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2
Journal of the
New York Entomological Society
published by
The New York Entomological Society
Contents Volume 93, 1985, Numbers 1-4
Number 1
The Coccinellidae (Coleoptera) of America North of Mexico Robert D. Gordon 1-912
Number 2
A revision of the rove beetle tribe Falagriini of America North of Mexico (Co-
leoptera: Staphylinidae: Aleocharinae) E. Richard Hoebeke 913-1018
Brood VI of 17-year periodical cicadas, Magicicada spp. (Hemiptera: Cicadidae):
New evidence from Connecticut, the hypothetical 4-year deceleration, and the
status of the brood Chris T. Maier 1019-1026
The natural history of Oncocnemis piffardi (Walker) (Lepidoptera: Noctuidae)
Tim L. McCabe 1027-1031
Number 3
Male territorial behavior in four species of the tribe Cercerini (Sphecidae: Phi-
lanthinae) Howard E. Evans and Kevin M. O’Neill 1033-1040
Orientation behavior of the slave-making ant Polyergus breviceps in an oak- wood-
land habitat H. Topoff, M. Pagani, M. Goldstein, and L. Mack 1041-1046
Cooperative colony foundation by females of the leafcutting ant Atta texana in
the laboratory Alex Mintzer and S. B. Vinson 1047-1051
General activity and reproductive behavior of Rhagoletis cornivora (Diptera: Te-
phritidae) flies in nature David Courtney Smith 1052-1056
Dictyla echii: seasonal history and North American records of an immigrant lace
bug (Hemiptera: Tingidae) A. G. Wheeler, Jr. and E. Richard Hoebeke 1057-1063
Scanning electron microscopic demonstration of bacteria on tarsi of Blattella
germanica Pasko Gazivoda and Durland Eish 1064-1067
Parasitism and mortality caused by field and laboratory strains of Brachymeria
intermedia (Nees) (Hymenoptera: Chalcididae)
Yuen-shaung Ng, James H. Lashomb, and Robert Chianese 1068-1072
Notes on the spider genus Eilica (Araneae: Gnaphosidae) Norman I. Platnick 1073-1081
Larval characters of a neotropical Scotocryptus (Coleoptera: Leiodidae), a nest
associate of stingless bees (Hymenoptera: Apidae) Quentin D. Wheeler 1082-1088
Species status and the hitherto unrecognized male of Papilio diaphora Staudinger
(1891), (Lepidoptera: Papilionidae)
Kurt Johnson, Rick Rozycki, and David Matusik 1089-1095
Revision of the Nearctic species of Hyssopus Girault (Hymenoptera: Eulophidae)
Michael E. Schauff 1096-1108
Antagonism of entomogenous fungal extracts to Dutch elm disease fungus, Cer-
atocystus ulmi J. N. Gemma, S. S. Wasti, and G. G. Hartmann 1109-1112
Melanism in Phigalia titea (Cramer) (Lepidoptera: Geometridae) in southern New
England: A response to forest disturbance? Theodore D. Sargent 1 113-1 120
Newly recognized synonyms, homonyms, and combinations in North American
Miridae (Heteroptera) Thomas J. Henry 1121-1136
Notes and Comments
Surface vibrational cues in the precopulatory behavior of whirligig beetles
Steven A. Kolmes 1 1 37-1 140
Number 4
Charles P. Alexander: a tribute, with emphasis on his boyhood in Fulton County,
New York, and his studies at Cornell University A. G. Wheeler, Jr.
A revision of the pentatomine genus Serdia StM, 1 860 (Pentatomidae: Hemiptera)
Donald B. Thomas, Jr. and L. H. Ralston
Names proposed and taxonomic publications by Herbert Ruckes (1895-1965)
L. H. Ralston and D. A. Rider
Revision of the platynine carabid genus Tanystoma Motschulsky (Coleoptera)
James K. Liebherr
Neotype designation for Scarites subterraneus Fabricius 1775 (Coleoptera: Ca-
rabidae: Scaritini) Stephen W. Nichols
Anthrenus pimpinellae F., a Palearctic dermestid established in eastern North
America (Coleoptera: Dermestidae)
E. Richard Hoebeke, A. G. Wheeler, Jr., and Richard S. Beal, Jr.
New host associations for Cerambycidae (Coleoptera) from selected species of
Leguminosae and Rutaceae Marlin E. Rice
New Ommatius Wiedemann (Diptera: Asilidae) from Cuba and the Bahamas
A. G. Scarbrough
Review of the Caribbean Geron Meigen (Diptera: Bombyliidae)
A. G. Scarbrough and Deborah A. Davidson
New family placement for the genus Cynipencyrtus (Hymenoptera: Chalcidoidea:
Tanaostigmatidae) John LaSalle and John S. Noyes
On the monophyly of the spider suborder Mesothelae (Arachnida: Araneae)
Norman L Platnick and Pablo A. Goloboff
1141-1164
1165-1172
1173-1181
1182-1211
1212-1215
1216-1222
1223-1225
1226-1239
1240-1260
1261-1264
1265-1270
Phytocoris adenostomae, a new mirine plant bug (Heteroptera: Miridae) from
southern California Gary M. Stonedahl 1271-1274
Notes and Comments
New water mite (Prostigmata: Parasitengona)— chironomid (Diptera) associations
from Otsego Lake, New York
Thomas W. Simmons
1275-1276
Book Reviews
Escarabajos. 20 Millones Afios de Evolucion
Brett C. Ratcliffe
1277-1278
The Spiders of Great Britain and Ireland
Norman I. Platnick
1279-1280
Population Biology and Evolution
John Jaenike
1280-1281
Chemical Ecology of Insects
Robert H. Hagen
1281-1285
Courtship Behaviors of the Hawaiian Picture-winged DrosophUa
David Grimaldi
1285-1287
Honorary, Life, and Sustaining Members
Reviewers for 1985
1288
1288
(
JANUARY 1985
No. 1
S' ^s, jom
Bnt
Vol. 93
Journal
of the
New York
Entomological Society
(ISSN 0028-7199)
Devoted to Entomology in General
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY
Editor: Randall T. Schuh, Department of Entomology, American Museum
of Natural History, Central Park West at 79th Street, New York, New
York 10024
Book Review Editor: Quentin D. Wheeler, Department of Entomology,
Cornell University, Ithaca, New York 14853
Publications Committee: Louis Trombetta, St. Johns University, Queens,
New York, Chairman; Alfred G. Wheeler, Jr., Pennsylvania State De-
partment of Agriculture, Harrisburg; Joseph M. Cerreta, St. Johns Uni-
versity, Queens, New York.
The New York Entomological Society
Incorporating The Brooklyn Entomological Society
President: Gerard Iwantsch, Department of Biological Sciences, Fordham
University, Bronx, New York 10458
Vice President: Henry M. Knizeski, Jr., Department of Biology, Mercy
College, Dobbs Ferry, New York 10522
Secretary: Irene E. Matejko, Science Department, The New Lincoln School,
New York, New York 10021
Assistant Secretary: Dennis J. Joslyn, Department of Biology, Rutgers
University, Camden, New Jersey 08102
Treasurer: Louis Sorkin, Department of Entomology, American Museum
of Natural History, New York, New York 10024
Trustees: Class of 79(54— Joseph Cerreta, St. Johns University, Queens,
New York; Durland Fish, Fordham University, Bronx, New York;
Class of 7955— Peter Chabora, Queens College, New York; Charles
Porter, Fordham University, Bronx, New York.
Annual dues are $18.00 for established professionals with journal, $10.00 without journal,
$10.00 for students with journal, $5.00 without journal. Sustaining memberships are $48.00
per year, institutional memberships are $120.00 per year, and life memberships are $300.00.
Subscriptions are $27.00 per year domestic and $30.00 foreign. All payments should be made
to the Treasurer. Back issues of the Journal of the New York Entomological Society, the Bulletin
of the Brooklyn Entomological Society, Entomologica Americana, the Torre- Bueno Glossary of
Entomology and other Society publications can be purchased from Lubrecht and Cramer, RD
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Meetings of the Society are held on the third Tuesday of each month (except June through
September) at 8 p.m. in the American Museum of Natural History, Central Park West at 79th
Street, New York, New York.
Mailed May 3, 1985
The Journal of the New York Entomological Society (ISSN 0028-7199) is published quarterly (January, April, July,
October) for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid at
New Brunswick, New Jersey and at additional mailing office.
Known office of publication: American Museum of Natural History, New York, New York 10024.
Journal of the New York Entomological Society, total copies printed 1,000, paid circulation 673, mail subscription 673, free
distribution by mail 7, total distribution 680, 320 copies left over each quarter.
THE COCCINELLIDAE (COLEOPTERA) OF
AMERICA NORTH OF MEXICO
ROBERT D. GORDON
Systematic Entomology Laboratory
IIBIII, Agricultural Research Service USDA,
% U.S. National Museum of Natural History,
Washington, D.C. 20560
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY
Volume 93, fascicle 1, pages 1-912
Price per copy $40.00
1
Plate 1.
Adalia bipunctata (L.) Cycloneda polita Casey
Axion tripustulatum (Degeer)
Paranaemia vittigera (Mannerheim) Hippodamia parenthesis (Say)
Epilachna borealis (F.)
Chilocorus stigma (Say) Coccinella trifasciata perplexa Mulsant
Plate 2.
Didion punctatum (Melsheimer) Nephaspis bioculatus (Blatchley)
Psyllobora vigintimaculata (Say)
Diomus terminalis (Say) Hyperaspis levrati Mulsant
Brachiacantha uteella Casey
Hyperaspis fastidiosa Casey Cephaloscymnus z. australis Gordon
iT
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2
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S’.
I
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]
THE COCCINELLIDAE (COLEOPTERA) OF
AMERICA NORTH OF MEXICO
ROBERT D. GORDON
i
n
' ■ !'■
(
/
4
(
Table of Contents
Introduction 3
Historical Resume 6
Methods 7
Terms 8
Acknowledgments 11
Biological Control and Coccinellidae in North America 12
Table 1 14
Table 2 29
Family Coccinellidae (Systematic Treatment) 33
Sticholotidinae 34
Micro weisini 36
Serangiini 58
Cephaloscymnini 66
Scymninae 74
Zilini 74
Stethorini 88
Scymnini 99
Selvadiini 347
Hyperaspini 352
Cryptognathini 599
Chilocorinae 602
Chilocorini 602
Coccidulinae 654
Coccidulini 655
Noviini 662
Exoplectrini 668
Azyini 671
Coccinellinae 678
Coccinellini 679
Psylloborini 851
Epilachninae 862
Epilachnini 863
Literature Cited 874
Index 893
1
Abstract,— T\iQ 57 genera and 475 species of Coccinellidae occurring in America north of
Mexico are treated taxonomically. Keys to all taxa, descriptions of the higher taxa, species
diagnoses, synonymies, and host records are included. Two new tribes, Cephaloscymnini and
Selvadiini, are erected for the genera Cephaloscymnus Crotch and Selvadius Casey. New species
are described as follows: Brachiacantha barberi; B. rotunda-, B. schwarzv, B. soltaur, B. stephani;
Exoplectra schaefferv, Gnathoweisea ferox-, G. hageni-, G. micula; G. texana-, Hyperaspidius
algodonus-, H. andrewsv, H. hardyi’, H. nanellus-, H. simulatus-, Hyperaspis caseyv, H. deludens;
H. dobzhanskyv, H. imitator, H. ornatella-, H. schaefferv, H. uteana-, Nephus (5”.) timberlakei;
Zagloba satana-, Zilus horni. A chapter on biological control involving the family Coccinellidae
includes discussions of the introduced species established in North America, and tables listing
all the species that have been introduced whether established or not.
1985
NORTH AMERICAN COCCINELLIDAE
3
Ladybird beetles (Coccinellidae) have been favorites of collectors and objects of
general popular interest for centuries because of the bright, contrasting red and black
colors of many of the species. Also, species tend to seek shelter in winter, hence are
commonly found in and around dwellings at a time of year when most insects are
not in evidence. Popular interest in the Ladybird (which in Europe is Coccinella
septempunctata) goes back at least to the fifteenth century and probably much farther.
The Ladybird is usually dedicated to the Virgin Mary; in Scandinavia it is called
Nyckelpiga, our Lady’s Key-maid, or Jung-fru Marias Gulhona, the Virgin Mary’s
Golden-hen. In Germany it is Frauen or Marien-Kafer, Ladybeetles of the Virgin
Mary, and in France it is known as Betes de la Vierge, Animals of the Virgin. Many
rhymes or verses stem from beliefs in the supernatural powers of the Ladybird, a
few of which are recited below (from Cowan, 1865).
From Vienna a superstition connected with the Ladybird’s ability to bring fine
weather:
Little birdie, birdie
Fly to Marybrunn
and bring us a fine sun
From the marsh of the Elbe comes a similar request:
May-cat
Fly away
Hasten away
Bring me good weather with you tomorrow
Northern Germany (Ploen) gives us a request based on the belief that the Ladybird
can foretell the harvest year; if the spots exceed 7, grain will be scarce, if there are
fewer than 7, there will be an abundant harvest:
Maerspart, fly to heaven
Bring me a sack full of biscuits, one for me, one for thee,
for all the little angels one
In northern Europe it is thought to be lucky when a young girl sees the Ladybird
in the spring, she lets it creep around her hand and says, “She measures me for
wedding gloves.” When it flies away the direction it takes is important because it
signifies from what direction her sweetheart will come. England provides us with this
rhyme:
This Ladyfly I take from off the grass,
whose spotted back might scarlet red surpass.
Fly, Ladybird, north, south, or east or west.
Fly where the man is found that I love best.
He leaves my hand, see to the west he’s flowen.
To call my true-love from the faithless town.
4
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Also from England (Norfolk) comes a similar wish in verse:
Bishop, Bishop Bamabee,
Tell me when my wedding be:
If it be tomorrow day,
Take your wings and fly away!
Fly to the east, fly to the west.
Fly to him that I love best.
In Scotland the Ladybird is a great favorite, and we have still more rhymes con-
cerning a sweetheart:
Lady, Lady Lanners
Lady, Lady Lanners,
Tak’ up your elowk about your head.
An’ flee awa’ to Planners (Flanders)
Flee ower firth and flee ower fell.
Flee ower pule and rinnan’ well.
Flee ower muir, and flee ower mead.
Flee ower livan, flee ower dead.
Flee ower com, and flee ower lee.
Flee ower river, flee ower sea.
Flee ye east, or flee ye west.
Flee till him that lo’es me best.
Or,— King, King collowa.
Up your wings and flee awa’
Over land and over sea;
Tell me where my love can be!
The Ladybird rhyme best known to children in England and America follows with
2 English versions:
Ladybird, ladybird, fly away home;
Your house is on fire your children’s at home.
All but one that ligs under the stone,—
Ply thee home, ladybird, ere it be gone.
From Yorkshire and Lancashire,—
Ladybird, ladybird, eigh thy way home;
Thy house is on fire, thy children all roam.
Except little Nan, who sits in her pan,
weaving gold laces as fast as she can.
The seemingly obscure meaning of the latter 2 rhymes is explained by the presence
of the Ladybird in large numbers among hop vines. The larvae feed on aphids of
the hop vine and fire was formerly used as a means of killing the aphids, thus
effectively killing the Ladybirds as well.
From “The Zoology of the English Poets” by Newell (1845) come 2 rather elegant
1985
NORTH AMERICAN COCCINELLIDAE
5
examples of verse concerning the Ladybird. First, from the tragedy of Sir Thomas
Moore by Hurdis:
Sir John.
What d’ye look at?
Cecilia.
A little animal, that round my glove.
And up and down to every finger’s tip.
Has travelled merrily, and travels still,
Tho’ it has wings to fly: what its name is
With learned men I know not; simple folk
Call it the lady-bird.
Sir John.
Save it.
Poor harmless thing!
Cecilia.
I would not hurt it for the world;
Its prettiness says. Spare me; and it bears
Armour so beautiful upon his back,
I could not injure it to be a queen:
Look, sir, its coat is scarlet dropp’d with jet.
Its eyes pure ivory.
Sir John.
Child, I am blind
To objects so minute: I know it well;
’Tis the companion of the waning year.
And lives among the blossoms of the hop;
It has fine silken wings enfolded close
Under that coat of mail.
Cecilia.
I see them, sir.
For it unfurls them now— ’tis up and gone.
And, from Southey’s “The Bumie-Bee”:
Back o’er thy shoulders throw thy ruby shards.
With many a tiny coal-black freckle deck’d;
My watchful eye thy loitering saunter guards.
My ready hand thy footsteps shall protect.
So shall the fairy train, by glowworm light.
With rainbow tints thy folding pennons fret.
6
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Thy scaly breast in deeper azure dight,
Thy burnish’d armour deck with glossier jet.
Some superstitions have existed about the Ladybird that don’t appear in verse,
such as the Ladybird as a cure for measles and colic (Newell, 1845); or as a cure for
the toothache when specimens are mashed and put into the hollow tooth. This latter
use of the Ladybird comes to us from Jaeger ( 1859) who states “I tried this application
in two instances, and the tooth-ache was immediately relieved; but whether the
remedy, or the faith of the patient, acted therapeutically, or the tooth ceased aching
of itself, I confess I do not pretend to know.”
Ladybird beetles are generally thought of as beneficial insects, predators of plant
pests; this is true for the most part, particularly in temperate regions. In tropical
regions, however, many are plant feeders, some economically significant. A few plant
feeders occur in temperate regions, the Mexican bean beetle being the prime North
American example.
Historically, the beneficial species have been classified as “Coccinellides Aphidi-
phages” (Chapuis, 1876) (aphid predators), and the plant feeders as “Coccinellides
phytophages” (Chapuis, 1876). This designation of beneficial species as aphid pred-
ators is accurate only in part. The beneficials actually divide into groups of species,
each of which has a preferred group of host species; as examples, species of Chilo-
corinae prey on scale insects, species of Stethorini on mites, and most species of
Coccinellinae are aphid predators. However, in the absence or scarcity of preferred
food, many species will feed on other insects such as the immature stages of Co-
leoptera, Lepidoptera, and Hymenoptera. Members of the genus Coleomegilla (Coc-
cinellinae) are able not only to survive on a variety of foods, but to complete de-
velopment when restricted to an unusual diet such as mites. Plant pollen also qualifies
as an essential food for Coleomegilla, and members of this genus are evidently the
most euryphagous of the Coccinellidae. The preferred food of another genus of
Coccinellinae, Neoharmonia, is evidently the larvae of a genus of Chrysomelidae.
Among the phytophagous Coccinellidae, most are typical leaf feeders, such as Epi-
lachna and Subcoccinella, but the Psylloborini have acquired the unusual habit of
feeding exclusively on fungal hyphae and spores.
Ladybirds are thus of considerable interest to naturalists, agriculturists, etc.; there-
fore a need exists for a comprehensive faunal treatment. The present volume is an
attempt to fill that need.
The purpose of this treatment is to provide the means to identify the species of
Coccinellidae occurring in America north of Mexico. To this end, keys, illustrations,
diagnoses, and synonymies are provided for all taxa known to occur in North Amer-
ica. A brief chapter on biological control importation efforts is included because of
the significance of many species as actual or potential control agents against plant
pests.
Historical Resume
Along with other animal groups, the classification of the Coccinellidae began with
Linnaeus in the mid 1700’s. Over the next 100 years it proceeded along the familiar
paths of insect classification, attended to by Fabricius, Degeer, Thunberg, Herbst,
1985
NORTH AMERICAN COCCINELLIDAE
7
etc. In 1850, Mulsant produced a monograph of the Coccinellidae on a world basis
that proved to be the foundation for modem classification and which still is an
indispensable tool in any coccinellid specialist’s shop. This treatment was so well
done that large portions of it remain unaltered by subsequent research. In 1853 and
1866, Mulsant published additional information as supplements to the 1850 volume.
The next major figure on the scene was George Robert Crotch, who again treated
the world Coccinellidae (1874). He changed Mulsant’s classification very little, but
added several new genera and many new species. Crotch was followed by Julius
Weise, who, although producing no single monumental work, succeeded in refining
coccinellid classification in a series of papers from 1 878 to 1930. Weise was a careful,
observant worker whose contributions were highly significant. He was also the first
coccinellid taxonomist to realize that male genitalia could be used to distinguish
species, although he did not pursue this to any extent. Korschefsky, a protege of
Weise, was an amateur coccinellid taxonomist of considerable ability. When Weise
died before writing the Coccinellidae portion of the Junk Catalogue, Korschefsky
proceeded to do the work which appeared in 1931 and 1932, and which remains the
single most useful taxonomic publication for coccinellid specialists anywhere in the
world. One of the most important contributions to coccinellid classification is rela-
tively recent; Sasaji (1968) published the “Phylogeny of the family Coccinellidae
(Coleoptera),” a thorough consideration of the relationships of the higher taxa of the
family. This publication has served as the basis for subfamily and tribal assignments
since 1968, and rightfully so; all morphological characteristics of adults and many
larval characteristics have been incorporated in the scheme in a logical fashion. I
regard this contribution as a landmark in coccinellid classification, to be compared
in significance with Mulsant’s classification of 1850.
Casey (1899) treated the Coccinellidae of the United States in their entirety, pro-
viding the foundation for taxonomic research in North America. The chief workers
in North American Coccinellidae since 1899 are Leng (1903-1920), Dobzhansky
(1931-1941), Chapin (1930-1966), Brown and de Ruette (1962), Brown (1962), and
Gordon ( 1 970-present). In addition, regional studies of Coccinellidae have been made
by Stehr (1930), Minnesota; Wingo (1952), Upper Mississippi Basin; Hatch (1961),
Pacific Northwest; J. Chapin (1974), Louisiana; and Belicek (1976), Western Canada
and Alaska.
Comprehensive publications on the biology, ecology, nutrition, metabolism, etc.,
are few. Some sources that contain literature reviews are Hagen ( 1 962), Hodek ( 1 966),
and Hodek (1967). Most recently Hodek (1973) has compiled much of this infor-
mation in a single source. Hodek’s book contains a short chapter on the taxonomy
and morphology of adults and an excellent chapter on the taxonomy and morphology
of the larvae. The bulk of the book is devoted to discussion of such biological
relationships as natural enemies, food sources, variability, and habitat.
Methods
In keeping with the primary purpose of this publication, to serve as an identification
manual, the systematics portion is kept as simple as possible. Thus, taxa above the
species level are fully described, but, except for new taxa, species are briefly diagnosed
8
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
rather than described. Illustrations are provided to facilitate identifications, and these
should be considered an essential part of the work and used accordingly.
Primary types. An effort has been made to locate and examine type material of all
authors included except the older European workers such as Linnaeus, Fabricius, etc.
When a species has been described from more than one specimen without designation
of a holotype, a lectotype is designated and, where possible, the remaining specimens
of the type series are designated as paralectotypes. The major type depositories for
North American Coccinellidae are the California Academy of Sciences, Museum of
Comparative Zoology, and U.S. National Museum; other institutions in North Amer-
ica that are type repositories are the Canadian National Collection and Purdue Uni-
versity. In several instances the type specimens have not been located and are either
known to be lost or are presumed to be. An example of the former is the Say collection;
examples of the latter are the Randall types and some of the Crotch and Melsheimer
types. When the type specimens are lost, not located, or not examined, the traditional
concept of the species has been accepted.
Locality records. Nearly all of the locality records listed in the text were taken from
specimens actually examined; published records were accepted only when the source
was unquestionably authoritative. Under “Distribution” only the specified locality
is given, plus county if stated on the label. Distribution data are given exactly as they
appear on the label except that obvious misspellings are corrected. For new species,
all information is given exactly as it apppears on the label. Distributions are presented
on maps with either symbols, shaded areas, or both. Shading is used when a species
is commonly collected; symbols are used when a species is rarely collected or when
the distribution pattern needs to be accurately defined. When a state record only is
available, “S” with the appropriate symbol appears in that state on the map.
Host data. Host data for members of each genus are listed in the generic discussion.
This is not intended to be a complete listing of all published host records; an ex-
haustive literature search has not been conducted, but all major sources of host
information have been consulted, additional records have been discovered in the
course of the study, and specimen label data have been included.
Terms
Most of the morphological terms used are germane to beetles in general, but some
are unique to ladybird beetles. To facilitate the use of the keys and descriptions, a
brief glossary follows (see Figs. 1, 2).
accessory gland, thin walled, saclike structure attached in basal Vi of spermathecal
capsule of female genitalia.
basal lobe, median apical projection of phallobase of male genitalia serving as a guide
for sipho.
basal piece, basal portion of phallobase of male genitalia to which the basal lobe,
paramere, and trabes attach.
bursa copulatrix, thin walled, saclike structure between infundibulum or sperm duct
and abdominal apex.
cornu, apical curved portion of spermathecal capsule of female genitalia.
cryptotetramerous (tarsus), tarsus composed of 4 segments, appearing 3 seg- mented
because 3rd segment minute, concealed between lobes of 2nd segment.
1985
NORTH AMERICAN COCCINELLIDAE
9
genital plates, sclerotized plates which are the divided 9th abdominal sternum in the
female.
infundibulum, sclerotized, funnel-like structure between sperm duct and bursa cop-
ulatrix of female genitalia.
nodulus, basal part of spermathecal capsule of female genitalia.
paramere, paired lateral apical projection of phallobase of male genitalia serving to
position and hold basal lobe in position during copulation.
phallobase, includes the basal piece, basal lobe, and paramere of male genitalia.
postcoxal line, the line on the 1st abdominal sternum posterior to hind coxa.
ramus, swelling or projection usually between cornu and nodulus of spermathecal
capsule of female genitalia.
sipho, sclerotized, curved rod which is inserted through the basal lobe and into the
female bursa copulatrix during copulation, corresponds to aedeagus or penis.
spermathecal capsule, part of the female genitalia composed of the cornu, ramus,
and nodulus (one or both of the latter may be absent).
10
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93( 1 )
Fig. 2. Male and female genitalia and postcoxal lines of Coccinellidae. a, b. Male genitalia,
c. Female, d-g. Postcoxal lines.
1985
NORTH AMERICAN COCCINELLIDAE
11
sperm duct, tube connecting spermathecal capsule to infundibulum or bursa copu-
latrix of female genitalia.
trabes, strut posterior to basal piece of male genitalia, connected by muscular at-
tachment to basal piece.
trimerous {tarsus), tarsus composed of 3 segments.
The postcoxal line on the 1 st abdominal sternum is a useful character for generic
discrimination. This line takes 4 major forms in the Coccinellidae; I refer to these
as the Pullus, Scymnus, Diomus, or Nephus types in the text without further expla-
nation. These types are illustrated in Figure 2, and are characterized as follows: Pullus
type-line complete, curved from base medially to base of sternum laterally; Scymnus
type-line incomplete, curved from base medially and forward toward base of sternum
laterally; Diomus type - line incomplete, extending down from base, joining apical
margin of sternum; and Nephus type - line incomplete, extending down from base
nearly to apical margin of sternum, extending parallel to apical margin toward lateral
margin.
Acknowledgments
For the loan of specimens of Coccinellidae I am indebted to the curators of the following
institutional collections (acronyms are those used in the text): (BMNH) British Museum (Natural
History), London, England; (CAS) California Academy of Sciences, San Francisco, California;
(CDA) California Department of Agriculture, Sacramento, California; (CM) Carnegie Museum
at Natural History, Pittsburgh, Pennsylvania; (CNC) Canadian National Collection, Ottawa,
Ontario; (DLM) Museum d’Histoire Naturelle (Dejean Collection), Lyon, France; (FSCA) Flor-
ida State Collection of Arthropods, Gainesville, Florida; (HSPA) Hawaiian Sugar Planters
Association, Honolulu, Hawaii; (INHS) Illinois Natural History Survey, Urbana, Illinois; (MCZ)
Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts; (NREA)
Naturhistoriska Riksmuseum, Entomologiska Avdelningen, Stockholm, Sweden; (PAS) Phila-
delphia Academy of Sciences, Philadelphia, Pennsylvania; (PM) Museum National d’Histoire
Naturelle, Paris, France; (PU) Purdue University, West Lafayette, Indiana; (UCCC) University
of Cambridge (Crotch Collection), Cambridge, England; (UCR) University of California, Riv-
erside, California; (UMMZ) University of Michigan Museum of Zoology, Ann Arbor, Michigan;
(UMZH) University Museum Helsinki, Finland; (USNM) United States National Museum,
Washington, D.C.; (WHN) William H. Nutting, Oakland, California; (ZMC) Zoologisk Museum
Copenhagen, Universitets Copenhagen, Denmark.
Assistance in the form of specimens, library materials, advice, encouragement, etc., was
rendered by the following individuals (institutional specimens were often involved, but acro-
nyms do not appear in the text): G. E. Ball, University of Alberta, Edmonton, Alberta; W. F.
Barr, University of Idaho, Moscow, Idaho; E. C. Becker, Biosystematics Research Institute,
Agriculture Canada, Ottawa, Ontario; J. M. Campbell, Biosystematics Research Institute, Ag-
riculture Canada, Ottawa, Ontario; J. B. Chapin, Louisiana State University, Baton Rouge,
Louisiana; J. A. Chemsak, University of California, Berkeley, California; J. T. Doyen, University
of California, Berkeley, California; H. Dozier, Clemson, South Carolina; W. A. Foster, Uni-
versity of Museum of Zoology, Cambridge, England; K. S. Hagen, University of California,
Division of Biological Control, Albany, California; T. J. Henry, USDA, Systematic Entomology
Laboratory, Washington, D.C.; H. F. Howden, Carleton University, Ottawa, Ontario; J. D.
Lattin, Oregon State University, Corvallis, Oregon; L, LeSage, Biosystematics Research Insti-
tute, Agriculture Canada, Ottawa, Ontario; W. H. Nutting, Oakland, California; R. D. Pope,
12
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
British Museum (Natural History), London, England; J. Smart, University Museum of Zoology,
Cambridge, England; A. G. Wheeler, Pennsylvania Department of Agriculture, Harrisburg,
Pennsylvania; R. E. Woodruff, Florida Department of Agriculture, Gainesville, Florida.
Host arthropod and plant names were verified by the following individuals: E. W. Baker, D.
R. Miller, and M. B. Stoetzel, USDA, Systematic Entomology Laboratory, Beltsville, Maryland;
D. C. Wasshausen, Smithsonian Institution, Washington, D.C.
The illustrations were prepared by contract illustrators Arthur Cushman, Criglersville, Vir-
ginia, Janine Higgins, Paris, Virginia, Britt Griswold, Washington, D.C., and Systematic Ento-
mology illustrators Linda Lawrence and Mary Lou Cooley. The color plates were prepared by
George Venable, Department of Entomology, Smithsonian Institution, Washington, D.C.
For reviewing the manuscript I am indebted to J. B. Chapin, Louisiana State University,
Baton Rouge, Lousiana; K. S. Hagen, University of California, Division of Biological Control,
Albany, California; R. D. Pope, British Museum (Natural History), London, England; W.
Steiner, Smithsonian Institution, Washington, D.C.; T. J. Henry, D. R. Whitehead, and E. W.
Baker, USDA, Systematic Entomology Laboratory, Washington, D.C.; J. R. Coulson, Beneficial
Insect Introduction Laboratory, Beltsville, Maryland.
Biological Control and Coccinellidae in North America
The history of biological control in North America has been well documented
beginning with Essig (1931). In addition to Essig, there have been several compre-
hensive reports on the subject which should be consulted for detailed information
and bibliographies. Chief among these are DeBach (1964), Hagen and Franz (1973),
and the articles in Huffaker and Messenger (1976). Clausen (1956b) discusses the
status of successfully established beneficial introductions prior to that date. Clausen
et al. (1978) present a broad view of the subject on a world wide basis.
The cottonycushion scale, a serious pest of citrus in California, precipitated the
first attempts at introducing foreign parasites and predators into North America. In
1888, Albert Koebele was sent to Australia to obtain natural enemies and sent back
to California several species of ladybird beetles, among which was the now famous
“vedalia” beetle, Rodolia cardinalis. This species proved to be an immediate and
spectacular success, and this success precipitated a wave of coccinellid introductions
which included 46 species between 1 89 1 and 1 892, all brought or sent from Australia
by Koebele (Hagen, 1974). Very few of these became established, and the interest in
predaceous coccinellids waned in favor of parasitic Hymenoptera and, later, pesti-
cides. In the 1960’s and 1970’s coccinellids were again introduced in significant
numbers with several useful establishments resulting. Table 1 gives a summary of
the species introduced into Canada and the United States, and is an attempt to list
all coccinellid species that have been introduced, whether established or not. This
attempt has not been completely successful because of ineffective record keeping
during much of the last 80 years, but is nearly complete for 1950-1983. Available
records show that 179 species have been intentionally imported into North America;
8 species have become established through accidental introductions, 5 of these had
been intentionally introduced but did not become established where released. A total
of 26 species of foreign Coccinellidae are now definitely or possibly established in
North America, 16 of these resulting from intentional releases. Following are sum-
maries of those species of Coccinellidae known to be established in North America
as a result of intentional or accidental introductions. Clausen (1956b), and Clausen
et al. (1978), and Tables 1 and 2 should be consulted for additional details.
1985
NORTH AMERICAN COCCINELLIDAE
13
Aphidecta obliterata (L.)
Aphidecta obliterata has been imported from Austria, Czechoslovakia, Germany,
Norway, and Sweden and released in Canada, the Pacific Northwest, and North
Carolina for control of the balsam woolly adelgid. The only release resulting in
establishment was from Germany into North Carolina in 1960-1963. It now occurs
in the Mt. Mitchell area only.
Azya orbigera orbigera Mulsant
There are no records of attempts made to introduce A. orbigera into Florida.
However, it is now definitely established in the Miami, Florida, area (1975 to date);
providing yet another example of an apparent accidental introduction. Woodruff and
Sailer (1 977) reviewed the history of the genus regarding biocontrol efforts in Florida.
Chilocorus bipustulatus (L.)
Attempts to establish Chilocorus bipustulatus in California were made in 1905,
1915, and 1927 from Israel and Italy for control of the black scale, citrus scale,
California red scale, etc. These attempts failed, but in 1951, C. bipustulatus was
imported from Israel and released for control of the olive scale, this time with
successful establishment. At present this species occurs in Fresno, Merced, and Ma-
dera counties, California, but the establishment may be tenuous.
Chilocorus kuwanae Silvestri
Introduced into the United States from Japan and China a number of times since
1 895 (as Chilocorus similis or kuwanae). Establishment resulted from an introduction
made in 1924-1925. White peach scale, California red scale, and San Jose scale were
the primary target hosts. At present C. kuwanae is known to occur in the vicinity of
Santa Barbara, California.
Cocci nella septempunctata L.
Attempts to establish C. septempunctata in the United States began in 1956 and
continued through 1971. Material was obtained from France, India, Italy, Norway,
and Sweden and released in several of the northeastern states, with an accompanying
rearing program that produced material sent to several other states as far west as
Arizona. All of these attempts apparently failed; however, specimens were collected
in Bergen County, New Jersey, in 1973 and 1974. The species is now known to be
established in several eastern states, but the origin of the New Jersey establishment
is unknown. Subsequent laboratory rearing and shipments of specimens have resulted
in establishment of C. septempunctata in Connecticut, Delaware, Georgia, Maine,
New York, Oklahoma, and Pennsylvania. Coccinella septempunctata was released
in New Brunswick in 1959-1960 without ensuing establishment; however, it is now
established in Quebec due either to an accidental introduction or spread northward
from Maine (Larochelle, 1979). New Jersey stock was also released in California, but
apparently did not become established there (K. Hagen, pers. comm.).
Table 1. Species of Coccinellidae intentionally introduced into North America.
14 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
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NORTH AMERICAN COCCINELLIDAE
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1985
NORTH AMERICAN COCCINELLIDAE
21
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22
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
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1985
NORTH AMERICAN COCCINELLIDAE
23
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1985
NORTH AMERICAN COCCINELLIDAE
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26
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
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punctata (L ) New Jersey L. (bouncing bet) Wheeler and Henry (1981);
Pope (1977)
30
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Coccinella undecimpunctata L.
First discovered in 1912 in Massachusetts, C. undecimpunctata has been subse-
quently reported from the northeastern United States and southern Canada, in the
vicinity of the St. Lawrence River and the Great Lakes, and as far north as New-
foundland; also from southern British Columbia. In 1965 it was discovered in the
Seattle area of Washington. The native distribution is Eurasian; the North American
populations are apparently the result of accidental introductions and subsequent
spread.
Cryptognatha nodiceps Marshall
The introductions were made in 1936 and 1938 from Puerto Rico and Trinidad
into south Florida (Miami) for use against the coconut scale. The species was re-
covered in 1940 and again in 1963. It is not certain whether it is actually established
or not, but if so, then the population is evidently very low and cannot be considered
as having any significant impact on pest populations at the present time. All available
records are from the Miami area.
Cryptolaemus montrouzieri Mulsant
The introductions took place in 1891-1892 and 1930 from Australia into Cali-
fornia, primarily for control of the citrus mealybug; but C montrouzieri is also a
predator of mealybugs of the genus Pseudococcus and will attack related genera such
as Phenacoccus and Ferrisia as well as the coccid genus Pulvinaria. Insectary reared
material from California was released in Florida where the species became estab-
lished, but attempts failed in Virginia in 1940-1941. A similar attempt also failed in
New Orleans, Louisiana, in 1908. C montrouzieri is presently established in Cali-
fornia and in central and southern Florida.
Diomus pumilio Weise
This Australian species has become established in California along the coast from
the San Francisco Bay area to San Diego, apparently as a result of releases made in
1975 and 1978, although it was first imported and released in 1892. Attempts have
been made to established it in eastern Canada (1958), North Carolina (1959), and
Washington (1959-1960), all without success.
Epilachna varivestis Mulsant
The Mexican bean beetle is a native of Mexico that probably migrated north as a
result of bean cultivation by Indians. It was first recorded in 1850 from the United
States (New Mexico) and later the beetle was discovered at Birmingham, Alabama,
in 1918. This latter introduction may have been a result of shipments of hay from
the west. It now occurs from Quebec south to Florida, west to Idaho and to the
Mexican border.
Exochomus flavipes (Thunberg)
Several attempts have been made to establish E. flavipes in California over the
years, but only an introduction from South Africa in 1978 succeeded, although
1985
NORTH AMERICAN COCCINELLIDAE
31
tenuously. The beetle was introduced for control of Pulvinariella mesembryanthemi
(Vallot) and Pulvinaria delottoi Gill, 2 species of scales on ice plant. E. flavipes now
occurs in the San Francisco Bay area.
Exochomus metallicus (Korschefsky)
This species is now established in Ventura County, California, from material in-
troduced from Eritrea, Ethiopia, in 1954 for control of the citricola and black scales.
Clausen et al. (1978) state that it is presently found in infestations of the citrus
mealybug on host plants other than citrus.
Exochomus quadripustulatus (L.)
First introduced into Massachusetts from Europe in 1905-1906 for control of
various lecaniine coccids; it was also imported into California from Italy in 1915,
and 1927-1928. No establishment resulted from three Massachusetts releases, but
it is now established in California where it feeds on several species of scale insects.
Halmus chalybeus (Boisduval)
This species was introduced into California as Orcus chalybeus from Australia in
1892 by Koebele and has been established in coastal southern California since. It
was released on the black scale and will develop on both lecaniine and diaspine
coccids. At present it is found mostly in infestations of the California red scale.
Harmonia dimidiata (F.)
The initial introduction as Leis dimidiata was from south China into California
in 1924. It was released in 1925 for aphid control, but did not become established.
A shipment sent to Florida from California in 1925 was released in 1926 with
establishment resulting. In 1 959 it was introduced into Oregon from India for control
of the balsam woolly adelgid but did not become established. At present H. dimidiata
occurs only in Florida.
Hyperaspis senegalensis hottentota Mulsant
This species was introduced into California from South Africa in 1978 for control
of scales on ice plant. It presently occurs in the San Francisco Bay area but “has only
a tenuous foothold” (Tassan et al., 1982).
Nephus {Sidis) binaevatus (Mulsant)
This species was introduced into California from South Africa in 1921 for control
of various mealybug species. It was released in 1922 and became established. At
present it occurs only in coastal and southern California.
Propylea quatuordecimpunctata (L.)
In 1968 a population of this palearctic species was discovered in the vicinity of
Montreal, Quebec, where it is apparently well established but still localized. Attempts
to establish this species in the United States from 1971 to 1982 were unsuccessful.
32
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Rhyzobius fores fieri (Mulsant)
This species has been misidentified as Rhyzobius ventralis Erichson, but Pope
(1981) corrected the long standing error. The true R. ventralis was among material
sent from Australia to California by Koebele in 1 889, but did not become established.
In 1892, another shipment thought to be “i?. ventralis''" was sent to California by
Koebele, this time the releases became established the same year. This second ship-
ment was composed of R. forestieri (Pope, 1951), a predator of the black scale on
various plants. The beetle is presently known only from California.
Rhyzobius lophanthae (Blaisdell)
Rhyzobius lophanthae (formerly Lindorus lophanthae) was first introduced from
Australia and established in 1892 for use against the black scale in California. It
preys on various species of Coccidae, but especially on diaspines. The present dis-
tribution throughout most of the southern United States may be a result of intro-
ductions from California stock or subsequent unrecorded releases. Cressman (1933)
recorded R. lophanthae from New Orleans, Louisiana, as an effective predator of
Chrysomphalus dictyospermi (Morgan); however, he gave no indication of introduc-
tion as being the population source. This species was also imported from South Africa
in 1959 and released in Texas, but it may already have been established there.
Rodolia cardinalis (Mulsant)
The vedalia beetle, Rodolia cardinalis, is the most famous introduced beneficial
insect in history. Koebele sent it from Australia to California in 1888-1889 for use
against the cottonycushion scale. It immediately became established and achieved a
startling success that led to the wholesale introduction of many other ladybird beetles
from Australia in ensuing years. At present it is established in California, south
Texas, Louisiana, and Rorida.
Scymnus {Pullus) impexus Mulsant
Scymnus impexus has been introduced into Canada and the United States from
Germany several times for control of the balsam woolly adelgid. It was released in
New Brunswick, Newfoundland, and Nova Scotia each year between 195 1 and 1960,
and persisted in small numbers through 1959. Early releases in British Columbia
apparently were unsuccessful, but success was achieved from releases in 1960 and
1961 in the Willamette Valley of Oregon. In North Carolina this beetle may have
become marginally established from releases in the early 1960’s, but at present this
cannot be documented.
Scymnus {Pullus) suturalis Thunberg
The first specimens reported from the United States were collected in Pennsylvania
in 1972, but misidentified as Scymnus {Pullus) coniferarum Crotch (Gordon, 1976b).
Subsequently the true identify was discovered (Gordon, 1982). The species is widely
distributed in Pennsylvania with additional records from New York, Michigan, and
Connecticut. The probable origin of this species is northern Europe possibly arriving
1985
NORTH AMERICAN COCCINELLIDAE
33
with imported nursery stock. S. (P.) suturalis was released in Michigan in 1961, but
whether the present Michigan populations result from that release cannot be docu-
mented.
Stethorus punctillum Weise
Stethorus punctillum, a European species of this mite-feeding genus, was first re-
ported in 1950 from Ontario and Massachusetts. It is now known to occur in eastern
North America from Massachusetts west to Wisconsin. In the west it occurs in western
Oregon, Washington, and Idaho.
Subcoccinella vigintiquatuorpunctata (L.)
This is one of 2 nonpredaceous (phytophagous) foreign coccinellids established in
North America. In the Old World S. 24-punctata is a serious pest of alfalfa. In North
America it apparently will not feed on alfalfa; instead it feeds primarily on bouncing
bet, Saponaria officinalis L. (Caryophyllaceae). The initial discovery of this species
was made in 1973, in Bergen Co., New Jersey. A subsequent survey in the eastern
states showed that the beetle was much more widespread and must have been es-
tablished long before 1973. It is now known from 7 states: Illinois, Maryland, New
Jersey, New York, Ohio, Pennsylvania, and West Virginia.
The following tables list all known introductions of foreign Coccinellidae. Table
1 deals with those intentionally imported, whether released or not, and Table 2 lists
those species accidentally introduced and established. The species are listed under
the currently accepted name or combination, with the name it was introduced under
in parentheses. The dates of importation and release are given if known with the
areas of release listed in the next column. The literature citations are those from
which the information presented for each species was gleaned. The tables were com-
piled from various literature sources beginning with Essig (1931). The major sources
utilized for the period from 1931 to date were the California Biological Control
Reports; Clausen (1956b); Canadian Insect Pest Review; Clausen et al. (1978). In
addition, I am indebted to the following individuals for information and assistance:
K. Hagen, University of California, Berkeley, Division of Biological Control, Albany;
R. Dysart and P. Schaefer, USDA Beneficial Insects Research Laboratory, Newark,
Delaware; J. Hall, Division of Biological Control, University of California, Riverside;
J. Coulson, USDA, Beneficial Insect Introduction Laboratory, Beltsville, Maryland;
R. Eye, USDA Yakima Agricultural Research Laboratory, Yakima, Washington; J.
Kelleher, Pesticide Information Liaison Section, Research Branch, Agriculture Can-
ada, Ottawa; R. Woodruff, Florida Department of Agriculture, Gainesville, Florida.
Systematic Treatment
Family Coccinellidae
Coccinellidae Latreille, 1807, p. 70— Westwood, 1839, p. 395— Crotch, 1874b, p.
53— Weise, 1885a, p. 3— Casey, 1899, p. 71— Mader, 1926, p. 1 — Korschefsky,
1931, p. 3-Wingo, 1952, p. 16-J. Chapin, 1974, p. 12-Belicek, 1976, p. 288.
Form usually oval to round, convex, sometimes elongate oval and weakly convex.
34
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Antenna usually 1 1 -segmented, often reduced to 10, 9, 8, or 7 segments, more or
less clubbed. Apical segment of maxillary palpus triangular (securiform), or parallel
sided, or conical. Elytron not truncate, not striate. Prostemal process distinctly sep-
arating transverse front coxae. Mesepimeron reaching middle coxal cavity. Abdomen
with 5 or 6 visible sterna, 7th rarely visible. First sternum of abdomen nearly always
with postcoxal line. Tarsus usually cryptotetramerous, often trimerous, rarely truly
tetramerous. Tibial spurs present or absent. Tarsal claw simple or toothed. Male
genitalia with sclerotized sipho (aedeagus), trilobed phallobase.
The cryptotetramerous tarsi and presence of postcoxal lines on the first abdominal
sternum will usually enable a coccinellid to be recognized as such. In those species
which lack postcoxal lines, the maxillary palpi are strongly securiform and the tarsi
are cryptotetramerous. The curved, sclerotized aedeagus (the sipho) is a certain char-
acter for family recognition.
Key to subfamilies of Coccinellidae north of Mexico
Clypeus expanded laterally, shelflike, partially dividing eye (Fig. 3a); dorsal surface
not pubescent Chilocorinae (p. 602)
Clypeus not expanded laterally, or if so, briefly and not shelflike; dorsal surface
pubescent or not 2
Mandible multidenticulate apically (Fig. 3c); antenna 1 1 -segmented, inserted dorsally
(Fig. 3b); dorsal surface pubescent; plant leaf feeders Epilachninae (p. 862)
Mandible rarely multidenticulate apically, if so, then length less than 3.0 mm; antenna
1 1 -segmented or not, insertion variable; dorsal surface pubescent or not; not plant
leaf feeders 3
Apical segment of maxillary palpus conical or elongate oval (Fig. 3d); mentum
narrowly articulated with submentum; length less than 3.0 mm; middle coxal cavities
broadly separated by articulation of meso- and metastema Sticholotidinae (p. 34)
Apical segment of maxillary palpus divergent apically (securiform) or nearly parallel
sided, rarely slightly convergent apically; mentum not narrowly articulated with
submentum; length often more than 2.0 mm; middle coxal cavities narrowly sepa-
rated except broadly separated in Scymninae 4
Antenna short, % or less as long as head width; apical segment of maxillary palpus
usually parallel sided or barrel shaped (Fig. 3d, e), rarely securiform; middle coxal
cavities broadly separated Scymninae (p. 74)
Antenna long, usually more than % as long as head width; apical segment of maxillary
palpus securiform (Fig. 3t); middle coxal cavities narrowly separated 5
Dorsal surface pubescent Coccidulinae (p. 654)
Dorsal surface glabrous Coccinellinae (p. 678)
Subfamily Sticholotidinae
Sticholotidinae Gordon, 1977, p. 186 (emendation).
Sticholotinae Weise, 1901, p. 430— Sasaji, 1967, p. 2— Sasaji, 1968, p. 19— J. Chapin,
1974, p. 13.
Small to medium-sized Coccinellidae; form hemispherical or elliptical. Functional
wings present or absent. Dorsally pubescent or not. Head with apical segment of
maxillary palpus more or less tapered, conical, barrel shaped or elongate oval; men-
1.
2(1).
3(2).
4(3).
5(4).
1985
NORTH AMERICAN COCCINELLIDAE
35
Fig. 3. a. Head of Chilocorus sp. b. Head of Epilachna sp. c. Epilachna mandible, d.
Microweisea maxillary palpus, e. Hyperaspis maxillary palpus, f. Coccinella maxillary palpus,
g. Coccidula antenna.
turn and submentum narrowly joined. Antenna usually inserted dorsally, with 7 to
1 1 segments, club with 1 to 5 segments. Pronotum sometimes with line or ridge
separating anterior angle from disc. Anterior coxal cavities open behind. Middle
coxal cavities broadly separated. Metendostemum with very broadly separated an-
terior tendons. Abdomen with 5 or 6 visible sterna; male 9th sternum flat. Tarsus
trimerous or cryptotetramerous. Female genital plate elongate, triangular.
The subfamily is principally characterized by the form of the terminal segment of
the maxillary palpus which is not securiform or distinctly broadened apically as is
typical of the rest of the Coccinellidae. The form of the maxillary palpus is an excellent
distinguishing character for members of the Serangiini, Microweisini, and Cephal-
oscymnini, but some mem^bers of tribes not occurring north of Mexico have that
segment more or less enlarged, approaching the typical coccinellid type. Members
of this subfamily are found throughout the tropical regions of the world with some
genera and species occurring also in temperate regions. The New World members of
Sticholotidinae were treated by Gordon (1977); see that paper for detailed discussion
of taxonomy, phylogeny and zoogeography. The Cephaloscymnini, new tribe, was
not recognized as belonging in this subfamily when that paper (1977) was prepared,
and therefore they were not included. The detailed study of the morphology of the
genus Cephaloscymnus required for this study showed that Cephaloscymnus and
36
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
related genera must be transferred to Sticholotidinae and a new tribe erected for
them. With this addition the tribes representing the subfamily north of Mexico are
Serangiini, Microweisini, and Cephaloscymnini.
Key to tribes of Sticholotidinae
1. Antennal club composed of a single knife-shaped or elongate-oval segment (Fig.
25b); femur broad, flat, fitting into depressions on ventral surface; prostemum greatly
expanded to conceal mouthparts (Fig. 25a) Serangiini
- Antennal club composed of more than a single segment or if only one, then segment
not knife-shaped; femur not broad or flat, ventral surface without depressions for
legs; prostemum not greatly expanded, not concealing mouthparts 2
2(1). Dorsal surface pubescent; head large, exposed, directed ventrally; eye large, narrow,
elongate (Fig. 30a) Cephaloscymini
Dorsal surface apparently glabrous; head small, at least slightly concealed under
pronotum, usually directed forward; eye small, round or oval (Fig. 4a) . . Microweisini
Tribe Microweisini
Microweisini Leng, 1920, p. 213 — Wingo, 1952, p. 19— J. Chapin, 1974, p. 14—
Sasaji, 1968, p. 20-Gordon, 1977, p. 200.
Pharini Casey, 1899, p. 1 10— Korschefsky, 1931, p. 209— Pope, 1962, p. 267 (in
part) (type-genus preoccupied).
Sticholotidinae with dorsal surface usually not pubescent; if so, then hairs of uni-
form length; size minute. Head capsule with prolonged frons and clypeus emarginate
around antennal insertion (Fig. 4a); eye small, facets ranging from extremely coarse
to fine. Mandible without apical or basal teeth. Apical segment of maxillary palpus
slender, tapered at apex (Fig. 4b). Antenna 7 to 1 0-segmented. Pronotum with oblique
anterolateral line inside anterolateral angle (Fig. 4c) (except Gnathoweisea schwarzi).
Intercoxal process of prostemum broad, with anterior lobe (Fig. 4d). Leg simple,
tibia unmodified. Tarsus cryptotetramerous or trimerous. Functional wing present.
Abdomen with 6 visible sterna; basal sternum with divided postcoxal line (Fig. 4g).
Male genitalia asymmetrical, phallobase with unpaired, basal apodeme (Fig. 8a).
Female spermathecal capsule bulbous (Fig. 8d).
This tribe is represented by 8 genera that occur from southern Canada to Chile
and Argentina and is apparently restricted to the Western Hemisphere. Microweisini
is a closely knit group of genera agreeing quite well in all essential characteristics.
The small size, characteristic habitus, the almost universal presence of an anterolateral
line on the pronotum, divided postcoxal line and broad T-shaped intercoxal process
of the prostemum serve to diagnose this tribe. See Gordon (1977) for a discussion
of all Western Hemisphere genera.
Key to genera of Microweisini
1 . Head entirely concealed beneath pronotum (Fig. 20d) Nipus Casey
- Head partially or not at all concealed 2
2(1). Head deeply inserted in prothorax, extremely elongate, slender (Fig. 15a)
Gnathoweisea Gordon
- Head not deeply inserted in prothorax, not elongate (Fig. 4a) 3
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NORTH AMERICAN COCCINELLIDAE
37
Fig. 4. Microweisea sp. a. Head. b. Maxillary palpus, c. Pronotum. d. Postemum. e. Antenna,
f. Leg. g. Postcoxal line.
3(2). Antenna with 3-segmented club (Fig. 4e) Microweisea Cockerell
- Antenna with 2-segmented club (Fig. 1 2a) Coccidophilus Brethes
Genus Microweisea Cockerell
Microweisea Cockerell, 1903, p. 38 (new name for Epismilia Cockerell, 1900)—
Wingo, 1952, p. 19-Pope, 1962, p. 637-J. Chapin, 1974, p. 15-Belicek, 1976,
p. 296— Gordon, 1970d, p. 207 — Gordon, 1977, p. 204. Type-species; Smilia
felschei Weise, by monotypy.
Smilia Weise, 1891, p. 288 (not Germar, 1833)— Horn, 1895, p. 82— Blatchley,
1910, p. 524.
Epismilia Cockerell, 1 900, p. 606 (not Fromental, 1861) (new name for Smilia Weise).
Pseudoweisea Schwarz, 1 904, p. 1 1 8 (name made available by accident).
Microweisini with form elongate, oval; dorsum glabrous. Head slightly prolonged
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anterior to antennal insertion (Fig. 4a); eyes separated by 4 times the width of an
eye. Apical segment of maxillary palpus elongate, slender, concical. Antenna with 7-
segmented scape, 3-segmented club (Fig. 4e). Prostemum with small anterior lobe.
Tarsus trimerous (Fig. 4f). Male genitalia asymmetrical, paramere reduced (Fig. 5a).
There are 5 species of this genus described from north of Mexico, and I am aware
of several undescribed species from Mexico and South America. Members of Mi-
croweisea are scale predators with available host records as follows; Lepidosaphes
beckii (Newman), Lepidosaphes sp., Melanaspis obscura (Comstock), Chionaspis pin-
ifoliae (Fitch), Pseudaonidia duplex (Cockerell), and Quadraspidiotus perniciosus
(Comstock). The North American species of Microweisea were taxonomically treated
by Gordon (1970d).
Key to species of Microweisea
1. Elytron with a transverse median yellow or yellowish red band (Fig. lOd)
coccidivora (Ashmead)
- Elytron without a transverse median yellow band 2
2(1). Species occurring in California 3
- Species not occurring in California 4
3(2). Elytron light yellowish brown, suture narrowly piceous; head and pronotum piceous;
surface of pronotum dull, strongly alutaceous suturalis (Schwarz)
- Elytron usually dark brown or piceous; unicolorous with head and pronotum; surface
of pronotum somewhat shiny, feebly alutaceous misella (LeConte)
4(2). Form extremely elongate (Fig. 1 Id); pronotum brown or yellowish brown, paler on
anterolateral angle than on disc; Florida ovalis (LeConte)
Form not extremely elongate; pronotum dark brown or piceous, anterolateral angle
not paler than disc; not restricted to Florida 5
5(4). Pronotum distinctly punctured; head strongly alutaceous; distributed from southern
Canada to Florida and Mexico misella (LeConte)
Pronotum without apparent punctures; head shiny, not or very feebly alutaceous;
Texas minuta (Casey)
Microweisea suturalis (Schwarz)
Fig. 5a-d; Map, Fig. 9
Pseudoweisea suturalis Schwarz, 1904, p. 118.
Microweisea suturalis: Leng, 1920, p. 213— Gordon, 1970d, p. 209.
Pentilia suturalis: Korschefsky, 1932, p. 225.
Diagnosis. Length 1.0 to 1.10 mm, width 0.90 to 0.95 mm. Color piceous; elytron
yellowish brown, elytral suture narrowly piceous (Fig. 5d), ventral surface brown.
Male genitalia as in Figure 5a-c.
Discussion. M. suturalis occurs only in California. The only species with which it
might be confused is M. misella, but M. misella usually has a uniformly dark dorsal
surface and the pronotum is mostly shiny and alutaceous; the pronotum of M.
suturalis is strongly alutaceous, dull. The holotype is a male specimen in the USNM
collection.
Type locality. Long Beach, California.
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Fig. 5. Microweisea suturalis.
Type depository. USNM (7936).
Distribution. Figure 9. CALIFORNIA: Long Beach; Los Angeles Co.
Microweisea minuta (Casey)
Fig. 6a-c; Map, Fig. 7
Smilia minuta Casey, 1899, p. 135.
Epismilia minuta: Cockerell, 1900, p. 606.
Microweisea minuta: Cockerell, 1903, p. 38 — Leng, 1920, p. 213— Gordon, 1970d,
p. 211.
Pentilia caseyi Korschefsky, 1931, p. 223 (unnecessary replacement name for minuta
Casey).
Diagnosis. Length 0.85 to 0.88 mm, width 0.55 to 0.60 mm. Color piceous, ventral
surface dark brown. Male genitalia as in Figure 6a-c. The small size will usually
distinguish this species; see remarks under M. misella. The type of M. minuta is a
unique male in the Casey collection which must be considered the holotype.
Type locality. Austin, Texas, on the Colorado River above Columbus.
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Fig. 6. Microweisea minuta.
Type depository. USNM (35241).
Distribution. Figure 7. TEXAS: Austin; Brownsville; San Diego, Sinton.
Microweisea misella (LeConte)
Fig. 8a-d; Map, Fig. 9
Pentilia misella LeConte, 1878a, p. 400— Korschefsky, 1931, p. 224.
Smilia misella-. Horn, 1895, p. 82— Casey, 1899, p. 135 — Blatchley, 1910, p. 524.
Epismilia misella: Cockerell, 1900, p. 606.
Microweisea misella: Cockerell, 1903, p. 38— Leng, 1920, p. 213— Wingo, 1952, p.
19-Gordon, 1970d, p. 211-J. Chapin, 1974, p. 15-Belicek, 1976, p. 297.
Diagnosis. Length 0.98 to 1 .45 mm, width 0.70 to 1 .05 mm. Color entirely piceous.
Male genitalia as in Figure 8a-c. Female genitalia as in Figure 8d.
Discussion. This is the most widely distributed member of the genus, having been
recorded from most areas of the United States and part of southern Canada. This
species and M. minuta are similar in appearance, but M. misella has distinct pronotal
punctures that are lacking in M. minuta, and nearly all specimens of M. misella are
obviously larger than the largest specimens of M. minuta. The male genitalia afford
a certain means of separating these 2 species. There are 8 specimens in the LeConte
collection that I consider types, the first of these, a male labeled “D.C./Type 6702
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NORTH AMERICAN COCCINELLIDAE
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(red paper)/pentilia misella Zim.”, I designate and label the lectotype. The remaining
7 specimens are designated as paralectotypes.
Type locality. Washington, D.C. (lectotype here designated).
Type depository. MCZ.
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Distribution. Figure 9. Southeastern Canada to Florida and east Texas, west to
British Columbia and northern California.
Microweisea coccidivora (Ashmead)
Fig. lOa-d; Map, Fig. 7
Hyperaspidius coccidivora Ashmead, 1880, p. 10.
Smilia coccidivora: Horn, 1895, p. 82— Casey, 1899, p. 135.
Epismilia coccidivora: Cockerell, 1900, p. 606.
Microweisea coccidivora: Cockerell, 1903, p. 38— Leng, 1920, p. 213— Gordon,
1970d, p. 212.
Pentilia coccidivora: Korschefsky, 1931, p. 223.
Diagnosis. Length 0.80 to 1.0 mm, width 0.60 to 0.70 mm. Color yellowish red;
elytral base and apex dark brown, transverse median area yellowish brown (Fig. 1 Od),
ventral surface and leg yellowish brown. Male genitalia as in Figure lOa-b.
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Fig. 9. Distribution. Microweisea misella (shaded); M. suturalis (dot).
Discussion. This is the only described species of the genus possessing a distinctive
dorsal color pattern which allows it to be easily recognized.
Type locality. Orlando, Florida (neotype designated by Gordon, 1970d).
Type depository. USNM (70409).
Distribution. Figure 7. Rorida; Georgia; South Carolina.
Microweisea ovalis (LeConte)
Fig. 1 1 a-d; Map, Fig. 7
Pentilia ovalis LeConte, 1878a, p. 400— Korschefsky, 1932, p. 225.
Smilia ovalis: Horn, 1895, p. 82.
Epismilia ovalis: Cockerell, 1900, p. 66.
Microweisea ovalis: Cockerell, 1903, p. 38 — Leng, 1920, p. 213— Gordon, 1970d, p.
213.
Smilia felscheiV^QisQ, 1891, p. 288 — Horn, 1895, p. 82.
Microweisea felschei: Leng, 1920, p. 13.
Diagnosis. Length 0.95 to 1.05 mm, width 0.50 to 0.63 mm. Form extremely
elongate (Fig. lid). Color brown; elytral suture piceous, anterior pronotal angle,
venter, and leg yellowish brown. Male genitalia as in Figure 1 la-c.
Discussion. The elongate form and pale pronotum distinguish M. ovalis from M.
suturalis which it most nearly resembles. LeConte had more than one type specimen,
but only one remains in his collection. This male labeled “Haulover, Fla, 11-10/977/
Type 6699(red paper)/Pentilia ovalis Lee.” is designated and labeled the lectotype.
Type specimen(s) of S. felschei have not been examined.
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Fig. 10. Microweisea coccidivora.
Type locality. Of ovalis, Haulover, Florida (lectotype here designated); of felschei,
Florida.
Type depository. Of ovalis, MCZ; of felschei, probably MNHUB (not examined).
Distribution. Figure 7. FLORIDA: Baldwin; Biscayne; Citrus City; Haulover; St.
Lucie; Tallahassee; Tampa. GEORGIA: Sapelo Island.
Genus Coccidophilus Brethes
Coccidophilus^vQXhQs, 1905, p. 76~Costa Lima, 1941, p. 409 — Pope, 1962,p. 628 —
Gordon, 1970d, p. 213— Gordon, 1977, p. 203. Type-species; Coccidophilus ci-
tricola Brethes, by monotypy and original designation.
Cryptoweisea Gordon, 1970d, p. 213— Gordon, 1977, p. 215. Type-species; Pentilia
marginata LeConte, by original designation.
Diagnosis. Microweisini with form elongate, oval; dorsum apparently glabrous.
Head slightly prolonged anterior to antennal insertion; eyes separated by 4 times the
width of an eye; frons often with 2 interocular depressions. Apical segment of max-
illary palpus elongate, conical (Fig. 12b). Antenna with 7-segmented scape and 2-
segmented club (Fig. 12a). Prostemum with small anterior lobe (Fig. 12c). Tarsus
trimerous. Male genitalia asymmetrical, paramere reduced (Fig. 1 2e).
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Fig. 1 1 . Microweisea ovalis.
Discussion. There are 4 described species in this genus and 2 of these occur north
of Mexico. It is difficult to separate Coccidophilus from Microweisea without counting
the antennal club segments, but species of Coccidophilus often have 2 depressions
on the frons between the eyes. These depressions are quite apparent in C. marginata,
but feeble and difficult to detect in C. atronitens. Members of Coccidophilus are scale
predators with available host records as follows: Chionaspis pinifoliae (Fitch), Lep-
idosaphes beckii (Newman); Aspidiotus sp.; Aonidiella aurantii (Masked); Pseudau-
lacaspis pentagona (Targioni-Tozzetti); Chrysomphalus aonidum (L.). The North
American species of Coccidophilus were taxonomically treated by Gordon (1970d)
under the generic name of Cryptoweisea.
Key to species of Coccidophilus
1 . Punctures on elytron coarse, dense, separated by the diameter of a puncture or less;
form slender, elongate (Fig. 14d); northern and eastern U.S. and southeastern Canada
marginata (LeConte)
- Punctures on elytron fine, separated by 2 or 3 times the diameter of puncture; form
oval (Fig. 12d); western United States atronitens (Casey)
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Fig. 1 2. Coccidophilus sp. a. Antenna, b. Maxillary palpus, c. Prostemum. d-h. Coccidophilus
atronitens.
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Fig. 13. Distribution. Coccidophilus atronitens (shaded); C. marginata (dot).
Coccidophilus atronitens (Casey)
Fig. 12d-h; Map, Fig. 13
Smilia atronitens Casey, 1899, p. 135.
Epismilia atronitens: Cockerell, 1900, p. 606.
Microweisea atronitens: Cockerell, 1903, p. 38— Leng, 1920, p. 213.
Pentilia atronitens: Korschefsky, 1931, p. 223.
Cryptoweisea atronitens: Gordon, 1970d, p. 215.
Coccidophilus atronitens: Gordon, 1977, p. 187.
Smilia reversa Fall, 1901, p. 231.
Microweisea reversa: Leng, 1920, p. 213.
Diagnosis. Length 1.10 to 1.20 mm, width 0.90 to 0.95 mm. Form oval (Fig. 12d).
Color dark brown; epipleuron and leg yellowish brown. Male genitalia as in Figure
1 2e-g. Female genitalia as in figure 1 2h.
Discussion. This species is smoother, more polished in appearance than C. mar-
ginata, and the dorsal punctation is very fine rather than coarse as in C. marginata.
The 2 species are strongly allopatric. There are 6 types of S. atronitens in the Casey
collection, all from the same locality. The first of these, a female, is here designated
and labeled the lectotype, the remainder are designated and labeled as paralectotypes.
Type specimens of S. reversa are in the Fall collection in the MCZ.
Type locality. Of atronitens Siskiyou Co., California (lectotype here designated);
of reversa, Lake Tahoe, San Bernardino Mts., California (lectotype not designated).
Type depository. Of atronitens USNM (35240); of reversa, MCZ.
Distribution. Figure 13. Colorado and Arizona to Oregon and California.
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Fig. 14. Coccidophilus marginata.
Coccidophilus marginata (LeConte)
Fig. 14a-d; Map, Fig. 13
Pentilia marginata LeConte, 1878a, p. 400— Korschefsky, 1931, p. 224.
Smilia marginata: Horn, 1895, p. 82— Casey, 1899, p. 135.
Epismilia marginata: Cockerell, 1903, p. 38.
Microweisea marginata: Cockerell, 1903, p. 38— Leng, 1920, p. 213— Wingo, 1952,
p. 27-Belicek, 1976, p. 297.
Cryptoweisea marginata: Gordon, 1970d, p. 215.
Coccidophilus marginata: Gordon, 1977, p. 203.
Diagnosis. Length 1.20 to 1.25 mm, width 0.70 to 1.00 mm. Form elongate (Fig.
14d). Color light brown; epipleuron yellowish brown. Male genitalia as in Figure
14a-b.
Discussion. The 2 interocular depressions on the frons are usually pronounced in
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NORTH AMERICAN COCCINELLIDAE
49
this species, feeble or absent in C atronitens', and C marginata is much more coarsely
punctured dorsally than C. atronitens (see remarks under C. atronitens). LeConte
apparently had one type specimen which must be considered the holotype. This male
in his collection is labeled “Marquette, Mich., 29-6/Type 6701(red paper)/Pentilia
marginata LeC.”
Type locality. Marquette, Michigan.
Type depository. MCZ.
Distribution. Figure 13. MAINE: Mt. Katahdin. MICHIGAN: Marquette. NEW
JERSEY: Anglesea; Burlington Co. NEW YORK: Ithaca; Mt. Whiteface. PENN-
SYLVANIA: Blair Co., Duncansville; Indiana Co., Shelocta; Philadelphia.
Genus Gnathoweisea Gordon
Gnathoweisea Gordon, 1970a, p. 47— Gordon, 1977, p. 204. Type-species; Smilia
planiceps Casey, by original designation.
Microweisini with form elongate, oval, pronotum partially covering head; dorsum
nearly glabrous, short, sparse pubescence present. Head elongate anterior to antennal
insertion, lateral border margined (Fig. 1 5a); eyes separated by 6 times the width of
an eye, very coarsely faceted. Apical segment of maxillary palpus elongate, slender,
conical. Antenna with 6-segmented scape, 3-segmented club (Fig. 15b). Prostemum
with or without anterior lobe. Postcoxal line as in Figure 15d. Tarsus trimerous.
Male genitalia asymmetrical, paramere somewhat reduced.
Two species have previously been placed in this genus, and four species are de-
scribed here. The extremely elongate head is the most obvious characteristic of
Gnathoweisea, but the 9-segmented antenna with a small, compact club is equally
distinctive within this tribe. The head is deeply inserted within the prothorax, the
intercoxal process is lobed anteriorly and protrudes ventrally except in G. schwarzi.
No host data is available, but members of this genus are undoubtedly scale predators,
probably on diaspine scales. The species of Gnathoweisea were reviewed by Gordon
(1970a), and the genus was discussed again by Gordon (1977).
Key to species of Gnathoweisea
1. Pronotum without oblique line across anterolateral angle; prostemum not lobed
anteriorly; anterior border of mesostemum raised (Fig. 15c) schwarzi Gordon
Pronotum with oblique line across anterolateral angle; prostemum lobed anteriorly;
anterior border of mesostemum flat 2
2(1). Length 1.20 mm or more hageni, n. sp.
Length 1.10 mm or less 3
3(2). Head extremely elongate, abmptly narrowed between hind margin of eye and an-
tennal insertion; Nevada ferox, n. sp.
- Head shorter, not abmptly narrowed; not known from Nevada 4
4(3). Dorsal color light brown; elytral punctures fine, lightly impressed; pronotal surface
feebly alutaceous micula, n. sp.
Dorsal color brown to black; elytral punctures coarse, distinctly
impressed; pronotal surface strongly alutaceous 5
5(4). Elytral punctures separated by a diameter or less; Arizona, California
Elytral punctures separated by more than a diameter; Texas
planiceps (Casey)
. . . texana, n. sp.
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Fig. 15. Gnathoweisea sp. a. Head. b. Antenna, c. Metastemum. d. Postcoxal line, e, f.
Gnathoweisea schwarzi.
Gnathoweisea schwarzi Gordon
Fig. 15c, e, f; Map, Fig. 17
Gnathoweisea schwarzi Gordon, 1970a, p. 50.
Diagnosis. Length 0.98 to 1.03 mm, width 0.70 to 0.75 mm. Color medium brown
except pronotum often dark brown or piceous. Male genitalia as in Figure 1 5e, f.
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51
Discussion. This species differs from the other members of the genus in having the
prostemum not at all lobed in front and the apex of the mesostemum raised to form
a ventrally directed ridge (Fig. 1 5c). The intercoxal process of the prostemum is also
much narrower than in the other 2 species.
Type locality. Williams, Arizona.
Type depository. USNM (70406).
Distribution. Figure 17. ARIZONA: type locality.
Gnathoweisea planiceps (Casey)
Fig. 16a-c; Map, Fig. 17
S mi Ha planiceps Casey, 1899, p. 135.
Microweisea planiceps: Cockerell, 1903, p. 38— Leng, 1920, p. 213.
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Pentilia planiceps: Korschefsky, 1932, p. 225.
Gnathoweisea planiceps: Gordon, 1970a, p. 50.
Diagnosis. Length 0.85 to 1.10 mm, width 0.72 to 0.78 mm. Color dark brown or
piceous. Male genitalia as in Figure 1 6a, b. Female genitalia as in Figure 1 6c.
Discussion. This species was previously known only from California, but I have
seen several specimens from Arizona that are apparently G. planiceps. For compar-
ative remarks see the discussion under G. ferox, n. sp. There are 2 type specimens
of S. planiceps in the Casey collection. The first of these, a male, is here designated
and labeled the lectotype; the other specimen is designated and labeled a paralec-
totype.
Type locality. Southern California (lectotype here designated).
Type depository. USNM (35242).
Distribution. Figure 17. ARIZONA: Bright Angel Camp; Huachucha Mts., Millers
Canyon; Hot Springs; Pima Co., Santa Rita Exp. Range; Santa Rita Mts.; Madera
Canyon. CALIFORNIA: Argus Mts.; Pomona; Riverside Co., Sage.
Gnathoweisea texana, new species
Map, Fig. 17
Description. Female, length 1.0 mm, width 0.72 mm. Form elongate, oval. Color
dark brown, head and pronotum nearly black. Head alutaceous, feebly shiny, nearly
impunctate; moderately prolonged anterior to eye, sides parallel. Pronotum dull,
alutaceous, meshes small, punctures fine, indistinct, separated by one to 3 times a
diameter. Elytron shiny, punctures coarse, separated by slightly more than a diameter.
Ventral surface smooth medially, lateral portion of metastemum and entire abdomen
alutaceous.
Holotype. Female. TEXAS: Bell Co., Co. Rd., 4 mi. E. Heidenheimer, Barton
Weems Farm, 26 Jun. 1978, coll. Robbins & Critchfield. USNM(101326).
This species closely resembles G. planiceps, but the elytral punctures are less dense
in planiceps. The only specimen examined is a female, therefore no genitalic com-
parisons are possible. The specific name refers to the state in which the holotype was
collected.
Gnathoweisea micula, new species
Map, Fig. 17
Description. Female, length 1.05 mm, width 0.80 mm. Form elongate, oval. Color
light brown; head, pronotum, and ventral surface slightly darker brown. Head shiny,
feebly alutaceous, impunctate; short anterior to eye and slightly widened. Pronotum
feebly alutaceous, somewhat shiny, punctures fine, indistinct, separated by less than
to twice a diameter. Elytron shiny, punctures feebly impressed, separated by one to
3 times a diameter. Ventral surface smooth medially, lateral portion of metastemum
and entire abdomen alutaceous.
Holotype. Female. NEW MEXICO: Deming, July 11-12, 4,300-4,400 ft., Wick-
ham. USNM (101327).
Paratypes. Total 2 (females) (Fig. 17). ARIZONA: Adamana, 7-V-03, HS Barber
collector; Walnut, Wickham. (USNM).
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NORTH AMERICAN COCCINELLIDAE
53
Fig. 17. Distribution. Gnathoweisea schwarzi (triangle); G. planiceps (dot); G. texana (open
circle); G. micula (star); G. hageni (circled star); G. ferox (square).
The pale color of G. micula and the feeble alutaceous sculpture on the head and
pronotum are diagnostic characters. The head anterior to the eye is very short, and
the sides are not parallel but slightly widened. The only other known species with
similar tendencies is G. hageni, n. sp. Only females of this species have been ex-
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Fig. 18. Gnathoweisea hageni.
amined. The specific epithet is from the Latin mica, meaning crumb, or morsel, and
refers to the small size.
Gnathoweisea hageni, new species
Fig. 18; Map, Fig. 17
Description. Female, length 1.50 mm, width, 1.0 mm. Form elongate, oval. Color
brown; head and pronotum dark brown. Head alutaceous, slightly shiny, punctures
fine, separated by 2 to 3 times a diameter; short anterior to eye and strongly widened.
Pronotum feebly alutaceous, shiny, punctures distinct, separated by one to 3 times
a diameter. Elytron shiny, punctures coarse, separated by 2 to 4 times a diameter.
Ventral surface smooth medially, lateral portion of metastemum and entire abdomen
alutaceous. Genitalia as in Figure 18.
Variation. Length 1.25 to 1.50 mm, width 0.90 to 1.0 mm.
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Fig. 19. Gnathoweisea ferox.
Holotype. Female. CALIFORNIA: Yuba Co., Bullards Bar, 11-1956, K. S. Hagen
Collector. USNM (101328).
Paratypes. Total 2 (females) (Fig. 17). CALIFORNIA: same data as holotype
(KSH).
This is the largest species of Gnathoweisea known. The head is short as in G.
micula, and the pronotal punctures are distinctly visible. All specimens examined
are females. The species is named for Kenneth Hagen, the collector, and one who
has contributed much to the biosystematics of Coccinellidae.
Gnathoweisea ferox, new species
Fig. 19a-c; Map, Fig. 17
Description. Male, length 1.0 mm, width 0.72 mm. Form elongate, oval. Color
dark brown; head and pronotum black; leg piceous; epipleuron yellowish brown.
Head dull, strongly alutaceous, nearly impunctate; extremely elongate, abruptly nar-
rowed between hind margin of eye and antennal insertion. Pronotum dull, alutaceous,
meshes very small, punctures very fine, indistinct, separated by one to 4 times a
diameter. Elytron shiny, densely, coarsely punctured, punctures separated by a di-
ameter or less. Ventral surface smooth medially, lateral portion of metastemum and
entire abdomen alutaceous. Genitalia as in Figure 1 9a-c.
Holotype. Male. NEVADA: Churchill Co., 6 mi. east of Frenchman, 16-VI-1973,
Stephen J. Chaplin. USNM(101329).
Paratypes. Total 9 (Fig. 17). Same data as holotype except 3 dated 22 Aug. 1972.
(USNM).
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Gnathoweisea ferox appears to have the head more strongly tapered (narrowed
from the hind margin of the eyes to the antennal insertion) than the other members
of the genus. The specific epithet is from the Latin meaning fierce, and refers to the
forbidding appearance of the head and mouthparts.
Genus Nipus Casey
NipusCsLsey, 1899, p. 132 — Leng, 1920, p. 213 — Korschefsky, 1931, p. 175— Gordon,
1970f, p. 71— Gordon, 1977, p. 208. Type-species; Nipus biplagiatus Casey, by
subsequent designation of Korschefsky, 1931.
Microweisini with form oval; dorsum glabrous or partially pubescent; pronotum
usually completely concealing head (Fig. 20d). Head strongly elongate anterior to
antennal insertion but not as elongate as in Gnathoweisea-, eyes separated by 3 times
the width of an eye. Apical segment of maxillary palpus elongate, somewhat conical.
Antenna with 7-segmented scape, 3-segmented club (Fig. 20a). Prostemum with
anterior lobe pronounced, semicircular, nearly concealing mouthparts ventrally (Fig.
20b). Postcoxal line as in Figure 20c. Tarsus trimerous. Male genitalia asymmetrical,
paramere reduced.
This genus contains 4 species occurring in the southwestern United States. Nipus
is readily distinguished from other genera of Microweisini because the head is almost
always completely concealed beneath the pronotum. The partially concealed head
found in Gnathoweisea is the only remotely similar condition known. The species
of Nipus were reviewed by Gordon (19701), and the genus was discussed again by
Gordon (1977). The only host record for this genus is that of N. biplagiatus preying
upon Ehrhornia cupressi (Ehrhom), but all members of the genus are undoubtedly
scale predators.
Key to species of Nipus
1 . Elytron with a pale red or yellow spot, or red or yellow band (Fig. 20d) 2
Elytron without pale spot or band 3
2(1). Form elongate, parallel sided (Fig. 20d); California biplagiatus Casey
Form elongate, oval, not parallel sided (Fig. 24); Arizona, Utah occiduus Gordon
3(1). Form narrow, elongate (Fig. 21); pronotum dull, strongly alutaceous; punctures on
elytron extremely coarse; California niger Casey
- Form oval (Fig. 23); pronotum shiny, feebly alutaceous; punctures on elytron fine;
Arizona, Colorado planatus Gordon
Nipus biplagiatus Casey
Fig. 20d-f; Map, Fig. 22
Nipus biplagiatus CdiSty, 1899, p. 133 — Leng, 1920, p. 213— Herbert, 1920, p. 18 —
Schilder, 1928, p. 237 — Korschefsky, 1931, p. 175— Gordon, 1970f, p. 72.
Diagnosis. Length 1.25 to 1.50 mm, width 0.75 to 0.82 mm. Form elongate,
parallel-sided, abruptly narrowed posteriorly (Fig. 20d). Color piceous; large median
area of elytron and anterior margin of pronotum yellow, ventral surface yellowish
brown. Male genitalia as in Figure 20e, f
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NORTH AMERICAN COCCINELLIDAE
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Fig. 20. Nipus biplagiatus. a. Antenna, b. Prostemum. c. Postcoxal line. d. Habitus; e, f.
male genitalia.
Discussion. This species is known only from California and resembles N. occiduus
which apparently does not occur in California. In addition to the key characters, N.
occiduus is smaller and not as coarsely punctured as N. biplagiatus. In the Casey
collection is a unique female which must be considered the holotype of N. biplagiatus.
Type locality. Los Angeles, California.
Type depository. USNM (35224).
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Distribution. Figure 22. CALIFORNIA: Contra Costa Co., Vine Hill; Los Angeles;
Los Gatos; Orange Co., Costa Mesa; San Bernardino; Upland.
Nipus niger Casey
Fig. 21; Map, Fig. 22
Nipus niger CnsQy, 1899, p. 133 — Leng, 1920, p. 2 1 3 — Korschefsky, 1931, p. 176 —
Gordon, 1970f, p. 73.
Diagnosis. Length 1.05 to 1.10 mm, width 0.70 to 0.74 mm. Form elongate, oval,
gradually narrowed posteriorly (Fig. 21). Color brownish piceous; anterior margin of
pronotum yellowish brown, ventral surface brown.
Discussion. No males of this species were available for study. Nipus niger is most
similar to N. planatus, but the key characters will readily separate the 2 species. A
unique female in the Casey collection must be considered the holotype of N. niger.
Type locality. Sonoma Co., California.
Type depository. USNM (35225).
Distribution. Figure 22. CALIFORNIA: Humboldt Co.; Los Gatos; Sonoma Co.
Nipus planatus Gordon
Fig. 23; Map, Fig. 22
Nipus planatus Gordon, 1970f, p. 74.
Diagnosis. Length 1.19 to 1.24 mm, width 0.81 to 0.84 mm. Form elongate, oval,
evenly narrowed anteriorly and posteriorly (Fig. 23). Color brown; anterior and lateral
borders of pronotum yellowish brown, mouthparts and leg yellowish brown.
Type locality. Salida, Colorado.
Type depository. USNM (70851).
Distribution. Figure 22. ARIZONA: Bright Angel Camp. COLORADO: Salida.
Nipus occiduus Gordon
Fig. 24; Map, Fig. 22
Nipus occiduus Gordon, 1970f, p. 75.
Diagnosis. Length 1.20 to 1.24 mm, width 0.75 to 0.78 mm. Form oval (Fig. 24).
Color piceous; elytron with yellow spot occupying Vi to % of elytron, anterior margin
of pronotum yellowish brown.
Discussion. See comparative remarks under N. biplagiatus.
Type locality. Wasatch, Utah.
Type depository. USNM (70852).
Distribution. Figure 22. ARIZONA: Chiricahua Mts.; Huachucha Mts., Millers
Canyon; Oracle; Santa Rita Mts; Williams. UTAH: Wasatch.
Tribe Serangiini
Sera/tgh/?/ Blackwelder, 1945, p. 450— Pope, 1962, p. 627— Sasaji, 1967, p. 2 — Sasaji,
1968, p. 20-Gordon, 1970e, p. 356-J. Chapin, 1974, p. 13-Belicek, 1976, p.
292-Gordon, 1977, p. 208.
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Fig. 2 1 . Nipus niger.
Sticholotidinae with form compact; dorsally pubescent or not. Head slightly pro-
longed anterior to antennal insertion, emarginate around insertion; eye coarsely fac-
eted (Fig. 25a). Apical segment of maxillary palpus either elongate and conical, or
short and barrel shaped. Antenna 8 or 9-segmented, club composed of a single
segment (Fig. 25b). Prostemum strongly lobed anteriorly, concealing mouthparts
(Fig. 25a), notched on each side for reception of antenna. Epipleuron with fovea for
reception of leg. Leg received in deep cavity on ventral surface; at least front femur
broad, flat, concealing tibia when leg retracted; at least front tibia angulate externally.
Tarsus cryptotetramerous or trimerous. Abdomen with 5 visible sterna. Postcoxal
line on first abdominal sternum complete (Fig. 25c). Male genitalia asymmetrical,
paramere reduced. Female genitalia lacking infundibulum.
This tribe presently contains 6 genera, 5 of which are native to the Old World.
Delphastus is the only native American representative of the tribe with 12 species
occurring from Canada to Argentina. Catana clauseni Chapin occurs in Cuba, but
was introduced from Indonesia in 1930 for biocontrol of the citrus blackfly, Aleu-
rocanthus woglumi Ashby. Serangiini is a closely knit group of genera, highly dis-
tinctive in appearance. The strongly lobed prostemum that conceals the mouthparts
and has a notch on each side for reception of the antenna is the most striking
characteristic; in addition, the ventral surface is deeply foveate for reception of the
legs, and at least the anterior leg is broad, flattened. See Gordon (1977) for further
discussion of the genera occurring in the Western Hemisphere.
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Fig. 22. Distribution. Nipus biplagiatus (dot); N. niger (star); N. planatus (square); N. oc-
ciduus (open circle).
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NORTH AMERICAN COCCINELLIDAE
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Fig. 23. Nipus planatus.
Genus Delphastus Casey
Delphastus CsLsey, 1899, p. Ill— Leng, 1920, p. 214— Korschefsky, 1931, p. 220—
Chapin, 1940, p. 264-Wingo, 1952, p. 22-J. Chapin, 1974, p. 13-Belicek, 1976,
p. 292— Gordon, 1970e, p. 357— Gordon, 1977, p. 209. Type-species; Oeneis
pusillus LeConte, by subsequent designation of Korschefsky, 1931.
Serangiini with form hemispherical, slightly elongate. Head with apical segment
of maxillary palpus slender, somewhat conical. Antenna 9-segmented (Fig. 25b).
Elytron without sutural line. Epipleuron not descending externally. Leg with femur
broad; tibia angulate externally. Tarsus trimerous.
There are only 3 species of this genus described from the area north of Mexico;
the remaining 9 described species occur from Mexico and the West Indies to Argen-
tina. Members of Delphastus are known as predators on whiteflies (Aleurodidae) with
available host records as follows: Aleurocanthus woglumi Ashby; Pelius kelloggi (Be-
mis); Trialeurodes floridensis (Quaintance); Dialeurodes citri (Ashmead); and Di-
aleurodes citrifolii (Morgan). However, a series of a species of Delphastus in the
USNM collection bears the host data “on Asterolecanium miliaris (Boisduval)”, a
pit scale. Kamiya (1966) records Serangium japonicum japonicum Chapin as feeding
on the soft scales Ceroplastes rubens Maskell and Ceroplastes japonicus Green in
Japan. It appears that members of the Serangiini feed on both whiteflies and scale
insects. The species of Delphastus were taxonomically treated by Gordon (1970e).
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Fig. 24. Nipus occiduus.
Key to species of Delphastus
1. Length less than 1.10 mm; color pale reddish yellow; Florida pallidus (LeConte)
Length more than 1.30 mm; color light reddish brown to black; not restricted to
Florida 2
2(1). Prostemal lobe densely, coarsely punctate; California catalinae (Rovn)
- Prostemal lobe smooth; not restricted to California pusillus (LeConte)
Delphastus pallidus (LeConte)
Fig. 25e, g; Map, Fig. 27
Oeneis pallidus LeConte, 1878a, p. 400.
Cryptognatha pallida: Horn, 1895, p. 83.
Delphastus pallidus: Casey, 1899, p. 112 — Leng, 1920, p. 214— Blatchley, 1924, p.
167-Korschefsky, 1931, p. 220-Gordon, 1970e, p. 360.
Diagnosis. Length 0.90 to 1.05 mm, width 0.70 to 0.80 mm. Color pale reddish
yellow except leg yellow. Male genitalia as in Figure 25e, g.
Discussion. This small, pale species is readily recognizeable by the key characters.
The type is a unique female in the LeConte collection labeled “Sand Pt, Fla, 18-2/
979/Type 6696(red paper)/Oeneis pallidus LeC.” which must be considered the
holotype.
Type locality. Sand Point, Florida.
Type depository. MCZ.
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NORTH AMERICAN COCCINELLIDAE
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Fig. 25. Delphastus pallidus. a. Head and prostemum. b. Antenna, c. Postcoxal line. d-g.
Male genitalia.
Distribution. Figure 27. FLORIDA: Homestead; Lake Alfred; Miami; Orlando;
Pasco Co.; Sand Point; Volusia Co.
Delphastus catalinae (Horn)
Fig. 26a-d; Map, Fig. 27
Cryptognatha catalinae Horn, 1895, p. 83.
Delphastus catalinae: Casey, 1899, p. 1 12 — Leng, 1920, p. 214— Korschefsky, 1931,
p. 220— Gordon, 1970e, p. 365.
Diagnosis. Length 1 .40 to 1 .50 mm, width 1 . 10 to 1.18 mm. Color medium reddish
brown, median area of pronotum slightly darker; legs and head of male pale yellowish
brown. Male genitalia as in Figure 26a-d.
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Fig. 26. Delphastus catalinae.
Discussion. The coarsely punctured prostemal lobe distinguishes this species from
other North American Delphastus. In addition, D. catalinae is broader and usually
paler in color than D. pusillus which it most closely resembles. The type is a unique
female in the Horn collection labeled “Catalina Cal., 7-21-94/Holotype 3169(red
paper)/Cryptognatha catalinae H.” which must be considered the holotype.
Type locality. Catalina, southern California.
Type depository. MCZ.
Distribution. Figure 27. CALIFORNIA: Catalina; Los Angeles Co., Oak Canyon,
Tanbark Flat; Pasadena, San Antonio Canyon; Santa Barbara.
Delphastus pusillus (LeConte)
Fig. 28a-c; Map, Fig. 27
Oeneis pusilla LeConte, 1852, p. 135 — Crotch, 1873, p. 377.
Cryptognatha pusilla: Crotch, 1874b, p. 207 — Horn, 1895, p. 83.
Delphastus pusillus: Casey, 1899, p. 1 12 — Blatchley, 1910, p. 519 — Leng, 1920, p.
214 — Wingo, 1952, p. 45— J. Chapin, 1974, p. 14— Korschefsky, 1931, p. 220—
Gordon, 1970e, p. 367.
Oeneis puncticollisTQCon\.Q, 1852, p. 135 — Crotch, 1873, p. 377 (as female of /7W5/7/^z).
Cryptognatha puncticollis: Crotch, 1874b, p. 207 — Horn,1895, p. 83.
Delphastus pusillus var. puncticollis: Casey, 1899, p. 1 12 — Korschefsky, 1931, p. 220.
Delphastus puncticollis: Gordon, 1970e, p. 367.
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NORTH AMERICAN COCCINELLIDAE
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Fig. 27. Distribution. Delphastus pallidus (dot); D. catalinae (open circle); D. pusillus (shad-
ed).
Delphastus sonoricus Casey, 1899, p. 1 12— Korschefsky, 1931, p. 221— Gordon,
1970e, p. 367.
Diagnosis. Length 1.40 to 1.60 mm, width 1.10 to 1.20 mm. Color black; pro-
stemum and leg yellow, male with head and lateral margin of pronotum yellow. Male
genitalia as in Figure 28a, b. Female genitalia as in Figure 28c.
Discussion. Delphastus pusillus is a widely distributed, variable species. The color
pattern described above was taken from a Maryland specimen which agrees quite
well with LeConte’s original description. The southwestern U.S. specimens are usually
dark brown rather than black and the males do not have lighter colored pronotal
margins, it was to this form that Casey gave the name D. sonoricus. The brown form
prevails south through Mexico and Central America with an occasional population
from a coastal locality exhibiting the color pattern of typical D. pusillus. Its range
appears to be continuous into South America at least as far as Peru.
I consider the three specimens of D. pusillus that remain in the LeConte collection
type material. The first of these, a male labeled “(orange disc)/ /Type 6697(red paper)/
Oe. pusilla LeC.”, I designate and label the lectotype. The second, bearing only an
orange disc, and the third bearing a pink disc are designated as paralectotypes.
LeConte apparently had only one example of O. puncticollis, and this specimen in
his collection labeled “(orange disc)/Type 6698/Oeneis puncticollis LeC.” must be
considered the holotype. Casey had 6 type specimens from southern Arizona and
southern California. I designate and label a male as the lectotype and the remainder
as paralectotypes.
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Fig. 28. Delphastus pusillus.
Type locality. Of pusillus, Georgia (lectotype here designated); of puncticollis,
“Southern States”; of sonoricus, Tucson, Arizona (lectotype here designated).
Type depository. Of pusillus and puncticollis, MCZ; of sonoricus, USNM (35230).
Distribution. Figure 27. Massachusetts to Florida, west to California.
Cephaloscymnini, new tribe
Sticholotidinae of small size, length less than 3.0 mm. Head prominent, exposed,
deflected ventrally; eye large, narrow, elongate, very finely faceted, inner margin
parallel or closer at posterior border of eye than at anterior border; apex of clypeus
truncate or subtruncate; gena with or without narrow extension onto eye. Antenna
inserted frontally at apex of eye, insertion exposed or not; antenna short, 8-10-
segmented, club 3-segmented. Apical segment of maxillary palpus long, slender,
conical or parallel sided. Mandible bidentate apically, or unidentate with feeble,
subapical tooth Pronotum short, deeply excavated for reception of head, lateral border
explanate, anterolateral angle strongly produced forward, extending nearly to apex
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Fig. 29. Cephaloscymnus zimmermani zimmermani.
of eye. Prostemum broad, bicarinate or not, produced anteriorly to partially conceal
mouthparts or not. Epipleuron broad or narrow, not foveate for reception of leg. Leg
slender, simple. Tarsus cryptotetramerous; tarsal claw without tooth. Abdomen with
5 visible sterna. Postcoxal line on 1st abdominal sternum complete (Fig. 30g). Male
genitalia symmetrical.
The group of genera here assigned to this tribe contain some of the most unusual
appearing Coccinellidae in the entire family. These genera have previously been
placed in the Scymninae, but examination of all morphological characters shows that
they belong in the subfamily Sticholotidinae. They are not closely related to members
of any presently established tribe; therefore, the establishment of the tribe Cephal-
oscymnini is deemed necessary. The included genera are Cephaloscymnus Crotch,
Prodilis Mulsant, Neaporia Gorham, Aneaporia Casey and Prodioloides Weise. The
genus Cephaloscymnus has been placed in the Scymnini or Ortaliini by authors, while
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Prodilis, Neaporia, and Prodiloides have been placed in the Ortaliini. Casey consid-
ered Aneaporia to belong to the Exoplectrini but this was an obviously incorrect
placement. All of these genera quite apparently share a common ancestry and must
be grouped together as done here. Examination of species of this group in existing
collections indicates that additional genera will have to be erected when a complete
study is completed. The combination of short antenna; large, ventrally directed head;
large, narrow, finely faceted eyes; and short, explanate pronotum readily separate
this tribe, not only from other tribes of Sticholotidinae, but from all other North
American Coccinellidae. The only genus in this tribe occurring north of Mexico is
Cephaloscymnus.
Genus Cephaloscymnus Crotch
Cephaloscymnus Croich., 1873, p. 382 — Horn, 1895, p. 81— Casey, 1899, p. 160 —
Blatchley, 1910, p. 524 — Leng, 1920, p. 214 — Korschefsky, 1931, p. 168— Wingo,
1952, p. 19— Gordon, 1970b, p. 66. Type-species; Cephaloscymnus zimmermanni
Crotch, by monotypy.
Cephaloscymnini with form elongate, slender (Fig. 29). Head broad between eyes,
frons 3 times the width of an eye; inner margin of eyes nearly parallel; apex of clypeus
subtruncate; gena not extending onto eye (Fig. 30a). Antennal insertion exposed;
antenna 9-segmented, club 3-segmented (Fig. 30b). Apical segment of maxillary
palpus slender, parallel-sided (Fig. 30c). Mandible unidentate apically, with feeble,
subapical tooth (Fig. 30d). Surface of head and pronotum deeply, densely punctured,
punctures contiguous or nearly so. Prostemum short, not produced anteriorly, with-
out carinae (Fig. 30e). Male metastemum with large, deep, pubescent pit (Fig. 30f).
Postcoxal line as in Figure 30g. Female genitalia without infundibulum; spermethecal
capsule simple, lacking cornu or ramus (Fig. 311).
The presence of a metacoxal pit in males and the short intercoxal prostemal process
lacking carinae or an anterior protuberance distinguish Cephaloscymnus from the
other genera of Cephaloscymnini. There are presently 7 species in this genus (Gordon,
1970b, 1974d), 2 of which are known only from Mexico and 4 from Mexico and the
United States. Cephaloscymnus bruchi Weise was described from Brazil; I have not
seen this species but suspect that it belongs in Prodilis. No host data is available for
members of this genus, but they are probably scale predators. Cephaloscymnus has
been revised by Gordon (1970b), with a subsequent paper (Gordon, 1974d) on
additional species from Mexico.
Key to species of Cephaloscymnus
1. Length 2.15 mm or more; eastern United States, Texas, Arizona, New Mexico, and
Mexico 2
- Length 2.15 mm or less; California, Arizona, Texas, Mexico 3
2(1). Pronotum and elytron piceous to black; eastern U.S
zimmermanni zimmermanni Crotch
Pronotum usually reddish, elytron piceous to brown; southwestern U.S. and north-
eastern Mexico zimmermanni australis Gordon
3(1). Ventral surface black (except legs and mouthparts) laevis Gordon
Ventral surface piceous or brown 4
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Fig. 30. Cephaloscymnus sp. a. Lateral view of head and pronotum. b. Antenna, c. Maxillary
palpus, d. Mandible, e. Prostemum. f. Metastemum. g. Postcoxal line.
4(3). Pronotum finely punctured, anterior angle feebly explanate occidentalis Horn
Pronotum coarsely punctured, anterior angle strongly explanate .... insulatus Gordon
Cephaloscymnus zimmermanni zimmermanni Crotch
Figs. 29, 31a-e; Map, Fig. 32
Cephaloscymnus zimmermanni Crotch, 1873, p. 382— Horn, 1895, p. 11— Casey,
1899, p. 161-Blatchley, 1910, p. 531-Leng, 1920, p. 214-Korschefsky, 1931,
p. 169-Wingo, 1952, p. 45.
Cephaloscymnus zimmermanni zimmermani: Gordon, 1970b, p. 67— Gordon, 1974d,
p. 45.
Diagnosis. Length 2.15 to 2.40 mm, width 1.30 to 1.45 mm. Form elongate (Fig.
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Fig. 31. a-e. Cephaloscymnus zimmermani zimmermani. f. C. z. australis.
29). Color piceous to black dorsally; ventral surface piceous, tarsus yellowish brown.
Male genitalia as in Figure 31a-d. Female genitalia as in Figure 31e. Crotch had
more than one type specimen, but only one female labeled “(yellow disc)/Type 8247/
Cephaloscymnus zimmermanni Crotch” remains in the LeConte collection. I des-
ignate and label that female the lectotype.
Type locality. “Central Valley” (Ohio, Illinois, etc.) (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 32. DISTRICT OF COLUMBIA. INDIANA (state record).
MARYLAND: Beltsville. NEW JERSEY: Montclair. SOUTH CAROLINA: (state
record). TENNESSEE: Oak Ridge. VIRGINIA: Falls Church; Winchester. WEST
VIRGINIA: Berkley.
Cephaloscymnus zimmermanni australis Gordon
Fig. 3 If; Map, Fig. 32
Cephaloscymnus zimmermanni australis Gordon, 1970b, p. 67.
Diagnosis. Length 2.20 to 2.36 mm, width 1.38 to 1.60 mm. Color piceous to
brown dorsally, pronotum red; venter black except leg, mouthparts and epipleuron
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Fig. 32. Distribution. Cephaloscymnus z. zimmermanni (dot); C z. australis (star); C. oc-
cidentalis (shaded, disjunct locality with circled star); C. laevis (open circle); C. insulatus (square).
yellowish brown. Male genitalia as illustrated for zimmermanni zimmermanni. Fe-
male spermathecal capsule as in Figure 3 If. See Gordon (1974d) for detailed dis-
cussion.
Type locality. Kerrville, Texas.
Type depository. USNM (70399).
Distribution. Figure 32. ARIZONA: Chiricahua Mts.; Cochise Co., Palmerlee;
Huachucha Mts., Millers Canyon. NEW MEXICO: Las Vegas. TEXAS: Kerrville;
Mountain Home.
Cephaloscymnus occidentalis Horn
Fig. 33a-e; Map, Fig. 32
Cephaloscymnus occidentalis Yiom, 1895, p. Ill— Casey, 1899, p. 161— Leng, 1920,
p. 214-Korschefsky, 1931, p. 169-Gordon, 1970b, p. 69-Gordon, 1974d, p.
46.
Diagnosis. Length 1 .85 to 2. 1 0 mm, width 1 . 1 0 to 1 .40 mm. Color brown dorsally,
pronotum reddish brown; venter piceous, leg brown. Male genitalia as in Figure 33a-
c. Female genitalia as in Figure 33e.
Discussion. Horn apparently had more than one specimen when he described C.
occidentalis, but only one specimen, a female labeled “425/Los Angeles Cal/Lectotype
3030/C. occidentalis Horn” remains in his collection, I designate and label this
specimen the lectotype.
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Type locality. Los Angeles, California (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 32. Arizona to California, also Texas (Brownsville).
Cephaloscymnus insulatus Gordon
Fig. 34a-d; Map, Fig. 32
Cephaloscymnus insulatus Gordon, 1970b, p. 69.
Diagnosis. Length 2.00 to 2. 10 mm, width 1 . 10 to 1 .30 mm. Color brown dorsally.
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Fig. 34. Cephaloscymnus insulatus.
pronotum reddish; venter piceous, legs, mouthparts, and epipleuron brown. Male
genitMia as in Figure 34a-d.
Type locality. Santa Rita Mts., Arizona.
Type depository. USNM (70400).
Distribution. Figure 32. ARIZONA: Oracle; Santa Rita Mts., Box Canyon.
Cephaloscymnus laevis Gordon
Fig. 35a-e; Map, Fig. 32
Cephaloscymnus laevis Gordon, 1970b, p. 70— Gordon, 1974d, p. 46.
Diagnosis. Length 2.00 mm, width 1.15 mm. Color light brown dorsally; venter
black, mouthparts, leg, and epipleuron light brown. Male genitalia as in Figure 35a-
d. Female genitalia as in Figure 35e.
Discussion. This species was originally described from a unique male from Nogales,
Arizona. Gordon (1974d) recorded 2 specimens of C. laevis from Hidalgo, Mexico.
Type locality. Nogales, Santa Cruz Co., Arizona.
Type depository. CAS.
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Distribution. Figure 32. ARIZONA: Pima Co.; Santa Rita Exp. Range. MEXICO:
Hidalgo.
Subfamily Scymninae
Scymninae Della Beffa, 1912, p. 168 — Sasaji, 1968, p. 23— J. Chapin, 1974, p. 15.
Coccinellidae with dorsal surface pubescent (Scymnini, Selvadiini, Blaisdelliana,
Zagloba) or glabrous (Hyperaspini, Zilus)\ size small. Antenna very short, usually %
or less the length of head, inserted ventrally. Terminal segment of maxillary palpus
not strongly securiform, usually parallel sided or barrel shaped. Mentum broadly
articulated with submentum. Epipleuron of elytron narrow, short. Middle coxae
broadly separated. Each femur nearly cylindrical, stout, occasionally flattened. Tarsus
cryptotetramerous or trimerous.
This subfamily contains the small, compact coccinellids as exemplified by members
of the genera Scymnus and Hyperaspis. Della Beffa (1912) was the first to group the
mostly pubescent Scymnini and usually glabrous Hyperaspini together, and this view
was recently reinforced by Sasaji ( 1 968). In America north of Mexico 5 tribes represent
this subfamily, one of which, the Selvadiini, is erected for the first time. Zilini is
provided as a replacement name for Scymnillini.
Key to tribes of Scymninae
1 . Abdomen with 5 visible sterna 2
- Abdomen with 6 or 7 visible sterna 3
2( 1 ). Prostemum with large anterior lobe concealing mouthparts; Florida . . Cryptognathini
- Prostemum unmodified, not concealing mouthparts; Florida and elsewhere Zilini
3(1). Surface of elytron pubescent 4
Surface of elytron glabrous Hyperaspini
4(3). Anterior margin of prostemum lobed, at least partially concealing mouthparts (Figs.
48c, 59c) 5
- Anterior margin of prostemum not lobed 6
5(4). Length less than 2.0 mm; pronotum black Stethorini
Length more than 3.0 mm; pronotum reddish yellow
Cryptolaemus montrouzieri Mulsant (Scymnini)
6(4). Head narrow, elongate in front of eye; apex of clypeus strongly emarginate, antero-
lateral angle produced forward (Fig. 292a) Blaisdelliana sexualis Casey (Hyperaspini/1)
- Head broad, not elongate in front of eye; apex of clypeus tmncate or nearly so,
anterolateral angle not produced 7
7(6). Form flattened, nearly parallel sided; eyes small, separated by
3 times the width of an eye; antennal club symmetrical (Fig. 287b) Selvadiini
Form usually convex, rounded; eyes large, separated by twice the width of an eye;
antennal club asymmetrical (Fig. 68a) Scymnini
Tribe Zilini, new name
Scymnillini Casey, 1899, p. 112— Leng, 1920, p. 2 1 4 — Korschefsky, 1931, p. 171 —
Blackwelder, 1945, p. 445-Sasaji, 1971, p. 58-J. Chapin, 1974, p. 46.
Scymninae of small size, usually less than 2.30 mm long; form round or elongate,
convex; dorsal surface either distinctly pubescent or apparently glabrous, head and
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Fig. 35. Cephaloscymnus laevis.
anterolateral pronotal angle always pubescent. Head partially inserted in pronotum.
Antenna short, compact, insertion exposed, 10-segmented, club symmetrical. Max-
illary palpus with apical segment cylindrical or slightly securiform. Pronotum deeply
emarginate anteriorly, lateral margin slightly explanate, anterolateral angle produced.
Prostemum with intercoxal process broad, flat, without carinae. Leg free, simple;
tarsus cryptotetramerous; tarsal claw with or without basal tooth. Abdomen with 5
visible sterna, sterna compact and tightly joined. Male genitalia symmetrical, form
simple. Female genitalia with sperm duct short; genital plate elongate, triangular.
There are 2 North American genera in this tribe, Zagloba and Zilus. The tribe is
strictly New World in distribution and forms a tightly knit group of genera and
species. The 5 -segmented abdomen, broad intercoxal process of the prostemum, and
partially concealed head distinguish this tribe from other tribes in the Scymninae.
Examination of species of Zilus and Scymnillus indicates that they are congeneric,
therefore Scymnillus is placed as a junior synonym of Zilus, and the tribal name
changed to Zilini.
Key to genera of Zilini
1 . Elytron apparently glabrous Zilus Mulsant
- Elytron densely pubescent Zagloba Casey
Genus Zilus Mulsant
Scymnus {Zilus) Mulsant, 1850, p. 958 — Korschefsky, 1931, p. 117. Type-species;
Scymnus {Zilus) fulvipes Mulsant, by monotypy.
Zilus: Blackwelder, 1945, p. 445.
Scymnillus Horn, 1895, p. 110— Casey, 1899, p. 114 — Leng, 1920, p. 214— Kor-
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schefsky, 1931, p. 171 — Blackwelder, 1945, p. 445. J. Chapin, 1974, p. 47. Type-
species; Scymnillus aterrimus Horn, by monotypy. New Synonymy.
Scymnillodes Sicard, 1922, p. 355 — Korschefsky, 1931, p. 221— Chapin, 1930, p.
490— Blackwelder, 1945, p. 445 (as synonym of Zilus). Type-species; Scymnillodes
viridimicans Sicard, by subsequent designation of Korschefsky, 1931.
Zilini with length less than 2.0 mm. Dorsal surface often with a metallic tint of
varied colors; pubescence usually limited to head and pronotum with occasional
sparse hairs present on elytron. Antenna extremely short, compact, club apparently
3-segmented (Fig. 36a). Apical segment of maxillary palpus slightly securiform (Fig.
36b). Gena extending onto eye. Tarsal claw with basal tooth (Fig. 36c). Postcoxal
line extending downward from base of first abdominal sternum, joining apex of
sternum nearly at lateral margin (Fig. 36d). Male genitalia simple, symmetrical.
Female genitalia with infundibulum slender, elongate (Fig. 36e).
The key characters will separate Zilus from Zagloba. In addition, Zilus often has
a metallic tint of green, violet, or blue, etc., on the dorsal surface, and the postcoxal
line extends in an arc from the base of the sternum to the posterolateral angle. Most
species of Zilus are neotropical with 4 species recorded from the United States. They
are apparently predators on various scale insects such as Lepidosaphes spp. and
Aspidiotus spp, but one species has been recorded on the whitefly Aleurocanthus
woglumi Ashby. The genus has not been treated taxonomically as a whole.
Key to species of Zilus
1. Length 1.0 mm or less eleutherae (Casey), n. comb.
- Length 1.20 mm or more 2
2(1). Dorsal surface with purple or blue tint; form broad; known only from Florida . . .
subtropicus (Casey), n. comb.
Dorsal surface black or brown; form somewhat elongate; not restricted to Florida 3
3(2). Dorsal surface reddish brown; western United States aterrimus (Horn), n. comb.
- Dorsal surface black; eastern United States horni, n. sp.
Zilus aterrimus (Horn), new combination
Fig. 37a-f; Map, Fig. 39
Scymnillus aterrimus Horn, 1895, p. 110— Casey, 1899,p. 115 — Leng, 1920, p. 214—
Korschefsky, 1931, p. 171— Hatch, 1961, p. 154.
Scymnillus cochisensis Nunenmacher, 1912, p. 451. New Synonymy.
Diagnosis. Length 1.25 to 1.60 mm, width 0.90 to 1.35 mm. Form elongate, oval
(Fig. 37f). Color reddish brown except antenna, mouthparts and leg yellowish brown.
Male genitalia as in Figure 37a-d. Female genitalia as in Figure 37e.
Discussion. I cannot separate Z. cochisensis (Nunenmacher) from Z. aterrimus;
therefore, I place Z. cochisensis as a junior synonym of Z. aterrimus. Nunenmacher
stated that he had 20 cotypes of S. cochisensis, 2 of which (male and female) are
now in the California Academy of Science. I here designate and label the female as
the lectotype and the male as a paralectotype. Horn had more than one specimen of
S. aterrimus, and there are 3 specimens now in his collection, the first of these, a
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c d
Fig. 36. Zilus sp. a. Maxillary palpus, b. Antenna, c. Tarsus, d. Postcoxal line.
female labeled “Oregon Koebele/40/Lectotype 3185/Scymnillus aterrimus Horn”, I
designate the lectotype, the remaining 2 are designated as paralectotypes.
Type locality. Of aterrimus, Oregon (lectotype here designated); of cochisensis,
Benson, Cochise Co., Arizona (lectotype here designated).
Type depository. Of aterrimus, MCZ; of cochisensis, CAS.
Distribution. Figure 39. Idaho and Washington to California and Arizona.
Zilus horni, new species
Fig. 38a-f; Map, Fig. 39
Scymnillus aterrimus: J. Chapin, 1974, p. 47 (not S. aterrimus Horn, 1895).
Description. Male, length 1.40 mm, width 1.0 mm. Form oval (Fig. 38f). Color
black except mouthparts, antenna, and leg yellowish brown. Head coarsely punctured,
punctures separated by less than a diameter. Pronotum with coarse punctures as on
head laterally, separated by a diameter or less, discal area finely punctured, punctures
separated by one to 2 times a diameter. Elytron finely punctured as on pronotal disc,
punctures separated by one to 3 times a diameter. Ventral surface smooth, finely
punctured medially, becoming dull with alutaceous sculpture and coarse punctures
laterally. Genitalia as in Figure 36a-d.
Female, similar to holotype except length 1.50 mm, width 1.10 mm. Genitalia as
in Figure 36e.
Variation. Length 1.40 to 1.60 mm.
Holotype. Male. MARYLAND: Piney Pt., Coll. Hubbard & Schwarz (USNM
101330).
Allotype. Female. MARYLAND: SI Java Farm Biol. Survey, 16:VII:1968, RE &
Jan White Collectors. (USNM).
Paratypes. Total 23. MARYLAND: same data as holotype; same data as allotype;
College Park, X-2-1960, P. J. Spangler. (USNM).
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Fig. 37. Zilus aterrimus.
Distribution. Figure 39. Maryland to Florida, west to Wisconsin. Disjunct localities:
LOUISIANA: Caddo Parish; East Baton Rouge Parish; Rapides Parish.
This eastern species has been confused with Z. aterrimus (Horn) although the
distributions are disjunct. In addition to differences in male and female genitalia,
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79
Fig. 38. Zilus horni.
these 2 species are also separable externally. Zilus horni is entirely black dorsally,
and the pronotum is finely punctate medially. Zilus aterrimus is reddish brown
dorsally, and the pronotum is closely, coarsely punctate throughout. The specific
epithet is in honor of George H. Horn.
Zilus eleutherae (Casey), new combination
Fig. 40a-e; Map, Fig. 39
Scymnillus eleutherae CdiSQy, 1899, p. 1 15 — Korschefsky, 1931, p. 171— Blatchley,
1920, p. 44.
Diagnosis. Length 0.90 to 1.0 mm, width 0.78 to 0.80 mm. Form round, convex
(Fig. 40e). Color purplish black; lateral pronotal border, ventral surface, and leg
(except tarsus) dark brown; antenna, mouthparts and tarsus yellow. Male genitalia
as in Figure 40a-c.
Discussion. This minute species was described from the Bahamas and first recorded
from Florida by Blatchley (1920). The size and muted purplish black dorsum char-
acterize Z. eleutherae in the North American fauna. There are 3 types in the Casey
collection, the first of which I designate and label the lectotype and the other 2 as
paralectotypes.
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Fig. 39. Distribution. Zilus aterrimus (shaded, western); Z. horni (shaded, eastern; disjunct
localities, star); Z. eleutherae (circled star); Z. subtropicus (dot).
Type locality. Eleuthera, Bahamas (lectotype here designated).
Type depository. USNM.
Distribution. Figure 39. FLORIDA: Cape Sable.
Zilus subtropicus (Casey), new combination
Fig. 41a-f; Map, Fig. 39
Delphastus subtropicus Casey, 1924, p. 170— Korschefsky, 1931, p. 221.
Scymnillodes subtropicus: Chapin, 1930, p. 493.
Diagnosis. Length 1.60 to 1.80 mm, width 1.28 to 1.42 mm. Form broad, oval
(Fig. 401). Color metallic purple or blue, pronotum often metallic green; ventral
surface yellow to reddish piceous, leg and mouthparts yellowish brown. Male genitalia
as in Figure 41a-d. Female genitalia as in Figure 41e.
Discussion. The metallic blue or purple dorsal color is very distinctive among
North American coccinellids but is shared with several other members of this genus
that occur in the West Indies. This species is apparently restricted to southern Florida
but may also occur in the West Indies. The type specimen is a unique female in the
Casey collection (holotype).
Type locality. Key West, Florida.
Type depository. USNM (35228).
Distribution. Figure 39. FLORIDA: Biscayne; Coral Gables; Davie; Florida City;
Fort Pierce; Hialeah; Key West; Miami; Paradise Key; Vero Beach.
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NORTH AMERICAN COCCINELLIDAE
81
Fig. 40. Zilus eleutherae.
Genus Zagloba Casey
Zagloba Casey, 1899, p. 113 — Leng, 1920, p. 214— Korschefsky, 1931, p. 172-
Hatch, 1961, p. 154 — Gordon, 1970g, p. 481. Type-species; Cephaloscymnus or-
natus Horn, by subsequent designation of Korschefsky, 1931.
Zilini with length usually less than 2.00 mm. Dorsal surface without metallic tint,
pubescence dense, mostly erect, present throughout. Antenna short, compact, club
distinctly 3-segmented (Fig. 42a). Apical segment of maxillary palpus not securiform,
sides nearly parallel, narrowed slightly at apex (Fig. 42b). Gena extending onto eye.
Tarsal claw with basal tooth (Fig. 42c). Postcoxal line complete or incomplete (Figs.
43f, 46f), never reaching apex of first abdominal sternum. Male genitalia simple,
symmetrical. Female genitalia with infundibulum usually large, flattened laterally,
sperm duct very short (Fig. 44e).
The described species of Zagloba occur from Venezuela and Colombia north to
Oregon and Pennsylvania with 3 known from the neotropics and 4 from the United
States. These species are not commonly collected and I have not seen host data for
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Fig. 4 1 . Zilus subtropicus.
any of the United States species. At least one Neotropical species, Z. obscura Gordon,
has been taken feeding on “scale insects” on banana and orange. We may presume,
therefore, that all species of Zagloba are likely to be scale predators. Zagloba has
not been taxonomically treated as a whole, but Gordon (1970g) reviewed the Central
and South American species.
Fig. 42. Zagloba sp. a. Antenna, b. Maxillary palpus, c. Tarsus.
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NORTH AMERICAN COCCINELLIDAE
83
Key to species of Zagloba
1 . Pronotum entirely yellowish red; elytron black or dark brown (Fig. 44f); Rorida
bicolor Casey
Pronotum entirely black or brown, at most with some lateral paler areas; elytron
uniformly dark or dark with yellow maculation; not known from Rorida 2
2(1). Elytron brown with yellow maculation; Pacific Coast, Arizona ornata (Horn)
- Elytron black or brown, immaculate; not known from the Pacific Coast 3
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Fig. 44. Zagloba bicolor.
3(2). Punctures within arc of postcoxal line coarse, distinct; pronotum paler than elytron
hystrix Casey
- Punctures within arc of postcoxal line fine, indistinct; pronotum and elytron con-
colorous satana, n. sp.
Zagloba ornata (Horn)
Fig. 43a-g; Map, Fig. 45
Cephaloscymnus ornatus Horn, 1895, p. 111.
Zagloba ornata: Casey, 1899, p. 114 — Leng, 1920, p. 214— Korschefsky, 1931, p.
172-Hatch, 1961, p. 154.
Zagloba laticollis Casey, 1899, p. 1 14 — Leng, 1920, p. 214. New Synonymy.
Zagloba orbipennis Casey, 1899, p. 114 — Leng, 1920, p. 214. New Synonymy.
Diagnosis. Length 1.75 to 2.00 mm, width 1.43 to 1.65 mm. Form elongate, oval.
Color dark brown to light brown; antenna, mouthparts, and leg yellowish brown;
pronotum often with yellowish brown lateral areas; elytron usually with 2 nebulous,
yellow spots feebly connected (Fig. 43g), but pattern variable as in Figure 43g. Po-
stcoxal line complete {Pullus type) in both sexes (Fig. 431). Male genitalia as in Figure
43a-d. Female genitalia with infundibulum small, elongate (Fig. 43e).
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85
Fig. 45. Distribution. Zagloba ornata (shaded); Z. bicolor (dot); Z. hystrix (open circle); Z.
satana (star).
Discussion. This species is unique among North American members of the genus
in having the postcoxal line complete in both sexes and in having a simple, reduced
infundibulum. Zagloba laticollis Casey and Z. orbipennis Casey are conspecific with
Z. ornata, and I place both names as junior synonyms. Both species were described
from unique specimens in the Casey collection which must be considered holotypes.
Zagloba ornata was described from several specimens, all from California, and I
designate and label as the lectotype a female in the Horn collection labeled “702/
Aug./Siskiyou Co., Cal./lectotype 3186(red paper)/C. omatus Horn.” Three other
type specimens from various California localities are designated as paralectotypes.
Type locality. Of ornata, Siskiyou Co., California (lectotype here designated); of
laticollis, California; of orbipennis, Healdsburg, Sonoma Co., California.
Type depository. Of ornata, MCZ; of laticollis (35234) and orbipennis (35233),
USNM.
Distribution. Figure 45. Southern Arizona and California, north to southwestern
Oregon.
Zagloba bicolor Casey
Fig. 44a-f; Map, Fig. 45
Zagloba bicolor Casey, 1899, p. 114— Leng, 1920, p. 214 — Korschefsky, 1931, p.
172.
Diagnosis. Length 1.65 to 1.85 mm, width 1.22 to 1.33 mm. Form elongate, oval
(Fig. 44f). Color pale yellowish brown; pronotum yellowish red; elytron black or dark
brown; meso- and metastemum and first abdominal sternum dark brown. Postcoxal
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line incomplete in both sexes. Male genitalia as in Figure 44a-d. Female genitalia
with infundibulum large, flattened laterally (Fig. 44e).
Discussion. This species is apparently restricted to Florida, and the color pattern
alone will distinguish it from other members of the genus. The type is a unique
(holotype) female in the Casey collection.
Type locality. Capron, Florida.
Type depository. USNM (35236).
Distribution. Figure 45. FLORIDA: Alachua Co., Gainesville; Dunedin; Jefferson
Co., Monticello; Miami; Tampa.
Zagloba hystrix Casey
Fig. 46a-g; Map, Fig. 45
Zagloba hystrix Casey, 1899, p. 114— Leng, 1920, p. 214— Korschefsky, 1931, p.
172.
Diagnosis. Length 1.45 to 1.75 mm, width 1.35 to 1.50 mm. Form rounded,
pronotum and elytron abruptly discontinuous in outline (Fig. 46g). Color medium
reddish brown; antenna, leg and mouthparts yellowish brown; elytron dark brown
to black. Postcoxal line incomplete in both sexes {Scymnus, s. str., type) (Fig. 46f).
Male genitalia as in Figure 46a-d. Female genitalia as in Figure 46e.
Discussion. This species is difficult to separate from Z. satana, n. sp., but the
pronotum is usually distinctly paler than the elytron in this species, and the abdominal
punctation is definitely coarser than in satana. There are 6 type specimens of hystrix
in the Casey collection, and the first of these, a female, is designated and labeled as
the lectotype. The other 5 types bear the same data and are designated as paralec-
totypes.
Type locality. Brownsville, Texas (lectotype here designated).
Type depository. USNM (35237).
Distribution. Figure 45. TEXAS: Brownsville; San Antonio; Zavalla Co., Nueces
River.
Zagloba satana, new species
Fig. 47a-h; Map, Fig. 45
Description. Male, length 1.65 mm, greatest width 1.32 mm. Form rounded (Fig.
47h), outline of pronotum and elytron strongly discontinuous. Color black; ventral
surface and lateral border of pronotum dark reddish brown; antenna, mouthparts,
and leg yellowish brown. Dorsum densely pubescent with grayish white hairs, hairs
erect on pronotum and elytron, appressed on head. Head densely, finely punctured,
punctures separated by a diameter or less. Pronotum the length of elytron; punc-
tures fine, separated by twice a diameter on disc, becoming contiguous along lateral
margin. Elytral punctation finer than on pronotum, punctures separated by less than
to twice a diameter. Metastemum smooth, nearly impunctate medially, punctures
becoming coarse and dense laterally. Abdominal punctation fine, punctures within
arc of postcoxal line indistinct; postcoxal line complete (Fig. 47g). Genitalia as in
Figure 47a-d.
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NORTH AMERICAN COCCINELLIDAE
87
Fig. 46. Zagloba hystrix.
Female, similar to holotype except length 1.60 mm, width 1.29 mm; postcoxal
line incomplete (Fig. 471); genitalia as in Figure 47e.
Variation. Length 1.45 to 1.75 mm, width 1.20 to 1.37 mm.
Holotype. Male. TEXAS: Devils River, V-2-07, E. A. Schwarz coll (USNM 101331).
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Allotype. Female. Same data as holotype except “on Pithecolobium”. (USNM).
Paratypes. (Fig. 45) Total 8. TEXAS: “Texas”; Devils River, V-4-07, FC Pratt
Collector; same data as holotype; Laredo, 28-5, Hubbard and Schwarz. (USNM).
This is the only known species of Zagloba exhibiting sexual dimorphism in the
shape of the postcoxal line. The punctation, both dorsal and ventral, is obviously
finer than in hystrix which satana most closely resembles. Genitalia should be ex-
amined in members of this group to ensure accurate determination. The specific
epithet refers to the type locality.
Tribe Stethorini
Stethorini Dobzhansky, 1924, p. 20— Korschefsky, 1931, p. 1 10— Kapur, 1948, 302—
Sasaji, 1968, p. 23— J. Chapin, 1974, p. 16.
Scymninae of small size, less than 2.0 mm; pubescent dorsally. Antenna 11 -seg-
mented; inserted between eye and clypeus, clypeus not emarginate around base.
Maxillary palpus with terminal segment convergent apically. Prostemum lobed an-
teriorly, partially concealing mouthparts; intercoxal process without carinae. Leg free,
simple; tarsus cryptotetramerous or trimerous. Abdomen with 6 visible sterna.
This tribe contains a single genus, Stethorus, which has usually been placed in the
tribe Scymnini. Dobzhansky (1924) erected the tribe Stethorini, but Korschefsky
(1931) synonymized Stethorini with Scymnini, and Kapur (1948) agreed with this
placement. Sasaji (1968) considered Stethorini a valid tribe and I concur with his
treatment.
Stethorini is easily separated from all other tribes of Scymninae because the clypeus
is not emarginate around the antennal bases, and the prostemum is arcuately pro-
duced in front, partly concealing the mouthparts.
Genus Stethorus Weise
Stethorus 1885a, p. 65 — Casey, 1899, p. 135 — Kapur, 1948, p. 300— Wingo,
1952, p. 19— J. Chapin, 1974, p. 16 — Belicek, 1976, p. 297— Gordon and Ander-
son, 1979, p. 61— Gordon and Chapin, 1983, p. 229. Type-species; Stethorus
punctillum Weise, by subsequent designation of Korschefsky, 1931.
NephopullusBYQX\\Qs, 1925, p. 167 — Kapur, 1948, p. 300. Type-species;
darwini Brethes, by subseqent designation of Korschefsky, 1931.
Body color black except antenna and mouthparts yellow, legs often yellow. Head
with moderately coarsely faceted eye; clypeus truncate anteriorly, anterolateral angle
rounded. Antenna short, 1 1 -segmented (Fig, 48a); inserted between eye and clypeus,
clypeus not emarginate around base. Maxillary palpus with apical segment oblong,
obliquely truncate and narrower toward apex (Fig. 48b). Prostemum without carinae,
produced anteriorly to partly conceal mouthparts (Fig. 48c). Tarsus trimerous or
cryptotetramerous; tarsal claw bifid (Fig. 48d), inner claw shorter in male than in
female. Abdomen with postcoxal line on basal sternum complete (Fig. 49e). Male
genitalia with basal lobe symmetrical or asymmetrical. Female spermathecal capsule
present or absent, genital plate small, not triangular (Fig. 48e).
There are 65 described species in this genus, and they are found in most parts of
the world; 6 species occur in America north of Mexico. Most coccinellids are pre-
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89
Fig. 47. Zagloba satana.
daceous on insects of the order Homoptera, and some are plant feeders, but species
of Stethorus feed almost exclusively on tetranychid mites. The western Hemisphere
species were treated by Gordon and Chapin (1983), see that publication for more
detailed information.
Key to species of Stethorus
1 . Postcoxal line not arched beyond middle of first abdominal sternum (figs. 54d); basal
abdominal sternum densely, coarsely punctured 2
Postcoxal line arched beyond middle of first abdominal sternum (Figs. 51e, 53e);
basal abdominal sternum sparsely, finely punctured 3
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2(1). Postcoxal line short, arc not reaching middle of first abdominal sternum; elytral
punctures larger than pronotal punctures punctillum Weise
- Postcoxal line usually arched to middle of first sternum or nearly so (Fig. 53e); elytral
and pronotal punctures equal in size punctum punctum (LeConte)
3(1). Leg (except tarsus) black or brown; punctures on abdominal sterna coarse, dense
(Fig. 53g) punctum picipes CasGy
Leg with at least tibia yellow; punctures on abdominal sterna fine, sparse (Fig. 49e)
4
4(3). Clypeal apex truncate; lateral pronotal punctures dense, contiguous
pinachi Gordon and Chapin
Clypeal apex emarginate; lateral pronotal punctures sparse, not contiguous 5
5(4). Elytral pubescence reddish brown; pronotal punctures fine, sparse
caseyi Gordon and Chapin
- Elytral pubescence yellowish white; pronotal punctures coarse, not sparse utilis (Horn)
Stethorus utilis (Horn)
Fig. 49a-e; Map, Fig. 52
Scymnus utilis Horn, 1895, p. 107.
Stethorus utilis: Casey, 1899, p. 136 — Korschefsky, 1931, p. 112— J. Chapin, 1974,
p. 17— Gordon and Chapin, 1983, p. 241.
Stethorus atomus Casey, 1899, p. 136 — Korschefsky, 1931, p. Ill— Gordon and
Chapin, 1983, p. 241.
Diagnosis. Length 1.0 to 2.0 mm, width 0.75 to 1.0 mm. Form elongate, oval.
Color black; antenna, mouthparts, and leg yellow except basal % of femur brown.
Dorsal pubescence moderately long, semierect, mostly yellowish white with traces
of brown. Head finely punctured, punctures separated by a diameter or less; pronotal
punctures coarse, slightly larger than elytral punctures, separated by about a diameter
on disc, less than a diameter laterally; elytral punctures shallow, separated by one to
2 times a diameter; metastemum with fine, dense punctures except nearly impunctate
on basomedian area; abdominal sterna finely, sparsely punctured. Arc of postcoxal
line extending % length of first abdominal sternum, angulate (Fig. 49e). Apex of 6th
abdominal sternum truncate. Male genitalia as in Figure 49a-c. Female spermathecal
capsule as in Figure 49d.
Discussion. This species is most easily confused with S. caseyi, but the dorsal
pubescence of S. caseyi is reddish brown. The male genitalia are similar in these 2
species, but the basal lobe in S. caseyi is more obviously triangular and shorter than
that of S. utilis.
Type locality. Of utilis, Barstow, Florida; of atomus, Columbus, Texas.
Type depository. Of utilis, MCZ; of atomus, USNM.
Distribution. Figure 52. North Carolina to Florida, west to east Texas.
Stethorus caseyi Gordon and Chapin
Fig. 50a-f; Map, Fig. 52
Stethorus Caseyi Gordon and Chapin, 1983, p. 241.
Diagnosis. Length 1.10 to 1.31 mm, width 0.75 to 1.05 mm. Form short, rounded
(Fig. 50f). Color black; antenna, mouthparts and leg except basal % of femur yellow.
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91
Fig. 48. Stethorus sp. a. Antenna, b. Maxillary palpus, c. Venter, d. Tarsus, e. Genital plates.
Dorsal pubescence long, nearly erect, reddish brown. Head shiny, finely punctured,
punctures separated by a diameter; pronotum with punctures coarser than on head,
punctures separated by one to 3 times a diameter; elytral punctation coarse, punctures
separated by about a diameter; metastemum with fine punctures medially, punctures
becoming coarse and dense laterally. Abdominal sterna with fine punctures sparse
on first sternum, dense on remaining sterna. Arc of postcoxal line extending Vt length
of first abdominal sternum, angulate (Fig. 50e). Apex of 6th abdominal sternum
feebly notched. Male genitalia as in Figure 50a-c. Female genitalia as in Figure 50d.
Discussion. The round form, reddish brown pubescence, and sparsely punctured
pronotum will separate S. caseyi from S. utilis which it most closely resembles.
Type locality. Devils River, Texas.
Type depository. USNM (10061).
Distribution. Figure 52. ARIZONA: Catalina Springs; Chiricahua Mountains; Or-
acle; Santa Rita Mountains. NEW MEXICO: Albuquerque. TEXAS: Brownsville;
Devils River; El Paso; Laredo; San Antonio; San Diego; Uvalde. UTAH: Leeds; St.
George.
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Fig. 49. Stethorus utilis.
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Fig. 50. Stethorus caseyi.
Stethorus pinachi Gordon and Chapin
Fig. 51a-e; Map, Fig. 52
Stethorus pinachi: Gordon and Chapin, 1983, p. 250.
Diagnosis. Length 1.25 to 1.40 mm, width 0.80 to 1.10 mm. Form elongate, oval.
Color black; antenna, mouthparts, and leg yellow except basal % of femur brown.
Dorsal pubescence short, semierect, yellowish white with traces of brown. Head shiny,
finely punctured, punctures separated by a diameter or more; pronotal punctation
coarser than on head, discal punctures separated by a diameter, lateral punctures
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Fig. 5 1 . Stethorus pinachi.
contiguous; elytral punctation very coarse, punctures separated by less than a di-
ameter. Metasternum coarsely and densely punctured laterally, punctures finer and
sparser medially; abdominal sterna with fine punctures sparse on first sternum, dense
on remaining sterna. Arc of postcoxal line extending more than length of first
abdominal sternum, rounded (Fig. 51e). Sixth sternum feebly emarginate apically.
Male genitalia as in Figure 51a-c. Female genitalia as in Figure 5 Id.
Discussion. This species is quite distinctive in the form of the male genitalia which
are most similar to those of S. punctum. The truncate apex of the clypeus will
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Fig. 52. Distribution. Stethorus utilis (shaded); S. caseyi (dot); S. pinachi (open circle).
distinguish S. chapini from S. utilis which it most closely resembles in external
appearance.
Type locality. Carrizo Springs, Dimmit Co., Texas.
Type depository. USNM (100664).
Distribution. Figure 52. TEXAS: type locality.
Stethorus punctum punctum (LeConte)
Fig. 53a-e; Map, Fig. 55
Scymnus punctum LeConte, 1852, p. 141— Horn, 1895, p. 106.
Stethorus punctum: Casey, 1899, p. 136 — Korschefsky, 1931, p. 112 — Wingo, 1952,
p. 27— J. Chapin, 1974, p. 17— Gordon and Chapin, 1983, p. 250.
Diagnosis. Length 1.35 to 1.55 mm, width 0.95 to 1.15 mm. Form elongate, oval.
Color black; antenna, mouthparts, and leg yellow except femur usually brown. Dorsal
pubescence short, semierect, yellowish white. Head finely punctured, punctures sep-
arated by more than a diameter; pronotum finely, densely punctured, punctures
separated by a diameter on disc, contiguous laterally; elytral punctures subequal in
size to those on pronotum, separated by a diameter or less; metastemum coarsely
punctured anteriorly and laterally; abdominal sterna with coarse, dense punctures
separated by less than a diameter. Arc of postcoxal line usually reaching middle of
basal abdominal sternum, sometimes shorter (Fig. 53e). Apex of 6th abdominal
sternum notched. Male genitalia as in Figure 53a-c. Female spermathecal capsule as
in Figure 53d.
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Discussion. This species is native to North America, but somewhat difficult to
distinguish from the introduced punctillum without examination of genitalia which
are highly distinctive in the males. The postcoxal line is shorter in S. punctillum than
in S. punctum, and the elytral punctures are distinctly larger than the pronotal punc-
tures in S. punctillum, nearly equal in S. punctum. The female of punctillum lacks a
spermathecal capsule.
Type locality. Ontario, northern shore of Lake Superior.
Type depository. MCZ.
Distribution. Figure 55. Southeastern Canada to North Carolina, west to Montana
and Colorado.
Stethorus punctum picipes Casey
Fig. 53f, g; Map, Fig. 55
Stethorus picipes Casey, 1899, p. 136 — Korschefsky, 1931, p. 112 — Hatch, 1961, p.
149 — Belicek, 1976, p. 298 — Gordon and Chapin, 1983, p. 252.
Stethorus brevis Casey , 1899, p. 1 36 — Korschefsky, 1931, p. 1 1 1 —Gordon and Chap-
in, 1983, p. 252.
Diagnosis. Description as for punctum except the leg black or dark brown (except
tarsus), the ventral punctation is noticeably more coarse and dense, and the postcoxal
line (Fig. 53g) extends beyond the middle of the basal abdominal sternum. Female
spermathecal capsule as in Figure 53f
Discussion. The male and female genitalia are identical in punctum and picipes,
but the 2 nominate forms can be distinguished on the basis of the characters men-
tioned above. I prefer to treat them as subspecies with punctum occurring from the
east coast to Colorado and Montana and picipes occurring from California and British
Columbia to Idaho and Alberta.
Type locality. Of picipes, Santa Rosa, Sonoma Co., California; of brevis, Siskiyou
Co., California.
Type depository. Of picipes and brevis, USNM.
Distribution. Figure 55. Idaho to British Columbia, south to southern California.
Stethorus punctillum Weise
Fig. 54a-d; Map, Fig. 56
Stethorus punctillum Weise, 1891, p. 391 {in Reitter et al.)— Casey, 1899, p. 136 —
Korschefsky, 1931, p. 112 — Kapur, 1948, p. 302 — Hatch, 1961, p. 149 — Belicek,
1976, p. 298— Gordon and Chapin, 1983, p. 270.
Coccinella minima Rossi, 1794, p. 89 (not Coccinella minima Muller, 1776).
Scymnus {Stethorus) minimus: Weise, 1885a, p. 74.
Coccinella pusilla Herbst, 1797, p. 346 (not Coccinella pusilla Muller, 1776).
Coccinella atra Illiger, 1798, p. 413 (not Coccinella atra Gmelin, 1790).
Stethorus at er: Korschefsky, 1931, p. 112.
Diagnosis. Length 1.35 to 1.57 mm, width 0.90 to 1.12 mm. Form elongate, oval.
Color black, antenna, mouthparts, and leg brownish yellow except basal Yt of femur
brown. Dorsal pubescence short, semierect, yellowish white. Punctation on head and
pronotum fine, pronotal punctures separated by about a diameter; elytral punctures
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Fig. 53. a-e. Stethorus punctum punctum. f, g. S. p. picipes
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Fig. 54. Stethoms punctillum.
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Fig. 55. Distribution. Stethorus punctum punctum (shaded, eastern); S. p. picipes (shaded,
western).
coarse, separated by less then a diameter; abdominal sterna with coarse, dense punc-
tures separated by less than a diameter. Arc of postcoxal line short, not reaching
middle of basal abdominal sternum, rounded (Fig. 54d). Male genitalia as in Figure
54a-c. Female genitalia lacking a spermathecal capsule and infundibulum.
Discussion, This species is apparently a European introduction, but not an inten-
tional one. Brown (1950) first reported it from North America (Framingham, Mass.;
Vineland Station and Leamington, Ontario) and gave a key to separate S. punct ilium,
S. punctum, and S. picipes. Stethorus punctillum is now known from several North
American localities and is often mixed with S. punctum in collections. The species
has become established on the west coast of the United States, but again it was not
intentionally introduced.
Type locality. Not stated.
Type depository. Type not examined.
Distribution. Figure 56. Eastern: southeastern Canada to Massachusetts, west to
Michigan and Wisconsin. Western: British Columbia (Vancover), to Oregon.
Tribe Scymnini
Scymnini Costa 1849, p. 9— Weise, 1895, p. 147— Casey, 1899, p. 133— Mader,
1924,p. 8-Leng, 1920, p. 213-Korschefsky, 1931, p. 110-Wingo, 1952, p. 19-
Sasaji, 1968, p. 23-J. Chapin, 1974, p. 18-Belicek, 1976, p. 295.
Scymninae of small size, usually less then 3.0 mm; form oval, rounded, or oblong;
dorsal surface and eye pubescent. Antenna 8 to 1 1 segmented, terminal segments
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forming distinct club. Maxillary palpus with apical segment cylindrical or securiform.
Leg free, simple, not expanded or enlarged; tarsus trimerous or cryptotetramerous,
tarsal claw simple or with basal tooth. Abdomen with 6 visible sterna, sterna usually
not fused medially, apex of 6th abdominal sternum of male modified. Male genitalia
vary from symmetrical to asymmetrical, form simple (Fig. 93b), or complex (Fig.
1 90a). Female genitalia with sclerotized infundibulum; genital plate long and narrow,
or short, nearly round.
Gordon (1976b) included the genera Selvadius and Blaisdelliana in this tribe. I
now consider Blaisdelliana a member of the Hyperaspini and erect the tribe Selvadiini
for Selvadius. There remain 6 North American genera in the Scymnini: one, Cryp-
tolaemus, is introduced; another, Didion, is apparently endemic; and the other 4 are
worldwide in distribution. Three of these genera (Scymnus, Nephus, and Diomus)
have previously been treated by Gordon (1976b); therefore, the descriptions of the
species are not included here, but the keys, illustrations, and synonymies are repeated.
Key to genera of Scymnini
1. Head with mouthparts directed postero-ventrad in repose, concealing prostemum;
basal antennal segment strongly enlarged (Fig. 57c) Nephaspis Casey
Head with mouthparts not concealing prostemum; basal antennal segment not strongly
enlarged 2
2(1). Prostemum enlarged, expanded, capable of concealing mouthparts in repose (Fig.
59c) Cryptolaemus Mulsant
Prostemum not enlarged, not concealing mouthparts 3
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Fig. 57. Nephaspis oculatus. a. Venter, b. Maxillary palpus, c. Antenna, d. Postcoxal line,
e-g. Male genitalia, h. Spermathecal capsule, i-1. Habitus and variations.
3(2). Prostemum with distinct carinae on intercoxal projection, carinae often reaching
anterior margin of prostemum (Fig. 68 c)
Prostemum without carinae, or at most with short ridges next to coxal cavities (Fig.
229g)
5
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4(3). Postcoxal line extending downward, joining hind margin of first abdominal sternum
(Fig. 257b), apex not recurved Diomus Mulsant
Postcoxal line complete or incomplete, not joining hind margin of first abdominal
sternum, apex recurved (Fig. 268e, g) Scymnus Kugelann
5(3). Postcoxal line complete, recurved to base of first abdominal sternum (Fig. 62e) . .
Didion Casey
- Postcoxal line extending nearly to lateral margin of first abdominal sternum, either
parallel to hind margin (Fig. 229j), or with apex curved forward (Fig. 229h)
Nephus Mulsant
Genus Nephaspis Casey
Nephaspis Casey, 1899, p. 168 — Casey, 1905, p. 161— Wingo, 1952, p. 44— Gordon,
1972b, p. 145— J. Chapin, 1974, p. 37 — Gordon, 1976b, p. 8. Type-species; Ne-
phaspis gorhami Casey, by subsequent designation of Gordon, 1972b.
Nephasis: Korschefsky, 1931, p. 168 — Blackwelder, 1945, p. 445 (error).
Scymnini with form elongate, somewhat oval; length less than 1.60 mm. Head with
mouthparts directed posteroventrally in repose, concealing prostemum (Fig. 57a);
clypeus extending beyond eye, anterolateral angle produced, rounded, anterior margin
truncate, lateral margin emarginate at antennal insertion; gena partially dividing eye.
Maxillary palpus with apical segment somewhat securiform (Fig. 57b). Antenna with
8-segmented scape, basal 2 segments enlarged, club 3-segmented (Fig. 57c). Pronotum
widest at posterolateral angle, narrowed apically. Prostemum short, only slightly
longer than anterior coxa, intercoxal process narrow, apex truncate. Metastemum
tumid. Front and middle femora slender, not enlarged; hind femur enlarged medially;
all tibiae slender; tarsus cryptotetramerous, claw simple, not toothed. Abdomen with
6 visible sterna; postcoxal line as in Scymnus {S. str.) (Fig. 57d). Male genitalia
symmetrical (Fig. 57e-g). Female genitalia with distinctly sclerotized spermathecal
capsule, infundibulum absent; genital plate long, slender.
The extremely large basal antennal segment, strongly tumid sternum and postero-
ventrally directed mouthparts characterize this genus. It is unlike any other Western
Hemisphere genus in these respects, being similar only to the Old World genus
Clitostethus.
The 4 known species are all entirely neotropical except N. oculatus which is es-
tablished in the United States. This species is probably native to Central America
and may have entered the West Indies and the United States on imported plant
materials. It is well established in Florida, and Wingo (1952) described it as N.
amnicola from specimens taken in Iowa. All available host data indicate that mem-
bers of this genus are predators on whiteflies of the family Aleurodidae. Specific host
records are: Aleurodicus dispersus Russell and A. cocois (Curtis). This genus was
revised by Gordon (1972b).
Nephaspis oculatus (Blatchley), new combination
Fig. 57e-l; Map, Fig. 58
Scymnus oculatus Blatchley, 1917, p. 140.
Nephaspis amnicola 1952, p. 44— Gordon, 1972b, p. 149— J. Chapin, 1974,
p. 37. New Synonymy.
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Diagnosis. Length 1.19 to 1.48 mm, width 0.79 to 1.00 mm. Color yellow; elytron
usually piceous to black along base and lateral margin, central area yellowish brown,
apex narrowly yellow (Fig. 57j); elytron in male is varied from completely black
(except apical yellow area) to black or piceous with red or yellow discal spot (Fig.
57i-l). Discal spot small and round, or elongate. Male genitalia as in Figure 57e-g.
Female genitalia as in Figure 57h.
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Fig. 59. Cryptolaemus montrouzieri. a. Antenna, b. Maxillary palpus, c. Venter, d. Tarsus,
e. Postcoxal line. f. Female genitalia.
Discussion. The name Scymnus oculatus Blatchley was overlooked during prepa-
ration of the revision of the genus Scymnus (Gordon, 1976b) and was brought to
my attention by Herbert Dozier. Examination of the holotype revealed that S. ocu-
latus is the same species later described as amnicola Wingo.
Type locality. Of oculatus, Dunedin, Florida; of amnicola, Iowa, Boone, Ledges
State Park.
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Type depository. Of oculatus, PU; of amnicola, USNM.
Distribution. Figure 58. FLORIDA: distributed throughout the state. IOWA: Boone,
Ledges State Park. LOUISIANA: Pointe Coupee Parish. NEW HAMPSHIRE: Web-
ster. TEXAS: Cameron Co., La Feria; Weslaco.
Genus Cryptolaemus Mulsant
Cryptolaemusyi\x\s2ini, 1853, p. 140— Crotch, 1874b, p. 204 — Leng, 1920, p. 214—
Korschefsky, 1931, p. 169— Wingo, 1952, p. 22 — Chapin, 1965, p. 198— J. Chapin,
1974, p. 38. Type-species; Cryptolaemus montrouzieri Mulsant, by monotypy.
Scymnini with length more than 3.00 mm; form oval, convex. Antenna with 7-
segmented scape, club 3-segmented, loose (Fig. 59a). Maxillary palpus with apical
segment securiform (Fig. 59b). Prostemum broadly rounded anteriorly and produced
to cover mouthparts and antenna (Fig. 59c); carinae weak, parallel, extending less
than halfway to anterior margin of prostemum. Tibial spurs absent; tarsus trimerous;
tarsal claw with broad basal tooth equal to half the length of claw (Fig. 59d). Abdomen
with postcoxal line complete, as in Scymnus (Pullus) (Fig. 59e). Male genitalia with
basal lobe symmetrical. Female genitalia with strong spermathecal capsule; sperm
duct short; infundibulum reduced to a small sclerite at head of bursa; genital plates
long, triangular (Fig. 59f).
Cryptolaemus is a small genus of the Indo-Australian region. The only species
present in the Western Hemisphere is C. montrouzieri which was introduced as a
scale predator. The expanded prostemum and large size readily separate Cryptolae-
mus from other genera of New World Scymnini. In my previous key (Gordon, 1 976b)
to the genera of Scymnini, I inadvertently omitted this genus. Specific host records
are as follows: Chloropulvinaria psidii (Maskell); Chrysomphalus /7z>2«w///^r (Maskell);
Coccus viridis (Green); Dactylopius confusus (Cockerell); Dactylopius opuntiae (Cock-
erell); Dactylopius tomentosus (Lamarck); Dysmicoccus boninsis (Kuwana); Dysmi-
coccus brevipes (Cockerell); Eriococcus araucariae (Maskell); Ferrisia virgata (Cock-
erell); Nipaecoccus aurilanatus (Maskell); Nipaecoccus filamentosus (Cockerell);
Nipaecoccus nipae (Maskell); Planococcus citri (Risso); Planococcus krauhniae (Ku-
wana); Planococcus vitis (Neidielski); Pseudococcus calceolariae (Maskell); Pseudo-
coccus comstocki (Kuwana); Pseudococcus crotonis (Green); Pseudococcus hirsutus
(Green); Pseudococcus longispinus (Targioni-Tozzetti); Pseudococcus maritimus (Ehr-
hom); Pseudococcus obscurus (Essig); Pulvinaria icerya (Guerin); Pulvinaria psidii
(Maskell); Rastrococcus iceryoides (Green); Saccharicoccus sacchari (Cockerell);
Trionymus insularis (Ehrhom). Ghorpade (198 1) recorded C. montrouzieri as feeding
on Aphis gossypii Glover in India.
Cryptolaemus montrouzieri Mulsant
Fig. 60a-e; Map, Fig. 61
Cryptolaemus montrousieri Mulsant, 1853, p. 140.
Cryptolaemus montrouzieri: Crotch, 1874b, p. 204 (emendation)— Leng, 1920, p.
214-Korschefsky, 1931, p. 169-Wingo, 1952, p. 45-Chapin, 1965, p. 199-J.
Chapin, 1974, p. 38.
Diagnosis. Length 3.40 to 4.50 mm, width 2.40 to 3.10 mm. Head, prothorax, tip
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Fig. 60. Cryptolaemus montrouzieri.
of elytron and abdomen reddish yellow; mesostemum and metastemum, leg and
elytron (except tip) black or blackish (Fig. 60e). Punctation of head and pronotum
dense, elytral punctation similar except on humeral callus which is shining, almost
devoid of punctures. Male genitalia as in Figure 60a-d.
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Fig. 61. Distribution. Cryptolaemus montrouzieri (shaded, disjunct localities dotted).
Discussion. This species is well established in California and south and central
Florida. There are 2 syntypes of montrouzieri, one in the UCCC collection, one in
the PM collection (R. D. Pope, pers. comm.).
Type locality. “Australia.”
Type depository. BMNH.
Distribution. Figure 6 1 . CALIFORNIA: San Francisco to San Diego. FLORIDA:
Clearwater. INDIANA: Lafayette (from Wingo, 1952). MISSOURI: Washington
(from Wingo, 1952).
Genus Didion Casey
Didion CdiSQy , 1899, p. 137 — Leng, 1920, p. 2 1 3 — Korschefsky, 1931, p. Ill— Gor-
don, 1976b, p. 8 — Belicek, 1976, p. 299. Type-species; Didion longulum Casey,
by subseqent designation of Korschefsky, 1931.
Scymnini with form elongate, oval; length less than 2.00 mm. Head short, eye
partially concealed by pronotum. Pronotum with lateral margin strongly convergent
apically (except D. nanum), base of pronotum distinctly narrower than base of elytra
(Fig. 63e). Antenna with scape 7-segmented, club 3 segmented, club segments uneven
on lower margin (Fig. 62a). Maxillary palpus with apical segment cylindrical, apex
oblique (Fig. 62b). Apex of prostemum truncate; intercoxal process flat, with a short
Carina next to each coxa (Fig. 62c). Tarsus cryptotetramerous; tarsal claw with strong
basal tooth (Fig. 62d). Abdomen with postcoxal line on basal sternum complete, as
in {Pullus) (Fig. 62e). Male genitalia with basal lobe somewhat triangular in ventral
view, shorter than paramere; paramere broad, apex rounded; trabes longer than
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Fig. 62. Didion sp. a. Antenna, b. Maxillary palpus, c. Prostemum. d. Tarsus, e. Postcoxal
line. f. Female genitalia.
phallobase (Fig. 63a). Female genitalia with spermathecal capsule bent near apex;
accessory gland present; sperm duct short, inserted at base of infundibulum; infun-
dibulum long, slender; coxal plate long, slender, with apical stylus (Figure 621).
Didion is apparently restricted to North America and is represented by 3 species.
No concrete information is available on host preferences of members of this genus,
but Wingo (1952) thought D. punctatum might be feeding on the two-spotted spider
mite. Belicek (1976) listed D. longulum as being on plants infested with spider mites.
Species of Didion are most likely to be confused with members of the genus
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109
Fig. 63. Didion punctatum.
Scymnus, subgenus Pullus, but Didion lacks complete prostemal carinae, has 10-
segmented antennae, usually has the lateral pronotal margin nearly straight and
strongly convergent anteriorly, and has the pronotal base distinctly narrower than
the elytral base.
Key to species of Didion
1. Elytron black with reddish orange discal spot (Fig. 63e) punctatum (Melsheimer)
Elytron immaculate 2
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2(1). Lateral margin of pronotum arcuate (fig. 67f); abdomen densely, coarsely punctured
nanum (LeConte)
Lateral margin of pronotum nearly straight, margins convergent apically (Fig. 63e);
abdomen not densely, coarsely punctured 3
3(2). Punctures on elytron large, deep, distinct; form extremely elongate, narrow (fig. 63e)
punctatum (Melsheimer)
Punctures on elytron small, shallow, indistinct; form oval longulum Casey
Didion punctatum (Melsheimer)
Fig. 63a-f; Map, Fig. 64
Scymnus punctatus Melsheimer, 1847, p. 80— Horn, 1895, p. 107 — Casey, 1899, p.
152-Leng, 1920, p. 214-Wingo, 1952, p. 27.
Scymnus (PuUus) punctatus: Korschefsky, 1931, p. 164— Wingo, 1952, p. 27.
Didion punctatum: Gordon, 1976b, p. 49 — Belicek, 1976, p. 300.
Diagnosis. Length 1.45 to 1.80 mm, width 0.90 to 1.25 mm. Form extremely
elongate, slender. Dorsal surface black or dark brown with anterior pronotal angle
pale; disc of elytron usually with reddish-orange spot (Figs. 63e, f)? occasionally
immaculate. Elytral punctures large, deep, distinct. Male genitalia as in Figure 63a-
d. Female genitalia as in Figure 62f.
Discussion. There are 8 specimens in the type series, all mounted in pairs on 4
points on the same pin bearing the labels “Melsh, punctatus/(a ragged piece of red
paper).” The top specimen nearest the tip of the point is here designated and labeled
as the lectotype, and the remaining 7 specimens as paralectotypes.
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111
Type locality. “Pennsylvania” (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 64. Quebec to Alabama, west to British Columbia and Cali-
fornia.
Didion longulum Casey
Fig. 65a-e; Map, Fig. 66
Didion longulum Casey, 1899, p. 137 — Leng, 1920, p. 213— Korschefsky, 1931, p.
111-Belicek, 1976, p. 299.
Didion parviceps Casey, 1899, p. 137 — Leng, 1920, p. 213— Korschefsky, 1931, p.
111. New Synonymy.
Scymnus {Pullus) occiduus Casey, 1899, p. 153 — Korschefsky, 1931, p. 163. New
Synonymy.
Diagnosis. Length 1.38 to 1.75 mm, width 0.95 to 1.20 mm. Form elongate, oval.
Dorsal surface black or piceous. Pronotum with surface alutaceous, punctures nearly
invisible except some northern specimens with fine but distinct punctures. Elytral
punctures fine, shallow. Abdominal sterna feebly punctured, mostly smooth. Male
genitalia as in Figure 65a-d. Female genitalia as in Figure 65e.
Discussion. This species and D. nanum are similar, but D. nanum has distinct,
coarse punctures on the pronotum and the surface between punctures is shiny. The
siphonal apices are also different in the 2 species (Figs. 65c, 67c).
In my revision of the subgenus Pullus (Gordon, 1976b), I had intended to point
out that Scymnus {Pullus) occiduus Casey belonged in the genus Didion, but failed
to do so. Therefore I now so indicate and also place occiduus as a junior synonym
of longulum. There are 2 types of occiduus (male and female) in the Casey collection,
I designate and label the female as the lectotype and the male as a paralectotype.
The types of D. longulum and D. parviceps are unique females (holotypes). I cannot
separate D. parviceps from D. longulum and consider them synonymous.
Type locality. Of longulum, California, Sonoma Co., Duncans Mills; of parviceps,
California, Sonoma Co.; of occiduus, Nevada, Reno (lectotype here designated).
Type depository. Of longulum (35247), parviceps (35248), and occiduus, (35249),
USNM.
Distribution. Figure 66. Alberta to British Columbia, south to California.
Didion nanum (LeConte)
Fig. 67a-f; Map, Fig. 66
Scymnus nanus LeConte, 1852, p. 141— Crotch, 1874b, p. 269— Horn, 1895, p.
107-Wingo, 1952, p. 28.
Scymnus {Pullus) nanus: Casey, 1899, p. 153 — Korschefsky, 1931, p. 163— Wingo,
1952, p. 28.
Didion nanum: Gordon, 1976b, p. 49 — Belicek, 1976, p. 300.
Diagnosis. Length 1.50 to 1.80 mm, width 1.15 to 1.40 mm. Form elongate, oval
(Fig. 67f). Color black; anterolateral angle of pronotum, mouthparts and leg dark
reddish brown. Punctation on head fine, punctures separated by a diameter or less.
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Pronotum with punctures equal in size to those on head, separated by less than to
twice a diameter; lateral margins not convergent anteriorly, rounded in apical ^4.
Elytron smooth, shiny, punctures coarser than on pronotum, separated by a diameter
or less; pubescence grayish white, arranged in S-curve. Postcoxal line nearly reaching
hind margin of hrst sternum. All abdominal sterna coarsely, densely punctured; 5th
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sternum feebly emarginate apically; 6th sternum broadly, deeply emarginate. Male
genitalia as in Figure 67a-d. Female genitalia as in Figure 67e.
Discussion. This species has often been identified as Scymnus {Pullus) tenebrosus
in collections. The form is broader than in other species of Didion, the pronotal
margins are not convergent anteriorly, and all abdominal sterna are coarsely, densely
punctured. Didion nanum resembles a typical Pullus in fascies more than it does
other species of Didion.
LeConte (1852) stated that he had 2 specimens of nanus. There are 2 specimens
now in his collection. The first of these, a female labeled “(pale green disc)/4698/
Type 6747(red paper)” is here designated and labeled the lectotype. The second
specimen, a male, bears a pale blue disc which denotes a Lake Superior locality; thus
I do not consider this specimen to be a type.
Type locality. “Missouri Territory” (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 66. IOWA: state record. ILLINOIS: Quincy. KANSAS: At-
chison. MASSACHUSETTS: Berlin; Boston. ONTARIO: Brockville; Pt. Pelee; Prince
Edward Co. PENNSYLVANIA: Dauphin Co., Harrisburg; Monroe Co., Canadensis;
Wind Gap.
Genus Scymnus Kugelann
-Seym Kugelann, 1794, p. 545. — Mulsant, 1846, p. 219. — Mulsant, 1850, p. 948.—
Mulsant 1853, p. 152.— Costa, 1849, p. 82. — LeConte, 1852, p. 130.— Crotch,
1874b, p. 239.-Chapuis, 1876, p. 211.-Weise, 1885a, p. 6, 67. -Horn 1895, p.
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Fig. 67. Didion nanum.
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NORTH AMERICAN COCCINELLIDAE
115
83.— Casey, 1899, p. 134. — Mader 1924, p. 8. — Korschefsky, 1931, p. 1 15.— Win-
go, 1952, p. 19. — Mader 1955, p. 869. — Fursch, 1958, p. 77. — Bielawski, 1959, p.
36.— Arnett, 1963, p. 809.— Chapin, 1965, p. 202.— J. Chapin, 1974, p. 18.—
Gordon, 1976b, p. 10. Type-species; Scymnus nigrinus Kugelann, by subsequent
designation of Korschefsky, 1931.
Scymnini with form varying from rounded to elongate, oval, widest at middle of
elytra unless otherwise stated. Antenna 1 0 or 11 -segmented, club of 4 or 5 segments,
lower margin of club segments uneven (Fig. 68a, b). Apical segment of maxillary
palpus cylindrical, apex obliquely truncate. Anterior margin of clypeus truncate or
slightly convex, clypeus extending slightly beyond eye, a narrow, short projection
extending onto eye at antennal insertion. Tarsus with 4 segments, tarsal claw of male
with inner claw larger than in female. Prostemum with distinct carinae (Fig. 68c).
Postcoxal line recurved toward base of first abdominal sternum, complete or incom-
plete. Female genitalia with sclerotized infundibulum; genital plate long, narrowly
triangular (Fig. 68d).
The genus Scymnus was revised by Gordon (1976b); therefore, only a^/ditional
locality records and some necessary corrections in synonymy are included for each
species herein, except for one introduced species not included in 1976b, Scymnus
{P.) suturalis Thunberg.
Key to subgenera of Scymnus
1 . Postcoxal line incomplete, apical end recurved, directed toward base of first sternum
(Fig. 68e) Scymnus Kugelann
- Postcoxal line complete, recurved, extending to base of first sternum (Fig. 68g)
Pullus Mulsant
Subgenus Scymnus Kugelann
Kugelann, 1794, p. 545. — Mulsant, 1846, p. 219. — Mulsant, 1850, p. 965.—
Casey, 1899, p. 138. — Leng, 1920, p. 213. — Korschefsky, 1931, p. 115. — Wingo,
1952, p. 27. — Mader, 1955, p. 929. — Fursch, 1958, p. 79. — Bielawski, 1959, p.
44.-Kamiya, 1961, p. 291. -Hatch, 1961, p. 151. -J. Chapin, 1974, p. 19.-
Gordon, 1976b, p. 10. — Belicek, 1976, p. 300. Type-species; Scymnus nigrinus
Kugelann, by subsequent designation of Korschefsky, 1931.
Antenna 10 or 11 -segmented (Fig. 68a, b); apical segment of maxillary palpus
cylindrical, obliquely truncate apically. Prostemum with 2 strong carinae nearly
always reaching anterior margin. Postcoxal line incomplete, curved forward apically
(Fig. 68e, f); male 5th and 6th abdominal sterna truncate or emarginate apically.
Female with distinct infundibulum (Fig. 69e).
Key to species of Scymnus {Scymnus)
1 . Species with elytron entirely pale or mostly pale with some dark areas, if mostly
dark then pale areas not restricted to apical third nor forming distinct median
spot on elytron 2
Species with elytron black or black with distinct, pale, median or apical spot . . 3
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Fig. 68. Scymnus sp. a, b. Antennae, c. Prostemum. d. Genital plates, e-g. Postcoxal lines.
2( 1 ). Form evenly tapered at both ends; average length less than 2.0 mm; elytron never
with distinct, irregular, dark spots difficilis Casey
Form broad in apical third, pronotum and elytron noticeably discontinuous; av-
erage length more than 2.0 mm.; elytron with distinct, irregular, dark spots at
least on California specimens nebulosus LeConte
3(1). Elytron black with pale antero-median spot circumspectus Horn
- Elytron black without pale antero-median spot 4
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NORTH AMERICAN COCCINELLIDAE
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4(3). Postcoxal line distinctly separated from hind margin of first abdominal sternum
(Fig. 68f) 5
Postcoxal line reaching hind margin of first abdominal sternum or approaching
it closely (Fig. 68e) 7
5(4). Species known only from east of the Mississippi River; postcoxal line approaching
hind margin of first sternum indianensis Weise
Species known only from west of the Mississippi River; postcoxal line not ap-
proaching hind margin of first sternum 6
6(5). Apex of elytron with pale area forming a spot occupying apical 1/4 or more . . .
coosi Hatch
Apex of elytron not or very feebly pale fended Malkin
7(4). Apical '/3 or more of elytron yellowish red; pronotum alutaceous .... opaculus Horn
Apex of elytron black or with narrow, pale yellow border; pronotum not alutaceous
(except caudnus) 8
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8(7). Pronotum alutaceous with punctures finer than on head; distribution mostly west
of Rocky Mountains caurinus Horn
- Pronotum not alutaceous, punctures usually larger than on head; distribution
mostly east of Rocky Mountains 9
9(8). Form extremely elongate, nearly parallel sided; known only from west of the
Mississippi River 10
- Form rounded, not parallel sided; known from both east and west of the Mississippi
River 11
10(9). Surface of elytron smooth, punctures distinctly coarser than on pronotum
apicanus pseudapicanus, new name
Surface of elytron distinctly micro-reticulate, punctures not or barely larger than
on pronotum paracanus linearis Gordon
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NORTH AMERICAN COCCINELLIDAE
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Fig. 71. Scymnus {S.) difficilis.
1 1(9). Basal lobe of male genitalia emarginate ventrally in lateral view (Fig. 82b); female
infundibulum slender, tapered at spermathecal end (Fig. 82e) . . americanus Mulsant
Male and female genitalia not as described above 12
12(1 1). Paramere of male genitalia short, strongly tapered from base to apex (Fig. 87b);
female infundibulum slender, sinuate toward spermathecal end (Fig. 87e)
paracanus paracanus J. Chapin
- Paramere of male genitalia elongate, not tapered toward apex (Fig. 84b); female
infundibulum short, broad at spermathecal end (Fig. 84e)
apicanus apicanus J. Chapin
Scymnus {Scymnus) nebulosus LeConte
Fig. 69a-e; Map, Fig. 70
Scymnus nebulosus LeConte, 1852, p. 137.— Crotch, 1874b, p. 262. — Horn, 1895,
p. 95. — Steinweden, 1929, p. 29.
Scymnus {Scymnus) nebulosus: Casey, 1899, p. 154. — Leng, 1920, p. 214.— Kor-
schefsky, 1931, p. 163.— J. Chapin, 1974, p. 22. — Gordon, 1976b, p. 13.
Scymnus infuscatus Boheman, 1859, p. 208. — Leng, 1920, p. 214. — Korschefsky,
1931, p. 160.— Gordon, 1976b, p. 15.
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Fig. 72. Scymnus (S.) coosi.
Scymnus phelpsii Crotch, 1874a, p. 77. — Horn, 1895, p. 96. — Gordon, 1976b, p. 15.
Scymnus {Scymnus) phelpsii. Casey, 1899, p. 1954. — Leng, 1920, p. 214.— Kor-
schefsky, 1931, p. 165. -Malkin, 1943b, p. 194.-Hatch, 1961, p. 153. -Gordon,
1976b, p. 15.-Belicek, 1976, p. 302.
Scymnus {Scymnus) harneyi WdiXch., 1961, p. 152. — Gordon, 1976b, p. 15.
For detailed description, and discussion see Gordon, 1976b, p. 13.
Scymnus {Scymnus) difficilis Casey
Fig. 71a-e; Map, Fig. 73
Scymnus {Scymnus) difficilis Casey, 1899, p. 154. — Leng, 1920, p. 214.— Kor-
schefsky, 1931, p. 157. — Gordon, 1976b, p. 19.
For detailed description, and discussion see Gordon, 1976b, p. 19.
Scymnus {Scymnus) coosi Hatch
Fig. 72a-d; Map, Fig. 73
Scymnus {Scymnus) coosi Hatch, 1961, p. 152. — Gordon, 1976b, p. 20.
For detailed description, and discussion see Gordon, 1976b, p. 20.
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121
Fig. 73. Distribution. Scymnus (S.) difficilis (dot); S. (S.) coosi (star).
Scymnus {Scymnus) fended Malkin
Fig. 74a-e; Map, Fig. 75
Scymnus {Scymnus) fended yiddkin, 1943a, p. 109. — Gordon, 1976b, p. 23.
For detailed description, and discussion see Gordon, 1976b, p. 23.
Scymnus {Scymnus) caudnus Horn
Fig. 76a-e; Map, Fig. 77
Scymnus caudnus Horn, 1895, p. 97.
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Scyrnnus {Scymnus) caurinus: Casey, 1899, p. 154. — Leng, 1920, p. 214.— Kor-
schefsky, 1931, p. 1 56. -Malkin, 1943b, p. 194.-Hatch, 1961, p. I51.-Belicek,
1976, p. 303. -Gordon, 1976b, 26.
Scymnus {Scymnus) aluticoUis Casey, 1899, p. 154. — Leng, 1920, p. 214.— Kor-
schefsky, 1931, p. 153. -Gordon, 1976b, p. 26.
For detailed description, and discussion see Gordon, 1976b, p. 26.
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124
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 76. Scymnus {S.) caurinus.
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NORTH AMERICAN COCCINELLIDAE
125
Fig. 77.
Distribution. Scymnus (S.) caurinus.
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Fig. 78. Scymnus (S.) indianensis.
Scymnus {Scymnus) indianensis Weise
Fig. 78a-e; Map, Fig. 79
Scymnus indianensis Weise, 1929, p. 33.
Scymnus {Scymnus) indianensis: Korschefsky, 1931, p. 160. — Wingo, 1952, p. 27.—
J. Chapin, 1973, p. 1071. -J. Chapin, 1974, p. 20.-Gordon, 1976b, p. 30.
Scymnus {Scymnus) rust icus Casey, 1899, p. 154 (not Weise, 1 895a). — Leng, 1920,
p. 214.
For detailed description, and discussion see Gordon, 1976b, p. 30.
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NORTH AMERICAN COCCINELLIDAE
127
Fig. 79. Distribution. Scymnus (S.) indianensis.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 80. Scymnus (S.) circumspectus.
Scyrnnus {Scymnus) circumspectus Horn
Fig. 80a-e; Map, Fig. 81
Scymnus circumspectus Horn, 1895, p. 96.
Scymnus {Scymnus) circumspectus: Casey, 1899, p. 153. — Leng, 1920, p. 214.—
Korschefsky, 193 1, p. 156. — Wingo, 1952, p. 27.— J. Chapin, 1974, p. 20. — Gordon,
1976b, p. 32.
For detailed description, and discussion see Gordon, 1976b, p. 32.
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NORTH AMERICAN COCCINELLIDAE
129
Fig. 8 1 . Distribution. Scymnus {S.) circumspect us.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 82. Scymnus (S.) americanus.
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NORTH AMERICAN COCCINELLIDAE
131
Fig. 83. Distribution. Scymnus (S.) americanus.
Scymnus {Scymnus) americanus Mulsant
Fig. 82a-e; Map, Fig. 83
Scymnus {Scymnus) americanus Mulsant, 1850, p. 965.— Casey, 1899, p. 153.—
Blatchley, 1910, p. 526.-Wingo, 1952, p. 27. -Gordon, 1976b, p. 35.
Scymnus americanus: LeConte, 1852, p. 137. — Crotch, 1874b, p. 262. — Horn, 1895,
p. 97. -Wilson, 1927, p. 170.
For detailed description, and discussion see Gordon, 1976b, p. 35.
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Fig. 84. Scymnus (S.) apicanus apicanus.
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NORTH AMERICAN COCCINELLIDAE
133
Fig. 85. Distribution. Scymnus {S.) apicanus apicanus (dot); S. a. pseudapicanus (star).
Scymnus {Scymnus) apicanus apicanus J. Chapin
Fig. 84a-e; Map, Fig. 85
Scymnus {Scymnus) apicanus 'S. Chapin, 1973, p. 1071.— J. Chapin, 1974, p. 20.—
Gordon, 1976b, p. 38.-Belicek, 1976, p. 301.
For detailed description, and discussion see Gordon, 1976b, p. 38.
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Fig. 86. Scymnus (S.) apicanus pscudapicanus.
Scymnus (Scymnus) apicanus pscudapicanus, new name
Fig. 86a-c; Map, Fig. 85
Scymnus (Scymnus) apicanus borealis Gordon, 1976b, p. 38, not Scymnus borealis
Hatch, 1961.
For detailed description, and discussion see Gordon, 1976b, p. 38.
Scymnus (Scymnus) paracanus paracanus J. Chapin
Fig. 87a-e; Map, Fig. 88
Scymnus (Scymnus) paracanus Chapin, 1973, p. 1071.— J. Chapin, 1974, p. 21.—
Gordon, 1976b, p. 41. — Belicek, 1976, p. 302.
For detailed description, and discussion see Gordon, 1976b, p. 41.
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NORTH AMERICAN COCCINELLIDAE
135
Fig. 87. Scymnus (S.) paracanus paracanus.
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Fig. 88. Distribution. Scymnus (5.) paracanus pamcanus (dot); S. p. linearis (star).
Scymnus {Scymnus) paracanus linearis Gordon
Fig. 89a-e; Map, Fig. 88
Scymnus {Scymnus) paracanus linearis Gordon, 1976b, p. 44.
For detailed description, and discussion see Gordon, 1976b, p. 41.
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137
Fig. 89. Scymnus (S.) pamcanus linearis.
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Scymnus {Scymnus) opaculus Horn
Fig. 90a-e; Map, Fig. 91
Scymnus opaculus Horn, 1895, p. 96. — Casey, 1899, p. 160.
Scymnus (Scymnus) opaculus: Leng, 1920, p. 214. — Korschefsky, 1931, p. 163.—
Hatch, 1961, p. 151. -Gordon, 1976b, p. 45.-Belicek, 1976, p. 302.
For detailed description, and discussion see Gordon, 1976b, p. 45.
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NORTH AMERICAN COCCINELLIDAE
139
Fig. 91. Distribution. Scymnus (S.) opaculus.
Subgenus Pullus Mulsant
Pw//w5 Mulsant, 1846, p. 241. — Mulsant, 1850, p. 976.— Weise, 1885a, p. 65.— Casey,
1899, p. 139.-Mader, 1924, p. 8.-Wingo, 1952, p. ll.-Fursch, 1958, p. 79.-
Bielawski, 1959, p. 37.— Arnett, 1963, p. 812.— J. Chapin, 1974, p. 22. — Gordon,
1976b, p. 48.— Belicek, 1976, p. 303. Type-species: Coccinella subvillosa Goeze,
by subsequent designation of Korschefsky, 1931.
Antenna 1 1 -segmented (Fig. 68b); apical segment of maxillary palpus cylindrical,
obliquely truncate apically. Prostemum with 2 strong carinae nearly always reaching
anterior margin. Tarsus cryptotetramerus. Postcoxal line complete, recurved apically,
reaching base of first abdominal sternum (Fig. 68g); male 5th and 6th abdominal
sterna moderately to strongly emarginate and impressed apically.
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Key to the species oe Scymnus {Pullus) of Region I
Map, Fig. 92
1 . Male with tubercle at center of first abdominal sternum; pronotum black or dark
with an obscure, narrow, pale anterior border; leg entirely black or piceous (see
postpictus Casey) marginicollis Mannerheim
Male lacking abdominal tubercle; pronotum variable but if black then with anterior
border also black; leg variable but rarely entirely dark 2
2( 1 ). Dorsal color pattern light reddish yellow with a dark median area extending from
basal portion of pronotum posteriorly along elytral suture, narrowed at apex of
elytron (Fig. 130e) (see nugator Casey) loewii Mulsant
Dorsal color pattern not as described above 3
3(2). Color entirely light yellowish brown; introduced into Eastern Canada and North
Carolina (see sutumlis Thunberg) impexus (Mulsant)
- Color not entirely yellowish brown, usually mostly black (except some forms of
brullei) 4
4(3). Species entirely black dorsally except head may be partly or entirely pale, apex of
elytron sometimes narrowly red or yellow, pronotal angle sometimes obscurely
paler than disc 5
- Species with at least anterior pronotal angle pale red or yellow, usually with
pronotum entirely pale or with a black, parabolic spot anterior to scutellum ... 9
5(4). Apex of elytron with a distinct yellow border pulvinatus Wingo
Apex of elytron black or barely perceptibly red . 6
6(5). Abdomen with median area of last 3 sterna distinctly pale, yellowish brown;
anterior pronotal angle with a relatively broad, obscure area noticeably paler than
disc; length less than 2.00 mm compar Casey
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141
7(6).
8(7).
9(4).
10(9).
11(10).
12(11).
13(12).
14(10).
15(14).
16(15).
17(16).
18(16).
19(18).
20(19).
Abdomen usually entirely black; anterior pronotal angle entirely black or with
very narrow paler area on margin; length more than 2.0 mm (except abbreviatus)
7
Length 1.90 to 2.05 mm; 1st abdominal sternum of male with a tri-angular, feebly
depressed, shining area medially abbreviatus LeConte
Length 2.0 mm or more, nearly always more than 2.20 mm; 1 st abdominal sternum
of male with a flattened median area, not depressed, somewhat rectangular .... 8
Average length 2.30-2.40 mm; form broad, robust; male genitalia as in Figure
138 tenebrosus MuXsdinX
Average length 2.10-2.20 mm; form elongate, slender; male genitalia as in Figure
211 lacustris
Elytron entirely light brown or with dark areas in basal V2 brullei Mulsant
Elytron with at least basal */2 black 10
Pronotum entirely red or yellow 11
Pronotum with at least median, basal projection black, usually with a large, black,
parabolic spot medially 14
Form rounded, sides of elytra not parallel; lateral pronotal margin continuous
with lateral margin of elytron; male with last sternum distinctly impressed .... 12
Form elongate, sides of elytra parallel at least medially; pronotum narrower than
elytra at base; male with last sternum barely perceptibly impressed . kansanus Casey
Elytron with large, red, apical spot, often occupying apical jhi 13
Elytron with apex narrowly red cervicalis Mulsant
Form robust, rounded; apex of elytral spot strongly arcuate .... nemorivagus Wingo
Form elongate; apex of elytral spot feebly arcuate (Fig. 108) semiruber Horn
Elytron with a distinct, pale area or spot in apical jhl 15
Elytron black, usually with a narrow, apical red or yellow border, always straight,
never taking the form of a defined spot 22
Form elongate, sides parallel medially; elytron with large, apical red spot extending
forward at suture (Fig. 120) festatus
Form rounded, sides not parallel, apical spot not extending forward at suture . . 16
Pronotum entirely red except basal median projection black; posterior third or
more of elytron red 17
Color combination not as above 18
Species known only from Missouri and Arkansas; male genitalia with basal lobe
heavily sclerotized, almost rectangular with a small median projection (Fig. 123)
nemorivagus Wingo
Species known only from extreme eastern United States, Massachusetts to Rorida;
male genitalia with basal lobe feebly sclerotized, slender, apex pointed (Fig. 108)
semiruber Horn
Length 2.00 mm or less, width less than 1.50 mm.; dorsal pubescence short,
appressed, grayish white; apical spot on elytron curved toward apex at suture (Fig.
116) rubricaudus Casey
Characters not all as above 19
Length more than 2.00 mm, width 1.40 mm or more; dorsal pubsecence long,
erect, yellowish white; apical spot on elytron as described for rubricaudus: prono-
tum entirely black except antero-lateral angle pale securus J. Chapin
Not entirely as described above; pronotum usually pale with black median spot
20
Elytron with red apical spot usually occupying at least apical '/4, red (Fig. 225);
last sternum of male deeply emarginate, lateral angle of emargination abrupt . .
brullei Mulsant
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
21(20).
22(14).
23(22).
24(23).
25(24).
26(25).
27(26).
28(27).
29(28).
30(29).
31(30).
Elytron with apical pale spot restricted to apical (Fig. 125); last sternum of
male feebly emarginate, feebly impressed 21
Male genitalia with dorsal processes of basal lobe convergent apically (Fig. 127)
louisianae J. Chapin
Male genitalia with dorsal processes of basal lobe widely separated apically (Fig.
125) fraternus
Male genitalia with ventral ala strongly fused to basal lobe, apex of basal lobe
bluntly triangular (Fig. 215) caudalis LeConte
Male genitalia not as described above 23
Last sternum of male deeply emarginate, lateral angle of emargination abrupt;
genitalia robust, heavily sclerotized, dorsal margin of paramere with long setae
(Fig. 225) bndlei Mulsant
Male genitalia not as described above 24
First sternum of male with a deep, elongate-oval pit surrounded by dense hairs,
often with a faint, longitudinal carina in middle of pit (Fig. 150); basal lobe of
genitalia with ventral projection blunt iowensis Casey
First sternum and genitalia not as described above 25
First sternum of male with a small, deep, triangular pit at apical margin; basal
lobe of genitalia with ventral apical projection feebly developed, an elongate-oval,
lightly sclerotized area on each side of middle in ventral view (Fig. 170)
consobrinus LeConte
First sternum and genitalia not as described above 26
Male genitalia with apex of basal lobe pointed, abruptly hooked downward in
lateral view (Fig. 198) uncus
Male genitalia not as described above 27
Male genitalia with basal lobe and ventral ala fused, apex of basal lobe projecting,
pointed, (Fig. 200) puncticollis LeConte
Male genitalia not as described above 28
Male genitalia with ventral projection of basal lobe tapered to a point, much longer
than dorsal projection (Fig. 136) LeConte
Male genitalia not as described above 29
Male genitalia heavily sclerotized, basal lobe broad, apex triangular in dorsal view,
ventral projection tapered to a blunt point in ventral view (Fig. 153)
majus, new name
Male genitalia not as described above 30
Male genitalia with apex of basal lobe bluntly rounded, fused to ventral ala,
margins of siphonal passage fused before apex (Fig. 218) creperus Mulsant
Male genitalia not as described above 31
Male genitalia with basal lobe fused to ventral ala, apex of basal lobe flattened,
triangularly spatulate, projecting beyond ventral ala (Fig. 217) . peninsularis Gordon
Male genitalia with basal lobe pointed apically; paramere slender, lower margin
produced medially (Fig. 187) wingoi Gordon
Key to the species of Scymnus (Pullus) of Region II
Map, Fig. 92
1. Elytron black with a large, median, reddish orange spot (Fig. 98) . . . pacificus Crotch
- Elytron without median spot 2
2(1). Form extremely elongate, parallel-sided; lateral margin of pronotum and elytron
strongly discontinuous (Fig. 93) coniferarum Crotch
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NORTH AMERICAN COCCINELLIDAE
143
3(2).
4(3).
5(4).
6(5).
7(6).
8(7).
9(8).
10(9).
11(9).
12(11).
13(12).
14(13).
15(14).
Form oval, not parallel-sided; lateral margin of pronotum and elytron not no-
ticeably discontinuous 3
Dorsal surface pale yellowish brown, elytron and pronotum unicolorous; length
2.00 mm or less; elytron distinctly alutaceous pallens LeConte
Dorsal surface dark, or with a contrasting color pattern, not entirely pale, if prono-
tum and elytron unicolorous then length more than 2.25 mm; elytron not aluta-
ceous 4
Dorsal color reddish brown, scutellum and sutural margin of elytron narrowly
black; length 2.00 mm or less nugator Casey
Dorsal color not as described above 5
Length less than 1.77 mm; pronotum pale yellowish brown, elytron dark reddish
brown; Texas, Big Bend pauculus Gordon
Length nearly always more than 1.75 mm; color pattern not as described above
or if so, then length more than 2.50 mm 6
Dorsal color pattern light brown with a dark median area extending from basal
portion of pronotum posteriorly along elytral suture, narrowed at apex of elytron
(Fig. 1 30e) loewii Mulsant
Color pattern not as described above 7
Form elongate, nearly parallel-sided; dorsal color pattern either entirely reddish
brown or with a median, black, pronotal spot flavescens Casey
Color pattern not as above, or, if so, then form distinctly rounded {brullei Mulsant)
Length 2.00 mm or less; dorsal color pattern light yellowish brown with basal
projection of pronotum and narrow sutural border dark brown to black, some
specimens also with a black lateral and anterior border on elytron; Texas, Big
Bend enochrus Gordon
Length usually more than 2.00 mm.; color pattern not as described above 9
Pronotum entirely red or yellow 10
Pronotum with at least median, basal projection black 11
Form rounded, sides of elytra not parallel; lateral pronotal margin continuous
with lateral margin of elytron; male with last sternum distinctly impressed ....
cervicalis Mulsant
Form elongate, sides of elytra nearly parallel, at least medially; pronotum narrower
than elytra at base; male with last sternum barely perceptibly impressed
kansanus Casey
Dorsal color entirely light reddish brown except some dark color on pronotum,
sometimes an obscure dark area present on basal jhi of elytron (Fig. 225); male
with last sternum strongly emarginate, angle of emargination abrup . brullei Mulsant
Dorsal color and male last sternum not as described above 12
Species with a large, definite pale area at apex of elytron 13
Species with apex of elytron black or with a more or less well-defined, pale apical
border, never a definite pale spot (see socer LeConte) 17
Punctures on elytron coarse, arranged in curved, transverse rows, giving a slightly
rugose appearance; male first sternum with tubercle medially postpictus Casey
Punctures on elytron fine, not arranged in rows; male first sternum without tubercle
14
Length less than 2.10 mm, form elongate, nearly parallel-sided (Fig. 116)
rubricaudus Casey
Length more than 2.10 mm; form rounded, not parallel-sided 15
Elytron with apical spot usually restricted to apical (4, yellow (Fig. 125); last
sternum of male feebly emarginate, feebly impressed
16
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
16(15).
17(12).
18(17).
19(18).
20(19).
21(20).
22(21).
23(22).
24(23).
25(24).
26(25).
27(24).
28(18).
29(28).
30(29).
Elytron with apical spot usually occupying at least apical V4 (Fig. 225); last sternum
of male deeply emarginate, lateral angle of emargination abrupt .... brullei Mulsant
Male genitalia with dorsal processes of basal lobe convergent apically (Fig. 127)
louisianae J. Chapin
Male genitalia with dorsal processes of basal lobe widely separated (Fig. 125) . .
fraternus LeConte
Postcoxal line reaching hind margin of first sternum; form elongate, almost par-
allel-sided; apex of elytron distinctly reddish yellow monticola Casey
Postcoxal line not reaching hind margin of first sternum; color and form variable
18
Pronotum entirely black or black with a very small, obscure, paler area at antero-
lateral angle 19
Pronotum mostly pale with a central dark area, or at least with antero-lateral angle
broadly, distinctly red or yellow (see horni Gorham) 28
Known from the eastern edge of Region II (see compar Casey) . . tenebrosus Mulsant
Known from Colorado westward (see weidti Casey) 20
Male genitalia with basal lobe slender, nearly as long as ventral ala, not fused to
ventral ala (Fig. 157) renoicus Casey
Male genitalia not as described above 21
Male genitalia with basal lobe much shorter than ventral ala, not fused to ala,
inner portion of ala lightly sclerotized, outer portion heavily sclerotized (Fig. 1 60)
mormon Casey
Male genitalia not as described above 22
Male genitalia pale, nearly transparent (Fig. 166) aridus Casey
Male genitalia darkened, definitely sclerotized 23
Male genitalia with apex of ventral projection of basal lobe robust, truncate (Fig.
151) Calaveras Casey
Male genitalia not as described above 24
Male genitalia with basal lobe and ventral ala fused, apex more or less pointed in
lateral view (Fig. 197) 25
Male genitalia with basal lobe and ventral ala fused, apex broadly rounded in
lateral view 27
Sclerotized area of ventral ala nearly truncate apically, apex of basal lobe bluntly
pointed (Fig. 197) papago Casey
Male genitalia not as described above 26
Sclerotized area of ventral ala deeply emarginate apically, apex of basal lobe
sharply pointed (Fig. 192) wickhami Gordon
Sclerotized area of ventral ala not emarginate, rounded apically, apex of basal
lobe sharply pointed (Fig. 184) impletus Gordon
Male genitalia with apex of basal lobe wide in lateral view, slightly enlarged before
apex (Fig. 211) lacustris LeConte
Male genitalia with apex of basal lobe narrow in lateral view, not enlarged before
apex (Fig. 213) tahoensis Casey
Elytron strongly alutaceous, feebly shining; length about 2.00 mm . . . uteanus Casey
Elytron not noticeably alutaceous, strongly shining; length usually more than 2.00
mm 29
Pronotum mostly yellow or red with a small parabolic spot medially at base, spot
not approaching anterior margin of pronotum 30
Pronotum mostly black with lateral margin and/or antero-lateral angle yellow or
red, anterior margin of pronotum black or very narrowly pale 35
Elytron with apical pale border wide, at least % of a mm; pronotal spot small.
1985
NORTH AMERICAN COCCINELLIDAE
145
usually confined to area just anterior to basal median projection (see uncus Wingo)
creperus Mulsant
Elytron with apical pale border narrow, less than 1/6 of a mm.; pronotal spot
usually extending ‘ the distance to anterior margin or more 31
31(30). Form rounded; length 2.40 mm garlandicus Casey
Form elongate; length 2.25 mm or less 32
32(31). Male genitalia with basal lobe and ventral ala not fused (Fig. 150) 34
Male genitalia with basal lobe and ventral ala fused (Fig. 174) 33
33(32). Male genitalia short, compact, feebly sclerotized (Fig. 174) cockerelli Casey
Male genitalia long, slender, lightly sclerotized (Fig. 215) caudalis LeConte
34(32). Male genitalia with ventral projection of basal lobe bluntly rounded, only slightly
longer than dorsal projection (Fig. 150); male with pit on first sternum deep,
elongate-oval, often with a fine, longitudinal carina at middle (Fig. 1 50)
iowensis Casey
- Male genitalia with ventral projection of median lobe pointed, distinctly longer
than dorsal projection (Fig. 186); median area of male first sternum glabrous,
slightly flattened, densely punctured simulans Gordon
35(29). Length 2.00 mm or less 36
- Length more than 2.10 mm 38
36(35). Male genitalia pale, nearly transparent 37
- Male genitalia darkened, definitely sclerotized (Fig. 156) utahensis Gordon
37(36). Male genitalia with apex of dorsal projection of basal lobe broad, truncate (Fig.
162) ardelio Casey
Male genitalia with apex of dorsal projection of basal lobe slender, no wider than
ventral projection (Fig. 166) aridus Casey
38(35). Dorsal pubescence at least partly yellowish brown 39
Dorsal pubescence entirely grayish or yellowish white 40
39(38). Black area of pronotum extending to anterior margin of pronotum, broad antero-
lateral angle pale barberi Gordon
- Black area of pronotum not quite reaching anterior margin medially, broad antero-
lateral angle and narrow anterior border pale solidus Casey
40(38). Male genitalia with basal lobe and ventral ala fused; male without pit on first
sternum 41
Male genitalia with basal lobe and ventral ala not fused, ventral projection of
basal lobe bluntly rounded (Fig. 1 50); male with deep elongate-oval pit on first
sternum (Fig. 150) iowensis Casey
4 1 (40). Male genitalia with central carinae of basal lobe divergent at apex, leaving a blunt,
triangular apical area (Fig. 215) caudalis LeConte
- Male genitalia with central carinae of basal lobe not divergent, apex of basal lobe
pointed in ventral view (Fig. 211) lacustris LeConte
Key to the species of Scymnus (Pullus) of Region III
Map, Fig. 92
1. Elytron with a large, median, reddish orange spot (Fig. 98) pacificus Crotch
Elytron without a median spot 2
2(1). Length nearly twice the width; lateral margin of pronotum strongly discontinuous
with lateral margin of elytron; elytron yellowish brown with suture and scutellum
usually black or dark brown (Fig. 93) coniferarum Crotch
Length much less than twice the width; lateral pronotal margin more or less
continuous with elytron; color variable 3
146
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
3(2).
4(3).
5(4).
6(4).
7(6).
8(7).
9(8).
10(9).
11(10).
12(3).
13(12).
14(13).
15(14).
16(15).
17(16).
Dorsal color primarily light brown or yellow, with or without a dark pattern . . 4
Dorsal color primarily black or dark brown, with or without pale areas 12
Postcoxal line on first sternum reaching hind margin of sternum 5
Postcoxal line on first sternum not reaching hind margin of sternum 6
Form elongate; pronotum entirely pale or with a median black area; postcoxal
line angulate flavescens Casey
Form rounded; pronotum always black at least medially; postcoxal line rounded,
not angulate nigricollis Gordon
Elytron distinctly alutaceous, feebly shining; dorsum entirely light yellowish brown;
form round; length less than 2.00 mm pallens LeConte
Elytron not alutaceous, shining; dorsum with or without a dark pattern, length
variable 7
Form elongate; dorsum entirely yellowish brown mimoides Gordon
Form not particularly elongate, dorsum not entirely yellowish brown 8
Length less than 1.75 mm, pronotum pale yellowish brown, elytron reddish brown;
Texas, Big Bend pauculus Gordon
Length more than 1.75 mm; color pattern not as described above 9
Length 2.00 mm or less; dorsal color pattern light yellowish brown with basal
projection of pronotum and narrow sutural border dark brown to black, some
specimens also with a black lateral and anterior border on elytron; Texas, Big
Bend enochms Gordon
Length usually more than 2.00 mm, color pattern not as described above .... 10
Dorsal color pattern light brown with a dark median area extending from basal
portion of pronotum posteriorly along elytral suture, narrowed at apex of elytron
(Fig. 1 30e) loewii Mulsant
Color pattern not as described above 11
Length 2.00 mm or slightly less; paramere of male genitalia as broad as basal
lobe (Fig. 177) nugat or CasQy
Length 1.75 mm or less; paramere of male genitalia narrower than basal lobe
(Fig. 179) neomexicanus Gordon
Pronotum entirely red or yellow cervicalis Mulsant
Pronotum black at least basally 13
Apical '/4 to V3 of elytron red (Fig. 1 34); punctures on elytron coarse, arranged in
curved, transverse rows, giving a slightly rugose appearance to elytron; male first
sternum with tubercle postpictus Casey
Species with all characters not as described above 14
Male with tubercle at middle of first sternum; elytron black except narrow apical
border pale; leg usually all black; pronotum with color variable but always with
at least a narrow anterior border pale marginicollis Mannerheim
Male without tubercle; leg usually pale or at least apical ‘ of tibia pale 15
Species with a distinct pale area on apex of elytron, or a pale, discal spot 16
Species with apex of elytron black or with a straight pale border, never with a
distinct pale spot 18
Species with a transversely oval, yellow spot restricted to apical % of elytron . 1 7
Species with apical '/t to ‘ of elytron red or with an elongate, median, red spot on
elytron (Fig. 130) loemi Mulsant
Pronotum black, anterolateral angle very narrowly yellow; male 1 st sternum not
depressed medially, coarsely, densely punctured caffer Gordon
Pronotum usually with median, parabolic, black spot, at least antero- lateral angle
broadly yellow or red; male 1st sternum depressed medially, finely, densely punc-
tured louisianae J. Chapin
1985
NORTH AMERICAN COCCINELLIDAE
147
18(15).
19(18).
20(19).
21(20).
22(21).
23(22).
24(18).
25(24).
26(25).
27(26).
28(27).
29(28).
30(29).
31(29).
32(26).
33(32).
34(33).
Pronotum entirely black (see tahoensis Casey) 19
Pronotum with at least anterolateral angle distinctly pale 24
Male genitalia with apex of ventral apical projection truncate in ventral view
(Fig. 151) Calaveras CdiSQy
Male genitalia not as described above 20
Male genitalia with basal lobe and ventral ala fused, apex of basal lobe sharply
pointed (Fig. 1 84) impletus Gordon
Male genitalia not as described above 21
Male genitalia with median area of ventral ala not sclerotized, and elongate-oval
area on each side of basal lobe also unsclerotized (Fig. 1 90) .... tenebricus Gordon
Male genitalia not as described above 22
Male genitalia with sclerotized area of anterolateral angle of ventral ala produced,
basal lobe with 2 median folds (Fig. 197) papago Casey
Male genitalia not as described above 23
Male genitalia with basal lobe and ventral ala strongly united, basal lobe slightly
shorter than ventral ala, apex rounded in lateral view (Fig. 202) weidti Casey
Male genitalia with basal lobe shorter than ventral ala, inner portion of ventral
ala membranous, extending beyond outer portion (Fig. 1 60) mormon Casey
Elytron strongly alutaceous, feebly shining; pronotum pale red with a small, black,
parabolic spot anterior to scutellum uteanus Casey
Elytron not noticeably alutaceous, strongly shining; pronotum variable 25
Elytron not black but a dark mahogany brown with sutural and lateral borders
obscurely black, apex of elytron with a wide, pale border; length usually less than
2.00 mm gilae Casey
Elytron black, apex of elytron with a narrow, pale border 26
Pronotum pale with a black, basal spot not approaching anterior margin of
pronotum 27
Pronotum mostly black, black area either reaching anterior border or very nar-
rowly separated from it 32
Pronotum with black area poorly defined, restricted to median, basal y^ of prono-
tum creperus Mulsant
Pronotum with black area parabolic, well defined, extending more than ‘ the
distance to apical margin 28
Male genitalia with apical ventral process of basal lobe long, stout, curved upward
in lateral view (Fig. 143) garlandicus Casey
Male genitalia not as described above 29
Male genitalia of the brullei type, basal lobe inflated (Fig. 223) . . hubbardi Gordon
Male genitalia not as described above 30
Male genitalia with basal lobe and ventral ala fused (Fig. 174) cockerel li Casey
Male genitalia not as described above 31
Male genitalia without ventral ala (Fig. 129) apithanus Gordon
Male genitalia with ventral ala (Fig. 159) horni Gorham
Pronotum black with anterolateral angle narrowly yellow; femur black except
apex pale aridoides Gordon
Pronotum with anterolateral angle broadly pale yellow; femur with at least apical
‘ pale 33
Male genitalia of the brullei type but with membranous lateral projection as in
figure 34
Male genitalia not of the brullei type, lacking membranous lateral projections . 35
Black area of pronotum separated from anterior margin by a narrow, yellow
border; Texas (Big Bend) howdeni Gordon
148
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
35(33).
36(35).
37(36).
38(37).
39(38).
40(38).
Black area of pronotum reaching anterior margin; Arizona huachuca Gordon
Male genitalia with apical ventral process of basal lobe long, stout, curved upward
in lateral view (Fig. 143) garlandicus Casey
Male genitalia not as described above 36
Male genitalia with basal lobe inflated in lateral view, a winglike lobe on each
side of siphonal aperture (Fig. 220) bryanti Gordon
Male genitalia not as described above 37
Male genitalia with apical ventral process of basal lobe long, broad at base, tapered
to blunt apex (Fig. 154) ignarus Gordon
Male genitalia not as described above 38
Male genitalia lightly sclerotized, nearly transparent 39
Male genitalia normally sclerotized 40
Male genitalia with apical dorsal projection of basal lobe broad, truncate at apex
(Fig. 162) ardelio Horn
Male genitalia with apical dorsal projection tapered to a point (Fig. 166)
aridus Casey
Male genitalia with inner border of ventral ala sclerotized (Fig. 142a) (see hum-
boldti Casey) solidus Casey
Male genitalia with inner border of ventral ala not sclerotized (Fig. 148)
barberi Gordon
Key to the species oe Scymnus (Fullus) of Region IV
Map, Fig. 92
1. Elytron with a large, median, reddish orange spot (Fig. 98) pacificus Crotch
Elytron without a median spot 2
2(2). Length nearly twice the width; lateral margin of pronotum strongly discontinous
with lateral margin of elytron; elytron yellowish brown with suture and scutellum
usually black or dark brown (Fig. 93) coniferarum Crotch
Length much less than twice the width; lateral pronotal margin more or less
continuous with elytron; color variable 3
3(2). Dorsal surface distinctly alutaceous, completely pale yellowish brown
pallens LeConte
- Dorsal surface not noticeably alutaceous, at least some dark areas present, usually
almost completely black 4
4(3). Dorsal color pattern light brown with a dark median area extending from basal
portion of pronotum posteriorly along elytral suture, narrowed at apex of elytron
(Fig. 1 30), lateral border may also be dark (Fig. 1 30) loewii Mulsant
Dorsal color not as described above 5
5(4). Apical 2/3 of elytron yellowish red, rest of elytron and pronotum except narrow
lateral border black (Fig. 140); California (Channel Islands) falli Gordon
- Color pattern not as described above 6
6(5). Dorsal color primarily pale yellowish brown with some dark marking 7
Dorsal color primarily black or dark brown, sometimes with pale marking .... 8
7(6). Form elongate, margins of elytra subparallel; sutural border of elytron narrowly
black, an obscure dark border on lateral margin mimoides Gordon
Form round, margins of elytra not parallel; sutural border of elytron narrowly
black but with no dark lateral border cockerelli Casey
8(6). Apical 'A to Vs of elytron red (Fig. 1 34); punctures on elytron coarse, arranged in
1985
NORTH AMERICAN COCCINELLIDAE
149
9(8).
10(9).
11(10).
12(11).
13(12).
14(11).
15(14).
16(15).
17(16).
18(17).
19(17).
20(19).
21(19).
22(21).
curved, transverse rows, giving a slightly rugose appearance to elytron; male 1 st
sternum with tubercle medially postpictus Casey
Apex of elytron black or narrowly pale; elytron not appearing rugose; male 1st
sternum not tuberculate (except marginicollis) 9
Male with median tubercle on 1st sternum; elytron black except narrow apical
border; legs usually all black or at least femora entirely black; pronotum varying
from almost entirely yellow to nearly all black but with at least apical border
narrowly pale marginicollis Mannerheim
Male 1st sternum without tubercle; legs usually pale but if black then at least apex
of femur pale 10
Pronotum entirely red or yellow (see cervicalis Mulsant) carri Gordon
Pronotum at least partly black 11
Pronotum mostly yellow or red with a black area medially anterior to scutellum,
black area not approaching anterior margin of pronotum 12
Pronotum mostly or entirely black, black area reaching anterior margin or very
narrowly separated from it 14
Postcoxal line reaching hind margin of 1st sternum; elytron distinctly micro-
reticulate (see uteanus Casey) nevadensis Weise
Postcoxal line not reaching hind margin of 1 st sternum; elytron not micro-retic-
ulate or feebly so 13
Length less than 2.00 mm.; 1st sternum of male densely punctured medially . .
erythronotum Gordon
Length 2.00 mm or more, 1 st sternum of male with a flattened, shining, impunctate
area medially (see garlandicus Casey) horni Gorham
Elytron dark mahogany brown with suture and lateral border black, apex of elytron
with a wide, pale border; length less than 2.00 mm gilae Casey
Elytron black or black with a pale apical border; length variable but usually more
than 2.00 mm 15
Pronotum entirely black 16
Pronotum with at least anterolateral angle pale 28
Large, robust, length usually 2.65 mm or more; dorsal pubescence yellowish brown;
dorsal surface entirely black except narrow apical margin pale calaveras Casey
Characters not all as described above 17
Male genitalia with basal lobe much shorter than ventral ala (Fig. 172) 18
Male genitalia with basal lobe as long as ventral ala or nearly so 19
Male genitalia with basal lobe extremely short (Fig. 172); 6th sternum of male
deeply, abruptly emarginate mendocino Casey
Male genitalia with basal lobe not extremely short (Fig. 1 94); male 6th sternum
normally emarginate elusivus Gordon
Basal lobe of male genitalia slender, not fused to ventral ala 20
Basal lobe of male genitalia robust, fused to ventral ala 21
Basal lobe of male genitalia with dorsal apical projection not wider than ventral
apical projection, dorsal margin of basal lobe sinuate (Fig. 157) renoicus Casey
Basal lobe of male genitalia with dorsal apical projection wider than ventral apical
projection, dorsal margin of basal lobe not sinuate (Fig. 145) jacobianus Casey
Male genitalia with a small, elongate sclerite medially at base of basal lobe, apex
bluntly pointed in lateral view, basal lobe and ventral ala fused (Fig. 202)
weidti Casey
Male genitalia not as described above 22
Apex of basal lobe of male genitalia rounded in lateral view (Fig. 207) 23
Apex of basal lobe of male genitalia pointed in lateral view (Fig. 1 84) 26
150
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
23(22).
24(23).
25(24).
26(22).
27(26).
28(15).
29(28).
30(29).
31(29).
32(31).
33(32).
34(33).
35(34).
36(33).
37(32).
38(37).
Apex of basal lobe slender, evenly rounded in lateral view (Fig. 213) 24
Apex of basal lobe broad, rounded dorsally, abruptly angled ventrally in lateral
view (Fig. 207) hesperius Gordon
Basal lobe with median ventral carinae slightly separated, ventral ala fused in
basal % (Fig. 213) tahoemis Casey
Basal lobe with median ventral carinae joined, ventral ala fused in more than
basal % (Fig. 21 1) 25
Apex of basal lobe as wide or wider than paramere in lateral view lacustris LeConte
Apex of basal lobe narrower than paramere in lateral view tahoemis Casey
Apex of ventral ala angulate, produced, or with a median membranous area ... 27
Apex of ventral ala not angulate or produced (Fig. 1 84) impletus Gordon
Apex of ventral ala simple, angulate (Fig. 192) wickhami Gordon
Apex of ventral ala divided by median membranous area, inner sclerotized area
angulate (Fig. 190) tenebricus Gordon
Male first sternum with a deep, elongate-oval pit, pit often with a median carina;
ventral apical projection of basal lobe of male genitalia rounded (Fig. 1 50) ....
iowensis Casey
Male first sternum without a pit, at most with a shallow depression; ventral apical
projection of basal lobe of male genitalia not as described above 29
Male genitalia feebly sclerotized, nearly transparent 30
Male genitalia distinctly sclerotized, darkened 31
Apex of dorsal projection of basal lobe truncate (Fig. 162) ardelio Horn
Apex of dorsal apical projection of basal lobe pointed (Fig. 166) aridus Casey
Basal lobe of male genitalia ovate in ventral view, ventral alae and paramere
curved inward (Fig. 181) mittingi Gordon
Male genitalia not as described above 32
Male genitalia with basal lobe and ventral ala fused 33
Male genitalia with basal lobe and ventral ala not fused 37
Basal lobe of male genitalia with apex broadly rounded in lateral view, no median
sclerite at base in ventral view 34
Basal lobe of male genitalia slender, a median, basal sclerite present in ventral
view 36
Median, ventral carinae of basal lobe distinctly separated at least basally (Fig.
208) luctuosus Casey
Median, ventral carinae of basal lobe united or nearly so from base to apex (Fig.
211) 35
Apex of basal lobe broad, broadly rounded in lateral view (Fig. 211)
lacustris LeConte
Apex of basal lobe narrow in lateral view (Fig. 213) tahoemis Casey
Basal lobe of male genitalia evenly tapered from base to apex (Fig. 204) Alberta
aquilonarius Gordon
Basal lobe of male genitalia narrowed before apex, apex slightly bulbous (Fig.
206); California martini Gordon
Basal lobe of male genitalia slender, tapered from base to apex (Fig. 156)
utahensis Gordon
Basal lobe of male genitalia not as described above 38
Male genitalia with inner margin of ventral ala sclerotized, basal lobe broad (Fig.
142) solidus Cc^sey
Male genitalia with inner margin of ventral ala not sclerotized, basal lobe slender
(Fig. 146) humboldti Casey
1985
NORTH AMERICAN COCCINELLIDAE
151
Scytnnus (Pullus) cajfer Gordon
Fig. 96a-d
Scymnus {Pullus) cajfer Gordon 1976b, p. 65.
For detailed description, and discussion see Gordon, 1976b, p. 65.
Scymnus {Pullus) coniferarum Crotch
Fig. 93a-f; Map, Fig. 95
Scymnus coniferarum Crotch, 1874a, p. 77. — Horn, 1895, p. 105.
Scymnus {Pullus) coniferarum: Casey, 1899, p. 152. — Leng, 1920, p. 214.— Kor-
schefsky, 1931, p. 157. — Hatch, 1961, p. 151.— Gordon, 1976b, p. 66. — Belicek,
1976, p. 305.
For detailed description, and discussion see Gordon, 1976b, p. 66, and Gordon
(1982).
Scymnus {Pullus) suturalis Thunberg
Fig. 94a-e; Map, Fig. 95
Scymnus suturalis Th.\xnhQY%, 1795, p. 106. — Korschefsky, 1931, p. 138.— Gordon,
1976b, p. 66. — Gordon, 1982, p. 250 (see Korschefsky, 1931, for complete syn-
onymy.
Diagnosis. Description as for S. {P.) coniferarum: Body slightly broader, less elon-
gate in appearance; punctures on elytron coarse, dense, separated by the diameter of
a puncture or less; basal lobe of male genitalia broad in ventral view, abruptly
narrowed in apical V4, apex in lateral view distinctly bent downward; apex of sipho
S-shaped (hgs. 94a-c); female genitalia with infundibulum slender, rodlike (hg. 94e).
Discussion. Gordon (1976b) included this species as S. {P.) coniferarum which is
primarily a California species. Subsequent investigation revealed that the Pennsyl-
vania and New York specimens were actually S. {P.) suturalis (Gordon, 1982). It
was introduced into Michigan from Germany in 1961, and has recently been collected
there, but whether this population is a result of the introduction or an accidental
establishment is not apparent (Hoebeke, in press).
Type locality. “Suecica”.
Type depository. Type not examined.
Distribution. Figure 95. CONNECTICUT: Middlesex Co., Clinton. MICHIGAN:
Saginaw Co., Saginaw. NEW YORK. PENNSYLVANIA: (see Gordon, 1982, for
specific localities).
Scymnus {Pullus) impexus Mulsant
Fig. 97a-d
Scymnus {Pullus) impexus Mulsant, 1850, p. 979. — Korschefsky, 1931, p. 127.—
Delucchi, 1954, pp. 243-278. -Gordon, 1976b, p. 70.
Scymnus {Pullus) abietisM\x\s?inX, 1846, p. 247 (not Paykull, 1 798). — Mulsant, 1850,
p. 979.
152
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 9 3( 1 )
Fig. 93. Scymnus {P.) coniferarum.
1985
NORTH AMERICAN COCCINELLIDAE
153
Fig. 94. Scymnus (P.) suturalis.
154
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 95. Distribution. Scymnus {P.) coniferarum (shaded, peripheral localities dotted); S.
{P.) suturalis (star); S. P. caffer (triangle).
For detailed description, and discussion see Gordon, 1976b, p. 70. Establishment
of this species has been effected in the Willamette Valley of Oregon following releases
made in 1960 and 1962.
Scymnus (Pullus) pacificus Crotch
Fig. 98a-e; Map, Fig. 99
Scymnus pacificus Crotch, 1874a, p. 77. — Horn, 1895, p. 100.
Scymnus {Pullus) pacificus: Casey, 1899, p. 152. — Leng, 1920, p. 214.— Korschefsky,
1931, p. 164.-Hatch, 1961, p. 151. -Gordon, 1976b, p. 72.
Scymnus stratus Horn, 1895, p. 100. — Gordon, 1976b, p. 72.
Scymnus {Pullus) stratus: Casey, 1899, p. 152. — Leng, 1920, p. 214.— Korschefsky,
1931, p. 166.
For detailed description, and discussion see Gordon, 1976b, p. 72.
Additional locality record: ARIZONA: Yavapai Co., 15 mi. S. Prescott.
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Fig. 96. Scymnus {P.) caffer.
Scymnus {Pullus) flavescens Casey
Fig. lOOa-d; Map, Fig. 101
Scymnus {Pullus) flavescens CdiSQy, 1899, p. 139. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 158. -Gordon, 1976b, p. 75.
For detailed description, and discussion see Gordon, 1976b, p. 75.
Additional locality records: ARIZONA: Apache Co., Chuska. UTAH: Wayne Co.,
Henry Mts.
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Fig. 97. Scymnus (P.) impexus.
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Fig. 98. Scymnus {P.) pacificus.
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Fig. 99. Distribution. Scymnus (P.) pacificus.
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NORTH AMERICAN COCCINELLIDAE
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Fig. 100. Scymnus (P.) flavescens.
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Fig. 101. Distribution. Scymnus (P.) flavescens (dot); S. (P.) nigricollis (star),
1985
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Fig. 102. Scymnus (P.) nigricollis.
Scymnus {Pullus) nigricollis Gordon
Fig. 102a-d; Map, Fig. 101
Scymnus {Pullus) nigricollis Gordon, 1976b, p. 78.
For detailed description, and discussion see Gordon, 1976b, p. 78.
1 62 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 103. Scymnus (P.) kansanus.
Scymnus {Pullus) kansanus Casey
Fig. 103a-d; Map, Fig. 104
Scymnus {Pullus) kansanus CdiSQy , 1899, p. 142. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 160.— Wingo, 1952, pp. 28.— Gordon, 1976b, p. 78.
For detailed description, and discussion see Gordon, 1976b, p. 78.
Additional locality record: NEW JERSEY : Fort Lee.
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Fig. 104. Distribution. Scymnus (P.) kansanus.
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Fig. 105. Scymnus (P.) pauculus.
Scymnus {Pullus) pauculus Gordon
Fig. 105a-d; Map, Fig. 106
Scymnus {Pullus) pauculus Gordon, 1976b, p. 81.
For detailed description, and discussion see Gordon, 1976b, p. 81. Additonal
locality record. ARIZONA: Oracle.
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165
Fig. 106. Distribution. Scymnus (P.) pauculus (star); S. {P.) pallens (dot).
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Fig. 107. Scymnus (P.) pallens.
Scymnus (Pullus) pallens LeConte
Fig. 107a-d; Map, Fig. 106
Scymnus pallens LeConte, 1852, p. 137. — Crotch, 1847b, p. 263. — Horn, 1895, p.
99.
Scymnus {Pullus) pallens: Casey, 1899, p. 140. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 164.-Hatch, 1961, p. 149.-Gordon, 1976b, p. 84.
For detailed description, and discussion see Gordon, 1976b, p. 84.
Additional locality record: TEXAS; Patricio Co., Martin, 12 mi. S.
1985 NORTH AMERICAN COCCINELLIDAE 167
Fig. 108. Scymnus (P.) semiruber.
Scymnus (Pullus) semiruber Horn
Fig. 108a-e; Map, Fig. 109
Scymnus semiruber Worn, 1895, p. 102.
Scymnus {Pullus) semiruber: Casey, 1899, p. 140.— Leng, 1920, p. 213.— Wilson,
1927, p. 169.-Korschefsky, 1931, p. 165. -Gordon, 1976b, p. 86.
Scymnus puritanus Casey, 1924, p. 174. — Leng, 1927, p. 33. — Korschefsky, 1931,
p. 165.-Gordon, 1976b, p. 86.
For detailed description and discussion see Gordon, 1976b, p. 86.
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Fig. 109. Distribution. Scymnus (P.) semiruber.
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Fig. 110. Scymnus (P.) gilae.
Scymnus (Pullus) gilae Casey
Fig. 1 lOa-d; Map, Fig. 1 1 1
Scymnus {Pullus) gilae Casey, 1899, p. 147. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 159. -Gordon, 1976b, p. 89.
Scymnus {Pullus) infans Casey, 1899, p. 149. — Leng, 1920, p. 213.— Korschefsky,
1931, p. I60.-Gordon, 1976b, p. 89.
Scymnus apiciventris Casey, 1924, p. 175. — Leng and Mutchler, 1927, p. 33. — Kor-
schefsky, 1931, p. 154.— Gordon, 1976b, p. 91.
For detailed description, and discussion see Gordon, 1976b, p. 89.
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Fig. 111. Distribution. Scymnus (P.) gilae.
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Fig. 112. Scymnus (P.) mimoides.
Scymnus (Pullus) mimoides Gordon
Fig. 112a-d; Map, Fig. 113
Scymnus {Pullus) mimoides Gordon, 1976b, p. 93.
For detailed description and discussion see Gordon, 1976b, p. 93.
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Fig. 1 13. Distribution. Scymnus (P.) mimoides.
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173
Fig. 1 14. Scymnus (P.) cervicalis.
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Fig. 115. Distribution. Scymnus {P.) cervicalis (disjunct localities dotted).
Scymnus (Pullus) cervicalis Mulsant
Fig. 114a-d; Map, Fig. 115
Scymnus {Pullus) cervicalis Mulsant, 1850, p. 984.— Casey, 1899, p. 142.— Leng,
1920, p. 213. — Korschefsky, 1931, p. 156. — Wingo, 1952, p. 29.— J. Chapin, 1974,
p. 28. — Gordon, 1976b, p. 95.
Scymnus cervicalis: LeConte, 1852, p. 139.— Crotch, 1874b, p. 266. — Horn, 1895,
p. 103. -Wilson, 1927, p. 169.
For detailed description, and discussion see Gordon, 1976b, p. 95.
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Fig. 116. Scymnus (P.) rubricaudus.
Scymnus (Pullus) rubricaudus Casey
Fig. 1 1 6a-e; Map, Fig. 1 1 7
Scymnus {Pullus) rubricauda Casey, 1899, p. 141. — Leng, 1920, p. 213.— Korschef-
sky, 1931, p. 165.
Scymnus {Pullus) texanus Casey, 1899, p. 141. — Leng, 1920, p. 213 (synonym of
creperus Mulsant). — Korschefsky, 1931, p. 157.— Wingo, 1952, p. 32.
Scymnus {Pullus) chromopyga Casey, 1899, p. 141. — Leng, 1920, p. 213.— Wingo,
1952, p. 32.
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Fig. 117. Distribution. Scymnus (P.) rubricaudus (peripheral and disjunct localities dotted).
Scymnus {Pul lus) canterius Casey, 1899, p. 142. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 156.-Gordon, 1976b, p. 100.
Scymnus {Pullus) rubricaudus’. Wingo, 1952, p. 32.— J. Chapin, 1974, p. 25.— Gor-
don, 1976b, p. 98.
Scymnus {Pullus) chromopygus: Korschefsky, 1931, p. 156.
For detailed synonymy, description, and discussion see Gordon, 1976b, p. 98.
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177
Scymnus (Pullus) enochrus Gordon
Fig. 118a-d; Map, Fig. 119
Scymnus {Pullus) enochrus Gordon, 1976b, p. 102.
For detailed description, and discussion see Gordon, 1976b, p. 102.
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Fig. 119. Distribution. Scymnus (P.) enochrus.
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Fig. 120. Scymnus {P.) festatus.
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Scymnus (Pullus) festatus Wingo
Fig. 120a-f; Map, Fig. 121
Scymnus {Pullus) festatus V^ingo, 1952, p. 31. — Gordon, 1976b, p. 103.
For detailed description, and discussion see Gordon, 1976b, p. 103.
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181
Fig. 122. Scymnus (P.) pulvinatus.
Scymnus (Pullus) pulvinatus Wingo
Fig. 122a-d; Map, Fig. 121
Scymnus {Pullus) pulvinatus Wingo, 1952, p. 34. — Gordon, 1976b, p. 106.
For detailed description, and discussion see Gordon, 1976b, p. 106.
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Fig. 123. Scymnus (P.) nemorivagus.
Scymnus (Pullus) nemorivagus Wingo
Fig. 123a-d; Map, Fig. 124
Scymnus {Pullus) nemorivagus 1952, p. 35. — Gordon, 1976b, p. 109.
For detailed description, and discussion see Gordon, 1976b, p. 109.
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183
Scymnus (Pullus) f rat emus LeConte
Fig. 125a-f; Map, Fig. 126
Scymnus fraternus LeConte, 1852, p. 138.— Crotch, 1874b, p. 264. — Horn, 1895, p.
lOl.-Stehr, 1930, p. 49.-Wingo, 1952, p. 31. -Chapin, 1973, p. 1072.
Scymnus haemorrhousl.QConXQ,\%51,^. 138. — Crotch, 1874b, p. 264. — Horn, 1895,
p. 101.
Scymnus {Pullus) creperus vdir, fraternus: Casey, 1899, p. 140. — Leng, 1920, p. 213.
Scymnus {Pullus) haemorrhous: Casey, 1899, p. 140.— Leng, 1920, p. 213.— Wilson,
1927, p. I70.-Korschefsky, 1931, p. 159.
Scymnus dentipes Fall, 1901, p. 234. — Fall, 1904, p. 176.
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Fig. 125. Scymnus (P.) fmternus.
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185
Fig. 126. Distribution. Scymnus (P.) fraternus (peripheral and disjunct localities dotted).
Scymnus (Pullus) fraternus: Wingo, 1952, p. 31.— J. Chapin, 1973, p. 1071.— J.
Chapin, 1974, p. 24.-Gordon, 1976b, p. 109.
For detailed description, and discussion see Gordon, 1976b, p. 109.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 127. Scymnus (P.) louisianae.
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187
Scymnus (Pullus) louisianae J. Chapin
Fig. 127a-d; Map, Fig. 128
Scymnus {Pullus) louisianae]. Chapin, 1973, p. 1071.— J. Chapin, 1974, p. 24.—
Gordon, 1976b, p. 115.
For detailed description, and discussion see Gordon, 1976b, p. 115.
Additional locality record: TEXAS: Hidalgo Co., Mac Allen.
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Scymnus {Pullus) apithanus Gordon
Fig. 129a-d; Map, Fig. 132
Scymnus {Pullus) apithanus Gordon, 1976b, p. 118.
For detailed description, and discussion see Gordon, 1976b, p. 118.
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Fig. 130. Scymnus (P.) loewii.
Scymnus {Pullus) loewii Mulsant
Fig. 130a-h; Map, Fig. 131
Scymnus {Pullus) loewii Mulsant, 1850, p. 908.— Gorham, 1897, p. 111.— Kor-
schefsky, 1931, p. 161. — Leng and Mutchler, 1933, p. 87.— J. Chapin, 1974, p.
27. -Gordon, 1976b, p. 119.
Scymnus loewii: Crotch, 1874b, p. 271.
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Fig. 132. Distribution. Scymnus (P.) apithanus (star); S. {P.) marginicollis (shaded, periph-
eral and disjunct localities dotted).
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Fig. 133. Scymnus (P.) marginicollis.
Scymnus cinctus LeConte, 1852, p. 137.-Crotch, 1874b, p. 263. — Horn, 1895, p. 99.
Scymnus (Pullus) cinctus: Gorham, 1897, p. 227. — Casey, 1899, p. 152.— Wilson,
1927, p. 169.-Leng, 1920, p. 214.-Korschefsky, 1931, p. 156.-Wingo, 1952,
p. 30.
Scymnus suturalis LeConte, 1852, p. 138 (not Thunberg, 1 795). — Crotch, 1874b, p.
264.
Scymnus lecontei Crotch, 1874b, p. 264. — Horn, 1895, p. 99.
Scymnus {Pullus) lecontei: Gorhdim, 1897,p. 227. — Casey, 1899,p. 152. — Leng, 1920,
p. 214. — Korschefsky, 1931, p. 161.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 134. Scymnus (P.) postpictus.
Scymnus flebilis Horn, 1895, p. 100.
Scymnus (Pullus) fJebilis: Casey, 1899, p. 160. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 158. -Gordon, 1976b, p. 120.
Scymnus (Pullus) sarpedon Casey , 1899, p. 152. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 165. -Gordon, 1976b, p. 120.
Scymnus (Pullus) nubes Casey, 1899, p. 151. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 163. -Gordon, 1976b, p. 120.
Scymnus scot ti Nunenmacher, 1934, p. 17. — Gordon, 1976b, p. 120.
For detailed description, and discussion see Gordon, 1976b, p. 119.
Additional locality record: TEXAS: Garza Co.; 2 mi. N. Justiceburg.
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Fig. 135. Distribution. Scymnus (P.) postpictus.
Scymnus (Pullus) marginicollis Mannerheim
Fig. 133a-e; Map, Fig. 132
Scymnus marginicollis Mannerheim, 1843, p. 313. — Mulsant, 1850, p. 1053.—
LeConte, 1852, p. I40.-Crotch, 1874b, p. 267. -Horn, 1895, p. 104.
Scymnus {Pullus) marginicollis: Casey, 1899, p. 142. — Leng, 1920, p. 213.— Kor-
schefsky, 1931, p. 161.— Wingo, 1952, p. 34. — Hatch, 1961, p. 150. — Gordon,
1976b, p. 125.-Belicek, 1976, p. 304.
Scymnus californicus Boheman, 1859, p. 207.
Scymnus {Pullus) californicus: Casey, 1899, p. 142. — Leng, 1920, p. 213.
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Fig. 136. Scymnus {P.) socer.
Scymnus (Pu/lus) desertorum Casey, 1899, p. 145. — Leng, 1920, p. 213.— Korschef-
sky, 1931, p. 157. -Gordon, 1976b, p. 125.
Scymnus (Pullus) marginicoUis borealis Hatch, 1961, p. 150. — Gordon, 1976b, p.
125.
For detailed description, and discussion see Gordon, 1976b, p. 125.
Scymnus {Pullus) postpictus Casey
Fig. 134a-e; Map, Fig. 135
Scymnus {Pullus) postpinctus Casey, 1899, p. 141 (lapsus). — Korschefsky, 1931, p.
164.
Scymnus {Pullus) postpictus Casey, 1908, p. 405 (emendation). — Leng, 1920, p. 213.—
Gordon, 1976b, p. 1 30. — Belicek, 1976, p. 304.
For detailed synonymy, description, and discussion see Gordon, 1976b, p. 130.
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Scymnus (Pullus) socer LeConte
Fig. 136a-d; Map, Fig. 137
Scymnus socer 'LQConlQ, 1852, p. 139. — Crotch, 1874b, p. 267. — Horn, 1895, p. 103.
Scymnus {Pullus) socer: Casey, 1899, p. 144. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 166.-J. Chapin, 1974, p. 29.-Gordon, 1976b, p. 133.
Scymnus {Pullus) kinzeli Casey, 1899, p. 143. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 160 {kinzelii).-]. Chapin, 1974, p. 30.
Scymnus {Pullus) innocens Casey, 1899, p. 145. — Leng, 1920, p. 213.— Korschefsky,
1931, p. I60.-Gordon, 1976b, p. 133.
For detailed description, and discussion see Gordon, 1976b, p. 133.
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Fig. 138. Scymnus (P.) tenebrosus.
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Fig. 139. Distribution. Scymnus {P.) tenebrosus.
Scymnus (Pullus) tenebrosus Mulsant
Fig. 138a-e; Map, Fig. 139
Scymnus {Pullus) tenebrosus Mulsant, 1850, p. 989. — Casey, 1899, p. 148.— Leng,
1920,p. 213.-Korschefsky, 1931, p. 166.-Wingo, 1952, p. 40.-J. Chapin, 1974,
p. 30.-Gordon, 1976b, p. 137.
Scymnus tenebrosus: LeConte, 1852, p. 140. — Crotch, 1874b, p. 268. — Horn, 1895,
p. 106.
For detailed description, and discussion see Gordon, 1976b, p. 137.
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Fig. 140. Scymnus (P.) falli.
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Fig. 141. Distribution. Scymnus (P.) falli (star); S. P. solidus (peripheral localities dotted).
Scymnus (Pullus) falli Gordon
Fig. 140a-e; Map, Fig. 141
Scymnus (Pullus) falli Gordon, 1976b, p. 140.
For detailed description, and discussion see Gordon, 1976b, p. 140.
Additional locality record: CALIFORNIA; Ventura Co., Santa Barbara Island.
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Scymnus (Pullus) solidus Casey
Fig. 142a-d; Map, Fig. 141
Scymnus {Pullus) solidus Casey, 1899, p. 145. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 166. -Gordon, 1976b, p. 143.
Scymnus {Pullus) blaisdelli Casey, 1899, p. 147. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 155. — Hatch, 1961, p. 150. — Gordon, 1976b, p. 143.
For detailed description, and discussion see Gordon, 1976b, p. 143.
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Fig. 143. Scymnus {P.) garlandicus.
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Fig. 144. Distribution. Scymnus {P.) garlandicus (star); S. {P.) jacobianus (dot).
Scymnus (Pul/us) garlandicus Casey
Fig. 143a-d; Map, Fig. 144
Scymnus (Pul/us) garlandicus CsLsey, 1899, p. 147. — Leng, 1920, p. 213.— Korschef-
sky, 1931, p. 159. -Gordon, 1976b, p. 145.
For detailed description, and discussion see Gordon, 1976b, p. 145.
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Fig. 145. Scymnus (P.) jacobianus.
Scymnus (Pullus) jacobianus Casey
Fig. 145a-d; Map, Fig. 144
Scymnus {Pullus) jacobianus CdiSty, 1899, p. 148. — Leng, 1920, p. 213.— Korschef-
sky, 1931, p. I60.-Gordon, 1976b, p. 148.
Scymnus {Pullus) jacinto CdiSQy, 1899, p. 148. — Leng, 1920, p. 213.— Korschefsky,
1931, p. I60.-Gordon, 1976b, p. 148.
Scymnus {Pullus) extricatus Casey , 1899, p. 148. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 158. -Gordon, 1976b, p. 148.
For detailed description, and discussion see Gordon, 1976b, p. 148.
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Scymnus (Pullus) humboldti Casey
Fig. 146a-d; Map, Fig. 147
Scymnus {Pullus) humboldti Casey, 1899, p. 146. — Leng, 1920, p. 213.— Korschef-
sky, 1931, p. 160. -Gordon, 1976b, p. 150.
For detailed description, and discussion see Gordon, 1976b, p. 150.
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Fig. 147. Distribution. Scymnus {P.) humboldti (disjunct locality dotted).
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Scymnus {Pul I us) barber i Gordon
Fig. 148a-d; Map, Fig. 149
Scymnus (Pullus) barberi Gordon, 1976b, p. 153.
For detailed description, and discussion see Gordon, 1976b, p. 153.
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Scyfnnus (Pullus) iowensis Casey
Fig. 150a-d; Map, Fig. 149
Scymnus {Pullus) iowensis C?iSQy , 1899, p. 143. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 160. — Wingo, 1952, p. 41. — Gordon, 1976b, p. 156.
Scymnus collaris Melsheimer, 1847, p. 180 (not Herbst, 1797). — LeConte, 1852, p.
I41.-Mulsant, 1856, p. 152.-Horn, 1895, p. 103.-Blatchley, 1910, p. 529.-
Weise, 1929, p. 33.
Scymnus {Pullus) collaris: Casey, 1899, p. 143. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 162.
Scymnus melsheimeri Weise, 1929, p. 33 (replacement name).
For detailed description, and discussion see Gordon, 1976b, p. 156.
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Fig. 150. Scymnus (P.) iowensis.
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Scymnus (Pullus) calaveras Casey
Fig. 51a-d; Map, Fig. 152
Scymnus {Pullus) calaveras Casey, 1899, p. 150. — Bowditch, 1902, p. 207.— Casey,
1910, p. llO.-Leng, 1920, p. 213.-Korschefsky, 1931, p. 155. -Malkin, 1943b,
p. 193. -Hatch, 1961, p. 151. -Gordon, 1976b, p. 159.-Belicek, 1976, p. 306.
Scymnus {Pullus) saginatus CdiS^Qy , 1899, p. 150. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 165.-Gordon, 1976b, p. 159.
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Fig. 152. Distribution. Scymnus {P.) calaveras (disjunct locality dotted).
Scymnus {PuUus) strenuus CsiSQy , 1899, p. 150. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 166. -Malkin, 1943b, p. 194.-Hatch, 1961, p. 151. -Gordon, 1976b,
p. 159.
Scymnus {Pullus) sty gicus CdiSQy, 1899, p. 151. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 166. — Gordon, 1976b, p. 159.
Scymnus {Pullus) tenuivestis Casey, 1899, p. 151. — Leng, 1920, p. 213.— Gordon,
1976b, p. 159.
Scymnus {Pullus) calaveras ab. tenuivestis: Korschefsky, 1931, p. 156.
For detailed description, and discussion see Gordon, 1976b, p. 159.
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Scymnus (Pullus) majus, new name
Fig. 153a-d
Scymnus {Pullus) majusculus Wingo, 1952, p. 40. — Gordon, 1976b, p. 163 (not
Scymnus {Pullus) majusculus Mader, 1950).
Diagnosis. Description and distribution. — See Gordon (1976b). It has been pointed
out to me by Herbert Dozier that the name majusculus Wingo, 1952, is a homonym
of majusculus Mader, 1950. I therefore propose the name majus, a Latin adjective
referring to the large size, for this species.
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Scymnus {Pullus) ignarus Gordon
Fig. 154a-d; Map, Fig. 155
Scymnus {Pullus) ignarus Gordon, 1976b, p. 163.
For detailed description, and discussion see Gordon, 1976b, p. 163.
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Fig. 155. Distribution. Scymnus {P.) ignarus (star); S. (P.) monticola (rectangle); S. {P.)
utahensis (dot).
Scymnus (Pullus) monticola Casey
Fig. 155
Scymnus {Pullus) monticola Casey, 1899,p. 146. — Leng, 1 920, p. 213.— Korschefsky,
1931, p. 162.-Gordon, 1976b, p. 165.
For detailed description, and discussion see Gordon, 1976b, p. 165.
214
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Scymnus (Pu/lus) utahensis Gordon
Fig. 156a-d; Map, Fig. 155
Scymnus {Pullus) utahensis Gordon, 1976b, p. 165.
For detailed description, and discussion see Gordon, 1976b, p. 165.
Additional locality records: OREGON: Harney Co.
1985
NORTH AMERICAN COCCINELLIDAE
215
Fig. 157. Scymnus (P.) renoicus.
Scymnus (Pullus) renoicus Casey
Fig. 157a-d; Map, Fig. 158
Scymnus {Pullus) renoicus Casey, 1899, p. 149. — Bowditch, 1902, p. 207.— Leng,
1920,p.213.-Korschefsky, 1931, p. 161. -Hatch, 1961, p. 151. -Gordon, 1976b,
p. 169.
For detailed description, and discussion see Gordon, 1976b, p. 169.
Additional locality records: IDAHO: Rupert. UTAH: Millard Co., Hawbush Dunes,
SE Delta.
216
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Fig. 158. Distribution. Scymnus (P.) renoicus (star); S. (P.) horni (shaded, peripheral local-
ities dotted).
Scymnus (Pullus) horni Gorham
Fig. 159a-d; Map, Fig. 158
Scymnus {Pullus) horni Gorham, 1897, p. 229. — Casey, 1899, p. 144. — Leng, 1920,
p. 213. — Korschefsky, 1931, p. 159. — Gordon, 1976b, p. 172.
For detailed description, and discussion see Gordon, 1976b, p. 172.
Additional locality records: TEXAS: Brewster Co., Marathon; Culberson Co., SE
Van Horn; Garza Co. 2 mi. N. Justiceburg; Hudspeth Co. 10 mi. S. Cornudas.
1985
NORTH AMERICAN COCCINELLIDAE
217
Fig. 1 59. Scymnus {P.) horni.
218
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Fig. 160. Scymnus (P.) mormon.
Scymnus (Pullus) mormon Casey
Fig. 160a-d; Map, Fig. 161
Scymnus {Pullus) mormon Casey, 1899, p. 150. — Leng, 1920, p. 213. — Casey, 1924,
p. 176.-Korschefsky, 1931, p. 162. -Gordon, 1976b, p. 175.
Scymnus {Pullus) subsimilis Casey, 1899, p. 150. — Casey, 1910, p. 109.— Casey,
1924, p. 176.
For detailed description, and discussion see Gordon, 1976b, p. 175.
1985
NORTH AMERICAN COCCINELLIDAE
219
Fig. 161. Y^isXrihviXion. Scymnus {P.) mormon.
220
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Scymnus (Pullus) ardelio Horn
Fig. 162a-d; Map, Fig. 163
Scymnus ardelio Horn, 1895, p. 105.
Scymnus {Pullus) ardelio: Casey, 1899, p. 148. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 154.-Hatch, 1961, p. 50.-Gordon, 1976b, p. 177.-Belicek, 1976, p.
305.
Scymnus {Pullus) apacheanus Casey, 1899, p. 146. — Leng, 1920, p. 213.— Kor-
schefsky, 1931, p. 154. — Gordon, 1976b, p. 177.
Scymnus {Pullus) decipiens Casey, 1899, p. 147 (not Weise, 1885). — Leng, 1920, p.
213.-Weise, 1929, p. 33. -Gordon, 1976b, p. 177.
Scymnus sanctus Weise, 1929, p. 33 (new name for decipiens Casey). — Korschefsky,
1931, p. 165.
For detailed description, and discussion see Gordon, 1976b, p. 177.
Additional locality records: IDAHO: Tuttle. NEW MEXICO: Lea Co., 8 mi. E.
Lovington. TEXAS: Culberson Co., 13 mi. W. Van Horn; Pecos Co., 6 mi. N. Pyote.
1985
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221
Fig. 163. T)isXri\)\xXion. Scymnus (P.) ardelio.
222
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Fig. 164. Scymnus (P.) erythwnotum.
Scymnus (Pullus) erythwnotum Gordon
Fig. 164a-d; Map, Fig. 165
Scymnus {Pullus) erythronotum Gordon, 1976b, p. 181.
For detailed description, and discussion see Gordon, 1976b, p. 181.
1985
NORTH AMERICAN COCCINELLIDAE
223
Fig. 165. Distribution. Scymnus {P.) erythronoturn.
224
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Scymnus (Pullus) aridus Casey
Fig. 166a-d; Map, Fig. 167
Scymnus {Pullus) aridus CdiSQy, 1899, p. 146. — Casey, 1924, p. 176. — Leng, 1920, p.
213. — Korschefsky, 1931, p. 154. — Gordon, 1976b, p. 184.
For detailed description, and discussion see Gordon, 1976b, p. 184.
Additional locality records: UTAH: San Juan Co., 26 mi. S. Hanksville.
1985
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225
Fig. 167. Distribution. Scymnus {P.) aridus.
226
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Scyrnnus {Pullus) aridoides Gordon
Fig. 168a-d; Map, Fig. 169
Scyrnnus {Pullus) aridoides Gordon, 1976b, p. 187.
For detailed description, and discussion see Gordon, 1976b, p. 187.
1985
NORTH AMERICAN COCCINELLIDAE
227
Fig. 169. Distribution. Scymnus {P.) aridoides.
Fig. 170. Scymnus {P.) consobrinus.
Scymnus (Pullus) consobrinus LeConte
Fig. 170a-d; Map, Fig. 171
Scymnus consobrinus l.QCon\Q, 1852, p. 139. — Horn, 1895, p. 103.— Crotch, 1874b,
p. 266.
Scymnus {Pullus) consobrinus: Mulsant, 1853, p. 153. — Casey, 1899, p. 142.— Leng,
1920, p. 213.-Korschefsky, 1931, p. 157.-Wingo, 1952, p. 42.-Gordon, 1976b,
p. 190.
For detailed description, and discussion see Gordon, 1976b, p. 190.
1985
NORTH AMERICAN COCCINELLIDAE
229
Fig. 171. Distribution. Scymnus (P.) consobrinus.
230
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Fig. 172. Scymnus (P.) mendocino.
Scymnus {PuUus) mendocino Casey
Fig. 172a-d; Map, Fig. 173
Scymnus (Pullus) mendocino Casey, 1899, p. 151. — Leng, 1920, p. 213.— Korschef-
sky, 1931, p. 162. — Gordon, 1976b, p. 193.
For detailed description, and discussion see Gordon, 1976b, p. 193.
1985
NORTH AMERICAN COCCINELLIDAE
231
Fig. 173. Distribution. Scymnus {P.) mendocino.
232
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Fig. 174. Scymnus (P.) cockerelli.
Scymnus {Pullus) cockerelli Casey
Fig. 174a-d; Map, Fig. 175
Scymnus {Pullus) cockerelli CsiSQy , 1899, p. 144. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 156. -Gordon, 1976b, p. 196.
Scymnus mimus Fall, 1901, p. 234. — Gordon, 1976b, p. 196.
Scymnus {Pullus) mimus: Leng, 1920, p. 213. — Korschefsky, 1931, p. 162.
For detailed description, and discussion see Gordon, 1976b, p. 196.
Additional locality record: UTAH: Leeds.
1985
NORTH AMERICAN COCCINELLIDAE
233
Fig. 175. Distribution. Scymnus (P.) cockerelli (dot); S. {P.) earn (star),
234
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig.. 1 76. Scymnus (P.) cam.
Scymnus (PuUus) cam Gordon
Fig. I76a-d; Map, Fig. 175
Scymnus {Pullus) carri Gordon, 1976b, p. 199. — Belicek, 1976, p. 304.
For detailed description, and discussion see Gordon, 1976b, p. 199.
1985
NORTH AMERICAN COCCINELLIDAE
235
Fig. 177. Scymnus (P.) nugator.
Scymnus (Pullus) nugator Casey
Fig. 177a-d; Map, Fig. 178
(Pw//w5) nugator Casey, 1899, p. 140. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 163. -Gordon, 1976b, p. 199.
For detailed description, and discussion see Gordon, 1976b, p. 199.
236
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 178. Distribution. Scymnus {P.) nugator.
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NORTH AMERICAN COCCINELLIDAE
237
Fig. 179. Scymnus (P.) neomexicanus.
Scymnus (Pullus) neomexicanus Gordon
Fig. 179a-d; Map, Fig. 180
Scymnus {Pullus) neomexicanus Gordon, 1976b, p. 203.
For detailed description, and discussion see Gordon, 1976b, p. 203.
238
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 1 80. Distribution. Scymnus (P.) neomexicanus (star); S. P. nuttingi (dot).
1985
NORTH AMERICAN COCCINELLIDAE
239
Fig. 181. Scymnus (P.) nuttingi.
Scymnus {Pul I us) nuttingi Gordon
Fig. 181a-d; Map, Fig. 180
Scymnus (PuUus) nuttingi Gordon, 1976b, p. 204.
For detailed description, and discussion see Gordon, 1976b, p. 204.
240
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 182. Scymnus (P.) compar.
Scymnus (Pullus) compar Casey
Fig. 182a-d; Map, Fig. 183
Scymnus {Pullus) compar CdiSQy , 1899, p. 148. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 156.— J. Chapin, 1974, p. 33. — Gordon, 1976b, p. 206.
Scymnus {Pullus) vicksburgicus Casey, 1924, p. 175. — Leng and Mutchler, 1927, p.
33. — Korschefsky, 1931, p. 167. — Gordon, 1976b, p. 206.
Scymnus {Pullus) impunctus 1952, p. 35.— J. Chapin, 1974, p. 33.— Gordon,
1976b, p. 207.
For detailed description, and discussion see Gordon, 1976b, p. 206.
1985
NORTH AMERICAN COCCINELLIDAE
241
Fig. 183. Distribution. Scymnus (P.) compar.
242
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 184. Scymnus (P.) impletus.
Scymnus (Pullus) impletus Gordon
Fig. 184a-d; Map, Fig. 185
Scymnus {Pullus) impletus Gordon, 1976b, p. 209.
For detailed description, and discussion see Gordon, 1976b, p. 209.
1985
NORTH AMERICAN COCCINELLIDAE
243
Fig. 185. Distribution. Scymnus (P.) impletus (shaded, peripheral localities dotted); S. (P.)
simulans (star); S. (P.) wingoi (rectangle).
244
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 186. Scymnus (P.) simulans.
Scymnus (Pullus) simulans Gordon
Fig. 186a, b; Map, Fig. 185
Scymnus (Pullus) simulans Gordon, 1976b, p. 214.
For detailed description, and discussion see Gordon, 1976b, p. 214.
1985
NORTH AMERICAN COCCINELLIDAE
245
Fig. 187. Scymnus (P.) wingoi.
Scymnus (Pullus) wingoi Gordon
Fig. 187a-d; Map, Fig. 185
Scymnus {Pullus) wingoi Gordon, 1976b, p. 215.
For detailed description, and discussion see Gordon, 1976b, p. 215.
246 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Scymnus (Pu/lus) abbreviatus LeConte
Fig. 188a-d; Map, Fig. 189
Scymnus abbreviatus 1852, p. 140. — Crotch, 1874b, p. 268. — Horn, 1895,
p. 104.
Scymnus {Pullus) abbreviatus: Casey, 1899, p. 153. — Leng, 1920, p. 213.— Kor-
schefsky, 1931, p. 153.— Gordon, 1976b, p. 216.
For detailed description, and discussion see Gordon, 1976b, p. 216.
1985
NORTH AMERICAN COCCINELLIDAE
247
Fig. 189. Distribution. Scymnus {P.) abbreviatus.
248
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 190. Scymnus (P.) tenebricus.
Scymnus (Pullus) tenebricus Gordon
Fig. 190a-d; Map, Fig. 191
Scymnus {Pullus) tenebricus Gordon, 1976b, p. 220.
For detailed description, and discussion see Gordon, 1976b, p. 220.
1985
NORTH AMERICAN COCCINELLIDAE
249
Fig. 191. Distribution. Scymnus (P.) tenebricus.
250
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Scymnus (Pullus) wickhami Gordon
Fig. I92a-d; Map, Fig. 193
Scymnus {Pullus) wickhami Gordon, 1976b, p. 223.
For detailed description, and discussion see Gordon, 1976b, p. 223.
1985
NORTH AMERICAN COCCINELLIDAE
251
Fig. 193. Distribution. Scymnus {P.) wickhami (dot); S. (P.) elusivus (star).
252
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 194. Scymnus (P.) elusivus.
Scymnus (Pullus) elusivus Gordon
Fig. I94a-d; Map, Fig. 193
Scymnus {Pullus) elusivus Gordon, 1976b, p. 226.
For detailed description, and discussion see Gordon, 1976b, p. 226.
Additional locality record: CALIFORNIA: San Diego Co., Julian.
1985
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253
Fig. 195. Scymnus (P.) uteanus.
Scymnus (Pullus) uteanus Casey
Fig. 195a-d; Map, Fig. 196
Scymnus {Pullus) uteanus CdiSQy , 1899, p. 144. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 167. -Gordon, 1976b, p. 226.
Scymnus {Pullus) rhesus Casey, 1899, p. 144. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 165.— Gordon, 1976b, p. 228.
For detailed description, and discussion see Gordon, 1976b, p. 226.
254
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 196. Distribution. Scymnus (P.) uteanus (dot); S. {P.) papago (star).
1985
NORTH AMERICAN COCCINELLIDAE
255
Scymnus (Pullus) papago Casey
Fig. 197a, b; Map,, Fig. 196
Scymnus {Pullus) papago CdiSQy , 1899, p. 151. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 164.-Gordon, 1976b, p. 230.
For detailed description, and discussion see Gordon, 1976b, p. 230.
256
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 198. Scymnus {P.) uncus.
Scymnus (Pullus) uncus Wingo
Fig. 198a-d; Map, Fig. 199
Scymnus {Pullus) uncus Wingo, 1952, p. 38.— J. Chapin, 1974, p. 32.— Gordon,
1976b, p. 232.
For detailed description, and discussion see Gordon, 1976b, p. 232.
1985
257
NORTH A
Ml
ERICAN COCCINELLIDAE
Fig. 199. Distribution. Scymnus (P.) uncus.
258
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 200. Scymnus (P.) puncticollis.
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NORTH AMERICAN COCCINELLIDAE
259
Scymnus {Pullus) puncticollis LeConte
Fig. 200a-d; Map, Fig. 201
Scymnus puncticollis LeConte, 1852, p. 139.— Crotch, 1874b, p. 266. — Horn, 1895,
p. 102.
Scymnus {Pullus) puncticollis: Casey, 1899, p. 160. — Leng, 1920, p. 133.— Kor-
schefsky, 1931, p. 165. — Wingo, 1952, p. 37.— J. Chapin, 1974, p. 31.— Gordon,
1976b, p. 232.
For detailed description, and discussion see Gordon, 1976b, p. 232.
260
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 202. Scymnus (P.) weidti.
Scymnus (Pullus) weidti Casey
Fig. 202a-d; Map, Fig. 203
Scymnus {Pullus) weidti Casey, 1899, p. 149. — Leng, 1920, p. 213.— Casey, 1924, p.
176.-Korschefsky, 1931, p. 167. -Gordon, 1976b, p. 237.
For detailed description, and discussion see Gordon, 1976b, p. 237.
1985
NORTH AMERICAN COCCINELLIDAE
261
Fig. 203. Distribution. Scymnus (P.) weidti (dot); S. (/*.) aquilonarius (star).
262
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 204. Scymnus (/*.) aquilonarius.
Scymnus (Pullus) aquilonarius Gordon
Fig. 204a-d; Map, Fig. 203
Scymnus {Pullus) aquilonarius Gordon, 1976b, p. 240. — Belicek, 1976, p. 305.
For detailed description, and discussion see Gordon, 1976b, p. 240.
1985
NORTH AMERICAN COCCINELLIDAE
263
Fig. 205.
Distribution. Scymnus (P.) martini (star); S. (P.) hesperius (dot).
264
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 206. Scymnus {P.) martini.
Scymnus {Pullus) martini Gordon
Fig. 206a-d; Map, Fig. 205
Scymnus {Pullus) martini Gordon, 1976b, p. 240.
For detailed description, and discussion see Gordon, 1976b, p. 240.
1985
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265
Fig. 207. Scymnus (P.) hesperius.
Scymnus (Pullus) hesperius Gordon
Fig. 207a-d; Map, Fig. 205
Scymnus {Pullus) hesperius Gordon, 1976b, p. 243.
For detailed description, and discussion see Gordon, 1976b, p. 243.
Additional locality record: CALIFORNIA: Eldorado Co., Pollock Pines.
266
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 208. Scymnus (P.) luctuosus.
Scyrnnus (Pullus) luctuosus Casey
Fig. 208a-d; Map, Fig. 209
Scymnus {Pullus) luctuosus CdiSQy , 1899, p. 146. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 161. -Gordon, 1976b, p. 245.
Scymnus {Pullus) sonomae CdiSQy , 1899, p. 147. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 166. — Gordon, 1976b, p. 245.
Scymnus {Pullus) advena Casey, 1899, p. 147. — Leng, 1920, p. 213.—
Korschefsky, 1931, p. 153. — Gordon, 1976b, p. 245.
For detailed description, and discussion see Gordon, 1976b, p. 245.
1985
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267
Fig. 209. Distribution. Scymnus {P.) luctuosus (dot); 5”. {P.) nevadensis (star).
268
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 210. Scymnus {P.) nevadensis.
Scymnus {Pullus) nevadensis Weise
Fig. 210a-d; Map, Fig. 209
Scymnus nevadensis Weise, 1929, p. 33. — Leng and Mutchler, 1933, p. 35.
Scymnus {Scymnus) nevadensis: Korschefsky, 1931, p. 163.— Gordon, 1976b, p. 248.
Scymnus {Scymnus) innocuus Casey, 1899, p. 154 (not Boheman, 1859).— Leng,
1920, p. 214.
For detailed description, and discussion see Gordon, 197613, p. 248.
Scymnus {Pullus) lacustris LeConte
Fig. 211a-d; Map, Fig. 212
Scymnus lacustris l^GConlQ, 1850, p. 239. — LeConte, 1852, p. 140.— Crotch, 1874b,
p. 268. -Horn, 1895, p. 105.
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269
Scymnus (Pullus) lacustris: Mulsant, 1850, p. 989. — Mulsant, 1853, p. 153.— Casey,
1899, p. 149.-Leng, 1920, p. 213.-Korschefsky, 1931, p. 160.-Hatch, 1961, p.
151. -Gordon, 1976b, p. 250.-Belicek, 1976, p. 305.
Scymnus {Pullus) cultratus Wingo, 1952, p. 38.— Gordon, 1976b, p. 250.
For detailed description, and discussion see Gordon, 1976b, p. 250.
270
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 212. Distribution. Scymnus {P.) lacustris.
Scymnus (Pullus) tahoensis Casey
Fig. 2I3a-h; Map, Fig. 214
Scymnus {Pullus) tahoensis CdiS^y , 1899, p. 150. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 166. -Gordon, 1976b, p. 253.-Belicek, 1976, p. 305.
For detailed description, and discussion see Gordon, 1976b, p. 253.
1985
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271
Fig. 213. Scymnus (P.) tahoensis.
272
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 214. Distribution. Scymnus (P.) tahoensis.
Scymnus {Pullus) caudalis LeConte
Fig. 215a-d; Map, Fig. 216
Scymnus caudalis LeConte, 1850, p. 238. — LeConte, 1852, p. 139. — Horn, 1895, p.
103.
Scymnus {Pullus) caudalis LeConte: Casey, 1899, p. 143. — Leng, 1920, p. 213.—
Korschefsky, 1931, p. 156. — Gordon, 1976b, p. 256.
Scymnus {Pullus) natchezianus Casey, 1899, p. 143. — Leng, 1920, p. 213.— Kor-
schefsky, 1931, p. 163.— Gordon, 1976b, p. 256.
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273
Fig. 215. Scymnus (P.) caudalis.
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Scymnus (Pullus) indut us Casey, 1899, p. 145. — Leng, 1920, p. 213.— Korschefsky,
1931, p. I60.-Gordon, 1976b, p. 256.
Scymnus {Pullus) agricola Casey, 1899, p. 145. — Leng, 1920, p. 213.— Korschefsky,
1931, p. 153. — Gordon, 1976b, p. 256.
For detailed description, and discussion see Gordon, 1976b, p. 256.
1985
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275
Fig. 217. Scymnus {P.) peninsularis.
Scymnus (Pullus) peninsularis Gordon
Fig. 217a-d; Map, Fig. 216
Scymnus {Pullus) peninsularis Gordon, 1976b, p. 259.
For detailed description, and discussion see Gordon, 1976b, p. 259.
276
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 218. Scymnus {P.) creperus.
Scymnus (Pullus) creperus Mulsant
Fig. 218a-d; Map, Fig. 219
Scymnus {Pullus) creperus Mulsant, 1850, p. 985. — Mulsant, 1853, p. 153.— Casey,
1899, p. 140.-Leng, 1920, p. 213.-Korschefsky, 1931, p. 1 57.- J. Chapin, 1974,
p. 28. -Gordon, 1976b, p. 260.
Scymnus creperus: LeConte, 1852, p. 139.— Crotch, 1874b, p. 265. — Horn, 1895, p.
101.
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111
Fig. 219. Distribution. Scymnus {P.) creperus (peripheral localities dotted).
Scymnus (Pullus) medionotans Casey, 1899, p. 143. — Leng, 1920, p. 213.— Kor-
schefsky, 1931, p. 162.— J. Chapin, 1974, p. 29.
Scymnus (Pullus) subtropicus Casey, 1899, p. 143. — Leng, 1920, p. 213.— Korschef-
sky, 1931, p. 162.— J. Chapin, 1974, p. 29.
Scymnus (Pullus) hortensis Wingo, 1952, p. 36.— J. Chapin, 1974, p. 29.
For detailed description, and discussion see Gordon, 1976b, p. 260.
Additional locality records: TEXAS: Garza Co., 2 mi. N. Justiceburg.
VIRGINIA: Virginia Beach.
278
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Scyrnnus (Pullus) bryanti Gordon
Fig. 220a-d; Map, Fig. 221
Scyrnnus {Pullus) bryanti Gordon, 1976b, p. 263.
For detailed description, and discussion see Gordon, 1976b, p. 263.
1985
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279
Fig. 221. Distribution. Scymnus {P.) bryanti (dot); 5'. {P.) howdeni (star); S. {P.) hubbardi
(rectangle); S. {P.) huachucha (open circle).
280
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Fig. 222. Scymnus {P.) howdeni.
Scymnus (Pullus) howdeni Gordon
Fig. 222a-d; Map, Fig. 221
Scymnus {Pullus) howdeni Gordon, 1976b, p. 265.
For detailed description, and discussion see Gordon, 1976b, p. 265.
Scymnus {Pullus) hubbardi Gordon
Fig. 223a-d; Map, Fig. 221
Scymnus {Pullus) hubbardi Gordon, 1976b, p. 268.
For detailed description, and discussion see Gordon, 1976b, p. 268.
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281
Fig. 223. Scymnus (/*.) hubbardi.
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Fig. 224. Scymnus (P.) huachucha.
Scymnus (Pullus) huachuca Gordon
Fig. 224a-d; Map, Fig. 221
Scymnus {Pullus) huachuca Gordon, 1976b, p. 269.
For detailed description, and discussion see Gordon, 1976b, p. 269.
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Fig. 225. Scymnus {P.) brullei.
Scymnus (Pullus) brullei Mulsant
Fig. 225a-h; Map, Fig. 226
Scymnus {Pullus) brullei M\x\^2ecii, 1850, p. 984.— Casey, 1899, p. 160.— Leng, 1920,
p. 213.-Korschefsky, 1931, p. 155.-Wingo, 1952, p. 33. -J. Chapin, 1974, p.
26.-Gordon, 1976b, p. 270.
Scymnus brullei: Crotch, 1874b, p. 264. — Horn, 1895, p. 101.
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Scymnus{Pullus)hemorrhous\2iY.divisusCdLSQy, 1899, p. 140.— Leng, 1920, p. 213.—
Korschefsky, 1931, p. 159. — Wingo, 1952, p. 33.
Scymnus {Pullus) hemorrhous var. laurenticus Casey, 1899, p. 140. — Leng, 1920, p.
2 13. -Korschefsky, 1931, p. 159. -Wingo, 1952, p. 33.
Scymnus {Pullus) hemorrhous var. subaeneus Casey, 1899, p. 141. — Leng, 1920, p.
2 13. -Korschefsky, 1931, p. 159. -Wingo, 1952, p. 33.
Scymnus {Pullus) lodi Stehr, 1946, p. 80. — Wingo, 1952, p. 30. — Gordon, 1970b, p.
270.
For detailed synonymy, description, and discussion see Gordon, 1976b, p. 270.
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Scymnus {Pullus) securus J. Chapin
Fig. 227a-b; Map, Fig. 228
Scymnus {Pullus) securus J. Chapin, 1973, p. 1072.— J. Chapin, 1974, p. 25.— Gor-
don, 1976b, p. 275.
For detailed description, and discussion see Gordon, 1976b, p. 275.
Additional locality record: ONTARIO: Kent Co., Tilbury.
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Genus Nephus Mulsant
Scymnus {Nephus) Mulsant, 1846, p. 237. — Mulsant, 1850, p. 958.— Casey, 1899,
p. 167. — Korschefsky, 1931, p. 116. — Mader, 1950, p. 60. — Wingo, 1952, p. 19.—
Bielewski, 1959, p. 49.— Arnett, 1963, p. 812.— J. Chapin, 1974, p. 33. Type-
species; Sphaeridium quadrimaculatum Herbst, by subsequent designation of Kor-
schefsky, 1931.
Nephus: Pope, 1957, p. 309.— Chapin, 1965, p. 200. — Gordon, 1976b, p. 276.—
Belicek, 1976, p. 306.
Scymnini with antenna 10 or 1 1 -segmented, basal 2 segments fused or not (Figs.
229a-f). Prosternum lacking intercoxal carinae except short carina often present
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287
adjacent to coxal cavity, not extending anterior to cavity (Fig. 229g). Abdomen with
6 visible sterna; postcoxal line on first sternum incomplete, nearly reaching lateral
margin, apical end either parallel to hind margin of sternum or recurved toward basal
margin (Fig. 229h-j). Tarsus 3-segmented. Female genitalia lacking infundibulum,
genital plate long, narrow, triangular (Fig. 229k); male genitalia simple, basal lobe
asymmetrical or symmetrical.
The genus Nephus was revised in detail by Gordon (1976b), therefore, only ad-
ditional locality records and some necessary corrections in synonymy are included
for each species, and one additional species is described.
Key to subgenera of Nephus
1. Postcoxal line on first abdominal sternum parallel to hind margin of sternum, at
most with only extreme apex curved forward (Fig. 229j); antenna 10-segmented,
basal 2 segments tightly joined (Fig. 229d) 2
- Postcoxal line not completely parallel to hind margin of sternum, definitely curved
forward apically (Fig. 229 h, i) 3
2(1). Body dorsoventrally flattened (Fig. 2551); antenna short, club oval (Fig. 2291) ....
Depressoscymnus Gordon
- Body not dorsoventrally flattened; antenna of normal length, club with inner margin
of segments discontinous (Fig. 229d) Scymnobius Casey
3( 1 ). Postcoxal line strongly curved forward along lateral border of first sternum, extending
onto basal half of sternum Sidis Mulsant
- Postcoxal line not extending onto basal half of sternum, gently curved forward
apically 4
4(3). Antenna 1 1 -segmented (Fig. 229b) Mulsant
Antenna 10-segmented (Fig. 229e) Turbocymnus Gordon
Subgenus Nephus Mulsant
Nephus Mulsant, 1846, p. 237. — Mulsant, 1850, p. 958.— Casey, 1899, p. 167.—
Wingo, 1952, p. 19. — Bielawski, 1959, p. 49.— Arnett, 1963, p. 812.— Gordon,
1976b, p. 278. — Belicek, 1976, p. 306. Type-species; Sphaeridium quadrimacu-
latum Herbst, by subsequent designation of Korschefsky, 1931.
Antenna 1 1 -segmented, basal 2 segments fused or at least tightly joined (Fig. 229b).
Abdomen with postcoxal line incomplete, distinctly curved forward apically, not
parallel to hind margin of first sternum (Fig. 229h).
Key to subspecies of Nephus {Nephus) ornatus (LeConte)
1. Elytron with irregular, elongate, yellow spot (Fig. 232); north of New England, west
and north of the Great Lakes ornatus naviculatus (Casey)
- Elytron with 2 large, yellow spots (Fig. 230f); New England west to Great Lakes ....
ornatus ornatus (LeConte)
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Fig. 229. a-f. Antennae, a. Nephus (Nephus) quadrimaculatus. b. Nephus (Nephus) ornatus
ornatus. c. Nephus (Sidis) binaevatus. d. Nephus (Scymnobius) sordidus. e. Nephus {Turboscyrn-
nus) georgei. f. Nephus (Depressoscymnus) schwarzi. g. Prostemum of Nephus sp. h. Postcoxal
line of Nephus {N.) ornatus ornatus. i. Postcoxal line of Nephus (Turboscymnus) georgei. j.
Postcoxal line of Nephus {Scymnobius sp.).
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Nephus (Nephus) ornatus ornatus (LeConte)
Fig. 230a-f; Map, Fig. 231
Scymnus ornatus 1850, p. 239. — LeConte, 1852, p. 135. — Crotch, 1874b,
p. 260.-Hom, 1895, p. 94.
Scymnus {Scymnobius) ornatus: Casey, 1899, p. 155. — Leng, 1920, p. 214.
Scymnus {Nephus) ornatus: Korschefsky, 1931, p. 164.
Scymnus {Scymnobius) sanguinifer C?iSQy , 1899, p. 155. — Leng, 1920, p. 214.
Scymnus {Nephus) sanguinifer: Korschefsky, 1931, p. 165.— Gordon, 1976b, p. p.
280.
Scymnus frosti Casey, 1924, p. 171. — Gordon, 1976b, p. 280.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93( 1 )
Fig. 23 1 . Distribution. Nephus {N.) ornatus ornatus (star); N. (N.) ornatus naviculatus (dot).
Nephus ornatus: Belicek, 1976, p. 307.
Nephus {Nephus) ornatus ornatus: Gordon, 1976b, p. 280.
For detailed description, and discussion see Gordon, 1976b, p. 280.
Nephus {Nephus) ornatus naviculatus (Casey)
Fig. 232; Map, fig. 231
Scymnus {Scymnobius) naviculatus Casey, 1899, p. 155. — Leng, 1920, p. 214.
Scymnus {Nephus) naviculatus: Korschefsky, 1931, p. 163.
Scymnus {Scymnus) kincaidi Hatch, 1961, p. 152. — Gordon, 1976b, p. 282.
Nephus {Nephus) ornatus naviculatus: Gordon, 1976b, p. 282.
For detailed description, and discussion see Gordon, 1976b, p. 282.
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NORTH AMERICAN COCCINELLIDAE
291
Fig. 232. Nephus {N.) naviculatus.
Subgenus Sidis Mulsant
Scymnus (Sidis) Mulsant, 1850, p. 975. — Korschefsky, 1931, p. 117.— Bielawski,
1959, p. 42. — Fursch, 1960, p. 305.— Gordon, 1976b, p. 282. Type-species: Scym-
nus (Sidis) binaevatus Mulsant, by subsequent designation of Korschefsky, 1931.
Nephus (Sidis): Gordon, 1976b, p. 282.
Antenna 1 0-segmented, large basal segment undivided (Fig. 229c); apical segment
of maxillary palpus cylindrical, obliquely truncate apically. Postcoxal line on 1st
abdominal sternum incomplete, nearly reaching lateral margin, curved forward par-
allel to lateral margin. Male genitalia with basal lobe asymmetrical (Fig. 233a);
spermathecal capsule of female divided into spindle-shaped nodulus and annulated
cornu, accessory gland opening at middle of nodus (Fig. 23 3e).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 233. Nephus {Sidis) binaevatus.
Nephus (Sidis) binaevatus (Mulsant)
Fig. 233a-e; Map, Fig. 234
Scymnus (Sidis) binaevatus Mulsant, 1850, p. 975. — Korschefsky, 1931, p. 150.—
Mader, 1950, p. 121. -Pope, 1957, p. 295.
Nephus (Sidis) binaevatus: Gordon, 1976b, p. 284.
For detailed description, and discussion see Gordon, 1976b, p. 284.
Additional locality records: CALIFORNIA: San Mateo Co., Daly City.
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293
Fig. 234. Distribution. Nephus (Sidis) binaevatus.
Subgenus Turboscymnus Gordon
Turboscymnus Gordon, 1976b, p. 287. Type-species: Scymnus georgei Weise, by
monotypy.
Antenna 1 0-segmented, one large basal segment present showing slight indication
of fusion (Fig. 229e); apical segment of maxillary palpus cylindrical, obliquely trun-
cate apically. Postcoxal line on 1st abdominal sternum curved throughout, apical
end approaching lateral border of sternum, curved forward (Fig. 229i). Male genitalia
with basal lobe asymmetrical; female spermathecal capsule feebly curved medially
(Fig. 235e).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 235. Nephus (Turboscymnus) georgei.
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NORTH AMERICAN COCCINELLIDAE
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Nephus (Turboscymnus) georgei (Weise)
Fig. 235a-e; Map, Fig. 236
Scymnus bisignatus Worn, 1895, p. 92. — (not Boheman, 1859).— Weise, 1929, p. 33.
Scymnus (Scymnobius) bisignatus: Casey, 1899, p. 160. — Leng, 1920, p. 214.
Scymnus georgei VsfQist, 1929, p. 33. — Korschefsky, 1931, p. 159.
Scymnus {Scymnus) bisignatus: Hatch, 1961, p. 153.
Nephus georgei: Belicek, 1976, p. 307.
Nephus {Turboscymnus) georgei (Weise): Gordon, 1976b, p. 287.
For detailed description, and discussion see Gordon, 1976b, p. 287.
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Subgenus Scymnobius Casey
Scymnus (Scymnobius) Casey, 1899, p. 139.— Weise, 1905, p. 220. — Leng, 1920, p.
213. — Korschefsky, 1931, p. 116. — Hatch, 1961, p. 153.— Arnett, 1963, p. 812.—
Gordon, 1976, p. 290. — Type-species; Scymnus Jlavifrons Melsheimer, by subse-
quent designation of Gordon, 1976b.
Nephus {Scymnobius): Gordon, 1976b, p. 290.
Antenna 10-segmented, basal 2 segments very tightly joined (Fig. 229d); apical
segment of maxillary palpus cylindrical, obliquely truncate apically. Postcoxal line
on 1st abdominal sternum running parallel to hind margin of sternum, not reaching
lateral margin, apex may be slightly curved forward. Male genitalia with basal lobe
symmetrical or asymmetrical; female spermathecal capsule bent or curved at ap-
proximately a right angle (Fig. 237e).
Key to species of Nephus {Scymnobius)
1 . Elytron entirely or mostly light yellow or brown 7
Elytron entirely black or at least dark, often with yellow areas or spots 2
2(1). Elytron entirely black; pronotum reddish yellow gordoni {Dozitv)
- Elytron not entirely black, or if so, then pronotum not entirely reddish yellow ... 3
3(2). Elytron completely black or with 2 yellow spots (Fig. 2431) or with one yellow spot
(Fig. 243g) or with 2 spots feebly connected (Fig. 243h) California and Oregon . . .
atramentarius (Boheman)
Elytron not as described above, or if so, not occurring in California or Oregon ... 4
4(3). Elytron with 2 yellow spots; Arizona quadrarius (Casey)
Elytron variable but never with 2 distinctly defined, yellow spots 5
5(4). Elytron with a more or less rounded, yellow or reddish yellow spot on apical half 6
- Elytron with 2 irregularly transverse yellow areas (Figs. 24 1 e, f), areas often obscurely
connected medially guttulatus (LeConte)
6(5). Form nearly round; pronotum entirely reddish yellow or with only a small, basal
area darkened; Florida bivulnerus (Horn)
- Form elongate; pronotum usually entirely black or with antero-lateral angle narrowly
pale; not restricted to Florida Jlavifrons (Melsheimer)
7(1). Elytron dark brown in basal ‘, apical ‘ paler brownish red (Fig. 252a)
timberlakei, n. sp.
- Elytron unicolorous or with only sutural margin darkened 8
8(7). Form short, rounded; pronotum distinctly paler than elytron wickhami Gordon
Form elongate, narrow, pronotum and base of elytron unicolorous 9
9(3). Occuring in eastern United States from Atlantic Coast to eastern Texas
intrusus (Horn)
- Occuring in western United States from Pacific Coast to western Texas, Colorado
and Idaho sordidus (Horn)
Nephus {Scymnobius) Jlavifrons (Melsheimer)
Fig. 237a-f; Map, Fig. 238
Scymnus Jlavifrons Melsheimer, 1847, p. 181. — LeConte, 1852, p. 136.— Crotch,
1874b, p. 261. -Horn, 1895, p. 93.-Blatchley, 1910, p. 526.
Scymnus {Scymnobius) Jlavifrons: Casey, 1899, p. 155. — Leng, 1920, p. 214.
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NORTH AMERICAN COCCINELLIDAE
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Fig. 237. Nephus {S.) flavifrons.
Scymnus {Nephus) flavifrons: Korschefsky, 1931, p. 158. — Wingo, 1952, p. 43.— J.
Chapin, 1974, p.34.
Scymnus {Nephus) bioculatus Mulsant, 1850, p. 960.
Scymnus bioculatus: LeConte, 1852, p. 136.— Crotch, 1874b, p. 261.
Scymnus flavifrons var. bioculatus: Horn, 1895, p. 93.
Scymnus {Scymnobius) bioculatus: Casey, 1899, p. 155.
Scymnus {Nephus) flavifrons ab. bioculatus: Korschefsky, 1931, p. 158.
Scymnus {Nephus) bioculatus guttiger Mulsant, 1850, p. 961.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 238. Distribution. Nephus (S.) flavifrons (peripheral and disjunct localities dotted).
Scymnus bioculatus var. guttiger: Horn, 1895, p. 93.
Scymnus {Nephus) flavifrons ab. guttiger: Korschefsky, 1931, p. 158.
Scymnus {Nephus) bioculatus marginellus Mulsant, 1850, p. 961.
Scymnus bioculatus var. marginellus: Horn, 1895, p. 93.
Scymnus {Nephus) flavifrons marginellus: Korschefsky, 1931, p. 158.
Scymnus ludovicianus Casey, 1924, p. 172. — Leng and Mutchler, 1927, p. 33.— J.
Chapin, 1974, p. 34. -Gordon, 1976b, p. 292.
Nephus {Scymnobius) flavifrons: Gordon, 1976b, p. 292.
For detailed description, and discussion see Gordon, 1976b, p. 292.
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Fig. 239. Nephus (S.) bivulnerus.
Nephus (Scymnobius) bivulnerus (Horn)
Fig. 239a-f; Map, Fig. 240
Scymnus bivulnerus Horn, 1895, p. 92.
Scymnus {Scymnobius) bivulnerus: Casey, 1899, p. 155. — Leng, 1920, p. 214.
Scymnus {Nephus) bivulnerus: Korschefsky, 1931, p. 155.
Nephus {Scymnobius) bivulnerus: Gordon, 1976b, p. 295.
For detailed description, and discussion see Gordon, 1976b, p. 295.
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Fig. 240. Distribution. Nephus (S.) bivulnerus.
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Fig. 241. Nephus (S.) guttulatus.
Nephus {Scymnobius) guttulatus (LeConte)
Fig. 241a-g; Map, Fig. 242
Scymnus guttulatus "LQConXQ, 1852, p. 136.— Crotch, 1874b, p. 261. — Horn, 1895,
p. 95.
Scymnus {Scymnobius) guttulatus: Casey, 1899, p. 155. — Leng, 1920, p. 214.
Scymnus {Nephus) guttulatus: Korschefsky, 1931, p. 159.
Scymnus coloradensis Horn, 1895, p. 94. — Leng, 1920, p. 214.— Gordon, 1976b,
298.
Scymnus {Scymnobius) coloradensis: Casey, 1899, p. 156.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 242. Distribution. Nephus (S.) guttulatus.
Scymnus {Nephus) coloradensis: Korschefsky, 1931, p. 156.
Scymnus (Scymnobius) scitus Casey, 1899, p. 156. — Leng, 1920, p. 214.— Korschef-
sky, 1931, p. 165. — Gordon, 1976b, p. 298.
Scymnus {Scymnobius) suavis Casey, 1899, p. 156. — Leng, 1920, p. 214.— Gordon,
1976b, p. 298.
Scymnus {Nephus) suavis: Korschefsky, 1931, p. 166.
Nephus {Scymnobius) guttulatus: Gordon, 1976b, p. 298.
For detailed description, and discussion see Gordon, 1976b, p. 299.
Additional locality records: CALIFORNIA: San Diego Co., Mouth of Tijuana R.;
San Luis Obispo Co., Oceano, Dune Lakes 3 mi. S.
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Fig. 243. Nephus (S.) atramentarius.
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Fig. 244. Distribution. Nephus (S.) atramentarius.
Nephus (Scymnobius) atramentarius (Boheman)
Fig. 243a-h; Map, Fig. 244
Scymnus atramentarius BohQm?in, 1859, p. 207. — Leng, 1920, p. 214.— Korschefsky,
1931, p. 154.
Scymnus {Scymnobius) maculatus Hatch, 1961, p. 153.— Gordon, 1976b, p. 302.
Nephus {Scymnobius) atramentarius: Gordon, 1976b, p. 301.
For detailed description, and discussion see Gordon, 1976b, p. 301.
Additional locality record: IDAHO: Caribou Co., Soda Springs.
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Fig. 245. Nephus {S.) quadrarius.
Nephus {Scymnobius) quadrarius (Casey)
Fig. 245a-e; Map, Fig. 246
Scymnus quadrarius Casey, 1924, p. 173.— Leng, and Mutchler, 1927, p. 33.
Scymnus schuberti^unQwvcvdichQY, 1934a, p. 17. — Gordon, 1976b, p. 303.
Nephus {Scymnobius) quadrarius: Gordon, 1976b, p. 303.
For detailed description, and discussion see Gordon, 1976b, p. 303.
Additonal locality record: ARIZONA: Santa Cruz Co., Madera Canyon.
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Fig. 246,
Distribution. Nephus (S.) quadrarius (dot); N. (S.) wickhami (star).
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Fig. 247. Nephus (S.) wickhami.
Nephus {Scymnobius) wickhami Gordon
Fig. 247a-d; Map, Fig. 246
Nephus {Scymnobius) wickhami Gordon, 1976b, p. 306.
For detailed description, and discussion see Gordon, 1976b, p. 306.
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Fig. 248. Nephus (S.) sordidus.
Nephus (Scyrnnobius) sordidus (Horn)
Fig. 248a-e; Map, Fig. 249
Scymnus sordidus Horn, 1895, p. 93.
Scymnus {Scyrnnobius) sordidus: Casey, 1899, p. 156. — Leng, 1920, p. 214.
Scymnus {Nephus) sordidus: Korschefsky, 1931, p. 166.
Scymnus {Scyrnnobius) intrusoides Hatch, 1961, p. 153. — Gordon, 1976b, p. 309.
Nephus sordidus: Belicek, 1976, p. 307.
Nephus {Scyrnnobius) sordidus: Gordon, 1976b, p. 309.
For detailed description, and discussion see Gordon, 1976b, p. 309.
Additional locality records: CALIFORNIA: Imperial Co., Glamis; Riverside Co.,
Rice Dunes; San Diego, Co., mouth of Tijuana R.
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Fig. 249. Distribution. Nephus {S.) sordidus.
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b
Fig. 250. Nephus (S.) intrusus.
Nephus {Scymnobius) intrusus (Horn)
Fig. 250a-b; Fig. 251c-e; Map, Fig. 253
Scymnus intrusus Horn, 1895, p. 92. — Blatchley, 1910, p. 526.
Scymnus {Scymnobius) intrusus: Casey, 1899, p. 156. — Leng, 1920, p. 214.
Scymnus {Nephus) intrusus: Korschefsky, 1931, p. 160.— Wingo, 1952, p. 43.— J.
Chapin, 1974, p. 34.
Scymnus {Scymnobius) inops: Casey, 1899, p. 156. — Leng, 1920, p. 214.— Gordon,
1976b, p. 312.
Scymnus {Nephus) inops: Korschefsky, 1931, p. 160.
Nephus {Scymnobius) intrusus: Gordon, 1976b, p. 312.
For detailed description, and discussion see Gordon, 1976b, p. 312.
Additional locality records: TEXAS: Jones Co., 2 mi. w. Noodle; Tom Green Co.,
16 mi. NE San Angelo.
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Fig. 251. Nephus (S.) intrusus.
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Fig. 252. Nephus (S.) timberlakei.
Nephus (Scymnobius) timberlakei, new species
Fig. 252a, b
Description. Female, length 1 .66 mm, width 1.22 mm. Oval, elongate. Color brown-
ish red, elytron dark brown in basal 1/2, dark brown area blended into brownish red
of apical ‘ (Fig. 252a), meso- and metasterna dark brown. Punctures on head and
pronotum extremely fine, nearly invisible. Elytron slightly dull, surface alutaceous,
coarsely punctured, punctures separated by one to 3 times a diameter; pubescence
yellowish white, semi-erect, arranged in S-curve. Postcoxal line parallel, widely sep-
arated from hind margin of 1st sternum, not approaching lateral margin. Apex of
6th sternum barely perceptibly emarginate. Female genitalia as in Figure 252b.
Variation. Length 1.66 to 2.0 mm; width 1.22 to 1.33 mm.
Holotype. Female. TEXAS: Brownsville, Apr. 21, ’15, Timberlake Coll., Salt Lake
Lab No. 9682. USNM (101332).
Paratypes. Total 3. Same data as holotype except collection dates Apr. 4, ’1 5; Apr.
20, ’15; Apr. 21, ’15. (UCR) (USNM).
This species most closely resembles N. intrusus, but N. timberlakei is larger and
has the elytron bicolored. No males were available for study, so the genitalic affinities
remain unknown. I name this species for P. H. Timberlake who first identified it as
an undescribed species, in recognition of his fine work in the Coccinellidae.
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Nephus {Scymnobius) gordoni (Dozier)
Fig. 254a-e; Map, Fig. 253
Scymnus {Scymnobius) gordoni Dozier, 1971, p. 87.
Nephus {Scymnobius) gordoni: Gordon, 1976b, p. 315.
For detailed description, and discussion see Gordon, 1976b, p. 315.
Additional locality record: SOUTH CAROLINA: 7 mi. NE Pickens.
Subgenus Depressoscymnus Gordon
Depressoscymnus Gordon, 1976b, p. 315. Type-species; Nephus {Depressoscymnus)
schwarzi, by monotypy.
Antenna short, 10-segmented, basal 2 segments tightly joined, club broad, oval
outer margin of segments nearly continuous (Fig. 2290; apical segment of maxillary
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NORTH AMERICAN COCCINELLIDAE
315
palpus distinctly securiform. Body flattened dorsoventrally (Fig. 25 5f). Postcoxal line
as in Scymnobius. Male genitalia symmetrical; spermathecal capsule of female of the
Scymnobius type.
Nephus {Depressoscymnus) schwarzi Gordon
' Fig. 255a-f; Map, Fig. 256
Nephus {Depressoscymnus) schwarzi Gordon, 1976b, p. 315.
For detailed description, and discussion see Gordon, 1976b, p. 315.
Additional locality record: ARIZONA: Pima Co., Molino Basin, Mt. Lemmon
Hwy., 4,400'.
Genus Diomus Mulsant
Scymnus {Diomus) Mulsant, 1850, p. 951.— Gorham, 1897, p. 226.— Casey, 1899,
p. 139.— Leng, 1920, p. 213. — Korschefsky, 1931, p. 1 16. — Mader, 1955, p. 955.—
Wingo, 1952, p. 19. -J. Chapin, 1974, p. 35.
Diomus: Weise, 1885a, p. 83. — Mader, 1924, p. 8.— Chapin, 1933, p. 95.— Pope,
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1957, p. 311. — Gordon, 1976b, p. 319. Type-species: Coccinella thomcicus Fa-
bricius, by subsequent designation of Korschefsky, 1931.
Head with clypeal margin truncate, gena extending onto eye beside antennal in-
sertion; antenna 10-segmented, 3rd segment as long as segments 4-6 combined (Fig.
257a); apical segment of maxillary palpus securiform. Prosternum with 2 fine, com-
plete carinae extending to anteror margin of prostemum. Tarsus 3-segmented. Po-
stcoxal line extending down and joining hind margin of 1st abdominal sternum (Fig.
257b). First abdominal sternum fused to 2nd medially; male with sterna 2-6 con-
tracted, 5th sternum broadly, feebly emarginate apically. Male genitalia with basal
lobe asymmetrical; sipho extremely long, slender, or short, evenly curved. Female
genitalia with genital plates short, rounded or truncate apically (Fig. 257c); sperm
duct long and tangled, or short, simple.
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NORTH AMERICAN COCCINELLIDAE
317
The genus Diomus was revised in detail by Gordon (1976b), therefore, only ad-
ditional locality records and some necessary corrections in synonymy are included
for each species. One imported species now established in California is included.
Key to species of Diomus
1.
2(1).
3(2).
4(3).
5(4).
6(5).
7(6).
8(7).
9(8).
10(9).
11(10).
12(11).
13(12).
14(11).
15(14).
Color completely pale yellow or yellowish brown; length less than 1 .40 mm . . .
debilis (LeConte)
Color not completely pale, or if so, then length more than 1 .40 mm 2
Elytron black with single, obliquely transverse, yellow band, anterior to middle
(Fig. 258a); Florida balteatus {l^&ConXQ)
Elytron not as described above; Florida and elsewhere 3
Elytron black except apical V4 yellow (Fig. 276e) terminatus (Say)
Elytron not as described above 4
Elytron black or at least dark with irregular yellow areas (Fig. 269d); California
taedatus (Fall)
Elytron not as described above; species not occurring in California 5
Elytron dark brown with 2 irregularly connected yellow spots (Fig. 268b); Browns-
ville, Texas pseudotaedatus Gordon
Elytron not as described above 6
Apex of elytron with a single yellow spot not reaching suture (Fig. 280d)
roseicollis (Mulsant)
Elytron not as described above 7
Pronotum yellow with large black median spot; elytron almost completely dark,
only narrow border pale, western Texas texanus Gordon
Pronotum and elytron not as described above; not known from western Texas . 8
Pronotum yellow; elytron black or dark with sutural and apical margins obviously
reddish brown, or entirely light yellowish brown; Arizona arizonicus Gordon
Dorsal color not as described above; not occurring in Arizona 9
Form depressed, oval; elytron dark with 2 rounded, yellow spots (Fig. 266d) . .
humilis Gordon
Form not depressed; elytron rarely with 2 yellow spots (except myrmidon) .... 10
Elytron entirely light reddish brown; Texas xanthaspis (Mulsant)
Elytron either entirely dark or dark with a distinct pale pattern 11
Elytron dark with some form of definite pale markings 14
Elytron entirely dark or with obscure, indefinite lightening toward apex, or with
apical border narrowly yellow 12
Elytron entirely black; coastal California (imported species) pumilio Weise
Elytron not entirely black, or if so, then not occurring in coastal California (native
species) 13
Elytron with apex distinctly, narrowly yellow; pronotum entirely reddish yellow
or at most with basal, median projection darkened xanthaspis (Mulsant)
Elytron entirely black or dark or with an obscurely paler area on apical half;
pronotum usually with at least basal half darkened, often entirely piceous
austrinus Gordon
Length 1.75 mm, or more; form robust, convex; elytron with 2 somewhat rounded,
yellow spots (Fig. 265d) myrmidon (Mulsant)
Length less than 1.80 mm; form not strongly robust or convex; elytron rarely with
2 rounded, yellow spots 15
Elytron with single yellow spot on disc, or with yellow spot on disc narrowly
connected to broad, apical, yellow area (Fig. 27 le, f); form strongly rounded . .
bigemmeus (Horn)
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93( 1 )
Fig. 257. Diomus sp. a. Antenna, b. Postcoxal line. c. Genital plates.
- Elytron with color pattern not as described above; form narrow, elongate, or form
short, broad, nearly truncate apically 16
16(15). Form elongate, narrow, parallel sided (Fig. 261 e-j) 17
Form short, broad, nearly truncate apically (Fig. 263d) liebecki (Horn)
17(16). Basal lobe of male genitalia robust, strongly asymmetrical in ventral view (Fig.
261a) amabilis (LeConte)
- Basal lobe of male genitalia slender, not strongly asymmetrical in ventral view
(Fig. 260a) floridanus {MulsanX)
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NORTH AMERICAN COCCINELLIDAE
319
Diomus balteatus (LeConte)
Fig. 258a-b; Map, Fig. 259
Scymnus balteatus LeConte, 1878a, p. 399. — Horn, 1895, p. 87.
Scymnus {Diomus) balteatus: Casey, 1899, p. 156. — Leng, 1920, p. 214.— Korschef-
sky, 1931, p. 155.
Diomus balteatus: Gordon, 1976b, p. 322.
For detailed description, and discussion see Gordon, 1976b, p. 322.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93( 1 )
Diomus floridanus (Mulsant)
Fig. 260a-e; Map, Fig. 259
Hyperapis floridana Mulsant, 1850, p. 1040. — Casey, 1899, p. 128.
Scymnus floridana: Crotch, 1873, p. 379. — Dobzhansky, 1941, p. 85.
Scymnus quadritaeniatus LeConXe, 1878a, p. 400. — Horn, 1895, p. 90. — Leng, 1920,
p. 214.-Korschefsky, 1931, p. 165. -Gordon, 1976b, p. 323.
Scymnus {Diomus) quadritaeniatus: Casey, 1899, p. 157.
Scymnus pellio Blatchley, 1927, p. 142. — Leng and Mutchler, 1933, p. 35.— Kor-
schefsky, 1931, p. 164. — Gordon, 1976b, p. 323.
Diomus floridanus: Gordon, 1976b, p. 322.
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NORTH AMERICAN COCCINELLIDAE
321
For detailed description, and discussion see Gordon, 1976b, p. 322.
Additional locality record: FLORIDA: Gainesville.
Diomus amabilis (LeConte)
Fig. 261a-e; Map, Fig. 262
Scymnus amabilis l.QCon\Q, 1852, p. 135. — Crotch, 1874b, p. 260. — Horn, 1895, p.
94.
Scymnus {Scymnobius) amabilis: Casey, 1899, p. 160. — Leng, 1920, p. 214.— Kor-
schefsky, 1931, p. 154.
Scymnus (Diomus) dulcis Casey, 1899, p. 159. — Leng, 1920, p. 214.— Korschefsky,
1931, p. 158. -Gordon, 1976b, p. 326.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 261. Diomus amabilis.
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NORTH AMERICAN COCCINELLIDAE
323
Scymnus {Diomus) emertoni Casey, 1924, p. 172. — Leng and Mutchler, 1927, p.
33.-Korschefsky, 1931, p. 158. -Gordon, 1976b, p. 326.
Scymnus {Diomus) amabilis: Wingo, 1952, p. 43.— J. Chapin, 1974, p. 35.
Diomus amabilis: Gordon, 1976b, p. 326.
For detailed description, and discussion see Gordon, 1976b, p. 326.
Additional locality record: MISSOURI: Columbia.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 263. Diomus liebecki.
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NORTH AMERICAN COCCINELLIDAE
325
Diomus liebecki (Horn)
Fig. 263a-e; Map, Fig. 264
Scymnus liebecki Horn, 1895, p. 89.— Blatchley, 1910, p. 527.
Scymnus {Diomus) liebecki: Casey, 1899,p. 157. — Leng, 1920, p. 214.— Korschefsky,
1931, p. 161.-Wingo, 1952, p. 43.
Scymnus {Diomus) adulans CdiSQy , 1899, p. 157. — Leng, 1920, p. 214.— Korschefsky,
1931, p. 153.-Gordon, 1976b, p. 329.
Scymnus {Diomus) ohioensis Stehr, 1946, p. 80.— Gordon, 1976b, p. 329.
Diomus liebecki: Gordon, 1976b, p. 329.
For detailed description, and discussion see Gordon, 1976b, p. 329.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 265. Diomus myrmidon.
Diomus myrmidon (Mulsant)
Fig. 265a-d; Map, Fig. 264
Scymnus {Diomus) myrmidon Mulsant, 1850, p. 954.— Crotch, 1874b, p. 261.—
Casey, 1899, p. 157. — Leng, 1920, p. 214. — Korschefsky, 1931, p. 162.
Scymnus myrmidon: LeConte, 1852, p. 136. — Horn, 1895, p. 89.
Diomus myrmidon: Gordon, 1976b, p. 331.
For detailed description, and discussion see Gordon, 1976b, p. 331.
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NORTH AMERICAN COCCINELLIDAE
327
Fig. 266. Diomus humilis.
Diomus humilis Gordon
Fig. 266a-d; Map, Fig. 267
Diomus humilis Gordon, 1976b, p. 333.
For detailed description, and discussion see Gordon, 1976b, p. 333.
Additional locality record: FLORIDA: Punta Gorda.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 267. Distribution. Diomus humilis (dot); D. pseudotaedatus (star).
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NORTH AMERICAN COCCINELLIDAE
329
Fig. 268. Diomus pseudotaedatus.
Diomus pseudotaedatus Gordon
Fig. 268a, b; Map, Fig. 267
Diomus pseudotaedatus Gordon, 1976b, p. 333.
For detailed description, and discussion see Gordon, 1976b, p. 333.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 269. Diomus taedatus.
Diomus taedatus (Fall)
Fig. 269a-d; Map, Fig. 270
Scymnus taedatus Fall, 1901, p. 233.
Scymnus {Diomus) taedatus: Leng, 1920, p. 213. — Korschefsky, 1931, p. 166.
Diomus taedatus: Gordon, 1976b, p. 335.
For detailed description and discussion see Gordon, 1976b, p. 335.
Additional locality record: ARIZONA: Santa Cruz Co., Madera Canyon.
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NORTH AMERICAN COCCINELLIDAE
331
Fig. 270. Distribution. Diomus taedatus.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Diomus bigem mens (Horn)
Fig. 271a-f; Map, Fig. 272
Scymnus bigemmeus Horn, 1895, p. 87. — Blatchley, 1918, p. 421.
Scymnus {Diomus) bigemmeus: Casey, 1899, p. 156. — Leng, 1920, p. 214.— Kor-
schefsky, 1931, p. 155. -J. Chapin, 1974, p. 36.
Scymnus {Diomus) stigma Casey, 1899, p. 158. (not Weise, 1898b). — Leng, 1920, p.
2l4.-Weise, 1929, p. 33. -Gordon, 1976b, p. 338.
Scymnus lunarisV^QiSQ, 1929, p. 33. — Korschefsky, 1931, p. 161. — Gordon, 1976b,
p. 338.
Diomus bigemmeus: Gordon, 1976b, p. 337.
For detailed description, and discussion see Gordon, 1976b, p. 337.
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NORTH AMERICAN COCCINELLIDAE
333
Fig. 272. Distribution. Diomus bigemmeus.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 273. Diomus austrinus.
Diornus austrinus Gordon
Fig. 273a-d; Map, Fig. 274
Diomus austrinus Gordon, 1976b, p. 341.
For detailed description, and discussion see Gordon, 1976b, p. 341.
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NORTH AMERICAN COCCINELLIDAE
335
Fig. 274. Distribution. Diomus austrinus.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 275. Diomus pumilio.
Diomus pumilio Weise
Fig. 275a-d; Map, Fig. 279
Diomus pumilio 1885b, p. 237. — Korschefsky, 1931, p. 148.
Scymnus flavifrons Blackburn, 1889, p. 95. — Blackburn, 1892, p. 250.
Scymnus (Scymnobius) pumilio: Hatch, 1961, p. 153.
Diagnosis. Length 1.35 to 1.60 mm, width 1.0 to 1.20 mm. Form oval, somewhat
oblong. Color black except male with anterior pronotal margin, head, propleuron,
mouthparts, and anterior leg reddish yellow. Male genitalia as in Figure 275a-c.
Female genitalia as in Figure 275d.
Discussion. This is an Australian species imported several times into the United
States and Canada, now known to be established only in coastal California (K. Hagen,
pers. comm.).
Type locality. Of pumilio and flavifrons, South Australia.
Type depository. Of pumilio and flavifrons, types not examined.
Distribution. Figure 279. CALIFORNIA: Marin Co. to San Diego (coastal).
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NORTH AMERICAN COCCINELLIDAE
337
Fig. 276. Diomus terminatus.
Diomus terminatus (Say)
Fig. 276a-e; Map, Fig. 277
Scymnus terminatus Say, 1835, p. 203.— LeConte, 1852, p. 136.— Crotch, 1874b, p.
259. -Horn, 1895, p. 90.
Scymnus {Diomus) terminatus: Mulsant, 1850, p. 952. — Casey, 1899, p. 158.— Leng,
1920,p. 214.-Korschefsky, 1931, p. 166.-Wingo, 1952, p. 43.- J. Chapin, 1974,
p. 36.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 277. Distribution. Diomus terminatus (peripheral localities dotted).
Scymnus femoralis LeConte, 1852, p. 136.— Crotch, 1874b, p. 260. — Horn, 1895,
p. 91.
Scymnus {Diomus) femoralis: Casey, 1899, p. 158. — Leng, 1920, p. 214.
Scymnus {Diomus) terminatus ab. femoralis: Korschefsky, 1931, p. 167.
Scymnus {Diomus) partitus Casey, 1899, p. 158. — Leng, 1920, p. 214.— Korschefsky,
1931, p. 164.-J. Chapin, 1974, p. 36.-Gordon, 1976b, p. 342.
Diomus terminatus: Gordon, 1976b, p. 341.
For detailed description, and discussion see Gordon, 1976b, p. 341.
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NORTH AMERICAN COCCINELLIDAE
339
Diomus texanus Gordon
Fig. 278a-c; Map, Fig. 279
Diomus texanus Gordon, 1976b, p. 346.
For detailed description, and discussion see Gordon, 1976b, p. 346.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 279. Distribution. Diomus texanus (dot); D. pumilio (shaded).
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NORTH AMERICAN COCCINELLIDAE
341
Diomus roseicollis (Mulsant)
Fig. 280a-d; Map, Fig. 28 1
Scymnus {Diomus) roseicollis 1853, p. 142. — Korschefsky, 1931, p. 165.
Scymnus roseicollis: Crotch, 1874b, p. 270. — Dimmock, 1906, p. 382.
Diomus roseicollis: Gordon, 1976b, p. 348.
For detailed description, and discussion see Gordon, 1976b, p. 348.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 28 1 . Distribution. Diomus wseicollis (star); D. xanthaspis (dot).
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NORTH AMERICAN COCCINELLIDAE
343
Fig. 282. Diomus xanthaspis.
Diomus xanthaspis (Mulsant)
Fig. 282a-d; Map, Fig. 281
Scymnus {Diomus) xanthaspis MuXsdinX, 1850, p. 952. — Horn, 1895, p. 90.— Casey,
1899, p. 160.-Leng, 1920, p. 214.-Korschefsky, 1931, p. 167.-Wingo, 1952,
p. 43. -J. Chapin, 1974, p. 37.
Scymnus xanthaspis: LeConte, 1852, p. 136. — Crotch, 1874b, p. 259.
Scymnus {Diomus) houstoni Casey, 1899, p. 158. — Leng, 1920, p. 214.— Korschef-
sky, 1931, p. I60.-Gordon, 1976b, p. 350.
Scymnus {Diomus) appalacheus Casey, 1899, p. 158. — Leng, 1920, p. 214.— Kor-
schefsky, 1931, p. 154.— Gordon, 1976b, p. 350.
Scymnus {Diomus) brunnescens CsLsey, 1899, p. 1 58 (not Motschulsky, 1866).— Leng,
1920, p. 214.-Weise, 1929, p. 33.-Korschefsky, 1931, p. 154.-Gordon, 1976b,
p. 350.
Scymnus caseyi Weise, 1929, p. 33 (not Brethes, 1924). — Korschefsky, 1931, p.
167, — Gordon, 1976b, p. 350.
Scymnus caseyianus Leng and Mutchler, 1933, p. 35.
Diomus xanthaspis: Gordon, 1976b, p. 348.
For detailed description, and discussion see Gordon, 1976b, p. 348.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 283. Diomus arizonicus.
Diomus arizonicus Gordon
Fig. 283a-d; Map, Fig. 284
Diomus arizonicus Gordon, 1976b, p. 353.
For detailed description, and discussion see Gordon, 1976b, p. 353.
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NORTH AMERICAN COCCINELLIDAE
345
Fig. 284. Distribution. Diomus arizonicus.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 285. Diomus debilis.
Fig. 286. Distribution. Diomus debilis.
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NORTH AMERICAN COCCINELLIDAE
347
Diomus debilis (LeConte)
Fig. 285a-d; Map, Fig. 286
Scymnus debilis LeConte, 1852, p. 137,— Crotch, 1874b, p. 263. — Horn, 1895, p.
91.
Scymnus {Diomus) debilis: Casey, 1899, p. 159.— Leng, 1920, p. 214.— Korschefsky,
1931, p. 157.
Scymnus {Diomus) pusio Casey, 1899, p. 159. — Leng, 1920, p. 214.— Korschefsky,
1931, p. 165. -Gordon, 1976b, p. 354.
Scymnus {Diomus) aeger CdiSQy, 1899, p. 159. — Leng, 1920, p. 214. — Wingo, 1952,
p. 43.— Gordon, 1976b, p. 354.
Scymnus {Diomus) molliculus Casey, 1924, p. 175. — Leng and Mutchler, 1927, p.
33.-Wingo, 1952, p. 43.-Gordon, 1976b, p. 354.
Scymnus {Diomus) minutissimus Casey, 1924, p. 176 (not de Villers, 1789).— Leng
and Mutchler, 1927, p. 33.— Gordon, 1976b, p. 354.
Scymnus {Diomus) minor , 1931, p. 162.
Diomus debilis (LeConte): Gordon, 1976b, p. 354.
For detailed synonymy, description, and discussion see Gordon, 1976b, p. 354.
Additional locality records: CALIFORNIA: El Centro; Inyo Co., Saline Valley.
TEXAS: Hudspeth Co., 10 mi. S. Comudas.
Selvadiini, new tribe
Hyperaspinae with form elongate, oblong, dorsoventrally flattened; dorsal surface
strongly pubescent. Antenna short, with fusiform club. Intercoxal process of pro-
stemum bicarinate. Epipleuron narrow, flat, not foveate for reception of femoral
apices. Leg free, simple, not enlarged or expanded; tarsus cryptotetramerous. Ab-
domen with 6 visible sterna; basal sternum broad, fused to 2nd sternum medially.
Male genitalia with basal lobe asymmetrical. Female genital plate short, transverse.
The genus Selvadius had previously been considered a member of the Scymnini
because of the obviously pubescent dorsal surface. However, the antenna is typically
hyperaspine, and the head is also hyperaspine in that it is broad apically, and partially
conceals the antennal insertions. The male and female genitalia do not particularly
resemble those of members of either the Hyperaspini or Scymini. Selvadius shows
some affinity to the genus Hyperaspidius (Hyperaspini), which lacks obvious dorsal
pubescence, and also resembles some members of the genus Diomus (Scymnini) in
the form of the female genital plate, sperm duct and male sipho. I prefer to erect a
new tribe for this genus rather than force it into either the Hyperaspini or Scymnini
because to do so would, in either case, cause an undesirable expansion of the tribal
limits.
Genus Selvadius Casey
Selvadius Casey, 1899, p. 137.— Leng, 1920, p. 213.— Korschefsky, 1931, p. 111.—
Gordon, 1970a, p. 45.— Gordon, 1976b, p. 8. Type-species; Selvadius rectus Casey,
by monotypy.
Diagnosis. Length less than 2.50 mm. Color pale yellowish brown to dark reddish
brown. Form elongate, oblong, dorsoventrally flattened. Head broad, surface convex,
width between eyes about 3 times the width of an eye; eye completely exposed;
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 287. Selvadius sp. a. Head. b. Antenna, c. Hind leg. d. Postcoxal line. e. Female genitalia.
clypeal apex broadly emarginate or nearly truncate, anterolateral angle abrupt, lateral
border emarginate at antennal insertion with flange partially covering antennal in-
sertion (Fig. 287a). Antenna 11-segmented (Fig. 287b). Apical segment of maxillary
palpus strongly securiform. Pronotum with intercoxal process raised, flat between
carinae which extend nearly to apex of prostemum. Tarsal claw with strong median
tooth (Fig. 287c). Postcoxal line on first abdominal sternum incomplete, of Scymnus
type (Fig. 287d). Male genitalia with basal lobe strongly curved in lateral view;
paramere broad, strongly narrowed in apical Vb; trabes longer than phallobase (Fig.
288); sipho with extremely long, attenuated apical portion (Fig. 290b). Female gen-
italia without definite spermathecal capsule; sperm duct long, coiled; genital plate
transverse, base narrowed, produced (Fig. 287e).
Discussion. Selvadius is apparently restricted to North America and is represented
there by 4 described species. No information is available on the biology of members
of this genus, but they may be similar to members of Hyperaspidius in this respect.
Species of both genera have been collected in grassland communities, very near the
ground, and Selvadius must feed on insects (almost certainly scale insects) associated
with grasses or herbs of that community. The type series of S. maderi was collected
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NORTH AMERICAN COCCINELLIDAE
349
Fig. 288. Selvadius rectus.
from oak {Quercus agrifolia). Because of the pubescent dorsal surface and incomplete
postcoxal lines, species of Selvadius are most likely to be confused with species of
Scymnus (Scymnus). The widely separated, completely exposed eyes and partially
concealed antennal insertions will distinguish Selvadius from members of the Scym-
nini.
Key to species of Selvadius
1. Length less than 1.50 mm 2
- Length 1.60 mm or more 3
2(1). Pronotal punctures coarse, nearly contiguous; Arizona, Texas rectus Casey
Pronotal punctures fine, not obvious, separated by the diameter of a puncture or
more; California maderi (Nunenmacher)
3(1). Length 1.60 to 2.0 mm; Colorado, Wyoming nunenmacheri Gordon
- Length 2.0 to 2.25 mm; Arizona, California, New Mexico megacephalus (Fall)
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 289. Selvadius maderi.
Selvadius rectus Casey
Fig. 288; Map, Fig. 293
Selvadius rectus Casey, 1899, p. 138. — Korschefsky, 1931, p. 111. — Gordon, 1970a,
p. 45.
Diagnosis. Length 1.40 mm, width 0.80 mm. Form elongate, not exactly parallel
sided, widest at middle of elytron. Postcoxal line short, widely incomplete. Male
genitalia as in Figure 288.
Discussion. This species resembles S. maderi in the small size and overall ap-
pearance. The only external difference that I have been able to detect is the size and
density of the pronotal punctures which are much coarser and closer together in S.
rectus than in 5’. maderi. However, I’ve seen only the type and 2 additional specimens
of S. rectus, so this character may not be constant. The type is a unique (holotype)
male in the Casey collection.
Type locality. Tucson, Arizona.
Type depository. USNM (35250).
Distribution. Figure 293. ARIZONA: Tucson. CALIFORNIA: Inyo Co., Eureka
Valley. TEXAS: El Paso.
Selvadius maderi (Nunenmacher), new combination
Fig. 289a, b; Map, Fig. 293
Scymnus maderi Nunenmacher, 1937, p. 183.
Scymnus quercus Nunenmacher, 1934, p. 18 (not Scymnus quercus Mulsant, 1850).
Diagnosis. Length 1.20 to 1.40 mm, width 0.75 to 0.90 mm. Similar to S. rectus
in all respects except pronotal punctures finer, less dense. Male genitalia as in Figure
289a, b.
Discussion. A male cotype is here designated as the lectotype, and 3 other type
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NORTH AMERICAN COCCINELLIDAE
351
Fig. 290. Selvadius nunenmacheri.
specimens designated as paralectotypes. I previously examined the types of this
species, and realized that it belonged to Selvadius, but I neglected to formally transfer
it although it was not included in the Scymnus revision (Gordon, 1976b).
Type locality. Vine Hill, Contra Costa Co., California (lectotype here designated).
Type depository. CAS.
Distribution. Figure 293. CALIFORNIA: type locality.
Selvadius nunenmacheri Gordon
Fig. 290a-c; Map, Fig. 293
Selvadius nunenmacheri Gordon, 1970a, p. 45.
Diagnosis. Length 1.55 to 2.35 mm, width 1.0 to 1.35 mm. Form varies from
parallel sided to slightly oval. Male genitalia as in Figure 290a, b. Female genitalia
as in Figure 290c.
Discussion. I regard the specimens recorded here as composing a single polymorphic
species. There is noticeable variation in size and some variation in the form of the
postcoxal line and body shape. The male genitalia, however, are constant throughout
and I cannot find a pattern in the observed variation. There are wide gaps in the
known distribution of S. nunenmacheri, and I expect that specimens from appropriate
localities will confirm the integrity of this species.
Type locality. Colorado Springs, Colorado.
Type depository. USNM (70401).
Distribution. Figure 293. COLORADO: Colorado Springs; Nunn, Pawnee Grass-
land. WYOMING: Cheyenne; Tipton.
Selvadius megacephalus (Fall)
Fig. 291a-c; Map, Fig. 293
Scymnus megacephalus Fall, 1901, p. 233. — Korschefsky, 1931, p. 162.
Selvadius megacephalus: Gordon, 1970a, p. 45.
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Fig. 291. Selvadius megacephalus.
Diagnosis. Length 2.0 to 2.25 mm, width 1.60 to 1.70 mm. Form parallel sided
to slightly oval (Fig. 291c). Dorsal surface reddish brown except narrow sutural
margin dark brown. Male genitalia as in Figure 291a, b.
Discussion. The type locality is Pasadena, California, and I have seen one additional
California specimen. Examples from Arizona and New Mexico appear to match the
type exactly; therefore, I refer them to this species. No solid external differences are
apparent that will separate S. megacephalus and S. nunenmacheri except size, and
there is a slight overlap even there. However, the male genitalia are noticeably
different, and I regard both species as valid. The type specimen is a unique (holotype)
female in the Fall collection.
Type locality. Pasadena, California.
Type depository. MCZ.
Distribution. Figure 293. ARIZONA: Santa Catalina Mts.; Santa Cruz Co., Mowry;
Tucson. CALIFORNIA: Cathedral City; Pasadena. NEW MEXICO: Hot Springs.
Tribe Hyperaspini
Hyperaspini Costa, 1849, pp. 9, 64.— Casey, 1899, p. 1 1 5. — Blatchley, 1910, p. 519.—
Korschefsky, 1932, p. 176.— Chapin, 1966, p. 278.— J. Chapin, 1974, p. 38.—
Belicek, 1976, p. 294.
Hyperaspidini, Wingo, 1952, p. 17.
Hyperaspiens Mulsant, 1850. p. 2.
Hyperaspidae Berg, 1874, p. 291.
Hyperaspides Crotch, 1873, p. 377.— Crotch, 1874b, p. 208.— Gorham, 1894, p. 183.
Hyperaspites Chapuis, 1876, p. 166.
Hyperaspidina Jacobson, 1916, p. 969.
Scymninae of small to medium size, 1.50 to 5.0 mm in length; form ranges from
elongate oval, depressed, to rounded, convex. Dorsal surface glabrous except in
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NORTH AMERICAN COCCINELLIDAE
353
Blaisdelliana. Antenna short, 9 to 1 1 -segmented; club elongate, fusiform, apical
segment small, recessed in preceding segment. Eye large, entire or weakly notched,
finely faceted, without pubescence. Maxillary palpus with apical segment securiform.
Scutellum usually large. Epipleuron of elytron narrow, usually flat, usually excavated
for reception of femoral apex except Hypemspidius and Blaisdelliana. Leg short;
femur grooved for reception of tibia; tarsus cryptotetramerous. Abdomen with 6
visible sterna in female, 7 sterna visible in male. Male genitalia asymmetrical. Female
coxal plate usually short, transverse, stylus reduced or absent.
I recognize 6 genera in America north of Mexico as belonging to this tribe. Five
of these have traditionally been placed here, and I now transfer Blaisdelliana Gordon
from the Scymnini to the Hyperaspini. Chapin (1966) was the first to critically study
the genera of Western Hemisphere Hyperaspini using internal characters as well as
external characters, and he succeeded in creating order from the chaos that previously
existed. El-Ali (unpubl. dissertation) further refined Chapin’s preliminary work to
provide a solid generic classification.
Key to genera of Hyperaspini
1 . Dorsal surface strongly pubescent Blaisdelliana Gordon
- Dorsal surface glabrous 2
2(1). Anterior tibia with external tooth or spine (Fig. 458c) Brachiacantha Chevrolat
Anterior tibia without external tooth or spine 3
3(2). Epipleuron of elytron not excavated for reception of middle and hind femoral apices
(Fig. 295b) Hyperaspidius CroXcYi
Epipleuron of elytron excavated for reception of middle and hind femora apices (Fig.
333d) 4
4(3). Epipleuron of elytron strongly descending externally; anterior tibia wide, angulate
or rounded anteriorly at external margin (Fig. 330b); elytron greenish black with red
spot behind middle Thalassa Mulsant
- Epipleuron of elytron flat or feebly inclined; anterior tibia slender or enlarged apically;
elytron not greenish black 5
5(4). Femur short, stout; tibia enlarged apically (Fig. 327b); elytron reddish brown, without
maculation (Fig. 328e); rare Helesius Casey
Femur slender; tibia slender, not enlarged apically (Fig. 3331); elytron usually black
or brown with pale maculation, rarely immaculate Hyperaspis Redtenbacher
Genus Blaisdelliana Gordon
Blaisdelliana Gordon, 1970a, p. 43. Type-species; Hyperaspis sexualis Casey, by
monotypy.
Hyperaspini with form broad, somewhat elongate, appearing almost rectangular;
length less than 2.0 mm; entire dorsal surface pubescent. Head elongate, inclined
downward. Antenna 10-segmented (Fig. 292b); antennal insertion exposed. Eye en-
tire, small, widely separated. Clypeus nearly parallel sided, apex emarginate (Fig.
292a). Lateral margin of pronotum rounded. Epipleuron of elytron narrow, flat, not
excavated for reception of middle or hind femoral apices. Prosternum with 2 faint
carinae extending nearly to apex. Posterior margin of metastemum nearly on equal
plane with abdomen between coxa and lateral margin. Leg long, slender; anterior
tibia simple; tarsal claw without basal tooth. Postcoxal line on first abdominal ster-
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Fig. 292. Blaisdelliana sexualis. a. Head. b. Antenna, c. Postcoxal lines, d-g. Male genitalia,
h. Habitus.
num incomplete, of Scymnus type (Fig. 292c). Apical abdominal sternum of male
truncate. Male genitalia with basal lobe slightly asymmetrical, paramere rooted in
phallobase (Fig. 292d). Female genitalia with compound spermatheca, basal portion
without appendix, coxal plate transverse (Fig. 292g).
Blaisdelliana is known only from the southwestern United States, and I recognize
only one species in the genus. Casey (1899) correctly placed sexualis in the Hyper-
aspini, Dobzhansky (1941) transferred it to the Scymnini, and Gordon ( 1970a) erected
the genus Blaisdelliana for it but retained it in the Scymnini. Present examination
of the female genitalia and antennae of this species has resulted in a reevaluation of
its position and I now consider it to belong in the Hyperaspini. The only morpho-
logical characteristic (albeit a most obvious one) that has caused B. sexualis to be
considered a scymnine is the presence of dorsal pubescence. In all other respects it
is a hyperaspine. Blaisdelliana is most similar to Hypemspidius but the dorsal pu-
bescence and produced anterolateral clypeal angles of Blaisdelliana will distinguish
that genus. No host data are available.
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NORTH AMERICAN COCCINELLIDAE
355
Fig. 293. Distribution. Selvadius rectus (star); S. maderi (open circle); S. nunenmacheri
(dot); S. megacephalus (square).
Blaisdelliana sexualis (Casey)
Fig. 292a-h; Map, Fig. 294
Hyperaspis sexualis C?iSQy , 1924, p. 167. — Korschefsky, 1931, p. 196.
Scymnus sexualis: Dobzhansky, 1941, p. 86.
Blaisdelliana vanduzeei Gordon, 1970a, p. 43.
Blaisdelliana sexualis: Gordon, 1974c, p. 209.
Diagnosis. Length 1.25 to 1.78 mm, width 0.84 to 1.25 mm. Dorsal surface black
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to brown (Fig. 292h); antenna, mouthparts, femoral apices and remainder of leg
yellow; male with clypeus and frons yellow (in California specimens narrow lateral
border of pronotum yellow). Male genitalia as in Figure 292d-f. Female genitalia as
in Figure 292g.
Discussion. The specimens from California are a little larger than those from Utah,
and the male has a yellow pronotal margin. The basal lobes of the male genitalia are
noticeably different when males from St. George, Utah, were compared with a male
from Fresno, California, but a male from Yuma, Arizona, exhibits an intermediate
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NORTH AMERICAN COCCINELLIDAE
357
form. I consider all specimens examined as conspecific on the premise that additional
material from intermediate localities will provide intergrading specimens.
Type locality. St. George, Utah (lectotype and paralectotypes previously designated
by Gordon (1974c).
Type depository. USNM (35158).
Distribution. Figure 294. ARIZONA: Yuma. CALIFORNIA: Fresno Co.; Indio;
Kings Co.; Rosamond. UTAH: St. George.
Genus Hyperaspidius Crotch
H y per aspidius Crotch, 1873, p. 382.— Casey, 1899, p. 130. — Leng, 1920, p. 212.—
Casey, 1924, p. 168. — Korschefsky, 1931, p. 199.— Wingo, 1952, p. 26. — Hatch,
1961, p. 155.— Chapin, 1966, p. 280. — Belicek, 1976, p. 308. Type-species; Hy-
peraspis vittigera LeConte (not Chrysomela trimaculatus L., 1767, of authors), by
original designation.
Hyperaspini with body usually elongate, subparallel to parallel sided, dorsoven-
trally compressed; dorsal surface glabrous. Head usually yellow in male, brown or
black in female. Antenna 10-segmented (Fig. 295a); antennal insertion exposed. Eye
entire. Scutellum small, wider than long. Epipleuron of elytron narrow, not descending
externally, not grooved medially, not excavated for reception of middle and hind
femoral apices (Fig. 295b). Prosternum with 2 carinae convergent anteriorly. Posterior
margin of metasternum nearly on equal plane with abdomen between coxa and lateral
margin. Leg with femur and tibia slightly compressed; anterior tibia simple; tarsal
claw without basal tooth. Postcoxal line on first abdominal sternum nearly complete,
similar to Pullus type, or incomplete, of Scymnus type. Apical abdominal sternum
in male feebly to strongly emarginate. Male genitalia with basal lobe asymmetrical,
paramere rooted in phallobase, of 3 distinct types (Figs. 295c, 298a, 309a). Female
genitalia with compound spermatheca, basal portion with appendix (Fig. 296d), coxal
plate always transverse.
The lack of epipleural depressions and simple tarsal claws will separate Hyper-
aspidius from other hyperaspine genera. Hyperaspidius is a New World genus con-
taining 26 species, none of which are known to occur south of Mexico. Crotch (1873)
designated Chrysomela trimaculata L. as the type species, however, the specimens
Crotch identified as trimaculata were North American specimens described as Hy-
peraspis vittigera by LeConte (1852). Since Chrysomela trimaculata L. is supposedly
a tropical American species. Crotch apparently misidentified the vittigera of LeConte,
therefore I recognize H. vittigera LeConte as the type-species of Hyperaspidius because
LeConte’s type was among other specimens actually seen by Crotch when he made
his type species designation. The type of Chrysomela trimaculata (L.) is missing from
the Linnean collection in London, therefore the exact identity of that species may
never be determined.
Host records for Hyperaspidius species are almost nonexistent. I have seen 2 females
from Phoenix, Arizona, labeled “on cottony cochineal scale of cactus.” El-Ali (unpubl.
dissertation) stated that specimens of H. comparatus Casey were collected feeding
on distichlis mealybug, Distichlicoccus salinus (Cockerell)?, and that they were reared
in the laboratory on the solanum mealybug, Phenacoccus solani Ferris. One species
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has been recorded in the literature as feeding on Dactylopius confusus (Cockerell) in
Texas, and “//. vittigerus"" was recorded on Antonina graminis (Maskell). Some
species of Hyperaspidius are prevalent in grasslands, others are known only from
sand dune areas. As a group they are rather uncommonly collected, perhaps because
they usually occur close to the ground where normal net sweeping will not reach
them. Pit traps are an effective method of collecting as evidenced by several large
series of specimens examined in the course of this study. Hyperaspidius is one of the
North American genera that needs to be studied further. I am not satisfied with many
of the conclusions reached herein, and biosystematic research will be needed as well
as the collection of many more specimens from critical areas in order to accurately
reflect the actual taxonomic picture.
Hyperaspidius was last treated in its entirety by Casey (1899). Since then, various
authors have published individual descriptions, and regional papers by Wingo (1952),
Hatch (1961), and Belicek (1976) have each included a few species.
The genus can be divided into 3 groups based on the form of the median lobe of
the male genitalia. I designate these as the comparatus, arcuatus, and vittigerus groups.
Morphological distinctions are discussed under each group heading.
Key to species of Hyperaspidius
1. Elytron vittate, always with yellow discal vitta which may be incomplete, and a
yellow vitta on lateral margin (Figs. 37 Id, 318d) 14
Elytron not appearing vittate, discal vitta absent, lateral margin vittate or not . 2
2(1). Species occurring east of Mississippi River 3
Species occurring west of Mississippi River 9
3(2). Elytron black with 4 yellow spots (Fig. 313) venustulus (Mulsant)
- Elytron never with 4 yellow spots as figured above 4
4(3). Elytron entirely yellow, immaculate (Fig. 298e) transfugatus Casey
Elytron yellow with dark maculation, or dark with yellow lateral border 5
5(4). Elytron yellow with dark maculation, humerus usually with small, elongate brown
spot (Fig. 299e) militaris (LeConte)
Elytron dark brown or black, with or without yellow lateral border 6
6(5). Elytron with complete yellow lateral border (Fig. 31 2d) marginatus (Gaines)
Elytron entirely brown or black, or dark with incomplete yellow border in humeral
area 7
7(6). Species known from Massachusetts (Fig. 310e) blatchleyi, n. name
Species known from North Carolina to Florida 8
8(7). Male pronotum reddish yellow except basal 1/3 with obscure, brown maculation;
female unknown; Florida flavocephalus Blatchley
- Male pronotum mostly brown or black; female pronotum dark brown or black
except anterior angle pale; North Carolina, Georgia nubilatus Casey
9(2). Elytron entirely yellow, immaculate (Fig. 308d) nanellus, n. sp.
Elytron brown or black, usually maculate 10
10(9). Elytron dark, with basal and lateral borders yellow (Fig. 303d) . . arcuatus (LeConte)
- Elytron not as described above 11
11(10). Pronotum entirely pale, yellow with reddish yellow maculation; elytron with discal
spot in apical V2 (Fig. 3 1 5c) insignis Casey
Pronotum mostly brown or black; elytron without discal spot 12
12(1 1). Elytron brown or black with complete yellow vitta on lateral margin (Fig. 31 2d);
Texas marginatus (Gaines)
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NORTH AMERICAN COCCINELLIDAE
359
Elytron brown or black, if yellow lateral vitta present, then vitta interrupted in
apical V4 with apical spot present; not occurring in Texas 13
1 3( 1 2). Elytron brown or black with yellow lateral vitta interrupted in apical ‘A with apical
spot present (Fig. 300d) tristis (LeConte)
- Elytron entirely brown or black, sometimes with faint maculation apparent on
humeral angle and/or disc (Fig. 302d) ploribundus (Nunenmacher)
14(1). Species occurring east of the 100th meridian, north of Texas (Fig. 317d)
wolcotti (Nunenmacher)
Species occurring west of the 100th meridian, or if east of the 100th meridian,
then only in Texas 15
15(14). Prostemum coarsely punctured, anterior margin nearly always broadly, feebly
emarginate 16
Prostemum impunctate or with fine, indistinct punctures, anterior margin truncate
24
16(15). . Female with 6th abdominal sternum abruptly narrowed to rounded apex; male
with 6th abdominal sternum strongly narrowed toward apex, apex strongly emar-
ginate; body form extremely elongate, tapered toward apex (Fig. 325d); Coral Pink
Sand Dunes, Utah andrewsi, n. sp.
- Female with 6th abdominal sternum gently narrowed to broadly rounded apex;
male with 6th abdominal sternum feebly narrowed toward apex, apex weakly
emarginate; not known from Coral Pink Sand Dunes 17
17(16). Species occurring in Texas 18
- Species not occurring in Texas 19
1 8( 1 7). Body elongate, parallel sided; dorsal maculation light brown, indistinct (Fig. 32 1 d)
shauli Nunenmacher
- Body broad, sides not appearing strongly parallel sided; dorsal maculation dark
brown or black (Fig. 3 1 8d) oblongus Casey
19(17). Pronotum yellow to reddish yellow, often yellow with indistinct reddish yellow
maculation 20
Pronotum mostly yellow or mostly black, if mostly yellow, then with some dark
brown or black maculation 21
20(19). Surface of pronotum dull, alutaceous; Colorado and Alberta insignis Casey
- Surface of pronotum shiny, polished; Algodones Dunes, Imperial Co., California
(Fig. 3 1 6d) algodonus, n. sp.
21(19). Female postcoxal line complete {Pullus type); area within postcoxal line of both
sexes smooth, polished, punctures scattered; inland sand dunes, southern Cali-
fornia hardyi, n. sp.
- Female postcoxal line incomplete (Scymnus type); area within postcoxal line of
both sexes dull, alutaceous, punctation often dense, usually coarse; not occurring
in southern California 22
22(21). Species known only from southern New Mexico; postcoxal line in both sexes
equally incomplete ingenitus Casey
- Species not known from New Mexico; postcoxal line nearly complete in male,
incomplete in female 23
23(22). Length less than 2.10 mm; female first abdominal sternum with punctures very
fine, female postcoxal line widely incomplete
(Fig. 322e) vittigerus (LeConte)
- Length more than 2.10 mm; female first abdominal sternum with punctures coarse,
dense, female postcoxal line narrowly incomplete (Fig. 323d) Hercules Belicek
24(1 5). Head of male dark brown with irregular yellow area adjacent to eye; male prono-
tum dark brown on anterior margin tristis (LeConte)
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
- Head of male always yellow except vertex usually brown or black; male pronotum
yellow on anterior margin (except arcuatus) 25
25(24). Basal lobe of male genitalia slender, without lateral projection in basal '/s (Fig.
295c) 26
- Basal lobe of male genitalia broad, with lateral projection in basal 'A (Fig. 304a)
27
26(25). Basal lobe of male genitalia longer than paramere (Fig. 295c) .... comparatus Casey
Basal lobe of male genitalia shorter than paramere (Fig. 296a) mimus Casey
27(25). Basal lobe of male genitalia with lateral projection in basal 'A pronounced, abruptly
rounded (Fig. 304a) simulatus, n. sp.
- Basal lobe of male genitalia with lateral projection in apical ‘A feeble, slightly
angulate or feebly rounded 28
28(27). Basal lobe of male genitalia triangular in apical % (Fig. 306a) bryanti Nunenmacher
Basal lobe of male genitalia not triangular in apical % 29
29(28). Basal lobe of male genitalia with lateral projection in apical V3 feebly rounded
(Fig. 305a); Arizona pallescens Casey
Basal lobe of male genitalia with lateral projection in apical 'A slightly angulate
(Fig. 303a); not known from Arizona arcuatus (LeConte)
comparatus group
Prosternum impunctate; male genitalia with basal lobe as long as, or longer than
paramere, slender, lacking lateral projection in basal ‘A (Fig. 295c).
Hyperaspidius comparatus Casey
Fig. 295a-h; Map, Fig. 287
Hyperaspidius comparatus 1899, p. 130. — Leng, 1920, p. 212.— Korschefsky,
1931, p. 199.
Hyperaspidius juniperus Nunenmacher, 1944, p. 145. New Synonymy.
Diagnosis. Length 1.40 to 2.10 mm; width 1.0 to 1.50 mm. Form oblong, lateral
margin of elytron feebly curved. Pronotum of male yellow with indistinct yellowish
brown maculation in basal ', or dark brown with anterior and lateral margins narrowly
yellow; pronotum of female yellowish brown with lateral margin narrowly yellow.
Elytron with 2 broad, yellow vittae often connected at apex (Fig. 295f-h), northern
specimens with vittae narrow, widely disconnected at apex. Postcoxal line complete
(male) or narrowly incomplete (female), area within line alutaceous, punctation barely
perceptible. Male genitalia as in Figure 295c-d.
Discussion. The male genitalia are the only certain criteria I can find that allow H.
comparatus to be recognized. The color patterns present in this species are also found
in several other species such as H. mimus and H. pallescens. I have included spec-
imens from New Mexico here because the genitalia appear to be identical with those
of California specimens. Since no specimens of H. comparatus have been seen from
Arizona, it is possible that we are dealing with 2 species. The holotype of H. juniperus
Nunenmacher is a typical example of H. comparatus, therefore I regard H. juniperus
as a junior synonym of H. comparatus. The type of H. comparatus is a unique female
(holotype).
Type locality. Of comparatus, Alameda Co., California; of juniperus, Tehachapi
Pass, Kern Co., California.
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NORTH AMERICAN COCCINELLIDAE
361
Fig. 295. Hyperaspidius sp. a. Antenna, b. Epipleuron. c-h. Hypemspidius compamtus.
Type depository. Of comparatus, USNM (35215); of juniperus, CAS.
Distribution. Figure 287. BRITISH COLUMBIA: Radium. CALIFORNIA: Ala-
meda Co.; Contra Costa Co., Pt. Molete Beach; Fresno Co., Fresno; Inyo Co., Owens
Lake; Kem Co., Bakersfield, Tehachapi Pass; Kings Co.; Lassen Co., Spaulding; Los
Angeles Co., Lancaster, Pasadena; Orange Co., Cypress; Paraiso Hot Springs; Riv-
erside Co., Temecula; Santa Barbara Co., county record, San Miguel Island. NEW
MEXICO: Bernallilo Co., Albuquerque; San Miguel Co., Las Vegas; Quay Co., Tuc-
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93( 1 )
Fig. 296. Hyperaspidius mimus.
umcari. UTAH: Iron Co., Buckskin Valley; Tooele Co., Skull Valley. WASHING-
TON: Benton Co., Rattlesnake Ridge; Doris.
Hyperaspidius mimus Casey
Fig. 296a-g; Map, Fig. 297
Hyperaspidius mimus Casey, 1924, p. 169. — Korschefsky, 1931, p. 199.
Hyperaspidius carri Nunenmacher, 1948, p. 6. New Synonymy.
Hyperaspidius coloradensis Nunenmacher, 1948, p. 7. New Synonymy.
Diagnosis. Length 1.40 to 1.80 mm; width 1.0 to 1.20 mm. Description as for H.
comparatus (Fig. 296e-g) except female pronotum sometimes with narrow, yellow
anterior margin. Male genitalia with basal lobe not longer than paramere (Fig. 296a-
c). Female genitalia as in Figure 296d.
Discussion. This species is very similar to H. comparatus, but has the basal lobe
of the male genitalia no longer than the paramere. Based on the specimens examined,
it can be said that the distributions of H. comparatus and H. mimus do not overlap,
or only narrowly so; but collections from additional localities may eliminate this
distinction. Hyperaspidius carri Nunenmacher and H. coloradensis Nunenmacher
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NORTH AMERICAN COCCINELLIDAE
363
Fig. 297. Distribution. Hyperaspidius comparatus (dot); H. mimus (star); H. transfugatus
(square); H. militaris (open circle).
are apparently identical in all respects to H. mimus, therefore I regard them as junior
synonyms.
Type locality. Of mimus, Boulder Co., Colorado; of earn. Medicine Hat, Alberta;
of color adensis, Colorado,
Type depository. Of mimus, USNM (35217); of carri and coloradensis, CAS.
Distribution. Figure 297. ALBERTA; Cypress Hills, Medicine Hat. SASKATCH-
EWAN: Swift Current. COLORADO: Boulder Co.; Douglas Co., Sedalia; El Paso
Co., Colorado Springs; Huerfano Co., La Veta; Lake Co., Leadville; Larimer Co.,
Fort Collins; Teller Co., Florissant; Weld Co., Pawnee National Grassland. IDAHO:
Cassia Co., Burley; Twin Falls Co., Hansen, Murtaugh. MONTANA: Lewis and
Clark Co., Helena. NEBRASKA: Scotts Bluff Co., Scottsbluff. UTAH: Salt Lake Co.,
Salt Lake. WYOMING: Carbon Co., Medicine Bow.
arcuatus group
Prostemum impunctate or minutely punctured; male genitalia with basal lobe not
longer than paramere, with feeble lateral projection in basal 'ri (Fig. 298a).
Hyperaspidius transfugatus Casey
Fig. 298a-e; Map, Fig. 297
Hyperaspidius transfugatus Casey, 1 899, p. 1 3 1 . — Leng, 1 920, p. 2 1 2.— Korschefsky,
1931, p. 200.-Wingo, 1952, p. 26.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Hyperaspidius pallidus Casey, 1924, p. 169. — Korschefsky, 1931, p. 200. New Syn-
onymy.
Hypemspidius horni Nunenmacher, 1934a, p. 19. New Synonymy.
Diagnosis. Length 1.90 to 2.10 mm, width 1.30 to 1.45 mm. Form oblong, lateral
margin of elytron feebly curved. Pronotum entirely yellowish brown. Elytron yellow
except sutural margin narrowly darkened (Fig. 298e). Postcoxal line narrowly incom-
plete (male), or widely incomplete (female) (Fig. 29 8d), area within line alutaceous,
punctation barely perceptible. Male genitalia as in Figure 298a-c.
Discussion. The pale dorsal color and eastern distribution combined make this an
easily recognized species. Hyperaspidius pallidus Casey and H. horni Nunenmacher
are junior synonyms of H. transfugatus. The types of both H. transfugatus and H.
pallidus are unique females (holotypes). I have seen one “cotype” of H. horni labeled
“Buena NJ/Coll. by C. Liebeck/Hyperaspidius horni Nun. Type” which I here des-
ignate and label the lectotype.
Type locality. Of transfugatus, Mt. Tom, Massachusetts; of pallidus. Southern
Pines, North Carolina; of horni, Buena, New Jersey (lectotype here designated).
Type depository. Of transfugatus (35221) and pallidus (35223), USNM; of horni,
CAS.
Distribution. Figure 287. MASSACHUSETTS: Plymouth Co., Marion; Worcester
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NORTH AMERICAN COCCINELLIDAE
365
Fig. 299. Hyperaspis militaris.
Co., Berlin. MINNESOTA: Jackson Co. NEW JERSEY: Atlantic Co., Buena. NORTH
CAROLINA: Moore Co., Southern Pines.
Hyperaspidius militaris (LeConte)
Fig. 299a-e; Map, Fig. 297
Hyperaspis militaris LeConte, 1852, p. 133. — Crotch, 1874b, p. 231.
Hyperaspidius militaris: Crotch, 1873, p. 382. — Schwarz, 1878, p. 448.— Casey,
1899, p. I31.-Blatchley, 1917, p. I40.-Korschefsky, 1931, p. 199.
Diagnosis. Length 1.90 to 2.50 mm, width 1.50 to 1.80 mm. Form oblong, lateral
margin of elytron feebly curved. Pronotum of male yellow with obscure brownish
yellow maculation in basal pronotum of female black with lateral border narrowly
yellow. Elytron yellow except broad sutural border brown or black, short vitta present
on humerus (Fig. 299e). Postcoxal line widely incomplete in both sexes (Fig. 299d),
area within line alutaceous, punctation barely perceptible. Male genitalia as in Figure
299a-c.
Discussion. No known species of Hyperaspidius has the elytral pattern of H. mil-
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Fig. 300. Hyperaspidius tristis.
itaris. The only other species that may occur in the southeastern United States are
H. nubilata, H. transfugatus, H. marginatus and H. flavocephalus, all of which possess
color patterns different from that of militaris. The unique female holotype of H.
militaris is labeled “(orange disc)/4655/Type 6726/Hyperaspis militaris Lee.”.
Type locality. Columbia, South Carolina.
Type depository. MCZ.
Distribution. Figure 287. ALABAMA: Barbour Co., Spring Hill. FLORIDA: Duval
Co., Jacksonville; Gadsden Co., Mt. Pleasant; Lee Co., Fort Myers, Estero; Pinellas
Co., St. Petersburg; Putnam Co., Crescent City; Volusia Co., Enterprise. SOUTH
CAROLINA: Richland Co., Columbia.
Hyperaspidius tristis (LeConte), new combination
Fig. 300a-d; Map, Fig. 301
Hyperaspis tristis TtConXQ, 1880, p. 188.— Casey, 1899, p. 128.— Korschefsky, 1931,
p. 198. — Dobzhansky, 1941, p. 85.
Hyperaspidius conspiratus Casey, 1899, p. 131. — Korschefsky, 1931, p. 199. New
Synonymy.
Diagnosis. Length 1.60 to 2.0 mm, width 1.0 to 1.50 mm. Form elongate, oval,
lateral margin of elytron definitely curved. Head of male brown or yellowish brown
with obscure yellow spot near eye; head of female dark brown. Pronotum of both
sexes dark brown with lateral margin narrowly yellow or yellowish brown. Elytron
typically dark brown with narrowly yellow lateral margin and apical yellow spot (Fig.
300d), apical spot and yellow lateral margin often feebly connected, or elytron almost
entirely immaculate. Postcoxal line narrowly incomplete in both sexes, area within
line alutaceous, punctation barely visible. Male genitalia as in Figure 300a-c.
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NORTH AMERICAN COCCINELLIDAE
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Discussion. The typical form of H. tristis is easily distinguished by the unique
elytral color pattern. However, reduction of this pattern occurs until only a small
yellow area on the humeral angle remains. There is also a tendency for a discal yellow
vitta to form which I have seen culminated in a complete discal vitta in 3 specimens
from Alameda Co., California. The rounded elytral margins and the mostly dark
male head and pronotum will usually distinguish specimens of H. tristis that do not
have the typical color pattern. The other species with a dark male head is H. plori-
bundus which has the lateral margins of the pronotum and elytron obviously dis-
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continuous, and an oblong body form. There are 2 examples under H. tristis in the
LeConte collection as stated by LeConte. The first of these, a female labeled “Cal./
Hardy/494/Type 6722(red paper)/H. tristis Lee.'" is here designated and labeled the
lectotype. The second type specimen is a male of Hyperaspis oculaticauda. There are
4 types of H. conspiratus in the Casey collection. The first specimen, a male, is here
designated and labeled the lectotype, the other 3 as paralectotypes. LeConte (1852)
listed the type locality of tristis as “Col.” (Colorado), but the specimen in his collection
is clearly labeled “Cal.” (California), and I regard the published type locality to be
erroneous.
Type locality. Of tristis, California (lectotype here designated); of conspiratus, Par-
aiso Hot Springs, Monterey Co., California (lectotype here designated).
Type depository, of tristis, MCZ; of conspiratus, USNM (35222).
Distribution. Figure 301. CALIFORNIA: Alameda Co.; Inyo Co., Saline Valley,
Saratoga Spring, Death Valley; Kern Co., Tehachapi Pass, Fort Tejon; Kings Co.;
Los Angeles Co., Altadena; Glendora; Modoc Co., Min Pass; Monterey Co., Paraiso
Hot Springs; Riverside Co., Riverside; San Timoteo Co.; San Bernardino Co., Cactus
Flat; Cajon Pass, Rte 395; Desert Springs; San Diego Co., Morena Lake; San Francisco
Co.; Tulare Co., Kaweah; Sequoia National Park.
Hyperaspidius ploribundus (Nunenmacher)
Fig. 302a-d; Map, Fig. 301
Hyperaspis ploribunda Nunenmacher, 1911, p. 74.
Hyperaspidius ploribunda: Leng, 1920, p. 212. — Korschefsky, 1931, p. 200.— Dob-
zhansky, 1941, p. 86. — Nunenmacher, 1944, p. 144.
Hyperaspidius immaculatus Hatch, 1961, p. 155. New Synonymy.
Hyperaspidius arcuatus: Hatch, 1961, p. 155. — Belicek, 1976, p. 309 (not arcuatus
LeConte).
Diagnosis. Length 1.50 to 1.80 mm, width 1.10 to 1.40 mm. Form oblong, outline
of pronotum and elytron abruptly discontinuous, lateral margin of elytron feebly
curved (Fig. 302d). Head and pronotum in both sexes typically dark brown, antero-
lateral angle of pronotum often yellowish brown. Elytron reddish brown, often with
humeral angle yellowish brown. Postcoxal line usually nearly complete in both sexes,
area within line alutaceous, punctation barely perceptible. Male genitalia as in Figure
302a-c.
Discussion. See comparative remarks under H. tristis. The few specimens examined
show a rather wide geographic range with large gaps present. It is possible that more
than one species is involved, but, based on the available evidence, I can identify only
one species. I consider H. immaculata Hatch a junior synonym of ploribundus. There
are 2 type specimens of H. ploribundus. One of these, a male labeled “Goldfield/
Esmeralda Co., Nev. VI-29-07/CoU’d by F. W. Nunenmacher/Type. Hyperaspis
ploribunda Nun.” is here designated and labeled the lectotype, the other specimen
is labeled a paralectotype.
Type locality. Of ploribundus. Goldfield, Esmeralda Co., Nevada (lectotype here
designated); of immaculatus, Redmond, Oregon.
Type depository. Of ploribundus and immaculatus, CAS.
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NORTH AMERICAN COCCINELLIDAE
369
Fig. 302. Hyperaspidius ploribundus.
Distribution. Figure 301. CALIFORNIA: Fresno Co.; Inyo Co., Independence;
Kem Co., Fort Tejon. IDAHO: Kootenai Co., Coeur d’Alene. NEVADA: Esmeralda
Co., Goldfield. OREGON: Deschutes Co., Redmond.
Hyperaspidius arcuatus (LeConte)
Fig. 303a-e; Map, Fig. 301
Hyperaspis arcuata LeConte, 1852, p. 133. — Crotch, 1874b, p. 232.
Hyperaspidius arcuata: Crotch, 1873, p. 382.— Casey, 1899, p. 131.— Leng, 1920,
p. 212.— Korschefsky, 1931, p. 199.
Hyperaspidius arcuatus: Belicek, 1976, p. 309.
Hyperaspidius rossi Nunenmacher, 1944, p. 145.— Hatch, 1961, p. 155. New Syn-
onymy.
Diagnosis. Length 1.60 to 2.0 mm, width 1.20 to 1.50 mm. Form oblong, outline
of pronotum and elytron abruptly discontinuous, lateral margin of elytron feebly
curved. Pronotum of male black with yellow lateral margin, or with anterior and
lateral margins yellow. Female not known. Elytron black with basal and lateral
margins yellow, yellow lateral margin reaching midpoint (Fig. 303d), or complete to
apex, apex often with yellow spot, often with discal vitta reaching apex (Fig. 303e).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93( 1 )
Postcoxal line complete (male), area within line feebly alutaceous, distinctly punc-
tured. Male genitalia as in Figure 303a-c.
Discussion. Females were not available for examination, and all remarks refer to
males. The forms with incomplete elytral maculation are outstanding in appearance
and readily recognized, but the form with complete discal vittae resembles several
other species of Hyperaspidius\ and male genitalia must be examined in these in-
stances. The female type of wssi appears to be an example of H. arcuatus and I
consider wssi to be a junior synonym of arcuatus. The unique male holotype of H.
arcuatus is labeled “(gold disc)/4657/Type 6727 (red paper)/H. arcuata Lee.”.
Type locality. Of arcuatus, “mouth of Gila River, California”; of rossi, Oregon.
Type depository. Of arcuatus, MCZ; of rossi, CAS.
Distribution. Figure 301. CALIFORNIA: Imperial Co., Algodones Dunes; Bard;
Gila River; Glamis; Olgiby; Riverside Co., Blythe; San Bernardino Co., 5 mi. N.
Buckmans Sp., sand dunes 10 mi. NW Kelso; San Diego Co., Borrego. NEVADA:
Washoe Co., Glendale. UTAH: Washington Co., St. George.
Hyperaspidius simuiatus, new species
Fig. 304a-d; Map, Fig. 307
Description. Male, length 1.65 mm, width 1.10 mm. Form somewhat oblong,
outline of pronotum and elytron slightly discontinuous, lateral margin of elytron
distinctly rounded. Head yellow except vertex brown; pronotum brown with narrowly
yellow lateral margin and broadly yellow anterior margin; elytron brown with broad
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NORTH AMERICAN COCCINELLIDAE
371
lateral and discal vittae connected apically and posteriorly (Fig. 304d). Punctures on
head fine, separated by a diameter or less; pronotal punctures fine, separated by one
or 2 times a diameter; punctures on elytron slightly coarser than on pronotum,
separated by 2 or 3 times a diameter. Metastemum with coarse, confluent punctures
laterally, punctures very fine, sparse medially. Abdominal sterna with fine, dense
punctures. Postcoxal line complete, area within line alutaceous, nearly impunctate.
Male genitalia as in Figure 304a-c.
Variation. Length 1.60 to 1.70 mm. Apical border of brown area on pronotum
may have a median, v-shaped indentation.
Holotype. Male. CALIFORNIA: Etiwanda, San Bernardino Co., VII-27-1972, Col-
lector E. L. Paddock USNM(101333).
Paratypes. Total 6 (Fig. 307). CALIFORNIA: same data as holotype. (USNM)
(CDA).
There are no external characteristics that will, with certainty, distinguish this species
from several other vittate species of Hyperaspidius. The character that must be seen
is the large, rounded lateral projection on the basal lobe of the male genitalia which
is unlike that of any other species examined. I have restricted the specimens desig-
nated as type material to the type locality, however, there are other specimens I
regard as this species from the following localities. Arizona: Coconino Co., Page;
Cochise Co; 7 mi. S. Picacho, Pinal Co.; Rillito River near Tucson. California: Kings
Co.; Riverside; Tulare Co. The specific name is from the Latin similis, referring to
the similarity in dorsal color pattern to several other species of Hyperaspidius.
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Hyperaspidius pallescens Casey
Fig. 305a-d; Map, Fig. 307
Hyperaspidius pallescens Casey, 1908, p. 420. — Leng, 1920, p. 212.— Korschefsky,
1931, p. 199.
Diagnosis. Length 1.60 to 2.10 mm, width 1.10 to 1.60 mm. Form oblong, lateral
margin of elytron distinctly curved. Pronotum of male yellow with indistinct yel-
lowish brown maculation; pronotum of female uniformly yellowish brown except
lateral margin narrowly yellow. Elytron with 2 broad, yellow vittae not connected,
or narrowly connected at apex (Fig. 30 5d). Postcoxal line complete in both sexes,
area within line alutaceous, punctation barely perceptible. Male genitalia as in Figure
305a-c.
Discussion. Thus far H. pallescens is known only from Arizona. The male and
female pronotal color pattern will distinguish this species from other presently known
Arizona species, and the feebly curved lateral projection on the basal lobe of the
male genitalia is diagnostic. The type of pallescens is a unique male (holotype).
Type locality. Nogales, Santa Cruz Co., Arizona.
Type depository. USNM (35220)
Distribution. Figure 297. ARIZONA: Cochise Co., Chiricahua Mountains; Hu-
achucha Mountains, Miller Canyon; Pinal Co., Oracle; Santa Cruz Co., Nogales;
Santa Rita Mountains, Madera Canyon.
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NORTH AMERICAN COCCINELLIDAE
373
Fig. 306. Hyperaspidius bryanti.
Hyperaspidius bryanti Nunenmacher
Fig. 306a-d; Map, Fig. 307
Hyperaspidius bryanti Nunenmacher, 1948, p. 7.
Diagnosis. Length 1.80 to 2.0 mm, width 1.10 to 1.45 mm. Form oblong, lateral
margin of elytron feebly curved. Pronotum of male yellow with basal 2/3 dark brown
except lateral margin yellow; female pronotum dark reddish brown except lateral
margin narrowly yellow. Elytron with lateral and discal vittae connected across base,
extending to posterior %, broadly separated from apical spot, or with discal vitta
reduced to discal spot (Fig. 306d). Postcoxal line narrowly incomplete in both sexes,
area within line alutaceous, distinctly punctured. Male genitalia as in Figure 306a-
c.
Discussion. The form with the discal vitta on the elytron reduced to a spot is very
distinctive, unlike any other species known from Arizona. The typical form is less
striking, but the widely separated apical spot is unusual. The male genitalia are quite
different from those of other species of Hyperaspidius in the triangular form of the
basal lobe.
Type locality. Santa Catalina Mts., Arizona.
Type depository. CAS.
Distribution. Figure 307. ARIZONA: Cochise Co., Dragoon. Coconino Co., Se-
dona. Pima Co., Tucson. Santa Cruz Co., Sonoita.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 307. Distribution. Hypemspidius simulatus (open circle); H. pallescens (dot); H. bryanti
(star); H. nanella (square); H. flavocephalus (circled star); H. blatchleyi (open star).
Hyperaspidius nanellus, new species
Fig. 308a-d; Map, Fig. 307
Description. Male, length 2.0 mm, width 1.15 mm. Form oblong, outline of prono-
tum and elytron slightly discontinuous, lateral margin of elytron feebly rounded.
Head yellow except vertex yellowish brown; pronotum yellow with obscure brownish
yellow maculation in basal 1/3; elytron entirely yellow (fig. 308d). Punctures on head
fine, separated by a diameter or less; pronotal punctures slightly coarser than on
head, separated by a diameter or less; punctures on elytron coarser than on pronotum,
separated by less than to twice a diameter. Metasternum with fine, dense punctures
laterally, nearly impunctate medially. Abdominal sterna with fine, dense punctures.
Postcoxal line complete (male) or widely incomplete (female), area within line alu-
taceous, nearly impunctate. Male genitalia as in Figure 308a-c.
Holotype. Male. TEXAS: Brownsville, VII, Wickham, Hyperaspis? cinctus?, Wick-
ham Collection USNM (101334).
Allotype. Female. TEXAS: Prairie 10 mi. NE Brownsville, 25.5.04, HS Barber
Collector. (USNM).
Paratype. Total 1 (Fig. 307). TEXAS: Burleson Co., 4/1 1/34, J.C. Gaines Collector
(WHN).
The entirely pale elytron of this species distinguishes it from all other species except
H. transfugatus which has no maculation on the pronotum and is known only from
east of the Mississippi River and Minnesota. The specific name is from the Latin
nanus, referring to the small size and generally insignificant appearance.
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NORTH AMERICAN COCCINELLIDAE
375
Fig. 308. Hyperaspidius nanella.
vittigerus group
Prosternum coarsely, often densely punctured; male genitalia with basal lobe not
longer than paramere, with strong, abrupt lateral angulation in basal Vs (Fig. 309a).
Hyperaspidius flavocephalus Blatchley
Fig. 309a-d; Map, Fig. 307
Hyperaspidius flavocephalus B\aich\ey, 1924, p. 167. — Korschefsky, 1931, p. 199.
Diagnosis. Length 2.0 mm, width 1.60 mm. Form oblong, lateral margin of elytron
nearly straight. Pronotum of male reddish yellow with irregular, obscure brown
maculation in basal Vs. Elytron black except humeral angle narrowly yellow on margin
from base to just beyond callus (fig. 309d). Postcoxal line narrowly incomplete, area
within line alutaceous, coarsely punctured. Male genitalia as in Figure 309a-c.
Discussion. The male holotype is the only specimen of this species examined. I
regard H. flavocephalus as a valid species, but it is very similar in appearance to H.
nubilatus and H. marginatus; the male genitalia of each species are apparently dis-
tinctive, and the dorsal color patterns are also different for each species. See remarks
under H. blatchleyi, n. name. The holotype of H. flavocephalus is labeled “Dunedin,
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93( 1 )
Fig. 309. Hyperaspidius flavocephalus.
Fla W.S.B. Coll. 3-27-18/822/Purdue Blatchley collection/Type(red paper)/Hyper-
aspidius flavocephalus Blatch.”.
Type locality. Dunedin, Florida.
Type depository. PU.
Distribution. Figure 307. FLORIDA: Pinellas Co., Dunedin.
Hyperaspidius blatchleyi, new name
Fig. 310a-e; Map, Fig. 307
Hyperaspidius flavocephalus Marshall, 1945, p. 177 (not flavocephalus Blatchley,
1924).
Diagnosis. Length 1.90 to 2.40 mm, width 1.30 to 1.60 mm. Form rounded, oval,
lateral margin of elytron definitely curved. Head and pronotum yellow, pronotum
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NORTH AMERICAN COCCINELLIDAE
377
Fig. 310. Hyperaspidius blatchleyi.
with median black area at base projecting forward at middle. Elytron black with
short, yellow vitta on lateral margin from humeral angle to middle (Fig. 3 1 Oe). Surface
of head alutaceous, finely punctured, punctures separated by 2 or 3 times a diameter;
surface of pronotum strongly alutaceous, punctures larger than on head, separated
by 2 or 3 times a diameter; surface of elytron smooth, punctures coarse, larger than
on pronotum, separated by 2 to 4 times a diameter. Metastemum coarsely, densely
punctured laterally, nearly impunctate medially. Abdominal sterna densely, coarsely
punctured; postcoxal line narrowly incomplete, area within line alutaceous, coarsely
punctured (Fig. 310d). Male genitalia as in Figure 310a-c.
Type locality. Berlin, Massachusetts.
Type depository. Location of allotype unknown.
Distribution. Figure 307. MASSACHUSETTS: Berlin; Natick; Wayland.
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Fig. 311. Hyperaspidius nubilatus.
Discussion. This species has been identified as H. flavocephalus Blatchley by pre-
vious authors. However, the male pronotal color patterns are quite different; the
body form of H. blatchleyi is rounded, that of H. flavocephalus is oblong. In addition,
the male genitalia are distinctive for each species. I name this species for W. S.
Blatchley. Marshall (1945) discussed this species under the name H. flavocephalus
Blatchley, and described an allotype and parallotypes.
Hyperaspidius nubilatus (Casey), new combination
Figs. 311a-e; Map, Fig. 314
Hyperaspis nubilata Casey, 1924, p. 166. — Korschefsky, 1931, p. 193.
Hyperaspis asphaltina Casey, 1924, p. 166. — Korschefsky, 1931, p. 184.— Dob-
zhansky, 1941, p. 83. New Combination.
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NORTH AMERICAN COCCINELLIDAE
379
Fig. 312. Hyperaspidius marginatus.
Diagnosis. Length 1.80 to 2.70 mm, width 1.30 to 2.0 mm. Form oblong, lateral
margin of elytron feebly curved. Pronotum of male black or dark brown with nebulous
black areas in basal ‘, always with lateral margin narrowly yellow; female pronotum
entirely black except anterior angle obscurely yellowish brown. Elytron black except
humeral angle obscurely yellowish brown, or with yellow vitta extending from hu-
meral angle to middle (Fig. 3 1 Id, e), occasional specimens with nebulus brown areas.
Postcoxal line narrowly incomplete (male) or widely incomplete (female), area within
line alutaceous, distinctly punctured. Male genitalia as in Figure 3 1 1 a-c.
Discussion. This species most closely resembles H. flavocephalus (see comments
under that species) and H. marginatus. In H. marginatus the lateral margin of the
elytron is yellow from the humeral angle to the apex. In H. nubilatus the margin is
either not yellow or yellow from the humeral angle to the midpoint. The type of H.
nubilatus is a unique female (holotype). There are 8 types of H. asphaltina, and I
here designate and label a male as the lectotype and the remainder as paralectotypes.
Type locality. Of nubilatus and asphaltina (lectotype here designated). Southern
Pines, North Carolina.
Type depository. Of nubilatus (35156) and asphaltina (35157), USNM.
Distribution. Figure 314. FLORIDA: Duval Co., Jacksonville. GEORGIA: Grady
Co., Beachton, Hutchison Place; Dodge Co., Chester. NORTH CAROLINA: Moore
Co., Southern Pines.
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Fig. 313. Hyperasidius venustulus.
Hyperaspidius marginal us (Gaines), new combination
Fig. 312a-d; Map, Fig. 314
Hyperaspis fimbriolata marginatus Gaines, 1933, p. 263.
Hyperaspis marginata: Dobzhansky, 1941, p. 58.
Diagnosis. Length 1.75 to 2.40 mm, width, 1.30 to 1.75 mm. Form oblong, lateral
margin of elytron feebly curved. Head yellow or brownish yellow in both sexes.
Pronotum reddish brown with nebulous brown maculation in male and some females,
other females with pronotum dark brown except lateral border broadly yellow. Ely-
tron dark brown or black with broad, yellow vitta on lateral margin from humeral
angle nearly to apex (Fig. 3 1 2d). Postcoxal line in both sexes widely incomplete, area
within line alutaceous, indistinctly punctured. Male genitalia as in Figure 3 1 2a-c.
Discussion. The complete yellow vitta on the lateral border of the elytron will
separate marginatus from any other species of Hyperaspidius. The resemblance of
//. marginatus to Hyperaspis fimbriolata is remarkable, and that resemblance is the
reason H. marginatus was described as a subspecies of fimbriolata in Hyperaspis.
Dobzhansky (1941) raised it to species level.
Type locality. College Station, Texas.
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NORTH AMERICAN COCCINELLIDAE
381
Fig. 314. Distribution. Hyperaspidius nubilatus (dot); H. marginatus (star); H. venustulus
(open circle); H. insignis (square); H. algodonus (circled star); H. wolcotti (open star).
Type depository. USNM (53746).
Distribution. Figure 314. TEXAS: Brazos Co., College Station; Colorado Co., Co-
lumbus; Refugio Co., Refugio; Robertson Co., Calvert; San Patricio Co., Sinton,
Welder Wildlife Refuge; Victoria Co., Victoria.
Hyperaspidius venustulus (Mulsant)
Fig. 313; Map, Fig. 314
Hyperaspis venustula Mulsant, 1850, p. 671.— Crotch, 1873, p. 381 (as a synonym
of Hyperaspis lugubris).— Crotch, 1874b, p. 235.— Weise, 1895a, p. 129.— Kor-
schefsky, 1931, p. 192. — Dobzhansky, 1941, p. 21.
Hyperaspidius venustulus: Gordon, 1974c, p. 210.
Diagnosis. Length 2.80 mm, width 1.85 mm. Form elongate, lateral margin of
elytron rounded in apical Pronotum yellowish red. Elytron black with 4 yellow
spots (Fig. 313). Postcoxal line reaching hind margin of first abdominal sternum,
narrowly incomplete, area within line alutaceous, nearly impunctate.
Discussion. I have seen only two specimens of this species, one of which is the
female lectotype, and another female from Georgia. The large size, elongate body,
and dorsal color pattern make H. venustulus an outstanding species that is unlike
any other North American species of Hyperaspini.
Type locality. “Amer. bor., LeConte” (lectotype designated by Gordon, 1974c).
Type depository. Dejean Collection (DLM).
Distribution. Figure 314. GEORGIA: Myrtle.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 315. Hyperaspidius insignis.
Hyperaspidius insignis Casey
Fig. 315a-f; Map, Fig. 314
Hyperaspidius insignis Casey, 1899, p. 131. — Leng, 1920, p. 212.— Korschefsky,
1931, p. 199.
Diagnosis. Length 2.25 to 3.20 mm, width 1.60 to 2.0 mm. Form oblong, lateral
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NORTH AMERICAN COCCINELLIDAE
383
Fig. 316. Hyperaspidius algodonus.
margin of elytron feebly curved. Head pale in both sexes. Pronotum in both sexes
yellow with reddish yellow maculation, often reddish yellow with reddish brown
maculation. Elytron black with complete yellow border on anterior and lateral mar-
gins, discal spot on apical ‘ sometimes connected to anterior border (Figs. 3 1 5e, 1).
Postcoxal line narrowly incomplete in both sexes (Fig. 3 1 5d), area within line alu-
taceous, distinctly punctured. Male genitalia as in Figure 3 1 5a-c.
Discussion. The pale pronotum and head in both sexes along with the usually large,
robust body form distinguish H. insignis from other members of the vittigerus group.
No known species occurring in the same geographic area is similar in appearance.
There are 2 type specimens of H. insignis in the Casey collection, I here designate
and label a male as the lectotype, and the other, a female, as a paralectotype.
Type locality. Colorado Springs, Colorado (lectotype here designated).
Type depository. USNM (35219).
Distribution. Figure 314. ALBERTA: Medicine Hat. COLORADO: Chaffee Co.,
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Fig. 317. Hyperaspidius wolcotti.
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NORTH AMERICAN COCCINELLIDAE
385
Buena Vista; El Paso Co., Colorado Springs. OKLAHOMA: Woodward Co., Wood-
ward.
Hyperaspidius aI*^odonus, new species
Fig. 316a-d; Map, Fig. 314
Description. Male, length 2.0 mm, width 1.40 mm. Form oblong, lateral margin
of elytron straight. Head and pronotum yellow, pronotum with faint, nebulous mac-
ulation in basal ‘. Elytron black with complete yellow vitta on anterior and lateral
borders connected to incomplete discal vitta (Fig. 316d). Punctures on head and
pronotum extremely fine, barely perceptible, surface of pronotum smooth, polished;
punctures on elytron fine, distinct, separated by one to 3 times a diameter. Meta-
stemum coarsely, densely punctured laterally, nearly impunctate medially. Abdom-
inal sterna with fine punctures separated by 2 or 3 times a diameter. Postcoxal line
narrowly incomplete, area within line alutaceous, nearly impunctate. Male genitalia
as in Figure 3 1 6a-c.
Holotype. Male. CALIFORNIA: Imperial Co., Algodones Dunes, 7 mi SE Glamis,
25°55'20"N, 114°59'14"W, Site 4, III-25-79 to IV-8-1979 (USNM 101335).
The holotype is the only example of this species I have seen. The dorsal color
pattern is like that of H. insignis, but the pronotal surface is smooth and polished
in H. algodonus, dull and alutaceous in H. insignis. The specific name refers to the
type locality.
Hyperaspidius wolcotti (Nunenmacher)
Fig. 317a-e; Map, Fig. 314
Hyperaspis wolcotti Nunenmacher, 1911, p. 73.
Hyperaspidius wolcotti: Leng, 1920, p. 212. — Korschefsky, 1931, p. 200.— Dob-
zhansky, 1941, p. 86.— Nunenmacher, 1944, p. 144. — Wingo, 1952, p. 26.
Diagnosis. Length 2.0 to 2.60 mm, width 1.40 to 1.80 mm. Form oblong, convex,
somewhat rounded, lateral margin of elytron curved. Pronotum of male yellow with
black maculation in basal ‘; female pronotum black, lateral margin broadly yellow.
Elytron black with complete yellow vitta on lateral margin, irregular, incomplete
discal vitta present (Fig. 317d, e). Postcoxal line narrowly incomplete in both sexes,
area within line alutaceous, densely, coarsely punctured. Male genitalia as in Figure
317a-c.
Discussion. The rounded, convex body form and dorsal color pattern will separate
H. wolcotti from any species presently known from the same geographic area. The
species most similar to H. wolcotti is H. Hercules, but the distribution patterns are
widely disjunct.
Type locality. Pine Barrens, Buffington, Indiana.
Type depository. CAS.
Distribution. Figure 314. INDIANA: Lake Co., Buffington, Pine Barrens; Hessville.
IOWA: Emmett Co., Estherville. KANSAS: Riley Co.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 318. Hyperaspidius oblongus.
Hyperaspidius oblongus Casey
Fig. 318a-d; Map, Fig. 320
Hyperaspidius oblongus Casey, 1908, p. 421.— Leng, 1920, p. 212.— Korschefsky,
1931, p. 199.
Hyperaspidius trimaculatus: Casey, 1899, p. 130 (not trimaculatus L., 1767).
Diagnosis. Length 1.70 to 2.30 mm, width 1.20 to 1.65 mm. Form oblong, some-
what convex, lateral margin of elytron straight. Pronotum of male yellow with ne-
bulous brown maculation in basal female pronotum black with lateral margin
narrowly yellow. Elytron black with complete yellow vitta on anterior and lateral
borders connected to incomplete discal vitta (Fig. 318d). Postcoxal line narrowly
incomplete in both sexes, area within line alutaceous, nearly impunctate. Male gen-
italia as in Figure 3 1 8a-c.
Discussion. This species, H. shauli, and H. ingenitus are very similar in appearance
with H. oblongus and H. ingenitus being extremely similar. The pronotal punctures
of H. ingenitus are definitely larger than the elytral punctures, the other two species
have the elytral punctures larger than the pronotal punctures. The body form of H.
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NORTH AMERICAN COCCINELLIDAE
387
Fig. 319. Hyperaspidius ingenitus.
shauli is distinctive because it is strongly dorsoventrally flattened and the elytra are
parallel sided, while H. oblongus is somewhat convex in lateral view and does not
appear extremely parallel sided. It is possible that H. oblongus and H. ingenitus are
conspecific; but there are differences in the male genitalia that I consider significant,
therefore I regard each as a valid species. There are 4 type specimens of H. oblongus,
I here designate and label a male as the lectotype, the remainder as paralectotypes.
Type locality. El Paso, Texas (lectotype here designated).
Type depository. USNM (35214).
Distribution. Figure 320. TEXAS: Colorado Co., Columbus; Duval Co., San Diego;
El Paso Co., El Paso.
Hyperaspidius ingenitus Casey
Fig. 319a-d; Map, Fig. 320
Hyperaspidius ingenitus Casey, 1899, p. 131. — Leng, 1920, p. 212.— Korschefsky,
1931, p. 199.
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Fig. 320. Distribution. Hyperaspdius oblongus (star); H. ingenitus (open circle); H. shauli
(square); H. vittigerus (dot).
Diagnosis. Length 1.75 to 2.50 mm, width 1.10 to 1.60 mm. Form oblong, lateral
margin of elytron feebly curved (Fig. 3 1 9d). Description as for H. oblongus except
area within postcoxal line coarsely, sparely punctured; male genitalia as in Figure
3 1 9a-c.
Discussion. For comparison of H. ingenitus to similar appearing species, see com-
ments under H. oblongus. The type of H. ingenitus is a unique male (holotype).
Type locality. Las Cruces, New Mexico.
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NORTH AMERICAN COCCINELLIDAE
389
Fig. 321. Hyperaspidius shauli.
Type depository. USNM (35218).
Distribution. Figure 320. NEW MEXICO: Dona Ana Co., Las Cruces.
Hyperaspidius shauli Nunenmacher
Fig. 321a-d; Map, Fig. 320
Hyperaspidius shauli ^xxnQnmdiQhQv, 1944, p. 145.
Diagnosis. Length 1.60 to 2.0 mm, width 0.90 to 1.10 mm. Form oblong, elongate,
extremely parallel sided, strongly dorsoventrally flattened. Pronotum of male yellow
with nebulous brown maculation in basal female pronotum mostly yellow with
median brown area. Elytron brown with complete yellow vitta on lateral and anterior
margins connected to complete or incomplete broad discal vitta (Fig. 32 1 d). Postcoxal
line complete in both sexes, area within line smooth, polished, with indistinct, coarse
punctures. Male genitalia as in Figure 321a-c.
Discussion. Some of the differences between H. shauli and similar species are
discussed under H. oblongus. In addition, the complete postcoxal line with area inside
of the line smooth are characters not shared with H. oblongus or H. ingenitus.
Type locality. Perryton, Texas.
Type depository. CAS.
Distribution. Figure 320. TEXAS: type locality.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
/ \ // C \
/ \ y w / \ X
V /
Fig. 322. Hyperaspidius vittigerus.
Hyperaspidius vittigerus (LeConte)
Fig. 322a-g; Map, Fig. 320
Hyperaspis vittigera LeConte, 1852, p. 133.— Crotch, 1874b, p. 231.
Hyperaspidius trimaculata: Crotch, 1873, p. 382 (not trimaculata L., 1767).
Hyperaspidius vittigera: Leng, 1920, p. 212. — Korschefsky, 1931, p. 200.
Hyperaspidius vittigerus: Wingo, 1952, p. 26. — Belicek, 1976, p. 309.
Diagnosis. Length 1.50 to 2.05 mm, width 1.25 to 1.50 mm. Form oblong, lateral
margin of elytron feebly curved. Pronotum of male yellow with black maculation in
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NORTH AMERICAN COCCINELLIDAE
391
basal female pronotum dark brown or black, lateral margin narrowly yellow. Elytron
black with complete yellow vitta on anterior and lateral borders, discal vitta incom-
plete or connected to lateral vitta at apex (Fig. 322g). Postcoxal line nearly complete
(male) or widely incomplete (female) (Fig. 322e, f), area within line alutaceous, finely,
densely punctured. Male genitalia as in Figure 322a-c. Female genitalia as in Figure
322d.
Discussion. The key characters and the differences in male genitalia are the only
characteristics I can find to distinguish H. vittigerus from H. Hercules (see comments
under H. Hercules). Superficially H. vittigerus resembles H. mimus even more than
it does H. Hercules, but H. vittigerus and H. mimus are in different groups within the
genus. LeConte (1852) had more than one type specimen, but only a single male in
his collection can now be identified as a type with certainty. I here designate and
label this specimen labeled “(green disc)/46 5 6/Type 6725 (red paper)/H. vittigera
LeC.” as the lectotype.
Type locality. “Missouri Territory” (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 310. ALBERTA: Edmonton; Medicine Hat. COLORADO: El
Paso Co., Colorado Springs; Larimer Co., Fort Collins; Weld Co., Pawnee National
Grassland; Yuma Co., Wray. IDAHO: St. Anthony sand dunes. MONTANA: Pe-
troleum Co., Winnett; Valley Co., Glasgow. NEW MEXICO: Roswell. NORTH
DAKOTA: Grant Co., Lake Tschida. SOUTH DAKOTA: Hutchinson Co., Tripp.
WYOMING: Teton Co., Grand Teton Park.
Hyperaspidius Hercules Belicek
Fig. 323a-f; Map, Fig. 324
Hyperaspidius Hercules Belicek, 1976, p. 308.
Diagnosis. Length 2.10 to 4.0 mm, width 1.50 to 2.20 mm. Form oblong, lateral
margin of elytron feebly curved. Pronotum of male yellow with black maculation in
basal ‘; female pronotum black, lateral margin narrowly, obscurely yellow. Elytron
black with complete yellow vitta on anterior and lateral borders, discal vitta incom-
plete or narrowly connected to lateral vitta at apex (Fig. 323e, !)• Postcoxal line
complete in male, narrowly incomplete in female (Fig. 323d), area within line alu-
taceous, densely punctured. Male genitalia as in Figure 323a-c.
Discussion. Most specimens of this species are large (more than 3.5 mm long), but
a few, usually males, are smaller and these are difficult to distinguish from vittigerus
without examining the male genitalia. The specimens of typical size are outstanding
on that characteristic alone.
Type locality. Medicine Hat, Alberta.
Type depository. CNC.
Distribution. Figure 324. ALBERTA: Medicine Hat. CALIFORNIA: Eureka Val-
ley. COLORADO: Denver Co., Denver. IDAHO: Cassia Co., Burley; Jefferson Co.,
Terreton; Twin Falls Co., Buhl, Twin Falls. MONTANA: Winnett. NEVADA:
Churchill Co., Sand Mountain; Humboldt Co. UTAH: Emery Co., 22 mi. n. Hanks-
ville; Tooele Co., Dugway Proving Ground. WYOMING: Goshen Co., Hell Gap
Camp; Laramie Co., Cheyenne; Teton Co., Grand Teton Park.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 323. Hyperaspidius Hercules.
Hyperaspidius andrewsi, new species
Fig. 325a-e; Map, Fig. 324
Description. Male, length 2.10 mm, width 1.40 mm. Form elongate, slender, nar-
rowed posteriorly, lateral margin of elytron feebly curved. Head and pronotum yel-
low, pronotum with base narrowly black medially, 4 maculae present in basal *.
Elytron yellow with sutural margin narrowly black, narrow, black vitta present me-
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NORTH AMERICAN COCCINELLIDAE
393
Fig. 324. Distribution. Hyperaspidius hercules (dot); H. andrewsi (star); H. hardyi (square).
dially, widely separated from basal and apical margins (Fig. 325d, e). Punctures on
head extremely fine, barely visible; pronotal punctures larger than on head, separated
by 2 or 3 times a diameter; surface of elytron dull, reticulate, punctures larger than
on pronotum, separated by a diameter or less. Metasternum coarsely, densely punc-
tured laterally,, nearly impunctate medially. Abdominal sterna densely, finely punc-
tured; postcoxal line complete, area within line alutaceous, densely punctured; apex
of 6th sternum with lateral angle abrupt, emarginate medially. Male genitalia as in
Figure 325a-c.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Female, length 2.35 mm, width 1.65 mm. Similar to holotype except head black;
pronotum black except lateral Vs and small median spot near apex yellow; 6th ab-
dominal sternum abruptly narrowed toward apex, apex rounded.
Variation. Length 1.80 to 2.65 mm, width 1.20 to 1.80 mm. The male pronotum
may lack some or all of the maculae in the basal ', and the female may have the
median pronotal yellow spot either entirely lacking, or expanded to reach apical
margin.
Holotype. Male. UTAH: Kane Co., Coral Pink Sand Dunes, VII- 16-75, Fred G.
Andrews, A. R. Hardy (USNM 101336).
Allotype. Female. Same data as holotype. (USNM).
Paratypes. Total 10 (Fig. 324). All with same data as holotype. (USNM) (CD A).
This species is the most striking and distinctive of all known species of Hyperas-
pidius. The elongate, apically tapered body, straw yellow color, and strongly modified
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NORTH AMERICAN COCCINELLIDAE
395
Fig. 326. Hyperaspidius hardyi.
6th sternum of both sexes are highly diagnostic. I name the species for one of the
collectors of the type series.
Hyperaspidius hardyi, new species
Fig. 326a-e; Map, Fig. 324
Description. Male, length 2. 10 mm, width 1 .65 mm. Form rounded, convex, lateral
margin of elytron definitely curved. Head and pronotum yellow, pronotum with
bilobed black area in basal ‘, each lobe with yellow spot present. Elytron yellow,
narrow sutural margin and broad median vitta dark brown (Fig. 326d, e). Punctures
on head extremely fine, barely perceptible; pronotal punctures larger than on head,
separated by one to 3 times a diameter; punctures on elytron equal in size to pronotal
punctures except on brown median vitta, there becoming coarse, separated by one
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
to 3 times a diameter. Metasternum coarsely, densely punctured laterally, nearly
impunctate medially. Abdominal sterna coarsely punctured; postcoxal line complete,
area within line shiny, polished, densely punctured. Genitalia as in Figure 326a-c.
Female, length 2.20 mm, width 1.75 mm. Similar to holotype except head brownish
yellow, vertex dark brown; pronotum dark brown except lateral ‘A yellow, midline
with faint yellowish brown vitta.
Variation. Length 2.10 to 3.0 mm, width 1.65 to 2.0 mm. The female head may
be yellow except vertex black, and the female pronotum may have the black area
reduced, as in the male.
Holotype. Male. CALIFORNIA: San Bernardino Co., Cadiz Dunes, IV-25-78, Alan
R. Hardy & F. G. Andrews, walking dunes at night (USNM 101337).
Allotype. Female. Same data as holotype. (USNM).
Paratypes. Total 6 (hg. 324). Five with same data as holotype; 1, 6 mi. N Palm
Springs Calif., VII-8-54, G. H. Nelson, Snow Creek, on Ephedra californica. (USNM)
(CDA) (WHN).
This species has the facies of a member of Hyperaspis rather than Hyperaspidius
because of the round, convex body shape. On that basis alone this species is quite
distinctive, also, the coarse punctures on the elytron are confined to the median
brown vitta, which I have not observed in any other Hyperaspidius species. I name
this species for one of the collectors of the type series.
Genus Helesius
Helesius Casey, 1899, p. 129. — Leng, 1920, p. 212. — Korschefsky, 1931, p. 202.—
Chapin, 1966, p. 280. Type-species; Helesius nubilans Casey, by subsequent des-
ignation of Korschefsky, 1931.
Hyperaspini with body elongate, oval, dorsoventrally convex; dorsal surface gla-
brous. Head and pronotum red, or head red, pronotum reddish brown; elytron brown
or black. Antenna 10-segmented (Fig. 327a); antennal insertion concealed. Eye entire.
Scutellum large, wider than long. Epipleuron of elytron narrow, obliquely inclined
toward outer margin, strongly excavated for reception of middle and hind femoral
apices. Prostemum with 2 parallel carinae not convergent anteriorly. Posterior margin
of metastemum abruptly descending between coxa and lateral margin. Leg with femur
and tibia compressed, apex of tibia thickened, excavated for reception of tarsal base
(Fig. 327b); hind femur extremely broad; tarsal claw without tooth. Postcoxal line
on first abdominal sternum incomplete, of Scymnus type (Fig. 327c). Apical abdom-
inal sternum of male feebly emarginate. Male genitalia with basal lobe asymmetrical,
paramere rooted in phallobase (Fig. 328a). Female genitalia with compound sper-
mathecal capsule (Fig. 327d); coxal plate transverse.
This genus is distinctive in the North American hyperaspine fauna because the
legs are compressed and the hind femur is extremely robust. Also, the apex of each
tibia is thickened and excavated. The genitalia (male and female) are of the type
possessed by the species in Section I of Hyperaspis, and Helesius is more closely
related to Hyperaspis than to any other genus of Hyperaspini. There are 3 species
presently described in Helesius, 2 of these are North American and one was described
from Colombia.
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NORTH AMERICAN COCCINELLIDAE
397
Fig. 327. Helesius sp. a. Antenna, b. Front leg. c. Postcoxal line. d. Spermathecal capsule.
A total of 9 specimens have been examined; no host data is on record for any
member of this genus.
Key to species of Helesius
1 . Punctures on head and elytron dense, Colorado, Montana nigripennis (LeConte)
- Punctures on head not apparent, barely perceptible, punctures on elytron fine, indistinct;
Texas nubilans Casey
Helesius nigripennis (LeConte)
Fig. 327d; Map, Fig. 329
Scymnus nigripennis LeConte, 1878b, p. 453.
Helesius nigripennis: Casey, 1899, p. 129. — Leng, 1920, p. 212.— Korschefsky, 1931,
p. 202.
Diagnosis. Length 2.45 to 3.0 mm, width 1.75 to 2.0 mm. Head red. Pronotum
red in lateral V3 with poorly defined reddish brown area medially. Elytron black or
dark brown. Female genitalia as in Figure 327d.
Discussion. The type in the LeConte collection labeled “8000 ft., Florissant, Col.,
Aug. 17-22, 1877/Type 6724(red paper)/S. nigripennis Lec./'is a Hyperaspis”, is a
holotype. I have not seen a male of this species.
Type locality. Colorado, Florissant, 8,000 feet.
Type depository. MCZ.
Distribution. Figure 329. COLORADO: Teller Co., Florissant. MONTANA: Lewis
and Clark Co., Helena.
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Fig. 328. Helesius nubilans.
Helesius nubilans Casey
Fig. 328a-e; Map, Fig. 329
Helesius nubilans C2iSQy , 1899, p. 129. — Leng, 1920, p. 212. — Korschefsky, 1931, p.
202.
Diagnosis. Length 2.80 to 3.0 mm, width 2.10 to 2.25 mm. Head red. Pronotum
dark red in lateral ‘A with poorly defined reddish brown area medially. Elytron black
or dark brown (Fig. 328e). Male genitalia as in Figure 328a-c. Female genitalia as
in Figure 328d.
Discussion. All specimens of H. nubilans examined, with one exception, have been
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NORTH AMERICAN COCCINELLIDAE
399
collected in the Brownsville, Texas area. The exception is a female (USNM) from
San Antonio, Texas. This specimen has an entirely red pronotum and is larger (3.5
mm long) than normal for H. nubilans. There is a good possibility that the specimen
represents an undescribed species, but a male is needed to ascertain this. There are
2 female type specimens of H. nubilans in the Casey collection, I here designate and
label one as the lectotype and the other a paralectotype.
Type locality Brownsville, Texas (lectotype here designated).
Type depository. USNM (35213).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Distribution. Figure 329. TEXAS: Bexar Co., San Antonio; Cameron Co., Browns-
ville.
Genus Thalassa
ThalassaMuXsdini, 1850, p. 511.— Crotch, 1874b, p. 209.— Gorham, 1894, p. 182.—
Leng, 1920, p. 212.— Korschefsky, 1931, p. 209.— Chapin, 1966, p. 280. Type-
species; Chilocorus pentaspilotus Chevrolat, by subsequent designation of Crotch,
1874b.
Hyperaspini with body rounded, convex; dorsal surface glabrous. Head yellow in
male, black or bluish black in female. Antenna 1 1 -segmented (Fig. 330a); antennal
insertion exposed. Eye entire. Epipleuron of elytron wide, strongly descending ex-
ternally, deeply excavated for reception of middle and hind femoral apices. Proster-
num with 2 parallel, incomplete carinae. Posterior margin of metastemum abruptly
descending between coxa and lateral margin. Leg compressed, anterior tibia flattened,
rounded or angulate at anterior part of outer margin; tarsal claw with large basal
tooth (Fig. 330b). Postcoxal line on first abdominal sternum incomplete, of Scymnus
type (Fig. 330c). Apical abdominal sternum in male feebly emarginate. Male genitalia
with basal lobe asymmetrical, paramere rooted in phallobase (Fig. 330d). Female
genitalia with compound spermatheca, basal portion with appendix, coxal plate trans-
verse (Fig. 331b).
The only species of this genus known to occur north of Mexico is Thalassa mon-
tezumae Mulsant, which can be recognized on body form and dorsal color pattern.
The primary diagnostic characteristics of the genus as a whole are the strongly de-
scending, deeply foveolate elytral epipleura, and the expanded tibial apices. Thalassa
is a New World genus containing 6 described species ranging from Arizona and Cuba
to Brazil. The only host record seen for Thalassa species is the soft scale, Toumeyella
mirabilis (Cockerell).
Thalassa montezumae Mulsant
Fig. 330a-f, 331a, b; Map, Fig. 332
Thalassa montezumae Mulsant, 1850, p. 512.— Crotch, 1873, p. 364.— Crotch, 1874b,
p. 209.— Gorham, 1894, p. 183. — Leng, 1903, p. 211. — Leng, 1920, p. 212.—
Korschefsky, 1931, p. 209.
Diagnosis. Length 4.50 to 5.80 mm, width 4.0 to 5.0 mm. Form rounded, convex.
Male pronotum bluish black with anterior and lateral margins narrowly yellow; female
pronotum entirely bluish black except anterolateral angle barely perceptibly yellow.
Elytron bluish black with reddish yellow spot in apical ‘ (Fig. 331a). Male genitalia
as in Figure 330d-f
Discussion. Two type specimens of montezumae exist in the Crotch collection, and
I here designate one of these labeled “Mexico/Type/” as the lectotype, the other
specimen as a paralectotype.
Type locality. “Mexique” (lectotype here designated).
Type depository. UCCC.
Distribution. Figure 332. ARIZONA: Cochise Co., San Bernardino Ranch; Douglas;
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NORTH AMERICAN COCCINELLIDAE
401
Fig. 330. Thalassa montezumae. a. Antenna, b. Front leg. c. Postcoxal line. d-f. Male
genitalia.
Graham Mts.; Huachucha Mts.; Oslar; Nogales; Palmerlee; Ruby; Santa Rita Mts.;
Tucson; Wilcox. TEXAS: Brownsville; Harlingen.
Genus Hyperaspis Redtenbacher
Redtenbacher, 1844, p. 8.— Mulsant, 1850, p. 649.— Costa, 1849, P. 64.—
Crotch, 1873, p. 379.-Crotch, 1874b, p. 224.-Gorham, 1894, p. 191. -Wick-
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 331. Thalassa montezumae.
ham, 1894, p. 299.— Casey, 1899, p. 120. — Blatchley, 1910, p. 521. — Korschefsky ,
1931, p. 177.-Dobzhansky, 1941, p. l.-Wingo, 1952, p. 17. -Chapin, 1966, p.
280.— J. Chapin, 1974, p. 39.— Belicek, 1976, p. 309. Type-species; Coccinella
reppensis Herbst, by subsequent designation of Crotch, 1874b.
Oxynychus LeConte, 1850, p. 238. — LeConte, 1852, p. 130. — Mulsant, 1850, p.
649.-Crotch, 1874b, p. 239.-Chapuis, 1876, p. 258.-Weise, 1890, p. 489.-
Casey, 1899, p. 1 16. — Korschefsky, 1931, p. 200. — Dobzhansky, 1941, p. 78.—
Bielawski, 1959, p. 54. Type-species; Oxynychus moerens LeConte, by monotypy.
(Korschefsky, 1931, incorrectly listed Coccinella erythrocephalus F. as the type-
species of Oxynychus.)
H y per aspis {Oxynychus): Mulsant, 1850, p. 694. — Mader, 1955, p. 850.— Miyatake,
1961, p. 154. — Kamiya, 1963, p. 79.
Hyperaspini with form elongate, oval, or rounded, dorsoventrally flattened or
hemispherical; dorsum glabrous. Head usually yellow in male, brown or black in
female; elytron usually with pale maculation on dark background, rarely immaculate.
Antenna 10 or 11 -segmented (Fig. 333a, b); antennal insertion exposed. Scutellum
large, wider than long. Epipleuron of elytron narrow, not descending externally, often
medially grooved, distinctly excavated for reception of middle and hind femoral
apices (Fig. 333d). Prostemum with 2 carinae convergent anteriorly (Fig. 333c).
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NORTH AMERICAN COCCINELLIDAE
403
Posterior margin of metastemum abruptly descending between coxa and lateral mar-
gin. Leg with femur and tibia slightly compressed; anterior tibia simple; tarsal claw
with or without basal tooth. Postcoxal line on first abdominal sternum incomplete,
of Scymnus (Scymnus) type (Fig. 333e). Apical abdominal sternum in male not or
very weakly modified. Male genitalia with median lobe asymmetrical, paramere
rooted in phallobase (Fig. 334a). Female genitalia with compound spermathecal
capsule (Fig. 334d), basal portion with appendix; coxal plate usually transverse.
Most members of Hyper aspis are easily recognized by the key characters, however.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 333. Hyperaspis sp. a, b. Antennae, c. Prostemum. d. Epipleuron. e. Postcoxal lines,
f. Front leg.
some of the species have the epipleural excavations reduced, and if not examined
closely, may be confused with species of Hyperaspidius. Hyperaspis is found world-
wide, but the bulk of the species of this huge genus are neotropical, with 103 species
and subspecies occurring in America north of Mexico.
I have not been able to locate type specimens of H. horni Crotch and H. subsignata
Crotch, nor can their identity be determined from the original description, therefore
they must remain unrecognized species. H. annularis Boheman does not occur north
of Mexico and is here deleted from the list of North American species. Four species
formerly placed in Hyperaspis, H. asphaltina Casey, H. nubilata Casey, H. marginata
Gaines, and H. tristis LeConte, are transferred to Hyperaspidius. Two species, H.
microsticta Casey and H. triplicans Casey are transferred to Brachiacantha. These
reassignments are based on examination of primary types.
Host records indicate that species of Hyperaspis prey only on families of Homop-
tera, and that many families within that order serve as hosts; the families Pseudo-
coccidae and Coccidae the most frequently attacked (El-Ali, unpubl. dissertation).
Specific host records are as follows: Scale insects; Amonostherium {=Erium) lichten-
siodes {CockQVQW), Antonina graminis (Maskell), Aspidiotus destructor {Signorti), Bod-
enheimera racheli (Bodenheimer), Ceroplastes sinensis {Ddi Guercio), Chrysomphalus
aonidum (Linn), Coccus hesperidum (L.), Coccus pseudomagnoliarum (Kuwana),
Dactylopius coccus Costa, Dactylopius confusus (Cockerell), Dactylopius opuntiae
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NORTH AMERICAN COCCINELLIDAE
405
(Cockerell), Dactylopius tomentosus (Lamarck), Dactylopius spp., Dysmicoccus brev-
ipes (Cockerell), Dysmicoccus {= Pseudococcus) boninsis (Kuwana), Dysmicoccus
{= Pseudococcus) ryani (Coquillett), Eriococcus quercus (Comstock), Ferrisia virgata
(Cockerell), Icerya purchasi (Maskell), ""Inglisia"' malvacearum (Cockerell), Lecan-
ium spp., Lepidosaphes beckii (Newman), Maconellicoccus (=Phenacoccus) hirsutus
(Green), Mesolecanium nigrofasciatum (Pergande), Metaceronema japonica (Mas-
kell), Neopulvinaria ineretina (Khazhibeili), Nipaecoccus Pseudococcus) aurilana-
tus (Maskell), Nipaecoccus {= Pseudococcus) filamentosus (Cockerell), Nipaecoccus
Pseudococcus) nipae (Maskell), Orthezia artemisiae (Cockerell), Orthezia urticae
(Linn), Orthezia spp., Parthenolecanium corni (Bouche), Phenacoccus acericola (King),
Phenacoccus colemani (Ehrhom), Phenacoccus gossypii (Townsend and Cockerell),
Phenacoccus helianthi (Cockerell), Phenacoccus pergandei (Cockerell), Physokermes
insignicola (Craw), Planococcus {= Pseudococcus) citri (Risso), Planococcus kenyae
(LePelly), Planococcus lilacinus (Cockerell), Protopulvinaria fukayai (Kuwana), Pro-
topulvinaria pyriformis (Cockerell), Pseudococcus calceolariae (Maskell), Pseudococ-
cus citriculus (Green), Pseudococcus comstocki (Kuwana), Pseudococcus longispinus
(Targioni-Tozzetti), Pseudococcus maritimus (Ehrhom), Pseudococcus spp., Pulvi-
naria acericola (Walsh and Riley), Pulvinaria aurantii (Cockerell), Pulvinaria citricola
(Kuwana); Pulvinaria floccifera (Westwood), Pulvinaria hazae Kuwana, Pulvinaria
hydrangeae (Steinweden), Pulvinaria innumerabilis (Rathvon), Pulvinaria torreyae
(Takahashi), Pulvinaria vitis (L.), Pulvinaria spp., Puto Pseudococcus) yuccae (Co-
quillett), Ripersia sp., Saccharicoccus {= Pseudococcus) sacchari (Cockerell), Sac-
charicoccus {=Trionymus) sacchari, Selenaspidus {=Pseudaonidia) articulatus (Mor-
gan), Sphaerolecanium prunastri (Boyer de Fonscolombe), Spilococcus {= Pseudococcus)
sequoiae {ColQmdcd), Takahashia japonica (Cockerell), Toumeyella liriodendri (Gme-
lin), Toumeyella mirabilis (Cockerell), Toumeyella parvicornis (Cockerell), Toume-
yella pini (King), Toumeyella pinicola (Ferris), Trionymus insularis (Ehrhom), Un-
aspis citri (Comstock). Aphids; Aphis craccivora Koch, Aphis fabae (Scopoli), Aphis
gossypii Glover, Aphis nerii Boyer de Fonscolombe, Aphis pomi Degeer, Cryptosi-
phum artemisiae (Buckton), Cryptosiphum gallarum (Kaltenbach), Macrosiphum eu-
phorbiae (Thomas), Melanaphis sacchari (Zehntner), Myzus malisuctus (Matsumura),
Rhopalosiphum maidis (Fitch), Schizaphis graminum (Rondani), Sipha flava (Forbes),
Toxoptera citricidus (Kirkaldy).
The North American species of Hyperaspis were taxonomically treated by Dob-
zhansky (1941), and the California species were recently the object of an excellent
dissertation (unpubl.) by El-Ali at the University of California, Berkeley. I have made
extensive use of El-Ali’s findings in preparing this section on Hyperaspis.
El-Ali was the first to realize that species of Hyperaspis possessed both 10 and 1 1-
segmented antennae, and he based his first major species grouping on this. He then
proceeded to define 1 9 minor groupings, modifying to a great extent Dobzhansky’s
(1941) grouping. The groups I recognize here differ to some extent from those of El-
Ali, principally for the following reasons: (1) El-Ali did not have available to him
many of the eastern species of Hyperaspis, and on examining these I find that some
of the criteria he used for group definition are rendered useless because a species
often possesses the external characteristics of one group and the internal character-
istics of another; (2) I believe that this type of informal grouping can be justified only
406
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
as long as it serves a practical purpose, therefore, I have combined many of El-Ali’s
groups because the definitions were tenuous, making them difficult to recognize and
use. The major division based on 1 1 -segmented or 1 0-segmented antennae is certainly
a valid one, and I refer to these as Section I and Section II, respectively. Within each
of these there are several groups based mainly on the type of genitalia and somewhat
on the body shape and color pattern.
Key to species of Hyperaspis
1 . Antenna 1 1 -segmented 2
- Antenna 1 0-segmented 53
2(1). Elytron with basal spot near scutellum (Fig. 373d), spot sometimes connected
to discal spot (Fig. 379) 3
Elytron without basal spot near scutellum 7
3(2). Elytron with 4 or 5 spots 4
Elytron with less than 4 spots, often appearing vittate 5
4(3). Pronotum mostly black, lateral border and/or anterior border yellow (Fig. 373d)
levrati Mulsant
- Pronotum yellow with irregular, median dark area (Fig. 395e)
longicoxitis Nutting
5(3). Elytron with irregular median vitta (connected spots) from base to apex (Fig.
382d); known only from Mississippi esclavium Dobzhansky
Elytron not as described above; species occurring west of the Mississippi River
6
6(5). Basal spot on elytron large, elongate oval, often connected to apical spot, discal
spot absent (Fig. 393d, e) fastidiosa Casey
Basal spot on elytron small, often narrowly connected to small discal spot, discal
spot often absent (Fig. 379d, e) revocans Casey
7(2). Elytron entirely black or brown, immaculate: California pluto Fall
- Elytron always with maculation; California and elsewhere 8
8(7). Elytron entirely red except narrow lateral border and broad basal border black
(Fig. 353d); known only from Florida nigrosuturalis Blatchley
Elytron not as described above; Florida and elsewhere 9
9(8). Elytron with humeral spot or with lateral vitta beginning at humeral angle .. 10
- Elytron without humeral spot or vitta 15
10(9). Elytron with lateral vitta extending from humeral angle beyond midpoint ... 11
- Elytron with humeral spot or short vitta not extending beyond midpoint .... 12
11(10). Surface of elytron dull, alutaceous; male pronotum almost entirely yellowish
red; discal and apical spots on elytron not connected lugubris (Randall)
- Surface of elytron shiny; male pronotum mostly black, or yellow with irregular,
median black area; discal and apical spots on elytron often connected, lateral
vitta often extending to apical spot (Fig. 393d, e) fastidiosa Casey
12(10). Elytron with 4 yellow spots, discal spot with anterior border emarginate (Fig.
347d) octonotata Casey
- Elytron with no more than 3 red or yellow spots, discal spot not emarginate
(often absent) 13
13(12). Discal spot on elytron broadly connected to lateral spot (Fig. 368d) . .excelsa Fall
- Discal spot on elytron present or absent, if present then not connected to lateral
spot (Fig. 365e, 367d) 14
14(13). Female pronotum entirely black; male pronotum narrowly yellow on lateral
and apical margin (Fig. 365e) lateralis Mulsant
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NORTH AMERICAN COCCINELLIDAE
407
15(9).
16(15).
17(16).
18(17).
19(18).
20(18).
21(20).
22(21).
23(22).
24(17).
25(24).
26(25).
27(26).
28(27).
29(28).
Female pronotum with yellow spot on lateral margin; male pronotum broadly
yellow laterally, narrowly yellow apically (Fig. 367d) pinguis Casey
Apex of clypeus truncate elytron with 2 small, apical spots, usually confluent
(Fig. 356d); coastal localities from Virgina to Georgia gemina LeConte
Apex of clypeus emarginate; elytron not as described above; not restricted to
eastern coastal localities 16
Body form nearly rectangular, elongate; elytron with large, median yellow spot
extending from lateral margin nearly to suture (Fig. 364d) eastern United States
lewisi Crotch
Body form oval or rounded, if somewhat rectangular then not from eastern
U.S.; pattern on elytron not as described above; eastern United States and
elsewhere 17
Elytron with 3 spots, one discal, one on lateral margin at or just posterior to
midpoint, one at apex (Figs. 374d) 18
Elytron with one to 4 spots, if 3 spots present, then not arranged as described
above 24
Species not known to occur west or south of Kansas and Missouri 19
Species known only from Texas, New Mexico and Arizona 20
Body form extremely convex, rounded (Fig. 374d); New England to Minnesota,
Kansas, and Missouri deludens, n. sp.
Body form elongate, oblong (Fig. 376); known only from “Missouri” and New
Jersey pratensis LeConte
Species occurring in Texas medialis Casey
Species occurring in Arizona and New Mexico 21
Lateral spot on elytron posterior to middle (Fig. 38 Id) triangulum Casey
Lateral spot on elytron at middle (Fig. 3 9 2d) 22
Male pronotum with anterior margin black medially (Fig. 392d); length 1.90
to 2.30 mm conspimns Casey
Male pronotum with anterior margin yellow medially, length 2.20 to 2.80 mm
23
Form oval, not strongly convex; apical spot on elytron heart shaped (Fig. 39 Id)
gemma Casey
Form rounded, strongly convex; apical spot on elytron round (Fig. 378d) . . .
aemulator Casey
Elytron with single apical spot (Fig. 357e) bigeminata (Randall)
Elytron with one or more spots, if only one spot present, then not located at
apex 25
Elytron with large, marginal red spot enclosing small yellow spot, and single
apical spot (Fig. 385d), apical spot may be absent, Utah uteana, n. sp.
Elytron not as described above; Utah and elsewhere 26
Elytron with 2 small spots, one at apex, one on lateral margin in apical '/s (Fig.
372e) chapini Dobzhansky
Elytron not as described above 27
Elytron with 4 spots (Fig. 335a); Brownsville, Texas weisei Schaeffer
Elytron not as described above 28
Elytron with discal spot and 2 apical spots (Fig. 334e) proba (Say)
Elytron not as described above 29
Elytron with irregular median vitta extending from base to apex (Fig. 3 8 2d);
known only from Mississippi esclavium Dobzhansky
Elytron not as described above; Mississippi and elsewhere 30
408
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
30(29).
31(30).
32(31).
33(32).
34(32).
35(34).
36(35).
37(31).
38(37).
39(37).
40(39).
41(40).
42(41).
43(42).
44(43).
45(44).
46(44).
Elytron with single yellow spot on lateral margin in apical spot often narrowly
elongated posteriorly (Fig. 383e); California osculans LeConte
Elytron not as described above; California and elsewhere 31
Elytron with 2 spots, one discal, one apical 32
Elytron with single spot in discal area or apical *, spot may be greatly expanded
37
Species occurring only in California 33
Species not occurring in California 34
Body form convex, rounded (Fig. 386d); anterior pronotal margin of male black
mckenziei Nutting
Body form broad, depressed (Fig. 34 1 d); anterior pronotal margin of male yellow
jovialis Fall
Body form very elongate, oval (Fig. 345d); mountains of northern New Mexico
haematosticta Fall
Body form rounded; not known from New Mexico 35
Body of female dorsoventrally flattened; basal lobe of male genitalia with lateral
projection on left side in ventral view (Fig. 354a) conviva Casey
Body of female not dorsoventrally flattened; basal lobe of male genitalia with
lateral projection on right side in ventral view 36
Basal lobe of male genitalia not strongly asymmetrical, apex rounded (Fig. 351a)
pistillata Watson
Basal lobe of male genitalia strongly asymmetrical, apex truncate (Figs. 348a)
signata signata (Olivier)
Species known only from California 38
Species not occurring in California 39
Spot on elytron distinctly separated from lateral margin, often with small,
subapical black spot enclosed (Fig. 34 Id) jovialis Fall
Spot on elytron touching or very narrowly separated from lateral margin (Fig.
342e) leachi Nunenmacher
Anterior margin of prostemum crenate southern Arizona oculifera Casey
Anterior margin of prostemum smooth; Arizona and elsewhere 40
Spot on elytron located in apical ‘ near apical margin (Fig. 370d); Florida . . .
ornatella, n. sp.
Spot on elytron located in discal area, or if in apical *, then not approaching
apical margin; Florida and elsewhere 41
Species known only from Arizona tuckeri Casey
Species not occurring in Arizona 42
Body form very elongate, oval (Fig. 345a); mountains of northern New Mexico
haematosticta Fall
Body form not extremely elongate; not known to occur in New Mexico 43
Length 2.0 mm or less; body extremely convex, rounded (Fig. 338e); Browns-
ville, Texas globula Casey
Length more than 2.0 mm; body not as described above; Texas and elsewhere
44
Species known only from Texas 45
Species not occurring in Texas 46
Discal spot on elytron posterior to middle (Fig. 359d) wickhami Casey
Discal spot on elytron on middle of disc signata bicentralis Casey
Female pronotum with yellow area on lateral margin; male pronotum with
lateral yellow area occupying % or more of pronotum
47
1985
NORTH AMERICAN COCCINELLIDAE
409
- Female pronotum entirely black; male pronotum with lateral yellow area oc-
cupying Vs or less of pronotum (except concavus male) 48
47(46). Discal spot on elytron on middle of disc, sometimes extended posteriorly (Fig.
337d) rivularis Dobzhansky
- Discal spot on elytron posterior to middle (Fig. 346d), occasionally greatly
enlarged inedita Mulsant
48(46). Body of female dorsoventrally flattened; basal lobe of male genitalia with lateral
projection on left side in ventral view (Fig. 354a) conviva Casey
- Body of female not dorsoventrally flattened; basal lobe of male genitalia with
lateral projection on right side in ventral view, or lacking lateral projection . 49
49(48). Basal lobe of male genitalia with apical angles rounded (Fig. 351a)
pistillata Watson
Basal lobe of male genitalia not as figured above 50
50(49). Paramere of male genitalia short, spatulate (Fig. 361a) 51
- Paramere of male genitalia long, slender, not spatulate 52
51(50). Pronotum of male with anterolateral angle and apical margin broadly yellow
(Fig. 361a) concavus Watson
Pronotum of male narrowly yellow on lateral and apical margins (Fig. 360d)
major Dobzhansky
52(50). Basal lobe of male genitalia slender, not strongly asymmetrical (Fig. 343a) . .
binotata (Say)
- Basal lobe of male genitalia broad, strongly asymmetrical (Fig. 348a)
signata signata (Olivier)
53(11). Head entirely pubescent 54
Head glabrous except often with sparse pubescence on apical border of clypeus
55
54(53). Elytron immaculate, or with lateral spot not extending forward to humeral angle
(Fig. 396e); Arizona, Utah, southern California significans Casey
Elytron always maculate, with the lateral spot usually extending forward to
humeral angle; Texas, New Mexico cruenta LeConte
55(53). Pronotum in both sexes with lateral yellow area large, more than 3/5 wider
than long; elytron with 2 spots, discal and apical, connected or not, apical spot
almost reaches hind margin of elytron 56
- Pronotum entirely black or with lateral yellow area twice as long as wide, if less
than twice as long as wide, then elytron not as described above 58
56(55). Elytron with discal and apical spots connected (Fig. 387e) . connectens (Thunberg)
- Elytron with discal and apical spots not connected 57
57(56). Apical spot on elytron not approaching suture (Fig. 388e); Texas . . rotunda Casey
Apical spot on elytron reaching suture or nearly so (Fig. 390d); Arizona ....
dobzhanskyi, n. sp.
58(55). Elytron immaculate 59
Elytron with at least one spot or vitta 64
59(58). Species occurring in the southeastern United States 60
- Species occurring west of the Mississippi River 61
60(59). Body elongate, somewhat flattened; male pronotum with lateral Va yellow (Fig.
410d) uniformis Casey
- Body oval, not distinctly flattened; male pronotum narrowly yellow on lateral
margin binaria Casey
6 1(59). Head black in both sexes; pronotum entirely black in both sexes; body slender,
elongate (fig. 454d); Alberta, Colorado, Wyoming jasperensis Belicek
410
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Head yellow in male; male pronotum narrowly yellow laterally; body oval; not
known from northern Rocky Mountains 62
62(6 1). Body dorsoventrally depressed, elongate oval (Fig. 452d); epipleuron not deeply
excavated for femoral apices; Arizona, Nevada simulans Casey
- Body convex, rounded; epipleuron deeply excavated for femoral apices; Ari-
zona, Nevada, and elsewhere 63
63(62). Pronotum narrowly yellow laterally in both sexes; western Washington and
Oregon immaculata Hatch
Pronotum black in female, narrowly yellow on lateral margin in male; not known
from western Washington and Oregon pleuralis Casey
64(58). Dorsal surface strongly alutaceous; elytron with postdiscal spot broadly con-
nected to complete lateral vitta (Fig. 456d); Illinois, Indiana, Kansas
bolteri LeConte
- Species not as described above, or if so, then not occurring east of the Mississippi
River 65
65(64). Elytron with marginal vitta extending from near scutellum around humeral
angle to apex, small, elongate discal spot present, often expanded posteriorly
to join marginal vitta (Fig. 457d-i); Texas, Arizona trifurcata Schaeffer
Elytron not as described above; Texas and elsewhere 66
66(65). Elytron with one basal spot in addition to lateral vitta or humeral spot 67
- Elytron without basal spot, with or without lateral vitta or humeral spot .... 69
67(66). Elytron with discal vitta (Fig. 443d) consimilis LeConte
- Elytron with postdiscal spot 68
68(67). Basal spot on elytron triangular, nearer humeral spot or lateral vitta than scu-
tellum (Fig. 43 5d) disconotata Mulsant
Basal spot on elytron round, nearer scutellum than humeral spot (Fig. 437d)
troglodytes Mulsant
69(66). Elytron vittate in appearance, one discal and one marginal vitta present .... 70
Elytron not appearing vittate 74
70(69). Dorsal surface dull, strongly alutaceous; Illinois, Indiana
brunnescens Dobzhansky
Dorsal surface shiny, lacking alutaceous sculpture; Iowa and west 71
71(70). Elytron with discal and marginal vitta usually broadly joined at apex (Fig.
428d)); California annexa LeConte
- Elytron with discal and marginal vittae not joined apically, or very narrowly
so; not known from California 72
72(71). Male pronotum black on anterior margin; occurring from Idaho east to Iowa,
south to New Mexico quadrivittata LeConte
Male pronotum yellow on anterior margin; Idaho, Washington, Oregon, north-
ern California 73
73(72). Basal lobe of male genitalia bisinuate on sclerotized side (Fig. 442a) occurring
principally from the Cascade Mountains to the Pacific Coast
borealis Dobzhansky
Basal lobe of male genitalia not bisinuate, emarginate in apical ‘ on sclerotized
side (Fig. 434a); occurring principally east of the Cascade Mountains
oregona Dobzhansky
74(69). Color pattern on elytron consisting only of complete vitta on lateral margin . 75
- Color pattern on elytron with or without complete vitta on lateral margin, if
vitta present, then additional maculation also present 81
75(74). Pronotum impunctate, surface dull, strongly alutaceous; marginal vitta on ely-
1985
NORTH AMERICAN COCCINELLIDAE
411
76(75).
77(76).
78(77).
79(78).
80(79).
81(74).
82(81).
83(82).
84(83).
85(83).
86(81).
87(86).
88(87).
iron '/4 or less the width of elytron; Pennsylvania to Rorida, Mississippi ....
fimbriolata Melsheimer
Pronotum punctate, surface usually shiny, not strongly alutaceous; not confined
to Atlantic and Gulf coast seaboards 76
Body form elongate, nearly parallel sided; apex of lateral vitta on elytron strongly
separated from margin (Fig. 449d); Arizona protensa Casey
Body form oval or rounded; apex of lateral vitta on elytron narrowly separated
from margin, if strongly separated then not occurring in Arizona; Arizona and
elsewhere 77
Elytron with vitta usually occupying 1/2 of elytron at midpoint (Fig. 442d);
California LeConte
Elytron with vitta occupying V3 or less of elytron at midpoint; California and
elsewhere 78
Basal lobe of male genitalia slender, rounded at apex, lateral projection rounded
(Fig. 398a); Santa Rita Mountains, Arizona sanctaeritae Dobzhansky
Basal lobe of male genitalia broad, apex acute, lateral projection not as figured
above; Arizona and elsewhere 79
Basal lobe of male genitalia with lateral projection near apex, blunt (Fig. 401a)
inflexa Casey
Basal lobe of male genitalia with lateral projection near midpoint, rounded (Fig.
403a) 80
Body form depressed dorsoventrally; marginal vitta on elytron occupying Va of
elytron, apex strongly separated from margin of elytron (Fig. 406d); known only
from San Diego, California limbalis Casey
Body form convex; marginal vitta on elytron occupying Vi of elytron, apex
narrowly separated from margin of elytron (Fig. 403d); not known from San
Diego caseyi, n. sp.
Species with single spot or short vitta on lateral margin of elytron in basal %,
spot never extended onto disc 82
Species with one or more spots on elytron, if only one, then in apical V3 or
extended onto disc from lateral margin 86
Species occurring in southeastern United States binaria Casey
Species occurring west of the Mississippi River 83
Elytron with lateral vitta extending from humeral angle beyond midpoint ... 84
Elytron with spot at humeral angle or at midpoint 85
Species occurring in southern Texas schaefferi, n. sp.
Species occurring west and north of New Mexico dissoluta nevadica Casey
Body form convex, apex slightly truncate; elytron with spot slightly posterior
to midpoint, (Fig. 413e) pleuralis Casey
Body form flattened dorsoventrally; elytron with spot slightly posterior to mid-
point, or with very narrow, elongate spot at humeral angle simulans Casey
Large robust species; elytron with triangular humeral spot and spot at apex (Fig.
42 Id) nunenmacheri Casey
Species not as described above 87
Elytron with marginal vitta from humeral angle beyond midpoint, and apical
spot 88
Elytron not maculate as described above 90
Female pronotum entirely black; male genitalia with strong lateral projection
in basal ‘ (Fig. 4 1 9a) dissoluta dissoluta Crotch
Female pronotum with lateral margin narrowly yellow 89
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
89(88). Basal lobe of male genitalia bisinuate on sclerotized side (Fig. 442a)
borealis Dobzhansky
- Basal lobe of male genitalia not bisinuate, emarginate in apical ‘ on sclerotized
side (Fig. 434a) oregona Dobzhansky
90(87). Elytron with single spot on apical Vs, may extend onto apical ‘ (also see sene-
galensis hottentota) 91
Elytron with more than one macula or spot not on apical Vs 95
91(90). Spot on elytron obliquely elongate, near suture (Fig. 448); occurring in Great
Lakes region moerens LeConte
- Spot on elytron not obliquely elongate; occurring west of the Mississippi River
92
92(9 1). Spot on elytron large, extending onto apical ‘, nearer lateral margin than suture
(Fig. 429d); Arizona arizonica Dobzhansky
Spot on elytron small, not extending onto apical ‘; not occurring in Arizona . 93
93(92). Spot on elytron narrow, elongate, nearer suture than apical margin of elytron
borealis Dobzhansky
- Spot on elytron rounded or wedge-shaped, very near apical margin of elytron,
distinctly removed from suture (Fig. 43 Id) 94
94(93). Length 2.40 mm or less; spot on elytron less than 2 times as far from suture
as from apical margin (Fig. 43 Id); basal lobe of male genitalia not longer than
paramere (Fig. 431a) oculaticauda Casey
Length 2.30 mm or more; spot on elytron usually more than 2 times as far from
suture as from apical margin (Fig. 42 3e); basal lobe of male genitalia longer
than paramere (Fig. 423a) postica LeConte
95(90). Discal spot on elytron broadly connected to lateral vitta or marginal spot (Fig.
417d, g) 96
- Discal spot on elytron absent, or if present, then discrete 97
96(95). Female pronotum black; surface of pronotum shiny; elytron with fine, indistinct
punctures, lateral vitta extending from humeral angle to just beyond midpoint
(Fig. 4 1 7e), or extending to apex of elytron (Fig. 4 1 7g), or interrupted just before
apex, leaving apical spot free taeniata LeConte
- Female pronotum yellow on lateral margin; surface of pronotum dull with
apparent alutaceous sculpture; elytron with coarse, dense punctures, lateral
yellow area not extending to humeral angle, or very narrowly so (Fig. 426e) .
quadrioculata (Motschulsky)
97(95). Elytron without discal spot 98
- Elytron with discrete discal spot 102
98(97). Elytron with large, oblong apical spot very near apical margin, and one small
spot on lateral margin (Fig. 433d) querquesi Nutting
- Elytron with spot pattern not exactly as described above 99
99(98). Elytron with 3 discrete marginal spots, humeral spot large, triangular (Fig. 433d)
100
Elytron with marginal vitta or with 2 or 3 marginal spots, if with 3 spots, then
humeral spot reduced to narrow, elongate streak 101
100(99). Female pronotum entirely black; male pronotum with yellow area on lateral
margin not extending to posterolateral angle psyche Casey
Female pronotum yellow on lateral margin; male pronotum with yellow area
on lateral margin extending to posterolateral angle (Fig. 426d)
quadrioculata (Motschulsky)
101(99). Basal lobe of male genitalia bisinuate on sclerotized side (Fig. 442a)
borealis Dobzhansky
1985
NORTH AMERICAN COCCINELLIDAE
413
102(97).
103(102).
104(102).
105(104).
106(105).
107(104).
108(107).
109(108).
110(107).
111(110).
112(110).
113(112).
Basal lobe of male genitalia not bisinuate, emarginate in apical ‘/s on sclerotized
side (Fig. 434a) oregona Dobzhansky
Humeral angle of elytron black 103
Humeral angle of elytron yellow 104
Discal spot on elytron wedge-shaped, large lateral spot present medially (Fig.
445d); body flattened dorsoventrally spiculinota Fall
Discal spot on elytron rounded or elongate, lateral spot absent or extremely
reduced (Fig. 426d); body not flattened quadrioculata (Motschulsky)
Body form slender, nearly parallel sided (Fig. 44 1 d) 105
Body form oval or rounded (Fig. 41 7d) 107
Species occurring along the Atlantic seaboard to Florida and Alabama
paludicola Schwarz
Species occurring west of the Mississippi River 106
Lateral margin of elytron with complete vitta (Fig. 407); Arizona . . . filiola Casey
Lateral margin of elytron with vitta extending from humeral angle beyond
midpoint, apical and discal spots present; Texas, South Dakota
punctata LeConte
Female pronotum entirely black; basal lobe of male genitalia with lateral pro-
jection in basal % (Fig. 411a) 108
Female pronotum with yellow area on lateral margin; basal lobe of male genitalia
bisinuate (Fig. 439a), or with broad emargination on sclerotized side (Fig. 425a),
or if with lateral projection, then projection near apex (Fig. 453a) 110
Elytron with lateral vitta strongly widened opposite discal spot (Fig. 417d) . .
taeniata LeConte
Elytron with lateral spots or vitta, if vitta present, then emarginate opposite
discal spot (Fig. 41 Id) 109
Basal lobe of male genitalia with lateral projection in apical ‘ (Fig. 411a) ....
bensonica Casey
Basal lobe of male genitalia with lateral projection in basal ‘ (Fig. 415a)
disrupta Dobzhansky
Surface of pronotum dull, with strong alutaceous sculpture Ill
Surface of pronotum shiny, polished, usually with alutaceous sculpture absent
or visible only under high magnification 112
Lateral yellow area on pronotum as wide or nearly as wide as humeral spot on
elytron (Fig. 439d); punctures on elytron much larger than pronotal punctures
undulata (Say)
Lateral yellow area on pronotum narrower than humeral spot on elytron, often
only ‘ as wide (Fig. 440d); punctures on elytron slightly larger than on pronotum
octavia Casey
Species known only from Texas imitator, n. sp.
Species occurring from Arizona and Montana to the Pacific coast 113
Elytron with discal spot wedge-shaped, narrow lateral vitta present from hu-
meral angle onto apical '/s (Fig. 425e); Idaho, Montana, Utah, Washington . .
simulatrix Dobzhansky
Elytron usually with discal spot round or elongate, rarely wedge-shaped, rarely
with lateral vitta, pattern extremely variable (Fig. 426d-f); California, Nevada
quadrioculata (Motschulsky)
414
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Section I
The definition of this Section is mainly based on the 1 1 -segmented antennae, but
there are some other criteria that apply to nearly all species in the Section as follows:
body shape usually rounded, convex; epipleuron of elytron broad, usually with me-
dian groove, excavations for femoral apices very deep; base of abdominal sternum
within the postcoxal arc usually strongly depressed with transverse fold or suture
evident. There are 4 species in the bigeminata group in which the first abdominal
sternum is similar to that described for Section II; H. osculans, H. revocans, H.
esclavium, and H. thangulum. These species also have the elytral epipleurae narrower
and less deeply excavated than the other species in Section I. I include 3 species with
10-segmented antennae in Section I, H. rotunda, H. connectens, and H. dobzhanskyi,
n. sp. These species have the male genitalia characteristic of members of the bige-
minata group and fit the criteria for Section I as outlined above. I consider the 10-
segmented antennae of these 3 species to be an independent reduction without phy-
letic significance.
proba group
Body very convex, round; female pronotum with large yellow area laterally; male
genitalia with paramere broad at base, tapered to slender, rounded process in apical
Vi, apex with tuft of short setae (Fig. 334a); female spermathecal capsule with appendix
nearly as long as basal portion, or longer (Fig. 334d).
Hyperaspis proba (Say)
Fig. 334a-e; Map, Fig. 336
Coccinella proba Say, 1826, p. 303.
Hyperaspis proba: Mulsant, 1850, p. 674. — Crotch, 1873, p. 380.— Crotch, 1874b,
p. 235.-LeConte, 1880, p. 188. -Wickham, 1894, p. 304.-Casey 1899, p. 123.-
Blatchley, 1910, p. 523.-Leng. 1920, p. 2 1 1 . - Korschefsky, 1931, p. 194.-Win-
go, 1952, p. 25.
Hyperaspis proba proba: Dobzhansky, 1941, p. 22.
Hyperaspis proba var. t rinif er , 1899, p. 123. — Dobzhansky, 1941, p. 23.
Hyperaspis proba ab. trinifer: Korschefsky, 1931, p. 194.
Diagnosis. Length 2.0 to 3.0 mm; width 1.60 to 2.50 mm. Form rounded, convex.
Pronotum of male with anterior margin and broad lateral area yellow; pronotum of
female with anterior margin black and lateral yellow area smaller than in male. Elytron
with 3 yellow or red spots (Fig. 334e). Postcoxal line evenly curved, not quite reaching
posterior margin of first abdominal sternum, area within line smooth, nearly im-
punctate. Male genitalia as in Figure 334a-c. Female genitalia as in Figure 334d.
Discussion. The elytral color pattern of H. proba is very distinctive and apparently
not variable, making this species one that is easily recognized. The type of trinifer
Casey is a unique female (holotype).
Type locality. Of proba, not stated; of trinifer. Las Vegas, New Mexico.
Type depository. Of proba, type lost; of trinifer, USNM (35163).
Distribution. Figure 336. Maine to South Carolina, west to South Dakota and west
Texas.
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Fig. 334. Hyperaspis proba.
Hyperaspis weisei Schaeffer
Fig. 335a, b; Map, Fig. 336
Hyperaspis weisei Schaeffer, 1908, p. 126. — Leng, 1920, p. 212.— Korschefsky, 1931,
p. 199.
Hyperaspis proba weisei: Dobzhansky, 1941, p. 23.
Hyperaspis kunzii Schaeffer, 1905, p. 145 (not kunzii Mulsant, 1850).— Schaeffer,
1908, p. 127.
Diagnosis. Length 2.25 mm, width 1.85 mm. Form rounded, convex. Description
as for proba except elytron with marginal spot behind humeral callus, spot extending
forward toward anterolateral angle (Fig. 335a). Female genitalia as in Figure 335b.
Discussion. Dobzhansky placed this species as a subspecies of H. proba with some
reservations. I have examined the female genitalia and find that both the spermathecal
capsule and appendix differ considerably from those of H. proba. Therefore I am
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Fig. 335. Hyperaspis weisei.
confident that H. weisei is a valid species, but is almost certainly a member of the
proba group. Schaeffer had 2 type specimens of this species, both females, one of
which I here designate and label as the lectotype. No other specimens have been
examined.
Fig. 336. Distribution. Hyperaspis proba (shaded, disjunct locality dotted); H. weisei (open
circle); H. rivularis (star).
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NORTH AMERICAN COCCINELLIDAE
417
Type locality. Brownsville, Texas (lectotype here designated).
Type depository. USNM.
Distribution. Figure 336. TEXAS: Brownsville.
Hyperaspis rivularis Dobzhansky
Fig. 337a-d; Map, Fig. 336
Hyperaspis rivularis Dobzhansky, 1941, p. 35.— Wingo, 1952, p. 26.
Diagnosis. Length 2.40 to 3.0 mm, width 1.90 to 2.50 mm. Form rounded, convex.
Color pattern as described for H. proba except elytron with single yellow or orange
discal spot either round or elongate, often very large (Fig. 337d). Postcoxal line on
first abdominal sternum and female genitalia as described for H. proba. Male genitalia
as in Figure 337a-c.
Discussion. Dobzhansky (1941) did not examine the male genitalia of this species
and therefore placed it near H. bigeminata. The male genitalia are of the proba type,
but the female pronotal pattern is like that of many species in the bigeminata group.
Type locality. Frankfort, Kentucky.
Type depository. USNM (54205).
Distribution. Figure 336. ILLINOIS: “southern.” KENTUCKY: type locality. MIS-
SOURI: Cliffcave; St. Louis.
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Fig. 338. Hypemspis globula.
Hyperaspis globula Casey
Fig. 338a-e; Map, Fig. 336
Hyperaspis globula Casey, 1899, p. 124. — Leng, 1920, p. 211. — Korschefsky, 1931,
p. 189. — Dobzhansky, 1941, p. 24.
Diagnosis. Length 1 .80 to 2.0 mm, width 1 .40 to 1 .60 mm. Form rounded, convex.
Pronotum with large yellow area laterally, in male anterior border narrowly yellow,
black in female. Elytron with single yellow discal spot (Fig. 338e). Postcoxal line
similar to that of H. proba, area within line polished, finely punctate. Male genitalia
as in Figure 338a-c. Female genitalia as in Figure 338d.
Discussion. The paramere of the male genitalia is not as strongly modified in H.
globula as it is in H. proba and H. rivularis, but they are similar enough to indicate
a common origin. The female spermathecal capsule does not have the appendix
longer than the capsule proper, but it is nearly as long as the capsule and very robust;
both the capsule and appendix are definitely of the proba type. Hyperaspis globula
and H. oculifera resemble each other externally, but the elytral spots are located more
posteriorly and the length is greater in H. oculifera. There are 2 type specimens in
the Casey collection, the first of these (female) is here designated and labeled as the
lectotype, and the other (male) as a paralectotype.
Type locality. Brownsville, Texas (lectotype here designated).
Type depository. USNM (35172).
Distribution. Figure 336. TEXAS: Brownsville.
tucker i group
Body robust, broad, slightly flattened dorsoventrally; male genitalia similar to proba
group but with inner membrane of phallobase extending well out of phallobase (Fig.
339a); female spermathecal capsule with appendix much longer than basal portion.
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419
Fig. 339. Hypemspis tucked.
This is a compact group of 3 species which are closely related to species of the
proba group. Examples of these species are extremely rare in collections, their habits
apparently are such that they are not taken by the usual collecting methods.
Hyperaspis tucked Casey
Fig. 339a-d; Map, Fig. 340
Hyperaspis tucked Csisey, 1924, p. 162.— Korschefsky, 1931, p. 198.— Dobzhansky,
1941, p. 36.
Diagnosis. Length 3.0 mm, width 2.45 mm. Form robust, elongate, broad. Prono-
tum of male with lateral ^4 yellow and apical margin broadly yellow. Elytron with
wide, elongate, red spot (Fig. 339d). Postcoxal line on first abdominal sternum as in
H. proba except with some fine punctures. Male genitalia as in Figsure339a-c.
Discussion. The unique male type and one other specimen are all I have seen. This
is a striking species because of the large, red elytral spots and broadly yellow pronotal
margins, and does not closely resemble any other Arizona species (see remarks under
H. jovialis).
Type locality. Tucson, Arizona.
Type depository. USNM (35164).
Distribution. Figure 340. ARIZONA: type locality; Globe.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 340. Distribution. Hyperaspis tucked (open circle); H. jovialis (dot); H. leachi (square).
Hyperaspis jovialis Fall
Fig. 341a-f; Map, Fig. 340
Hyperaspis jovialis Fall, 1925, p. 3 1 1 . — Korschefsky, 1931, p. 190.— Dobzhansky,
1941, p. 80.
Hyperaspis californica Dobzhansky, 1941, p. 81. New Synonymy.
Hyperaspis taeniata perpallida Dobzhansky, 1941, p. 44. New Synonymy.
Diagnosis. Length 2.40 to 2.80 mm, width 1.70 to 2.0 mm. Form robust, elongate.
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NORTH AMERICAN COCCINELLIDAE
421
Fig. 341. Hyperaspis jovialis.
broad. Pronotum of male with lateral Va yellow and apical margin either broadly
yellow or with yellow median area (Fig. 341d-f); pronotum of female entirely black
except lateral margin narrowly yellow. Color pattern on elytron variable from black
with 2 orange spots (Fig. 34 le) to mostly orange with black border and enclosed
black spot (Fig. 34 Id). Postcoxal line not reaching hind margin of first abdominal
sternum, flattened along hind margin, outer V3 straight, area within line smooth, with
some vague punctures. Male genitalia as in Figure 341a-c. Female genitalia as figured
for H. leachi.
Discussion. This species, H. leachi, and H. tuckeri, are very similar in both external
and internal characteristics. On the basis of specimens examined, I presently regard
them all as valid species, but very few specimens exist in collections, and it is possible
that all of these names are synonyms. However, the extent of character variation
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 342. Hyperaspis leachi.
cannot be determined at present. El-Ali (unpubl. dissertation) considered H. cali-
fornica Dobzhansky a junior synonym of H. jovialis, an opinion with which I agree.
Hyperaspis taeniata perpallida Dobzhansky is a pale variant and junior synonym of
H. jovialis. The unique type (holotype) of jovialis is a female labeled “Kem Co. Cal./
Havilah VI-5-13/jovialis Type/M. C.Z. Type 24542/Fall collection.”
Type locality. Of jovialis, Havilah, Kem Co., California; of californica; Mount San
Jacinto, California; of perpallida, Sacramento, Co., Grand Island, California.
Type depository. Of jovialis, MCZ; of californica (54220) and perpallida (54206),
USNM.
Distribution. Figure 340. CALIFORNIA: Bishop; Claremont; Fresno Co; L. Ar-
rowhead; San Bernardino Co., Forest Home; S. Jacinto Mts.; Tulare Co.; Ventura
Co., Lockwood Valley; Yolo Co., Davis. NEVADA: Carlin. WASHINGTON: Soap
Lake.
Hyperaspis leachi Nunenmacher
Fig. 342a-e; Map, Fig. 340
Hyperaspis leachi '\^unQnm?ichQr, 1934, p. 19. — Dobzhansky, 1941, p. 31.
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NORTH AMERICAN COCCINELLIDAE
423
Diagnosis. Length 2.40 to 3.10 mm, width 1.80 to 2.60 mm. Description as for
H. jovialis except pronotum of male mostly black, lateral and narrow apical margin
yellow; female pronotum black except lateral margin narrowly yellow; elytron with
large, discrete orange spot nearly reaching lateral margin (Fig. 342e). Male genitalia
as in Figure 342a-c. Female genitalia as in Figure 342d.
Discussion. Hyperaspis leachi is very similar to H. tuckeri and H. jovialis (see
remarks under H. jovialis), but can be separated from either of those species by the
pronotal and elytral color pattern. Males of both H. jovialis and H. tuckeri have the
apical margin of the pronotum broadly yellow and the lateral ‘A yellow. Males of H.
leachi have the apical margin narrowly yellow and the lateral or less yellow. The
orange spot on the elytron nearly reaches the lateral margin in H. leachi, but is clearly
separate from the margin in both H. jovialis and H. tuckeri. Nunenmacher (1925)
designated 2 primary types (male and female). I here designated and label the male
labeled “Riverside Co. Cal. III-25-18 E. R. Leach Coll./male sign/ Hyperaspis leachi
Nun.” as the lectotype. The female is designated as a paralectotype.
Type locality. Riverside Co., California (lectotype here designated).
Type depository. CAS.
Distribution. Figure 340. CALIFORNIA: Inyo Co., Independence; Kem Co.; San
Bernardino Co., Hesperia; Jacumba; San Diego; Los Angeles Co.; Sonoma Co.; Tulare
Co., Isabella.
bi not at a group
Male genitalia with paramere slender, slightly narrowed toward apex, basal lobe
slender, nearly parallel-sided, apex obliquely truncate (Fig. 343a); female spermathe-
cal capsule rounded, appendix very small.
Hyperaspis inedita has the bigeminata pronotal color pattern in the female, but
the male genitalia are of the binotata type. All members of this group have the elytron
black with one or two red or yellow spots except H. octonotata which has 4 yellow
spots on each elytron but possesses the binotata type of genitalia.
Hyperaspis binotata (Say)
Fig. 343a-d; Map, Fig. 344
Coccinella binotata Say, 1826, p. 302.
Hyperaspis binotata: Crotch, 1873, p. 380.— Casey, 1899, p. 124. — Blatchley, 1910,
p. 523. — Leng, 1920, p. 21 1. — Korschefsky, 1931, p. 196.— Dobzhansky, 1941, p.
27.-Wingo, 1952, p. 25. -Watson, 1960, p. 232.-Watson, 1969, p. 370.-J.
Chapin, 1974, p. 41.
Coccinella normata Say, 1826, p. 302.
Hyperaspis normata Crotch, 1873, p. 380.
Coccinella Randall, 1838b, p. 50. — Mulsant, 1850, p. 1051.
Hyperaspis leucopsisMoishQimQv, 1847, p. 179.— Crotch, 1873, p. 380.
Hyperaspis paspalis Watson, 1960, p. 233.— Watson, 1969, p. 370. New Synonymy.
Diagnosis. Length 2.40 to 4.50 mm, width 1 .90 to 3.70 mm. Form rounded, convex.
Pronotum of male narrowly yellow on lateral margin, often narrowly yellow on
anterior margin; pronotum of female black. Elytron black with single red spot (Fig.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 343. Hyperaspis binotata.
343d), rarely with apical spot (see remarks under H. signata). Postcoxal line not
reaching hind margin of first abdominal sternum, briefly flattened along hind margin,
area within line distinctly punctate. Male genitalia as in Figure 343a-c.
Discussion. Hyperaspis binotata is a common, widespread species recognized with
certainty only if the male genitalia are examined. I have examined the holotype and
several paratypes of H. paspalis Watson, and am unable to separate H. paspalis from
H. binotata. The key character used by Watson to distinguish H. paspalis concerns
the prostemal carinae which are supposed to be parallel, not joining anteriorly in H.
paspalis, convergent and joined anteriorly in H. binotata. This character is variable
in any long series of H .binotata, and even some of the paratypes of H. paspalis have
convergent carinae. The male genitalia are also somewhat variable and those of H.
paspalis vary within the range exhibited by H. binotata. Watson (1960) illustrated
the male genitalia of H. paspalis in ventral view and those of H. binotata in dorsal
view which, because of the asymmetrical basal lobe, presents a somewhat confusing
picture.
Type locality. Of binotata and normata, not stated; of affinis, “vicinity of Boston”;
of leucopsis, Pennsylvania; of paspalis. Iron Bridge, Ontario.
Type depository. Of binotata and normata, types lost; of affinis, not located; of
leucopsis, not located; of paspalis, CNC.
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NORTH AMERICAN COCCINELLIDAE
425
Fig. 344. Distribution. Hyperaspis binotata (shaded); H. haematosticta (star); H. inedita
(dot); H. octonotata (open circle).
Distribution. Figure 344. Maine and Quebec to North Carolina, west to North
Dakota and Louisiana. Peripheral locality; DeFuniak Spring, Florida.
Hyperaspis haematosticta Fall
Fig. 345a-e; Map, Fig. 344
Hyperaspis haematosticta Fall, 1907, p. 222.— Leng, 1920, p. 211.— Korschefsky,
1931, p. 190. — Dobzhansky, 1941, p. 29.
Diagnosis. Length 2.70 to 3.80 mm, width 1 .90 to 2.70 mm. Form elongate, convex.
Pronotal color pattern as in H. binotata. Elytron with single red discal spot or with
discal spot plus apical spot (Fig. 345d, e). Postcoxal line reaching hind margin of
first abdominal sternum, flattened along margin, outer Vz abruptly angled forward,
area within line smooth with scattered coarse punctures. Male genitalia as in Figure
345a-€.
Discussion. The combination of elongate body and elytral color pattern makes H.
haematosticta a reasonably distinctive species in the geographic region in which it
occurs. Fall had 3 types of this species but I have seen only a single female type
labeled “Santa Fe N.M. 8.97/haematosticta TYPE/head, front and sides of tho. pale/
M.C.Z. Type 24541 (red paper)/H.C. Fall Collection/Hyperaspis haematosticta Fall”
which I designate and label the lectotype.
Type locality. Santa Fe, New Mexico (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 344. ARIZONA: Chiricahua Mts.; Williams. NEW MEXICO:
Las Vegas.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 345. Hyperaspis haernatosticta.
Hyperaspis inedita Mulsant
Fig. 346a-d; Map, Fig. 344
Hyperaspis inedita Mulsant, 1850, p. 684.— Crotch, 1873, p. 380. — Crotch, 1874b,
p. 238. -Casey, 1899, p. 124.-Leng, 1920, p. 21 l.-Korschefsky, 1931, p. 190.-
Dobzhansky, 1941, p. 28.
Hyperaspis regalis Casey, 1899, p. 123. — Leng, 1920, p. 21 1.— Korschefsky, 1931,
p. 195. — Dobzhansky, 1941, p. 31. New Synonymy.
Hyperaspis nigropennis B\di\ch\Qy, 1924, p. 167. — Korschefsky, 1931, p. 192.— Dob-
zhansky, 1941, p. 84. New Synonymy.
Hyperaspis pinorum Casey, 1924, p. 162. — Korschefsky, 1931, p. 194.— Dobzhan-
sky, 1941, p. 28.— J. Chapin, 1974, p. 43. New Synonymy.
Hyperaspis centralis plagiata Dobzhansky, 1941, p. 34. New Synonymy.
Diagnosis. Length 2.65 to 3.0 mm, width 1.90 to 2.30 mm. Form elongate oval,
convex. Pronotum of male with anterior margin and broad lateral area yellow; prono-
tum of female with anterior margin black, broad lateral area yellow. Elytron with
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427
Fig. 346. Hyperaspis inedit a.
single red spot of variable size, rarely immaculate (Fig. 346d). Postcoxal line reaching
hind margin of first abdominal sternum, flattened along margin, then evenly curved
forward in outer 1/3, area within line smooth with scattered, coarse punctures. Male
genitalia as in Figure 346a-c.
Discussion. The broadly yellow pronotal margin of the female causes this species
to resemble members of the bigeminata and proba groups. The elongate shape,
pronotal color pattern, and geographic distribution will usually allow H. inedita to
be recognized, but genitalia should be examined when males are available. A single
female of H. inedita exists in the Dejean collection labeled “Amer. bor. LeConte,”
designated a lectotype by Gordon (1974c). Hyperaspis pinorum Casey is a junior
synonym of inedita as suggested by Dobzhansky (1941). There are 4 types of pinorum
in the Casey collection; and I here designate and label the first of these (male) as the
lectotype and the remainder as paralectotypes. The holotype of H. nigropennis is a
female lacking elytral spots. Rarely does a specimen of H. inedita lack these spots,
but in all other characteristics H. nigropennis and H. inedita appear to be conspecific,
therefore I consider H. nigropennis to be a junior synonym. Hyperaspis regalis is an
example of H. inedita with greatly enlarged elytral spots. The type is a unique female
(holotype) in the Casey collection.
Type locality. Of inedita, “L’Amerique septentrionale” (lectotype here designated);
of regalis, Jacksonville, Florida; of pinorum. Southern Pines, North Carolina (lec-
totype here designated); of nigropennis, Dunedin, Florida; of centralis plagiata, 2.3
miles east of Piney Point, Maryland.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 347. Hyperaspis octonotata.
Type depository. Of inedita, DLM; of regalis (35165), pinorum (35173), and cen-
tralis plagiata (54204), USNM; of nigropennis, PU.
Distribution. Figure 344. FLORIDA: Gainesville; Lake City; Navarre; Perry.
GEORGIA: Bamsville; Milner. LOUISIANA: Baton Rouge; Ouachita Parish; St.
Tammany Parish. MISSOURI: Jackson Co., Lee Summit. NORTH CAROLINA:
Southern Pines.
Hyperaspis octonotata Casey
Fig. 347a-d; Map, Fig. 344
Hyperaspis 8 -not at a Casey, 1899, p. 121.
Hyperaspis octonatata: Leng, 1920, p, 2 1 1 . — Korschefsky, 1931, p. 193.— Dob-
zhansky, 1941, p. 7.
Diagnosis. Length 2.30 to 3.50 mm, width 1 .90 to 2.80 mm. Form rounded, convex.
Pronotum of male with anterior margin and broad lateral area yellow; pronotum of
female with anterior margin black, lateral area broadly yellow. Elytron with 4 yellow
or red spots (Fig. 347d), anterior margin of discal spot obliquely truncate or emar-
ginate. Postcoxal line not reaching hind margin of first abdominal sternum, evenly
curved throughout. Male genitalia as in Figure 347a-c.
Discussion. The color pattern alone is sufficient for recognition of H. octonotata,
the arrangement of the elytral spots and the shape of the discal spot are unique in
the North American fauna. The type in the Casey collection is a unique female
(holotype).
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NORTH AMERICAN COCCINELLIDAE
429
Type locality. Arizona.
Type depository. USNM (35148).
Distribution. Figure 344. ARIZONA: Benson; Cochise Co., Palmerly; Douglas;
Graham Co.; Huachucha Mts.; Sta. Catalina Mts.; Tucson; Thatcher; Wilcox. CAL-
IFORNIA: Ft. Tejon; Lebec; Los Gatos; Palo Alto; Redwood City; Santa Barbara;
Santa Maria; Sequoia Nat. Pk.; Tulare Co. COLORADO: Canon City. TEXAS:
Brownsville; Presidio; Sanderson. UTAH: St. George.
signata group
Female pronotum always entirely black; male genitalia with paramere slender,
slightly narrowed toward apex, basal lobe narrow at base, wide at apex, apex truncate
or obliquely truncate (Fig. 348a); female spermathecal capsule rounded, appendix
very small (Fig. 348d).
This group contains those members of Hyperaspis most difficult to separate from
each other; in most cases male genitalia must be examined to correctly identify a
species. The shape is mostly oval and convex; the dorsal color is black with one or
2 red or yellow spots on each elytron; adequate external structural characters are
lacking.
Hyperaspis signata signata (Olivier)
Fig. 348a-f; Map, Fig. 349
Coccinella signata Olivier, 1808, p. 1047.
Hyperaspis signata Mulsant, 1850, p. 683.— Crotch, 1873, p. 380. — Crotch, 1874b,
p. 234. — LeConte, 1880, p. 187. — Casey, 1899, p. 122.— Leng, 1920, p. 211.—
Korschefsky, 1931, p. 196. — Dobzhansky, 1941, p. 28.— Wingo, 1952, p. 25.— J.
Chapin, 1974, p. 41.
Hyperaspis taedata hcConiQ, 1880, p. 187. — Leng, 1920, p. 212. — Korschefsky, 1931,
p. 197.— Dobzhansky, 1941, p. 20. New Synonymy.
Diagnosis. Length 2.60 to 4.0 mm, width 1.90 to 3.20 mm. Form oval, convex.
Pronotum of male with anterior and lateral margins narrowly yellow. Elytron with
one or 2 yellow or red spots (Fig. 348e, f), rarely with discal spot enlarged to humeral
angle. Postcoxal line not reaching hind margin of first abdominal sternum, evenly
curved except outer Va straight, area within line polished with scattered, coarse punc-
tures. Male genitalia as in Figure 348a-c. Female genitalia as in Figure 348d.
Discussion. This species is very similar to H. pistillata Watson. The male genitalia
are quite distinctive for both species, in addition, the distal capsule of the femle
spermathecae differ where the connecting ducts enter. In H. signata the connecting
duct and process of the capsule merge smoothly while in H. pistillata the duct is
noticeably more slender than the process of the capsule. Hyperaspis signata is a
common eastern species which has undoubtedly been mixed with several other species
in collections of any size for the past 100 years. I have not seen the Olivier type
material and therefore am not certain of the exact identity of H. signata. However,
I am arbitrarily assigning the name H. signata to the species described here as did
Dobzhansky (1941). The type series of H. taedata consists of 2 specimens, a female
with the typical signata color pattern labeled “Baldwin June 1, Fla/963/Type 671 1/
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 348. Hyperaspis signata signata.
H. taedata LeC,” and a male with an odd color pattern which I designate the lectotype
labeled “/Sand Pt. Fla./ 18.2/964.” The color pattern of the lectotype is very unusual
for this species. A unique male in the USNM collection, also from Florida, has an
identical color pattern but is a specimen of H. binotatal In both instances the male
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NORTH AMERICAN COCCINELLIDAE
431
Fig. 349. Distribution. Hyperaspis signata signata (shaded, disjunct localities dotted); H. s.
bicentralis-, (star).
genitalia are the criteria for placement, and I consider H. taedata a junior synonym
of H. signata.
Type locality. Of signata, “Elle se trouve la Caroline”; of taedata, Sand Point,
Florida (lectotype here designated).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 350. Hypemspis signal a bicentralis.
Type depository. Of signata, type not seen; of taedata, MCZ.
Distribution. Figure 349. Massachusetts to Florida, west to Wisconsin and east
Texas. Peripheral localities; Beaver Dam and Marinette Co., Wisconsin.
Hyperaspis signata bicentralis Casey, new status
Fig. 350a-e; Map, Fig. 349
Hyperaspis bicentralis CdiSty, 1899, p. 124. — Korschefsky, 1931, p. 184.
Hyperaspis bicentralis bicentralis: Dobzhansky, 1941, p. 32.
Diagnosis. Length 2.60 to 3.25 mm, width 2.20 to 2.70 mm. Description as for H.
signata except average size smaller; discal spot on elytron large, apical spot lacking
(Fig. 350e); basal lobe of male genitalia longer and narrower (Fig. 350a). Female
genitalia as in Figure 350d.
Discussion. In view of the differences between H. signata and H. bicentralis listed
above, I prefer to maintain H. bicentralis as a subspecies rather than a synonym of
H. signata. Two specimens from College Station, Texas, have been examined which
are intermediate between H. signata and H. bicentralis, therefore I regard H. bicen-
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433
Fig. 351. Hyperaspis pistillata.
trails as a well characterized subspecies of H. signata. The type of H. bicentralis is
a unique female in the Casey collection (holotype).
Type locality. Colorado River above Columbus, Austin, Texas.
Type depository. USNM (35170).
Distribution. Figure 349. OKLAHOMA: Mountain Pk. TEXAS: Austin; Dallas;
Kerrville; Lavaca Co.; Otey; Paris; Victoria.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Hyperaspis pistil lata Watson
Fig. 351a-e; Map, Fig. 352
Hyperaspis pistillata Watson, 1969, p. 369.
Diagnosis. Length 2.75 to 4.0 mm, width 2.20 to 3.10 mm. Form oval, convex.
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NORTH AMERICAN COCCINELLIDAE
435
Fig. 353. Hyperaspis nigrosuturalis.
Pronotum of male with anterior and lateral margins yellow; pronotum of female
entirely black. Elytron with one or 2 yellow or red spots (Fig. 35 le), as in H. signata.
Postcoxal line not reaching hind margin of first abdominal sternum, more or less
evenly curved, area within line smooth, coarsely punctured except depressed apical
portion strongly alutaceous. Male genitalia as in Figure 351a-c. Female genitalia as
in Figure 35 Id.
Discussion. This species closely resembles H. signata\ see comparative remarks
under H. signata.
Type locality. Dunedin, Florida.
Type depository. UMMZ.
Distribution. Figure 352. Massachusetts to Florida and Louisiana.
Hyperaspis nigrosuturalis Blatchley
Fig. 353a-d; Map, Fig. 352
Hyperaspis nigrosuturalis BXdtichXty, 1918, p. 420. — Leng, 1920, p. 212.— Blatchley,
1930, p. 43.— Korschefsky, 1931, p. 192. — Dobzhansky, 1941, p. 32.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 354. Hypemspis conviva.
Diagnosis. Length 3.40 mm, width 2.70 mm. Form oval, convex. Head of male
yellow except vertex black, entirely black in female. Pronotum of male with lateral
margin narrowly yellow; pronotum of female entirely black. Elytron red except mar-
gins narrowly black (Fig. 353d). Postcoxal line not reaching hind margin of first
abdominal sternum, evenly curved except outer ‘/a slightly angulate, area within line
rough, coarsely, densely punctured. Male genitalia as in Figure 353a-c. Female gen-
italia as figured for H. pistillata.
Discussion. I have seen only the holotype and one other specimen of this distinctive
species. The male genitalia are very similar to those of H. pistillata, but H. nigro-
suturalis would not normally be associated with H. pistillata because the dorsal color
patterns are so different.
Type locality. Lakeland, Florida.
Type depository. PU.
Distribution. Figure 352. FLORIDA: Lake Alfred.
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NORTH AMERICAN COCCINELLIDAE
437
Hyperaspis conviva Casey
Fig. 354a-e; Map, Fig. 355
Hyperaspis conviva CasQy, 1924, p. 163. — Korschefsky, 1931, p. 186.
Hyperaspis insolens CdiSQy , 1924, p. 164. — Korschefsky, 1931, p. 190.
Hyperaspis binotata: Dobzhansky, 1941, p. 27 (in part).
Hyperaspis congressis Watson, 1960, p. 231.— Watson, 1969, p. 370. J. Chapin, 1974,
p. 42. New Synonymy.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Diagnosis. Length 2.70 to 3.80 mm, width 2.0 to 2.60 mm. Form elongate, oval,
usually somewhat parallel sided and dorsoventrally flattened, particularly in female.
Color pattern as in H. signata (Fig. 354d, e). Postcoxal line not reaching hind margin
of first abdominal sternum, somewhat flattened along margin. Male genitalia as in
Figure 354a-c. Female genitalia as described for H. signata.
Discussion. This is the most easily recognized species in the signata group because
of the dorsoventrally flattened, elongate form which is characteristic of most females
and many of the males. The male genitalia are very peculiar because the asymmetry
is reversed, presenting nearly a mirror image of the H. signata genitalia. The genitalia
are also reversed in the abdomen, lying on the right side of the abdomen (in ventral
view from apex) rather than the left side as they do in all other species of Hyperaspis
that I have seen. Hyperaspis conviva is extremely widespread, and partially because
of this, has been named 3 times. Dobzhansky (1941) placed H. conviva Casey and
H. insolens Casey as junior synonyms of H. binotata. Watson (1960) described H.
congressis as a new species because he had not seen the type of either H. insolens or
H. conviva; I here place H. congressis as a junior synonym of H. conviva. The types
of both conviva (female) and insolens (male) are uniques (holotypes) in the Casey
collection.
Type locality. Of conviva. Southern Pines, North Carolina; of insolens. Grayling,
near Bay City, Michigan; of congressis, Savanne, Ontario.
Type depository. Of conviva (35174) and insolens (35176), USNM; of congressis,
CNC.
Distribution. Figure 355. MANITOBA: Beausejour; Fairford; Pine Falls; Reynolds;
Sandilands Forest Reserve; Stead; Victoria. ONTARIO: Agawa; Fort William; Ger-
man; Gogama; Hawk Lake; Lost Bay; McIntosh; Savanne; Walford. SASKATCH-
EWAN: Holbein; Hudson Bay; Prince Albert. ALABAMA: Mobile. DISTRICT OF
COLUMBIA: Washington; Woodridge. FLORIDA: Pensacola; Tallahassee. LOUI-
SIANA: East Baton Rouge Parish; Ouachita Parish; West Feliciana Parish. MAINE:
Mt. Katahdin. MARYLAND: Beltsville; Priest Br. MICHIGAN: Bay City; Roscom-
mon. NEW JERSEY: Clementon; Lakehurst; Pemberton; Riverton; Warren Co.;
Westville. NEW YORK: Brooklyn; Top of Mt. Whiteface; Seneca Co., Willard.
NORTH CAROLINA: Tryon. PENNSYLVANIA: Harrisburg. VIRGINIA: Gum
Spring; Rosslyn; Wallops Id. WEST VIRGINIA: White Sulphur Springs.
bigeminata group
Male genitalia with paramere broad, almost spoon-shaped, basal lobe deeply con-
cave on one side, angulate on other side (Fig. 356a); female spermathecal capsule
with appendix short (Fig. 357d).
The bulk of the species in Section I belong to this group which is well characterized
by the type of male genitalia.
Hyperaspis gemina LeConte
Fig. 356a-d; Map, Fig. 358
Hyperaspis gemina LeConte, 1880, p. 188. — Casey, 1899, p. 128. — Leng, 1920, p.
212. — Korschefsky, 1931, p. 189. — Dobzhansky, 1941, p. 37.
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NORTH AMERICAN COCCINELLIDAE
439
Fig. 356. Hyperaspis gemina.
Diagnosis. Length 2.80 to 4.0 mm, width 2.10 to 3.0 mm. Form oval, convex.
Apical margin of clypeus truncate. Head yellow, pronotum with broad lateral area
yellow in both sexes. Elytron with 2 narrowly connected yellow spots at apex (Fig.
356d). Postcoxal line evenly curved, except outer Vs slightly angulate, nearly reaching
hind margin of first abdominal sternum, area within line smooth, impunctate. Male
genitalia as in Figure 356a-c.
Discussion. This rarely collected species is unusual in having the clypeal apex
truncate and the head yellow in both sexes. There are 2 types of H. gemina in the
LeConte collection, one of these labeled “Ga./Type 67 1 3(red paper)///, gemina Lec.’’\
I here designate and label as the lectotype. The other specimen labeled “Tex.” is
designated and labeled as a paralectotype.
Type locality. Georgia (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 358. NORTH CAROLINA: Bell Island; Wenona. SOUTH
CAROLINA: Myrtle Beach. VIRGINIA: Cape Henry; Ft. Monroe.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93( 1 )
Fig. 357. Hyper aspis bigeminata.
Hyperaspis bigeminata (Randall)
Fig. 357a-e; Map, Fig. 358
Coccinella bigeminata Randall, 1838a, p. 32. — Mulsant, 1850, p. 1050.
Hyperaspis bigeminata heContc, 1852, p. 135.— Crotch, 1873, p. 380.— Crotch, 1874b,
p.234.-LeConte, 1880, p. 188. -Wickham, 1894, p. 304.-Casey, 1899, p. 122.-
Blatchley, 1910, p. 523. — Korschefsky, 1931, p. 185. — Dobzhansky, 1941, p. 36.—
Wingo, 1952, p. 26. -J. Chapin, 1974, p. 43.
Hyperaspis guexi Mulsant, 1850, p. 687.— Crotch, 1873, p. 380.
Diagnosis. Length 2.40 to 3.35 mm, width 2.0 to 2.70 mm. Form oval, convex.
Pronotum of male with anterior margin and broad lateral area yellow; pronotum of
female with anterior margin black, broad lateral area yellow. Elytron with single
yellow or red apical spot (Fig. 357e). Postcoxal line reaching hind margin of first
abdominal sternum, evenly curved, area within line smooth, distinctly punctured.
Male genitalia as in Figure 357a-c. Female genitalia as in Figure 357d.
Discussion. The apical position of the spot on the elytron is usually sufficient to
distinguish H. bigeminata from similar appearing species of Hyperaspis. A single
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NORTH AMERICAN COCCINELLIDAE
441
Fig. 358. Distribution. Hypemspis gemina (circled dot); H. bigeminata (shaded; H. wick-
hami (circled star); H. major (star).
male type of H. guexi exists in the Dejean collection labeled “Ameri. bor., LeConte.”
I here designate and label this specimen as the lectotype. The type of H. bigeminata
has not been located and may be either lost or not recognizable.
Type locality. Of bigeminata. Blue Mountains, Maine; of guexi, “Ameri. bor.”
(lectotype were designated).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 359. Hyperaspis wickhami.
Type depository. Of bigeminata, not located; of guexi, DLM.
Distribution. Figure 358. Maine to Florida, west to Michigan and east Texas.
Hyperaspis wickhami Casey
Fig. 359a-obz\\?ins\iy, 1941, p. 72. — Belicek, 1976, p. 315.
Diagnosis. Length 2.20 to 2.70 mm, width 1.60 to 1.85 mm. Form elongate, oval.
Elytron with yellow marginal vitta from base to apical 3/5, apical spot transversely
oval, discal spot elongate, angulate (Fig. 425e). Postcoxal line extending to hind
margin of hrst abdominal sternum, flattened along margin, outer ‘/a angulate, area
within line alutaceous, indistinctly punctured. Male genitalia as in Figure 425a-c.
Female genitalia as in Figure 425d.
Discussion. The dorsal color pattern of H. simulatrix is similar to that of H.
quadrioculata ifidelis form) and H. spiculinota, both of which have allopatric distri-
butions. The male genitalia are practically identical to those of H. postica, but the
female genitalia of the 2 species are distinctive for each, and H. postica never has a
marginal vitta or an apical spot on the elytron.
Type locality. Oakley, Idaho.
Type depository. USNM (54216).
Distribution. Figure 424. OREGON: Harney Co., Tencent Lake. WASHINGTON:
Smyrna.
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NORTH AMERICAN COCCINELLIDAE
521
Hyperaspis quadrioculata (Motschulsky)
Fig. 426a-f; Map, Fig. 427
Exochomus quadrioculatus Motschulsky, 1845b, p. 383.
Hyperaspis undulata var. 4-oculata\ Crotch, 1873, p. 381.
Hyperaspis quadrioculata: Crotch, 1874b, p. 231. — LeConte, 1880, p. 188.— Casey,
1899, p. 128.-Korschefsky, 1931, p. 194.
Hyperaspis quadrioculata quadrioculata: Dobzhansky, 1941, p. 68.
Hyperaspis notatula Casey, 1899, p. 121.— Casey, 1908, p. 413. — Leng, 1920, p.
212. — Korschefsky, 1931, p. 193. New Synonymy.
Hyperaspis quadrioculata notatula: Dobzhansky, 1941, p. 69.
Hyperaspis fidelis CdiSQy , 1908, p. 418.— Leng, 1920, p. 212. — Korschefsky, 1931, p.
188. New Synonymy.
Hyperaspis quadrioculata scotti Dobzhansky, 1941, p. 70. New Synonymy.
Hyperaspis quadrioculata fidelis: Dobzhansky, 1941, p. 70.
Diagnosis. Length 2.70 to 4.0 mm, width 1.50 to 2.25 mm. Form elongate, oval.
Color pattern on elytron variable as in Figure 426d-f. Postcoxal line not reaching
first abdominal sternum, evenly curved throughout, area within line alutaceous,
indistinctly punctured. Male genitalia as in Figure 426a-c.
Discussion. The variable color pattern of this species has caused several names to
be proposed, either as species, or subspecies of H. quadrioculata. The name fidelis
Casey was proposed for the color pattern in Figure 426e, which intergrades with the
pattern of notatula (Figure 426d) frequently and at so many localities that it is
apparent that fidelis must be a junior synonym. The patterns of typical quadrioculata
(Fig. 426f) and notatula do not intergrade as frequently nor at as many localities, but
the distribution of both forms and their intergrades indicates that they cannot be
maintained as subspecies, therefore notatula becomes a synonym of quadrioculata.
The form called quadrioculata scotti by Dobzhansky I also regard as a junior synonym
of quadrioculata. Most of the localities from which I have seen scotti are also localities
where ""notatula'' occurs, and the scotti color pattern intergrades with that of notatula
in most instances. There are also other unnamed color forms such as in a series from
Fort Tejon, California, which has the typical ""notatula" pattern except that the discal
spot on the elytron is missing in most of the specimens, however, 3 of these specimens
show distinct traces of the discal spot. A series from Mill Creek, San Bernardino
Mts., has most examples lacking the humeral spot of "" notatula" , and were identified
as triangulum Casey, but 4 examples have a small humeral spot present (this is the
intermediate form discussed by El-Ali below). El-Ali (unpubl. dissertation) hybridized
the forms discussed here and found that crosses attempted produced progeny. His
results are excerpted as follows:
“1) the quadrioculata form had an F generation which were all quadrioculata-like
with regard to elytral spots. In subsequent generations the same results were obtained.
2) the scotti form had an F generation which were all scotti-\ike in appearance.
Most progeny had the humeral spot reduced or disappeared. Marginal and discal
spots enlarged and merged together. Subsequent generations gave only this scotti-
form.
3) the notatula form had an F generation of different elytral patterns: the largest
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 426. Hyperaspis quadrioculata.
number of individuals and the notatula-Mke elytral pattern, followed by the individ-
uals that had the intermediate form of elytral pattern, and a few individuals were
like the quadrioculata-iorm. Some of the notatula-Xike forms and the intermediate
forms had the discal and marginal spots large, but not connected. Selection of these
types and crossing them manifested in their progeny a few individuals that were
scotti-\\kQ and some individuals showed maculations of the previously mentioned
forms of the F generation.
4) the intermediate form of the F generation produced individuals having the
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NORTH AMERICAN COCCINELLIDAE
523
Fig. 427. Distribution. Hyperaspis quadrioculata (shaded, disjunct locality dotted).
patterns of notatula, quadrioculata, scotti and some of the intermediate elytral form.
The notatula and the intermediate forms were the most numerous.”
The type specimens of fidelis (female) and notatula (male) are unique (holotypes)
in the Casey collection. The type of quadrioculata has not been located.
Type locality. Of quadrioculata, type not examined; of notatula, Reno, Nevada; of
fidelis, Los Angeles, California; of scotti, San Joaquin Co., California.
Type depository. Of quadrioculata, not located; of notatula (35209), and fidelis
(35199), USNM; of scotti, CAS.
Distribution. Figure 427. CALIFORNIA: central and southern.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 428. Hyperaspis annexa. ■
Hyperaspis annexa LeConte
Fig. 428a-d; Map, Fig. 430
Hyperaspis annexa l^tConXQ, 1852, p. 133. — LeConte, 1880, p. 188.— Crotch, 1873,
p. 381. -Crotch, 1874b, p. 232. -Casey, 1899, p. 128.-Leng, 1920, p. 212.-
Korschefsky, 1931, p. 184. — Dobzhansky, 1941, p. 74. Wingo, 1952, p. 26.
Diagnosis. Length 2.0 to 2.75 mm, width 1.40 to 2.0 mm. Form elongate, oval,
flattened dorsoventrally. Male pronotum usually widely yellow on anterior margin,
but often narrowly yellow, occasional black. Elytron black with 2 yellow vittae on
disc and lateral margin (Fig. 428d). Postcoxal line reaching hind margin of first
abdominal sternum, flattened along margin, outer */3 angulate, area within line alu-
taceous, indistinctly punctured. Male genitalia as in Figure 428a-c.
Discussion. The distinctive vittate appearance of most examples of H. annexa
makes it one of the most easily recognized species of Hyperaspis. However, I suspect
that H. annexa and H. quadrioculata may not be specifically different. The male
genitalia of H. annexa are essentially identical to those of H. quadrioculata, and I
have seen intergrading color patterns in 2 instances. The lectotype of H. annexa has
the apical margin of the pronotum entirely black, the only instance of this observed,
all other males having a distinct, usually broad, yellow apical margin. A series from
Mill Creek, San Bernardino Mts., exhibits all degrees of intergradation of the elytral
pattern from the annexa type to the quadrioculata type (notatula form). Thus far
these are the only 2 obvious instances of apparent intergradation, therefore I prefer
to consider H. annexa a valid species for the present, but point out the distinct
possibility it may be synonymous with H. quadrioculata. Hyperaspis annexa was
associated in a group with H. quadrivitta by Dobzhansky, however they have little
in common except a vittate dorsal appearance. LeConte (1852) states that he had a
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NORTH AMERICAN COCCINELLIDAE
525
pair of types, and there are 2 specimens now in his collection. I here designate and
label the male labeled “gold disc/male sign/Type 6718(red paper)H. annexa Lee.”
as the lectotype, and the female bearing only a gold disc as the paralectotype.
Type locality. San Francisco, California (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 430. CALIFORNIA: Cajon Pass; Compton; Fillmore; Kem
Co., Tehachapi Pass; La Canada; Mt. Wilson; Paraiso Hot Springs; Pasadena; Riv-
erside; San Bernardino; San Diego Co.; San Francisco Co.; San Mateo Co.; Santa
Maria; Santa Paula; Tulare Co., Isabella.
Hyperaspis arizonica Dobzhansky
Fig. 429a-d; Map, Fig. 430
Hyperaspis biornata arizonica Dobzhansky, 1941, p. 53.
Diagnosis. Length 2.30 to 3.0 mm, width 1.70 to 2.20 mm. Form elongate, oval.
Elytron black with large orange spot on outer margin in posterior Vi (Fig. 429d).
Postcoxal line not reaching hind margin of first abdominal sternum, evenly curved
throughout, area within line alutaceous, densely, coarsely punctured. Male genitalia
as in Figure 429a-c.
Discussion. The single large spot occupying most of the posterior 1/2 of an elytron
makes H. arizonica an easily recognizable species. This species may resemble some
color forms of H. taeniata, but females of the latter species have entirely black
pronota.
Type locality. Bright Angel, Arizona (Grand Canyon Nat. Park).
Type depository. USNM (54211).
Distribution. Figure 430. ARIZONA: Bright Angel; South Rim Grand Canyon.
CALIFORNIA: Bishop.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 430. Distribution. Hyperaspis annexa (dot); H. arizonica (star); H. oculaticauda (open
circle); //. oregona (square); H. immaculata (circled star); H. querquesi (open star).
Hyperaspis oculaticauda Casey
Fig. 431a-d; Map, Fig. 430
Hyperaspis oculaticauda Casey, 1899, p. 127. — Leng, 1920, p. 212.— Korschefsky,
1931, p. 193. — Dobzhansky, 1941, p. 41. — Hatch, 1961, p. 160.
Hyperaspis ejficta Casey, 1899, p. 127. — Leng, 1920, p. 2 12. — Korschefsky, 1931, p.
1 88. — Dobzhansky, 1941, p. 41. — Hatch, 1961, p. 160. New Synonymy.
Hyperaspis suhdcprcssa Casey, 1899, p. 127. — Leng, 1920, p. 212.— Korschefsky,
1931, p. 197. — Dobzhansky, 1941, p. 42. New Synonymy.
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NORTH AMERICAN COCCINELLIDAE
527
Diagnosis. Length 1.80 to 2.40 mm, width 1.40 to 1.80 mm. Form elongate, oval.
Elytron black with single, apical spot (Fig. 43 Id). Postcoxal line as described for H.
postica. Male genitalia as in Figure 431a-c.
Discussion. This species is a minature edition of H. postica, having the same dorsal
color pattern and same general type of male genitalia. It is much smaller than H.
postica, the maximum observed length of H. oculaticauda being 2.40 mm, the min-
imum observed length of H. postica being 2.30 mm. The basal lobe of the male
genitalia of H. oculaticauda is laterally emarginate in the apical ‘/2, the basal lobe in
H. postica is emarginate in apical % or %. Hypemspis effeta Casey is a junior synonym
of H. oculaticauda, the original description was based on a teneral male. I also regard
H. subdepressa as a junior synonym of H. oculaticauda although the elytral punctation
is slightly coarser than typical H. oculaticauda. The type series of H. oculaticauda is
composed of a male and 4 females. I here designate and label the male as the lectotype,
and the females as paralectotypes. The type of H. effeta (male) and the type of H.
subdepressa (female) one uniques (holotypes).
Type locality. Of oculaticauda, Humboldt or Siskiyou Co., California (which county
not stated) (lectotype here designated); of effeta. Placer Co., California, of subdepressa,
Alameda Co., California.
Type depository. Of oculaticauda (35197), effeta (35191), and subdepressa (35 1 90),
USNM.
Distribution. Figure 430. CALIFORNIA: Alameda Co., Livermore, Bay Farm Is.;
Cisco; Eldorado Co., Eldorado; Lassen Co., Hallelujah Junction; Modoc Co., Lake
City; Marin Co., Lagunitas; Monterey Co., Carmel; Nevada Co., Boca; Pleasanton;
Sacramento; Shasta Springs; Sierra Co., Sierraville; Trinity Co., Carrville; Tuolumne
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 432. Hyperaspis immaculata.
Co., Chipmunk Flat, Strawberry; Yreka. NEVADA: Reno; Washoe Co., Pyramid.
OREGON: Baker Creek; Curry Co., Cape Sebastian St. Pk.; Jackson Co., Siskiyou
Summit; Klamath Falls; Geary Ranch; Lakeview; McMinnville; Woods.
Hyperaspis immaculata Hatch
Fig. 432; Map, Fig.430
Hyperaspis immaculata Hatch, 1961, p. 161.
Diagnosis. Length 2.0 mm. Form elongate, oval. Elytron entirely black. Pronotum
narrowly yellow in both sexes. Postcoxal line not reaching hind margin of first ab-
dominal sternum, evenly curved throughout, area within line alutaceous, impunctate
or with fine, nearly imperceptible punctures. Male genitalia as in Figure 432 (sipho
lost).
Discussion. The only species with entirely black elytra with which H. immaculata
could be confused is H. pleuralis which is not known to occur in western Washington
and Oregon where H. immaculata is found. In addition, the body form of H. pleuralis
is more rounded and convex than that of H. immaculata.
Type locality. Olympic Hot Springs, Washington.
Type depository. USNM.
Distribution. Figure 430. OREGON: Alston Mt.; McMinnville.
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NORTH AMERICAN COCCINELLIDAE
529
Fig. 433. Hyperaspis querquesi.
Hyperaspis querquesi Nutting
Fig. 433a-d; Map, Fig. 430
Hyperaspis querquesi Nutting, 1980, p. 263.
Diagnosis. Length 2.55 to 2.70 mm, width 1.60 to 2.0 mm. Form elongate, oval.
Elytron black with large, apical yellow spot and small lateral spot on margin behind
middle (Fig. 433d). Postcoxal line not reaching hind margin of first abdominal ster-
num, evenly curved throughout, area within line alutaceous, impunctate or with fine,
nearly imperceptible punctures. Male genitalia as in Figure 433a-c.
Discussion. The elytral color pattern is like that of H. postica except for the small
lateral spot, and like that of the form of H. quadrioculata having the lateral spot
except for the presence of a discal spot. The male genitalia of H. querquesi are unlike
either of these species because the lateral emargination of the basal lobe is in the
apical 1/2, as in H. oculaticauda.
Type locality. Bird Observation Station, Marin Co., California.
Type depository. CAS.
Distribution. Figure 430. CALIFORNIA: Mendocino Co., Inglenook Fen.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 434. Hyperaspis oregona.
Hyperaspis oregona Dobzhansky
Fig. 434a-f; Map, Fig. 430
Hyperaspis oregona'Dobzh^insky, 1941, p. 76. — Hatch, 1961, p. 157. — Belicek, 1976,
p. 314.
Hyperaspis lanei Hatch, 1961, p. 159. New Synonymy.
Diagnosis. Length 2.0 to 2.60 mm, width 1.50 to 1.78 mm. Form elongate, oval.
Elytron black with oblique discal vitta, transverse apical spot, and narrow marginal
vitta extending from base to apical ^4 (Fig. 4341), maculation often reduced to remnant
of discal vitta or loss of discal vitta (Fig. 434d, e). Postcoxal line not reaching hind
margin of first abdominal sternum, flattened along hind margin, outer '/3 angulate,
area within line alutaceous, feebly punctured. Male genitalia as in Fig. 434a-c.
Discussion. Typical examples of this species resemble examples of H. quadrivittata,
from which H. oregona differs in having the elytral vittae not approaching the elytral
base, and the marginal vittae not reaching the apical spots. Hyperaspis lanei Hatch
has male genitalia nearly identical to those of oregona and I consider lanei a color
variant of H. oregona, therefore a junior synonym. The tendency toward the dis-
appearance of the discal vittae in any long series of typical H. oregona is apparent.
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NORTH AMERICAN COCCINELLIDAE
531
and the complete loss of these vittae would result in the form described as lanei (Fig.
434d). Hyperaspis oregona has previously been placed in a group with H. quadrivit-
tata, but the male genitalia indicate that H. oregona belongs in the postica group.
Type locality. Of oregona, Harney Co., Oregon; of lanei. Bead Lake, Washington.
Type depository. Of oregona, CAS; of lanei, USNM.
Distribution. Figure 430. ALBERTA: Banff. BRITISH COLUMBIA: Creston; Fer-
nie; Royal Oak. CALIFORNIA: Mono Co., Bridgeport; Siskiyou Co., Bartle. IDAHO:
Cassia Co., Rock Creek Can.; Centerville; Hayden’s L. OREGON: Blue Mts.; Cot-
tonwood Creek; Harper. WASHINGTON: Ellensberg; Yakima R. Canyon; Entiat;
Granger; Moxee; Neppel; Pullman; Soap Lake; Wenatchee. WYOMING: “Nat. Park.”
disconotata group
Female pronotum entirely black; form rounded, convex; base of hrst abdominal
sternum within postcoxal line depressed, with transverse suture; male genitalia of
the undulata type without bisinuate lateral margin of basal lobe.
The affinities of the 2 species included in this group appear to be with members
of the undulata group, but the female pronotum is entirely black and the basal lobe
of the male genitalia is not bisinuate, therefore I do not include them in the undulata
group.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Hyperaspis disconotata Mulsant
Fig. 435a-d; Map, Fig. 436
Hyperaspis disconotata Mulsant, 1850, p. 653.— Crotch, 1873, p. 380.— LeConte,
1880, p. 187. -Casey, 1899, p. 127.-Leng, 1920, p. 212.-Korschefsky, 1931, p.
187.-Wingo, 1952, p. 26.
Hyperaspis disconotata disconotata: Dobzhansky, 1941, p. 61.
Diagnosis. Length 2.30 to 2.80 mm, width 1.75 to 2.0 mm. Form elongate, oval,
moderately convex. Elytron with 5 yellow spots (Fig. 43 5d), median basal spot oblique
and close to humeral spot. Postcoxal line distinctly removed from hind margin of
first abdominal sternum, slightly flattened parallel to hind margin, area within line
dull, strongly alutaceous, barely perceptibly punctured. Male genitalia as in Figure
435a-c.
Discussion. This species and H. troglodytes have been variously considered as
synonyms or varieties of each other by authors. Dobzhansky (1941) considered H.
troglodytes to be a subspecies of H. disconotata based on a presumed difference in
distribution. I have seen both H. disconotata and H. troglodytes from the same
locality, Sherborn, Massachusetts. Hyperaspis disconotata is elongate, moderately
convex in form, the pronotal punctures are very fine, indistinct in some specimens.
1985
NORTH AMERICAN COCCINELLIDAE
533
and the 2 basal spots on each elytron almost touch. Hyperaspis troglodytes is rounded,
convex in form, the pronotal punctures are distinct, and the 2 basal spots on each
elytron do not approach each other. For these reasons I consider both species valid.
There are 2 female types in the LeConte collection, one of which, labeled “(pale blue
disc, clipped)/4660/H. disconotata Muls.”, I here designate and label as the lectotype.
The other specimen bearing only a pale blue, clipped disc, I designate a paralectotype.
Type locality. Lake Superior (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 436. ALBERTA: Edmonton. QUEBEC: Duparquet. MAS-
SACHUSETTS: Northboro; Sherbom. MINNESOTA: Duluth; Garrison. NEW
YORK: Mt. Marcy. WISCONSIN: Oneida Co.
534
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 437. Hyperaspis troglodytes.
Hyperasp is troglodytes Mulsant
Fig. 437a-d; Map, Fig. 438
Hyperaspis troglodytes Mulsant, 1853, p. 91. — Casey, 1899, p. 127. — Leng, 1920, p.
212.-Korschefsky, 1931, p. 198.
Hyperaspis disconotata troglodytes: Dobzhansky, 1941, p. 62.
Hyperaspis discreta LeConte, 1880, p. 187. — Casey, 1899, p. 127.
Hyperaspis troglodytes ab. discreta: Korschefsky, 1931, p. 198.
Hyperaspis novascotiae 1955, p. 153. New Synonymy.
Diagnosis. Length 2.0 to 2.75 mm, width 1 .60 to 2.40 mm. Form rounded, convex.
Color pattern of elytron as described for H. disconotata except basal spots on elytron
widely separated (Fig. 437d). Postcoxal line as described for H. disconotata. Male
genitalia as in Figure 437a-c.
Discussion. The external differences between H. troglodytes and H. disconotata
have been discussed under the latter species. In addition, the male genitalia are distinct
in each case. I have not located a type of H. troglodytes. The type of H. discreta is
a unique (holotype) male labeled “Cambr. 20.2.74/Type 6710 (red paper)///, discreta
LeC.’’’’ The male holotype of novascotiae lacks the median basal spot and the genitalia
are slightly different than those of troglodytes, but I regard it as synonymous with
troglodytes. Hyperaspis troglodytes is so rare in collections that the possible range of
variation cannot be assessed.
Type locality. Of troglodytes. United States; of discreta, Cambridge, Massachusetts;
of novascotiae, Bridgewater, Crescent Beach, Nova Scotia.
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NORTH AMERICAN COCCINELLIDAE
535
Fig. 438. Distribution. Hyperaspis troglodytes (star); H. undulata (dot); H. octavia (open
star); H. paludicola (open circle).
Type depository. Of troglodytes, type not located; of discreta, MCZ; of novascotiae,
CNC.
Distribution. Figure 438. CONNECTICUT: New Haven. INDIANA; Orange Co.
IOWA: Mt. Pleasant. MASSACHUSETTS: Berlin; Framingham; Sherborn. MIN-
NESOTA: Fillmore Co.; Houston Co.; Mille Lacs Co.; Plummer. PENNSYLVANIA:
Mt. Alta.
undulata group
Female pronotum narrowly yellow on lateral border; form elongate or rounded;
male genitalia with basal lobe bisinuate on lateral margin.
The bisinuate basal lobe of the male genitalia is the striking characteristic of this
group. The body form is generally elongate, sometimes depressed, in general more
convex than in members of the quadrivittata group.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 439. Hyperaspis undulata.
Hyperaspis undulata (Say)
Fig. 439a-d; Map, Fig. 438
Coccinella undulata Say, 1824, p. 92. — Mulsant, 1850, p. 1049.
Hyperaspis undulata: Crotch, 1873, p. 381.— Wickham, 1894, p. 304.— Weise, 1895a,
p. 128.-Casey, 1899, p. 128.-Blatchley, 1910, p. 521.-Korschefsky, 1931, p.
198.-Dobzhansky, 1941, p. 65.-Wingo, 1952, p. 26.-J. Chapin, 1974, p. 43.-
Belicek, 1976, p. 314.
Hyperaspis maculifera Melsheimer, 1847, p. 179. — LeConte, 1880, p. 189.
Hyperaspis elegansMwXsdLnX, 1850, p. 658. — LeConte, 1852, p. 134. — LeConte, 1880,
p. 187.
Hyperaspis elegans var. guttifera Weise, 1895a, p. 128.
Hyperaspis undulata ab. guttifera: Korschefsky, 1931, p. 199.
Diagnosis. Length 1.80 to 2.75 mm, width 1.50 to 2.0 mm. Form elongate, oval,
moderately convex. Elytron black with complete, irregular, lateral vitta (Fig. 439d),
or with vitta broken to form apical spot. Postcoxal line not reaching hind margin of
first abdominal sternum, evenly curved throughout. Male genitalia as in Figure
439a-c.
Discussion. With the exception of H. octavia, H. undulata is readily recognized by
the dorsal color pattern over most of the area in which it occurs. There is no good
1985
NORTH AMERICAN COCCINELLIDAE
537
external character that will distinguish H. undulata from H. octavia, but H. octavia
is usually larger, the body form less elongate, and the surface of the pronotum less
strongly alutaceous than in H. undulata. The type of undulata no longer exists and
the type of maculifera could not be located. The type of elegans Mulsant is a female
labeled “Amer. bor., LeConte” which I designate and label as the lectotype.
Type locality. Of undulata, “Missouri”; of maculifera, Pennsylvania; of elegans,
“I’Amerique boreale”; of guttifera, not stated.
Type depository. Of undulata, type lost; of maculifera, type not located; of elegans,
DLM; of guttifera, not located.
Distribution. Figure 438. ALBERTA: Calgary; Coaldale; Medicine Hat. BRITISH
COLUMBIA: Creston; Lulu Is; Sumas; Summerland; Wynndel. ONTARIO: Mus-
kota Co., Gull Lake; Pr. Edw. Co. CONNECTICUT: Lakeville. COLORADO: Eckert.
ILLINOIS: Macon Co.; Urbana. INDIANA: Dunes Park Beach. IOWA: Ames. KAN-
SAS: Atchison; Manhattan; Mission. LOUISIANA: Cameron Parish; Orleans Parish;
Webster Parish. MARYLAND: Upper Marlboro. MASSACHUSETTS: Ashbumhas;
Framingham; Natick; Northboro. MICHIGAN: Birmingham; Charlevoix Co.; Kew-
eenaw Co.; East Lansing; Rochester; Royal Oak; Washtenaw Co. MINNESOTA:
Clay Co., Buffalo River. NEW JERSEY: Haddon Hts. NEW YORK: Geneva; Long
Beach; Onondaga Co.; Seneca Co., Willard. NORTH DAKOTA: Billings Co.; Bot-
tineau Co.; Grant Co., Lake Tschida; Lake Tewaukon; Richland Co., Mirror Pool;
Wahpeton. OHIO: Champaign Co.; Green Co., John Bryan St. Pk.; Highland Co.,
Rocky Ford L; Licking Co. OREGON: Klamath Falls; Redmond. PENNSYLVANIA:
Mt. Alta. TENNESSEE: Chattanooga. UTAH: Provo. WASHINGTON: Benton Co.,
Hanford Works. WISCONSIN: Dane Co., Madison.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Hyperaspis octavia Casey
Fig. 440a-d; Map, Fig. 438
Hyperaspis octavia Casey, 1908, p. 419. — Korschefsky, 1931, p. 193.— Dobzhan-
sky, 1941, p. 65.-Wingo, 1952, p. 26.
Diagnosis. Length 2.20 to 2.80 mm, width 1.70 to 2.10 mm. Form broadly oval,
convex. Elytron black with pattern variable from that described for H. undulata to
having the lateral vitta broken into 3 spots (Fig. 440d). Postcoxal line as described
for H. undulata. Male genitalia as in Figure 440a-c.
Discussion. This species will normally be confused only with H. undulata (see
comments under H. undulata). There are 3 types of H. octavia in the Casey collection,
one male and 2 females. I here designate and label the male as the lectotype, the
females as paralectotypes.
Type locality. Vicksburg, Mississippi (lectotype here designated).
Type depository. USNM (35204).
Distribution. Figure 438. QUEBEC: St. Hilaire. MICHIGAN: Detroit; Oakland
Co.
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NORTH AMERICAN COCCINELLIDAE
539
Hyperaspis paludicola Schwarz
Fig. 441a-d; Map, Fig. 438
Hyperaspis paludicola Schwarz, 1878, p. 362. — LeConte, 1880, p. 188.— Casey,
1899, p. 128.— Leng, 1920, p. 212. — Korschefsky, 1931, p. 193. — Dobzhansky,
1941, p. 66.
Diagnosis. Length 1.70 to 2.10 mm, width 1.10 to 1.40 mm. Form elongate, nearly
parallel sided, depressed. Elytron black with narrow, complete lateral vitta and one
discal spot (Fig. 44 1 d). Postcoxal line not reaching hind margin of first abdominal
sternum, evenly curved throughout, area within line alutaceous, barely perceptibly
punctured. Male genitalia as in Figure 44 1 a-c.
Discussion. The dorsal color pattern is identical to that of typical H. undulata, but
the parallel sided depressed form of H. paludicola is quite different from the oval,
moderately convex appearance of H. undulata. These 2 species are apparently al-
lopatric. I have seen 5 type specimens of paludicola labeled “Type”, of these I select
and label a male labeled “Capron Fla. 3.4/Coll Hubbard & Schwarz/Type No. 4519
U.S.N.M.” as the lectotype. The other 4 specimens from Capron and Lake Ashby I
designate as paralectotypes.
Type locality. Capron, Florida (lectotype here designated).
Type depository. USNM (4519).
Distribution. Figure 438. ALBERTA: Mobile. FLORIDA: Kissimmee; Sanford;
Steinhatchee R. GEORGIA: Dade Co., Grant’s Blowing Spring. MISSISSIPPI: Pas-
cagula. SOUTH CAROLINA: Myrtle Beach; Sassafras Mtn.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 442. Hyperaspis borealis.
Hyperaspis borealis Dobzhansky
Fig. 442a-e; Map, Fig. 444
Hyperaspis oregona borealis Dobzhansky, 1941, p. 76.
Hyperaspis oregona: Belicek, 1976, p. 314.
Hyperaspis borealis: Hatch, 1961, p. 76.
Hyperaspis obscura Malkin, 1943a, p. 1 10. — Hatch, 1961, p. 159. New Synonymy.
Hyperaspis simuloides Hatch, 1961, p. 159. New Synonymy.
Hyperaspis schuhi Hatch, 1961, p. 160. New Synonymy.
Hyperaspis elali Nutting, 1980, p. 262. New Synonymy.
Diagnosis. Length 2. 10 to 3.0 mm, width 1 .50 to 2.0 mm. Form elongate, somewhat
oval. Elytron black with discal and lateral vittae joined at apex (Fig. 442d), pattern
variable as in Figure 442d, e. Postcoxal line nearly reaching hind margin of first
abdominal sternum, flattened along margin, area within line alutaceous, distinctly
punctured. Male genitalia as in Figure 442a-c.
Discussion. The fully marked form resembles H. quadrivittata and H. oregona but
so many color variants occur in this species that male genitalia are the only positive
criteria for recognizing H. borealis. Dobzhansky (1941) described borealis as a sub-
species of oregona, but H. borealis belongs in the undulata group while H. oregona
is in the postica group. The names I place in synonymy are all color variants of H.
borealis, and the male genitalia form the basis of this synonymy.
Type locality. Of borealis. Lake Cle-Elum, Washington; of obscura. Lake of the
Woods, Klamath Co., Oregon; of simuloides, Almota, Washington; oi schuhi, Sprague
1985
NORTH AMERICAN COCCINELLIDAE
541
Fig. 443. Hyper aspis consimilis.
River, 5 mi. E. Ely, Oregon; of elali, Yosemite National Park, Tuolumne Co., Cal-
ifornia.
Type depository. Of borealis, obscura, and elali, CAS: of simuloides and schuhi,
USNM.
Distribution. Figure 444. BRITISH COLUMBIA: Vancouver; Yale. CALIFOR-
NIA: Alameda Co., Hayward; Trinity Co., Trinity River; Tuolomne Co., Yosemite
Nat. Park. OREGON: Corvallis; Harper; Klamath Falls; McMinnville. WASHING-
TON: Coupeville; Sunnyside; Fidalgo I; King Co., Seattle; Kittitas; North Bend;
Thurston Co., Offut L.; Olympia; Olympic N. F., Hurricane Rdge; Port Angeles;
Pullman; Puyallup; Seaview; Skagit Co., Clear L.; Snohomish Co., Chase L.; Sultan.
Hyperaspis consimilis LeConte
Fig. 443a-d; Map, Fig. 444
Oxynchus consimilis l^tConiQ, 1852, p. 134. — Korschefsky, 1931, p. 202.
Hyperaspis consimilis: Crotch, 1873, p. 381.— Crotch, 1874b, p. 233.— LeConte,
1880, p. 189.-Casey, 1899, p. 128.-Dobzhansky, 1941, p. 78.
Hyperaspis disconotata canadensis Dobzhansky, 1941, p. 63. New Synonymy.
Hyperaspis moerens: Wingo, 1952, p. 26.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 444. Distribution. Hyperaspis borealis (dot); H. consimilis (star); H. spiculinota (open
star); H. quadrivittata (open circle); H. brunnescens (circled star).
Diagnosis. Length 2.30 to 2.70 mm, width 1.70 to 1.90 mm. Form elongate, oval,
moderately convex. Elytron with narrow, complete lateral vitta, one median, basal
spot, one oblique discal vitta extending from disc nearly to apical vitta (Fig. 443d).
Postcoxal line nearly reaching hind margin of first abdominal sternum, slightly flat-
tened along margin, area within line alutaceous, finely punctured. Male genitalia as
in Figure 443a-c.
Discussion. The dorsal color pattern of H. consimilis is most like that of H. trog-
lodytes, but the male genitalia are so dissimilar in these 2 species that I place them
in different groups. Crotch (1873) first considered H. consimilis to be a junior syn-
onym of H. moerens LeConte, and this opinion has been followed by authors ever
since. In fact, H. moerens is much more closely allied to H. quadrivittata than to H.
consimilis, and I regard H. consimilis as a valid species. Because of this confusion
as to the true identity of H. consimilis, Dobzhansky (1941) described as new the
form he called H. disconotata canadensis, which is identical in all respects to H.
consimilis. The type of H. consimilis is a unique (holotype) female labeled “(blue
clipped disc)/4661/Type 6720 (red paper)/H. consimilis Lee.”.
Type locality. Of consimilis. Lake Superior; of canadensis, Whitford Lake, Alberta.
Type depository. Of consimilis, MCZ; of canadensis, CAS.
Distribution. Figure 444. ALBERTA: Whitford Lake. QUEBEC: Duparquet. NEW
YORK: Cascade.
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NORTH AMERICAN COCCINELLIDAE
543
Fig. 445. Hyperaspis spiculinota.
Hyperaspis spiculinota Fall
Fig. 445a-d; Map, Fig. 444
Hyperaspis spiculinota Fall, 1901, p. 232. — Leng, 1920, p. 212.— Korschefsky, 1931,
p. 197. — Dobzhansky, 1941, p. 72.
Diagnosis. Length 2.40 to 3.0 mm, width 1.70 to 2.10 mm. Form elongate, feebly
oval, dorsoventrally flattened. Elytron with wedge shaped discal spot, spot on lateral
margin in apical half, and transverse apical spot sometimes touching lateral spot (Fig.
445d). Postcoxal line nearly reaching hind margin of first abdominal sternum, slightly
flattened along margin, area within line alutaceous, finely punctured. Male genitalia
as in Figure 445a-c.
Discussion. The dorsal color pattern is like that of a variation of H. quadrioculata,
but the strongly flattened form and the wedge shaped discal spot on each elytron are
characteristic of H. spiculinota. The type of male genitalia causes me to place this
species in the undulata group rather than the postica group where it would appear
to belong on the basis of external characters. A single male type remains in the Fall
collection labeled “Pom Cal 10/ 19/9 5/Type spiculinota/MCZ Type 24544 (red pa-
per)/ H.C. Fall collection”. Since Fall had more than one type specimen, I designate
this male as the lectotype.
Type locality. Pomona, California (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 444. CALIFORNIA; Claremont; Pasadena; Santa Barbara Co.;
Santa Paula.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 446. Hyper aspis brunnescens.
quadrivittata group
Female pronotum with narrow yellow border (except H. brunnescens and H. jas-
perensis); form elongate, flattened; color pattern of elytron often vittate; male genitalia
with strong lateral projection near apex of basal lobe.
Of the species included here, H. jasperensis is atypical in that the male head is
black (the only species of Hyperaspis known to have that characteristic) and the
female pronotum is entirely black. The body form and male genitalia of H. jasperensis
are of the quadrivittata type, therefore I include it in the quadrivittata group in spite
of the aberrant color pattern.
Hyperaspis brunnescens Dobzhansky
Fig. 446a-c; Map, Fig. 444
Hyperaspis brunnescens Dobzhansky, 1941, p. 77. — Wingo, 1952, p. 26.
Diagnosis. Length 2.30 to 2.50 mm, width 1.60 to 1.80 mm. Form elongate, oval,
subdepressed. Pronotum of male mostly dull yellow; pronotum of female brownish
black with indistinct, yellow lateral border. Elytron brownish black with 2 vittae,
one complete marginal vitta and oblique discal vitta (Fig. 446c). Surface of elytron
dull, strongly alutaceous. Postcoxal line not reaching hind margin of first abdominal
sternum, slightly flattened along margin, area within line alutaceous, barely percep-
tibly punctured. Male genitalia as in Figure 446a, b.
Discussion. The dull dorsal surface and vittate color pattern of the elytron are
distinctive for H. brunnescens. Hyperaspis quadrivittata has the same elytral pattern
but is shiny on the dorsum.
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NORTH AMERICAN COCCINELLIDAE
545
Fig. 447. Hypemspis quadrivittata.
Type locality. Illinois.
Type depository. USNM (54219).
Distribution. Figure 444. ILLINOIS: State record. IOWA: Clarke Co.
Hyperaspis quadrivittata LeConte
Fig. 447a-e; Map, Fig. 444
Hyperaspis quadrivittata LeConte, 1852, p. 133.— Crotch, 1873, p. 381.— Crotch,
1874b, p. 233.-LeConte, 1880, p. 188. -Casey, 1899, p. 128.-Blatchley, 1910,
p. 522.-Leng, 1920, p. 212.-Korschefsky, 1931, p. 195.-Wingo, 1952, p. 26.-
Belicek, 1976, p. 313.
Hyperaspis quadrivittata quadrivittata: Dobzhansky, 1941, p. 74.
Hyperaspis tetraneura Casey, 1908, p. 420. — Leng, 1920, p. 212. — Belicek, 1976, p.
313.
Hyperaspis quadrivittata variety Xetraneura: Dobzhansky, 1941, p. 75.
Diagnosis. Length 2.0 to 2.70 mm, width 1.30 to 1.80 mm. Form elongate, oval,
subdepressed. Elytron black with vittae as described for H. brunnescens except some
specimens with lateral vitta incomplete (Fig. 447d, e). Postcoxal line as described
for H. brunnescens except area within line strongly punctured. Male genitalia as in
Figure 447a-c.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 448. Hyperaspis moerens.
Discussion. This is one of the most easily recognized species of Hyperaspis because
of the strongly vittate appearance and elongate form (see comments under H. brun-
nescens). Hyperaspis oregona of the postica group has the same basic color pattern,
but the male genitalia of the 2 specimens are not at all alike and they are nearly
allopatric.
The unique (holotype) female type of quadrivittata is labeled “(green disc)/4659/
Type 6719 (red paper)/H. quadrivittata LeC.” LeConte (1852) stated that the type
was “found near Long’s Peak.” The type of tetraneura in the Casey collection is a
unique female (holotype).
Type locality. Of quadrivittata. Long’s Peak, Colorado; of tetraneura, Boulder,
Colorado.
Type depository. Of quadrivittata, MCZ; of tetraneura, USNM (35207),
Distribution. Figure 444. ALBERTA: Cypress Hills; Edmonton; Medicine Hat;
Waterton. ARIZONA: Chiricahua Mts.; IDAHO: Boville. NORTH DAKOTA: Bil-
lings Co.; Grant Co., Lake Tschida. OREGON: Brothers; Harney Co., Tencent Lake;
Prineville. WASHINGTON: Moses Canyon. WYOMING: Lonetree.
Hyperaspis moerens (LeConte)
Fig. 448; Map, Fig. 450
Oxynychus moerens 1850, p. 238.— Crotch, 1874b, p. 239.— Korschefsky,
1931, p. 202.
Hyperaspis {Oxynychus) moerens: Mulsant, 1850, p. 694.
Hyperaspis moerens: LeConte, 1880, p. 189. — Dobzhansky, 1941, p. 78.
Diagnosis. Length 2.25 mm, width 1.70 mm. Form oval, subdepressed, pronotum
abruptly narrower than elytral base, narrowed apically. Elytron black with obscure.
1985
NORTH AMERICAN COCCINELLIDAE
547
Fig. 449.
obliquely triangular spot on apical declivity (Fig. 448). Postcoxal line nearly reaching
hind margin of first abdominal sternum, slightly flattened along margin, area within
line alutaceous, finely punctate. Male genitalia not known.
Discussion. I have seen only the type series (females) and one other female specimen
of this species. The abruptly narrowed form of the pronotum is not shared by any
other species of North American Hyperaspis, and I have no doubt that this is a valid
species. No males were available, therefore the placement of moerens in this group
is based on an external similarity to quadrivittata, which may be superficial. Two
types presently exist in the LeConte collection, one of which, labeled “(blue clipped
disc)/4671/Tyepe 6721 (red paper)/Oxynychus moerens Lee. (Hyper. Muls.)”, I des-
ignate and label as the lectotype, the other as a paralectotype.
Type locality. Lake Superior (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 450.
Hyperaspis protensa Casey
Fig. 449a-d; Map, Fig. 450
Hyperaspis protensa Casey, 1908, p. 417.-Leng, 1920, p. 212. — Korschefsky, 1931,
p. 194.— Dobzhansky, 1941, p. 57.
Diagnosis. Length 1.50 to 2.20 mm, width 1.0 to 1.50 mm. Form elongate, parallel
sided. Elytron black with complete vitta on lateral margin (Fig. 449d), vitta often
incomplete. Postcoxal line not approaching hind margin of first abdominal sternum,
evenly curved throughout, area within line dull, alutaceous, distinctly punctured.
Male genitalia as in Figure 449a-c.
Discussion. The color pattern of this southwestern species is similar to that of H.
inflexa and H. caseyi, but the elongate, narrow form of H. protensa is completely
unlike the short, rounded form of the other 2 species. Thus far H. protensa is known
548
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 450. Distribution. Hyperaspis moerens (star); H. protensa (dot); H. punctata (open
circle); H. simulans (square); H. imitator (open star).
only from Arizona, and it is not likely to be confused with any other species of
Hyperaspis occurring there. The type is a unique female (holotype) in the Casey
collection.
Type locality. Nogales, Arizona.
Type depository. USNM (35185).
Distribution. Figure 450. ARIZONA: Dragoon Mts., Stronghold. Graham Mts.;
Huachucha Mts., Copper Canyon; Pajarito Mts., Sycamore Canyon; Pinaleno Mts.;
Santa Catalina Mts.; Santa Cruz Co., Parker Canyon; Santa Rita Mts., Box Canyon,
Madera Canyon.
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NORTH AMERICAN COCCINELLIDAE
549
Hyperaspis punctata LeConte
Fig. 451a-b; Map, Fig. 450
Hyperaspis punctata LeConte, 1880, p. 188. — Casey, 1899, p. 128.— Korschefsky,
1931, p. 194. — Dobzhansky, 1941, p. 67.— Wingo, 1952, p. 26.
Diagnosis. Length 1.70 to 2.30 mm, width 1.25 to 1.80 mm. Form elongate, nearly
parallel sided, subdepressed. Elytron black with discal spot, apical spot, and irregular
vitta on lateral margin from base past midpoint (Fig. 45 Id). Postcoxal line removed
from hind margin of first abdominal sternum, evenly curved throughout, area within
line dull, alutaceous, distinctly punctured. Male genitalia as in Figure 451a-c.
Discussion. This species and H. protensa are practically identical in body form and
color pattern except that H. punctata has a discal spot on the elytron. The genitalia
are similar but not identical, however, it is conceivable that these are forms of a
single species. Material from northern Mexico is needed to ascertain the actual dis-
tribution of both punctata and protensa. A female in the LeConte collection labeled
“Tex./ 184/Type 6717 (red paper)/H. punctata Lec'\ is here designated and labeled
as the lectotype. An additional female is designated a paralectotype.
Type locality. Texas (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 450. TEXAS: Bexar Co., Salado Creek, Fort Sam Houston;
Flatonia; Perryton; Sanderson.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Hyperaspis simulans Casey
Fig. 452a-d; Map, Fig. 450
Hyperaspis simulans 1899, p. 128. — Leng, 1920, p. 212. — Korschefsky, 1931,
p. 197. — Dobzhansky, 1941, p. 79.
Diagnosis. Length 2.30 to 2.70 mm, width 1.60 to 1.90 mm. Form regularly oval,
moderately convex. Elytron entirely black except vague yellow area may be present
on humeral angle (Fig. 4 5 2d). Postcoxal line nearly reaching hind margin of first
abdominal sternum, slightly flattened along margin, area within line alutaceous,
distinctly punctured. Male genitalia as in Figure 452a-c.
Discussion. The regularly oval form and nearly black, immaculate appearance
characterize H. simulans externally, and enable it to be separated from other south-
western species of Hyperaspis. I have seen a single female from Reno, Nevada, which
I regard as H. simulans, but this specimen has a small spot just posterior to the
middle of the elytron on the lateral margin. Casey’s type of simulans is a unique
female (holotype).
Type locality. Arizona.
Type depository. USNM (35208).
Distribution. Figure 450. ARIZONA: Huachucha Mts.; Santa Cruz Co., Adobe
Canyon. NEVADA: Reno.
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NORTH AMERICAN COCCINELLIDAE
551
Fig. 453. Hyperaspis imitator.
Hyperaspis imitator, new species
Fig. 453a-d; Map, Fig. 450
Description. Male, length 2.0 mm, width 1.60 mm. Form oval, moderately convex.
Pronotum black with lateral area yellow; elytron black with discal spot and lateral
yellow vitta from base to apex (Fig. 453d). Punctures on head fine, separated by a
diameter or less; pronotal punctures coarser than on head, separated by a diameter
or less; punctures on elytron slightly coarser than on pronotum, separated by less
than to twice a diameter. Metastemum coarsely punctured laterally, punctures be-
coming fine, sparse toward midline. Abdominal sterna with fine, dense punctures
except basal 2 sterna coarsely punctured. Postcoxal line not approaching hind margin
of first abdominal sternum, evenly curved throughout, area within line alutaceous,
sparsely punctured. Male genitalia as in Figure 453a-c.
Female, similar to male except length 2.25 mm, width 1.75 mm; head black; lateral
pronotal border not as broadly yellow.
Holotype. Male. TEXAS: San Antonio, Olmos Park, 28-VI-1947, B. E. White Coll.,
sweeping Ceanothus sp. (CAS).
Allotype. Female. Same data as holotype. (CAS).
The male genitalia of H. imitator place it in the H. quadrivittata group; the dorsal
color pattern of the male is exactly like that of H. bensonica males, but the female
pronotum is yellow laterally, entirely black in H. bensonica. Because of the similarity
in color pattern and also body form, males of H. imitator will probably be mixed
with males of H. bensonica unless genitalia are examined. The holotype and allotype
are the only specimens of H. imitator examined. The specific name refers to the
resemblance of H. imitator to H. bensonica.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 454. Hyperaspis jasperensis.
Hyperaspis jasperensis Belicek
Fig. 454a-d; Map, Fig. 455
Hyperaspis jasperensis Belicek, 1976, p. 316.
Diagnosis. Length 1.50 to 2.00 mm, width 1.0 to 1.50 mm. Form elongate, slender,
oblong. Head brownish black in both sexes. Pronotum brownish black in both sexes
except lateral margin slightly paler. Elytron entirely brownish black (Fig. 454d).
Postcoxal line nearly reaching hind margin of first abdominal sternum, slightly flat-
tened along margin, area within line alutaceous, barely punctured. Male genitalia as
in Figure 454a-c.
Discussion. The black female head, oblong form, and immaculate elytron char-
acterize this little species. In addition to material from the type locality and the
Northwest Territories, I regard 2 specimens from Colorado and one from Wyoming
as being H. jasperensis. The latter 3 specimens are more flattened dorsoventrally
than Alberta specimens, but the male genitalia are so similar that I consider them
conspecific.
Type locality. Alberta, Jasper National Park, Bald Hills.
Type depository. CNC.
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NORTH AMERICAN COCCINELLIDAE
553
Fig, 455. Distribution. Hyperaspis jasperensis (star); H. bolteri (square); H. trifurcata (open
circle).
Distribution. Figure 455. NORTHWEST TERRITORIES: Dempster, Richardson
Mts. COLORADO: Argentine Pass. WYOMING: Yellowstone Nat. Park.
Hyperaspis species not assigned to groups
Two species, H. bolteri LeConte and H. trifur at a Schaeffer, are included here
because they seem to have no affinities with other Hyperaspis, either North American
species or species from south of the United States. I prefer to treat them in this way
rather than create a probably meaningless “group” for each.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Hyperaspis bolteri LeConte
Fig. 456a-d; Map, Fig. 455
Hyperaspis bolteri LeContQ, 1880, p. 186.— Casey, 1899, p. 121. — Blatchley, 1910,
p. 522.-Leng, 1920, p. 2 1 1 . - Korschefsky, 1931, p. 185.-Wingo, 1952, p. 26.
Diagnosis. Length 3.0 to 3.25 mm, width 2.10 to 2.65 mm. Form regularly oval,
convex. Male pronotum with narrow yellow area laterally; female pronotum entirely
black; surface of pronotum dull, strongly alutaceous, punctures nearly invisible. Ely-
tron dull black with yellowish orange lateral vitta broadly expanded onto disc medially
(Fig. 456d). Postcoxal line widely separated from hind margin of first abdominal
sternum, flattened along margin, area within line depressed basally, alutaceous, finely
punctured. Male genitalia as in Figure 456a-c.
Discussion. Specimens of H. bolteri have been seen only from Illinois, Indiana,
and Kansas. The highly distinctive color pattern and extremely dull pronotal surface
characterize this species. A female in the LeConte collection labeled “111. /Type 6706
(red paper)/Hyperaspis bolteri Lee.''' is here designated and labeled the lectotype.
Type locality. Illinois (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 455. INDIANA: Pine (Lake Beach). KANSAS: State record.
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NORTH AMERICAN COCCINELLIDAE
555
Fig. 457. Hypemspis trifurcata.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Hyperaspis trifurcata Schaeffer
Figs. 457a-i; Map, Fig. 455
Hyperaspis trifurcata Schaeffer, 1905, p. 143.— Casey, 1908, p. 420.— Leng, 1920, p.
211.-Korschefsky, 1931, p. 198.-Dobzhansky, 1941, p. 63.
Hyperaspis durangoensis CdiSQy , 1924, p. 167.— Korschefsky, 1931, p. 187.— Dob-
zhansky, 1941, p. 64. New Synonymy.
Hyperaspis disjunctus Casey, 1924, p. 168.— Korschefsky, 1931, p. 187.— Dob-
zhansky, 1941, p. 64. New Synonymy.
Diagnosis. Length 2.30 to 3.0 mm, width 1.80 to 2.40 mm. Form oval, convex.
Head sparsely pubescent except pubescence on clypeus dense, long. Pronotum of
both sexes black with narrow, reddish yellow, lateral border. Elytron black with
variable red or yellow pattern (Fig. 457e-i). Postcoxal line not reaching hind margin
of first abdominal sternum, evenly curved throughout, area within line alutaceous,
densely punctured. Male genitalia as in Figure 457a-c.
Discussion. The elytral color pattern is somewhat variable, but always distinctive
for this species. No other North American species of Hyperaspis has a comparable
color pattern. The labrum and clypeus are extremely hairy, although there is no
pubescence on the frons. Comparison of the genitalia of H. trifurcata with those of
H. durangoensis has shown them to be identical, therefore I regard H. durangoensis
as a junior synonym of H. trifurcata. Hyperaspis disjunctus is only a color variant
of trifurcata from the same type locality as H. durangoensis, and is also a junior
synonym. There are 2 types of H. durangoensis in the Casey collection and I here
designate and label a male as the lectotype, the other specimen as a paralectotype.
The type of H. disjunctus is a unique female (holotype). I have seen 2 type specimens
of H. trifurcata and here designate and label a male labeled “Tex. /trifurcata type/
Cotype No. 42550 U.S.N.M.” as the lectotype, the other specimen as a paralectotype.
Type locality. Of durangoensis (lectotype here designated) and disjunctus, Durango
City, Durango, Mexico; of trifurcata, Texas (lectotype here designated).
Type depository. Of durangoensis (35 1 68), disjunctus (35 1 69), and trifurcata (42550),
(USNM).
Distribution. Figure 455. ARIZONA: Pima Co., Tucson. TEXAS: S. of Alamo;
Alice; Brownsville; College Station; Corpus Christi; El Paso; Falfurrias; Floresville;
Isabel; Kerrville; Laredo; Refugio Co., Tivoli; Sabinal; San Antonio; San Diego;
Seguin; Uvalde; Victoria.
Genus Brachiacantha
Brachiacantha Dejean, 1837, p. 458. — Melsheimer, 1847, p. 178.— LeConte, 1852,
p. 130.-Belicek, 1976, p. 317.
Brachyacantha: Chevrolat, 1842, p. 704 (unjustified emendation). — Mulsant, 1850,
p. 520.-Crotch, 1873, p. 377. -Crotch, 1874b, p. 210.-Chapuis, 1876, p. 228.-
Weise, 1885, p. 5. -Gorham, 1894, p. 184.- Wickham, 1894, p. 299. -Casey,
1899, p. I16.-Blatchley, 1910, p. 520.-Leng, 1911, p. 281. -Wheeler, 1911, p.
169. -Leng, 1920, p. 2 12. -Korschefsky, 1931, p. 202.-Wingo, 1952, p. 18.-
Hatch, 1961, p. 161.— J. Chapin, 1974, p. 44. Type-species; Coccinella dentipes
F., by subsequent designation of Crotch, 1873, p. 377.
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NORTH AMERICAN COCCINELLIDAE
557
Fig. 458. Brachiacantha sp. a. Antenna, b, c. Front tibiae, d. Abdomen, e. Female genitalia.
Hyperaspini with body elongate oval to rounded, strongly convex; dorsal surface
glabrous except head indistinctly pubescent. Head usually entirely yellow in male,
at least with clypeus brown or black in female. Antenna 1 1 -segmented (Fig. 458a);
antennal insertion concealed. Eye narrowly emarginated by expansion of epistoma.
Scutellum wider than long. Epipleuron of elytron narrow, not descending externally,
strongly excavated for reception of middle and hind femoral apices. Prostemum with
or without 2 carinae, if carinae present, then only slightly convergent apically. An-
terior tibia grooved or flanged, with spine at about basal % (Fig. 458b, c); tarsal claw
with large, basal, quadrate lobe. Abdomen with 7 apparent sterna in male, 6 in
female; sexual modification present on sterna 3-6 in male (depending on group).
Postcoxal line on first abdominal sternum incomplete {Scymnus type) (Fig. 458d).
Male genitalia with basal lobe symmetrical or asymmetrical; paramere not rooted in
phallobase, longer than phallobase (Figs. 459a, 471a); sipho strongly sclerotized with
fan-like membranous lobes in apical ‘A (except indubitabilis and lepida groups) (Fig.
459c). Female genitalia with simple spermathecal capsule, infundibulum present,
coxal plate transverse (Fig. 45 8e).
The spine on the anterior tibia and the emarginate eyes will distinguish Brachia-
cantha from all other hyperaspine genera. Brachiacantha is a New World genus
containing approximately 50 species and subspecies which range from Canada to
Argentina. The original spelling of the generic name, Brachiacantha, was published
by Dejean (1837) without a generic description, but the name was validated by the
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
inclusion of several previously described species. Chevrolat ( 1 842) emended the name
to Brachyacantha, an unjustified emendation which has been used by all subsequent
authors except Melsheimer (1847), LeConte (1852), and Belicek (1976).
The larvae of members of this genus for which data are available feed on Coccidae
in ant nests. This was well documented by Wheeler (1911), who found larvae of B.
quadripunctata quadripunctata in nests oi""Lasius umbratus var. aphidicola"" at Great
Blue Hill near Boston, Massachusetts. Wheeler also mentioned the recording of
similar observations by Smith (1886), Schwarz (1890), and Mann (1911). Adult host
preferences are almost completely unknown. Hosts mentioned in the literature are
“mealybugs,” “root coccids and root aphids;” Pemphigus sp.; Toumeyella parvicornis
(Cockerell); Dysmicoccus brevipes (Cockerell).
This genus was revised by Leng (1911), and his classification has remained un-
changed since then except for descriptions of some new species. The classification
proposed herein is based on Leng’s, but there are a number of changes in synonymy.
Twenty six species and subspecies are here considered to occur north of Mexico.
North American Brachiacantha can be divided into 4 groups based on the type of
male genitalia and modifications of the male abdominal sterna. I designate these as
the dentipes, ursina, lepida and indubitabilis groups. Morphological distinctions are
discussed under each group heading. The male genitalia of Brachiacantha species are
not diagnostic in many instances, and other, often less satisfactory characteristics
such as color, body form, etc., must be used to recognize species.
Key to species of Brachiacantha
1 . Anterior tibia with arcuate flange on outer margin (Fig. 458c); male abdomen with
3rd sternum bicuspid (Fig. 460d) 2
Anterior tibia not noticeably flanged, or if so, then flange not arcuate (Fig. 458b);
male abdomen with 3rd sternum lacking cusps (Fig. 458d) 15
2(1). Elytron with transverse median band only (Fig. 470e); Arizona, Texas
subfasciata Mulsant
Elytron with transverse median band or not, if median band present, then addi-
tional maculation also present; Arizona, Texas and elsewhere 3
3(2). Species occurring east of the Mississippi River 4
Species occurring west of the Mississippi River 6
4(3). Form oval to rounded; elytron with median band composed of connected spots
(Fig. 459d); Florida decora Casey
- Form distinctly oblong; elytron with median band usually appearing entire .... 5
5(4). First abdominal sternum with area within postcoxal line yellow or yellowish
brown; male pronotum yellowish orange except feeble dark maculae at base (Fig.
46 3e); 3rd abdominal sterna of male with cusps reduced (Fig. 463d); Mississippi
soltaui, n. sp.
- First abdominal sternum with area within postcoxal line black, brown, or only
partially yellow; male pronotum mostly black or dark brown, 3rd abdominal
sternum of male with cusps prominent (Fig. 462d); Mississippi and elsewhere .
dentipes (F.)
6(3). Species occurring in California, Nevada, Oregon (Fig. 464) . blaisdelli Nunenmacher
Species occurring east of Nevada and Oregon 7
7(6). Specimens from Arizona 8
- Specimens not from Arizona 10
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NORTH AMERICAN COCCINELLIDAE
559
8(7). Punctures on head fine, about equal in diameter to eye facet; median 'Z? of 3rd
and 4th abdominal sterna with punctures fine, nearly absent; elytron with single,
median, orange band (Fig. 470e) subfasciata Mulsant
Punctures on head coarse, diameter larger than eye facet; median Vs of 3rd and
4th abdominal sterna distinctly, densely punctured; elytron never with single,
median, orange band 9
9(8). Male head entirely yellow; 6th abdominal sternum of male strongly concave me-
dially; Arizona and elsewhere tau LeConte
Male head black except vertex and frons with orange spot; 6th abdominal sternum
of male nearly flat; Santa Rita Mts., Arizona stephani, n. sp.
10(7). Length less than 3.50 mm; male abdomen with cusps on 3rd abdominal sternum
reduced, 4th sternum with equally reduced cusps (Fig. 468d); south Texas
barberi, n. sp.
Length more than 3.50 mm; male abdomen with prominent cusps on 3rd abdom-
inal sternum, 4th sternum without cusps, or if present, reduced; south Texas and
elsewhere 11
11(10). Form oval to rounded; elytron with median band composed of connected spots,
apical spot present (Fig. 459d), all maculation yellow; south Texas . . . decora Casey
- Form oblong; elytron with median band more or less straight, apical spot present
or not, or elytron mostly yellow, all maculation orange or yellowish orange except
some northern specimens of tau with maculation yellow; south Texas and else-
where 12
12(1 1). Male abdomen with cusps on 3rd sternum separated by the width of a cusp or
more (Fig. 460d) 13
Male abdomen with cusps separated by V2 the diameter of a cusp, or cusps con-
nected by an intermediate ridge (Fig. 466d) 14
13(12). Length 4.20 mm or more; punctures on head coarse, diameter larger than eye
facet; median V3 of 3rd and 4th abdominal sterna distinctly, densely punctured;
Montana and Idaho to west Texas tau LeConte
Length 4.0 mm or less; punctures on head fine, about equal in diameter to eye
facet; median V3 of 3rd and 4th abdominal sterna with punctures fine, nearly
absent; south Texas subfasciata Mulsant
14(12). Male abdomen with cusps on 3rd sternum distinctly, strongly connected by a
ridge, ridge feebly, triangularly depressed medially (Fig. 466d); elytron usually
with single apical spot (Fig. 466f), pattern often variable (Fig. 466e, f); south and
central Texas quadrillum LeConte
Male abdomen with cusps on 3rd sternum distinctly separated (Fig. 462e), if
slightly connected, then intermediate ridge strongly, arcuately depressed medially;
elytron never with single apical spot; Kansas, Colorado, New Mexico . . dentipes (F.)
15(1). Elytron yellow with 2 median black spots, plus 2 black spots on suture confluent
with spots on opposite elytron (Fig. 493d) lepida Mulsant
Elytron not as described above 16
1 6( 1 5). Anterior tibia with prominent, rounded tooth on outer margin near tarsal insertion,
tooth angled inward (Fig. 479d); Arizona arizonica Schaeffer
- Anterior tibia with feeble tooth on outer margin near tarsal insertion, tooth not
angled inward (Fig. 458b) Arizona and elsewhere 17
17(16). Elytron black with large basal and apical spots (Fig. 48 2d); Rorida 18
- Elytron not as described above; Florida and elsewhere 19
18(17). Form oval (Fig. 483d); spots on elytron orange, anterior spot entire
querceti Schwarz
- Form round (Fig. 482d); spots on elytron yellow, anterior spot with hind margin
partially divided schwarzi, n. sp.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
19(17).
20(19).
21(20).
22(20).
23(22).
24(23).
25(23).
26(25).
27(26).
28(27).
29(27).
30(29).
Elytron yellow, suture and 2 spots black (Fig. 489d), spots sometimes feebly
confluent with suture; Manitoba and Alberta to Colorado, Nebraska (also see
illustris) albifrons (Say)
Elytron black with yellow spots, or mostly yellow with variable black maculation,
not as described above 20
Elytron black, maculation variable but always lacking discal spot, species occurring
east of Texas 21
Elytron black with discal spot, or mostly yellow with variable black maculation;
Texas and elsewhere 22
Elytron with basal and apical spots, male with additional feeble anterolateral spot
(Fig. 486d) quadripunctata quadripunctata (Melsheimer)
Elytron with basal and apical spots, plus median spot on lateral margin, male
with additional anterolateral spot sometimes connected to basal spot (Fig. 487d)
quadripunctata flavifrons Mulsant
Maculation on elytron consisting of 3 spots, 2 median, one apical, without basal
spot (Fig. 49 Id); spine on anterior tibia slender; occurring east of Mississipi River
indubitabilis Crotch
Maculation on elytron not as described above, or if so, then occurring west of
Mississippi River; spine on anterior tibia usually broad, triangular 23
Elytron with 5 large yellow spots, spots may be partially confluent or confluent
to the extent that the elytron is mostly yellow; length less than 3.0 mm; Texas
and Louisiana 24
Elytron with maculation variable, if with 5 yellow spots, then spots reduced in
size, or species not occurring in Texas or Louisiana 25
Outline of spots on elytron indistinct, often partly confluent or mostly confluent
(Fig. 485d, e) bollii Crotch
Outline of spots on elytron distinct, spots not confluent (Fig. 480d) . . . testudo Casey
Elytron with spots confluent in apical '/a, humeral and basal spot present (Fig.
484d); Florida floridensis Blatchley
Elytron not as described above; Florida and elsewhere 26
Length less than 3.0 mm; form oval; head coarsely, densely punctured; elytron
with 5 small, yellow spots (Fig. 474d) decempustulata (Melsheimer)
Length usually more than 3.0 mm (except felina)', form oblong oval or round;
head finely, indistinctly punctured; elytron with color pattern variable, but usually
with 5 yellow spots 27
Form round (Fig. 47 Id) 28
Form elongate oval (Fig. 478d) 29
Length 3.0 mm or less; male pronotum with anterior margin of median black area
irregular, slightly emarginate at middle (Fig. 4 7 2d) felina (F.)
Length 3.0 mm or more; male pronotum with anterior margin of median black
area straight, not emarginate at middle (Fig. 47 Id) rotunda, n. sp.
Elytron with 5 large, yellow spots (Fig. 476d); eastern Canada and Virginia to
Manitoba and Iowa ursina (F.)
Elytron with 5 yellow spots, or with variable maculation; occurring mostly west
of 100th meridian 30
Elytron usually black with 5 yellow spots (Fig. 476d), or with variable maculation
(Fig. 476e, f); South Dakota to New Mexico, west to British Columbia and northern
California uteella Casey
Elytron yellow with sutural margin and 2 spots black, black areas often confluent;
Manitoba to Colorado, west to Alberta and Idaho 31
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NORTH AMERICAN COCCINELLIDAE
561
31(30). Black spots on elytron usually not confluent (Fig. 489d); form slender, elongate;
distribution mostly Great Plains albifrons (Say)
- Black spots on elytron nearly always confluent (Figs. 490d-f); form oval, robust;
high altitudes, Idaho, Montana, Wyoming illustris Casey
dentipes group
Anterior tibia with arcuate flange on margin, widest just before spine (Fig. 458c);
abdomen of male with 3rd sternum bicuspid (Fig. 460d), 4th and 5th sterna modified;
basal lobe of male genitalia asymmetrical, apex usually abruptly bent to the left in
ventral view (Fig. 459a), sipho with fan-like membranous lobes (Fig. 459c).
This group contains Leng’s (1911) group 1 and group 2 which I have combined
because the features of the tibia, male abdomen, and male genitalia are essentially
the same in both groups. Leng used the body shape and color pattern to distinguish
his “groups” from each other, however, I do not consider these characteristics sig-
nificant at the group level.
Brachiacantha decora Casey
Fig. 459a-e; Map, Fig. 461
Brachyacantha decora Casey, 1899, p. 1 19.— Bowditch, 1902, p. 206.
Brachyacantha bistripustulata var. decora’. Leng, 191 1, p. 298. — Leng, 1920, p. 213.
Brachyacantha bistripustulata ab. decora’. Korschefsky, 1931, p. 203.
Brachyacantha bistripustulata var. minor Leng, 1911, p. 298. — Leng, 1920, p. 213.
New Synonymy.
Brachyacantha bistripustulata ab. minor. Korschefsky, 1931, p. 203.
Diagnosis. Length 3.0 to 4.20 mm, width 2.30 to 3.20 mm. Form oval to rounded.
Pronotum of male mostly yellow with median black area not reaching anterior margin;
pronotum of female black except broad lateral area yellow. Elytron black with yellow
apical spot and median band composed of 2 connected spots (Fig. 459d). Postcoxal
line angulate. Male genitalia as in Figure 459a-c.
Discussion. The yellow maculation on the elytron and oval to rounded body form
will distinguish B. decora from other species occurring in south Texas. This name
may be only a synonym or a subspecies of B. bistripustulata F. (Fig. 459e) but I
prefer to regard B. decora as a valid species until the Mexican and Central American
species are examined in detail. There are 5 type specimens in the Casey collection,
and I here designate and label a male as the lectotype, the remainder as paralectotypes.
The type of B. bistripustulata var. minor Leng is apparently a depauperate example
of B. decora having the median band on the elytron composed of separated spots.
The type locality is Brownsville, therefore the name minor cannot be maintained in
subspecific status. The type specimen was not located.
Type locality. Of decora and minor, Brownsville, Texas (lectotype of decora here
designated).
Type depository. Of decora, USNM (35574); of minor, not located.
Distribution. Figure 461. ARIZONA: Phoenix. FLORIDA: Broward Co., Andy-
town; Dade Co., Matheson Hammock; Miami; Monroe Co., Key Largo; Palm Beach
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 459. Brachiacantha decora.
Co., Lake Worth; West Palm Beach. TEXAS: Beeville; Big Bend National Park;
Brownsville; Del Rio; Devils River; Edna; Harlingen; Mission; New Braunfels; Rich-
mond; Sabinal; San Antonio; Uvalde; Victoria; Webb Co., 30 mi. E. Laredo.
Brachiacantha tau LeConte
Fig. 460a-g; Map, Fig. 461
Brachiacantha tau LeConte, 1859d, p. 28.
Brachyacantha tau: Crotch, 1873, p. 378.— Crotch, 1874b, p. 212. — Leng, 1911, p.
305.-Leng, 1920, p. 21 3.-Korschefsky, 1931, p. 207.
Diagnosis. Length 4.40 to 5.50 mm; width 3.0 to 3.80 mm. Form oblong. Pronotum
of male mostly yellow with small, black, basal area in northern specimens, mostly
black with narrow anterior margin and broad lateral area yellow in southern speci-
mens; pronotum of female mostly black except broad lateral area yellow, anterior
margin narrowly yellow in most northern specimens. Color pattern on elytron vari-
able from mostly yellow or orange in northern specimens (Fig. 460e), to a pattern
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NORTH AMERICAN COCCINELLIDAE
563
Fig. 460. Brachiacantha tau.
resembling that of B. decora in southern specimens (Figure 460f, g). Postcoxal line
angulate; male abdominal cusps separated by more than the width of a cusp (Fig.
460d). Male genitalia as in Figure 460a-c.
Discussion. The pale northern specimens are easily recognized, but the usually
darker southern specimens can be confused with other species of the dentipes group
unless the male abdominal cusps are compared as indicated by statements in the key
to species. LeConte (1859) had a single male type specimen (holotype) labeled “(green
disc)/male sign/Type 6703(red paper)/B. tau Led\
Type locality. Fort Riley, Kansas.
Type depository. MCZ.
Distribution. Figure 461. ARIZONA: Huachucha Mts.; Patagonia; Williams. COL-
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Fig. 46 1 . Distribution. Brachiacantha decora (dot); B. tau (star); B. dentipes (shaded).
ORADO: Boulder; Valmont Butte; Fort Collins; Wray. IDAHO: Hansen; Idaho Falls.
MONTANA: state record. NEW MEXICO: Rio Arriba Co. TEXAS: Alpine. UTAH:
Cache Co., Cornish; Logan Canyon; Manti; Millard Co., Parowan; Provo Canyon;
Torrey; Utah Co.
Brachiacantha dentipes (F.)
Fig. 462a-h; Map, Fig. 461
Coccinella dentipes F., 1801, p. 381. — Olivier, 1808, p. 1051. — Say, 1835, p. 202.
Brachyacantha dentipes: Mulsant, 1850, p. 525.— Crotch, 1873, p. 378.— Casey,
1899, p. 120.— Nunenmacher, 1909, p. 162. — Leng, 1911, p. 300. — Korschefsky,
1931, p. 204.-Wingo, 1952, p. 27. -J. Chapin, 1974, p. 44.
Brachyacantha socialis Casey, 1899, p. 119.— Wingo, 1952, p. 27.
Brachyacantha dentipes socialis Leng, 1911, p. 301.
Brachyacantha dentipes ab. socialis: Korschefsky, 1931, p. 204.
Brachyacantha dentipes var. separata Leng, 1911, p. 31.— Wingo, 1952, p. 27.
Brachyacantha dentipes ab. separata: Korschefsky, 1931, p. 204.
Diagnosis. Length 4.75 to 6.30, width 3.60 to 4.60 mm. Form oblong, sometimes
slightly oval. Pronotum of male mostly black except narrow anterior margin and
wide lateral area yellow or orange; pronotum of female similar to male except anterior
margin black. Elytron black with orange or yellow apical spot and irregular median
band varying in width and shape (Fig. 462f-h). Postcoxal line angulate; male ab-
dominal cusps separated by about Vi the diameter of a cusp (Fig. 462d, e). Male
genitalia as in Figure 462a-c.
Discussion. This is the most widely distributed member of the dentipes group and
can usually be recognized by the characters given in the key.
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NORTH AMERICAN COCCINELLIDAE
565
Fig. 462. Brachiacantha dentipes.
I must include 2 previously proposed names, separata and socialis, with B. dentipes
because I cannot find stable characters to differentiate species or subspecies. The
northern, and particularly the western specimens have noticeably more yellow or
orange on the elytra than do those from Missouri, North Carolina, etc., southward.
This is not constant however, occasional southern specimens are as pale as any
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
western specimens. The size and deptth of separation of the cusps on the male 3rd
abdominal sternum seem to differ. The cusps are reduced in size, and not distinctly
separated in most northern and western specimen (Fig. 46 2e) but large and deeply
separated in most southern specimens (Fig. 462d). This character also varies because
specimens in long series from Mobile, Alabama, and St. Louis, Missouri, have most
of the males with large cusps,. A few males in each series have the small cusps
characteristic of northern and western specimens. The type of socialis is a unique
male (holotype) in the Casey collection, the type of dentipes is also apparently unique,
but was not examined. The type of separata was not located; a female labeled as a
paratype from southern Illinois is in the USNM collection.
Type locality. Of dentipes, “Habitat in Carolina”; of socialis, Kansas, of separata,
Virginia.
Type depository. Of dentipes, ZMC; of socialis, USNM (35575); of separata, not
located.
Distribution. Figure 46 1 . New England and Ontario to Florida, west to Colorado
and New Mexico.
Brachiacantha soltaui, new species
Fig. 463a-e; Map, Fig. 465
Description. Male, length 5.0 mm, width 3.80 mm. Form oblong, som^ what oval.
Head and pronotum yellow except pronotum with small, irregular, basal brown area.
Elytron black with humeral angle, median basal, and apical spot orange (Fig. 46 3e).
Ventral surface dark brown except metepistemum yellow, leg reddish yellow, area
within postocxal line and most of abdomen yellowish brown. Punctures on head fine,
dense, separated by a diameter or less; pronotal punctures coarser then on head,
separated by a diameter or less; punctures on elytron equal in size to pronotal punc-
tures, separated by less than to twice a diameter. Metastemum coarsely, densely
punctured, punctures nearly contiguous laterally. Abdominal sterna coarsely, densely
punctured, punctures contiguous laterally, nearly contiguous medially. Postcoxal line
on first abdominal sternum rounded, area within line polished, coarsely, densely
punctured; cusps on third abdominal sternum reduced, connected by arcuate ridge
(Fig. 463d). Sterna 5 and 6 deeply concave medially, sternum 4 weakly concave
medially. Genitalia as in Figure 463a-c.
Female, length 5.30 mm, width 4.0 mm. Similar to holotype except head black
with large, median orange spot; pronotum black except large lateral area reddish
yellow.
Variation. Length 5.0 to 5.30 mm, width 3.80 to 4.0 mm.
Holotype. Male. MISSISSIPPI: Southern Mississippi, June 1 1, 1893, Collection H.
Soltau (USNM 101343).
Allotype. Female. Same data as holotype. (USNM).
Paratypes. Total 7 (Fig. 465). All with same data as holotype. (USNM).
Males of this species are easily recognized because of the nearly all yellow pronotum
and reduced cusps on the third abdominal sternum. Females may be confused with
females of B. dentipes, however, the entirely yellow or yellowish brown area within
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NORTH AMERICAN COCCINELLIDAE
567
Fig. 463. Bmchiacantha soltaui.
the postcoxal line on the first abdominal sternum of B. soltaui is a constant character
in all specimens of the type series. In B. dentipes, the same area is usually entirely
black or brown, or only partially yellowish brown. The species is named for H.
Soltau, the collector.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 464. Brachiacantha blaisdelli.
Brachiacantha blaisdelli Nunenmacher
Fig. 464a-f; Map, Fig. 465
Brachyacantha Nunenmacher, 1909, p. 162. — Leng, 191 1, p. 304.— Leng,
1920, p. 212. — Korschefsky, 1931, p. 204.
Diagnosis. Length 4.40 to 5.0 mm, width 3.0 mm to 3.40 mm. Form oblong,
narrow, elongate. Pronotum of male black except anterior margin broadly yellow,
with large reddish yellow area laterally; pronotum of female black except large lateral
area orange. Elytron black with orange maculation varying from a form with irregular
median band and apical spot to a form with median band and apical spot obscurely
connected (Fig. 464e, f). Postcoxal line angulate; male abdominal cusps widely sep-
arated, large, prominent (Fig. 464d). Male genitalia as in Figure 464a-c.
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NORTH AMERICAN COCCINELLIDAE
569
Fig. 465. Distribution. Bmchiacantha soltaui (circled star); B. blaisdelli (star); B. quadrillum
(dot); B. stephani (triangle).
Discussion. This species is the only member of the dentipes group with an extreme
western distribution. The body form is more narrow and elongate than in any other
member of the group. The male abdominal cusps are widely separated as in B. tau,
but the cusps are much more prominent than those of B. tau. Nunenmacher (1909)
stated that he had 2 type specimens, I here designate and label one of these (male)
as the lectotype, the other (female) as a paralectotype.
Type locality. Goldfield, Esmeralda Co., Nevada (lectotype here designated).
Type depository. CAS.
Distribution. Figure 465. CALIFORNIA: Humboldt Co.; Mendocino Co.; Modoc
Co., Lake City; Napa Co.; Sacramento Co., Galt; Siskiyou Co., Etna; Sonoma Co.,
Rio Nido; Tehachapi Pass. NEVADA: Esmeralda Co., Goldfield. OREGON: Grant’s
Pass.
Brachiacantha quadrillum LeConte
Fig. 466a-g; Map, Fig. 465
Brachiacantha quadrillum LeConte, 1858, p. 89.
Brachyacantha quadrillum: Crotch, 1873, p. 378.— Crotch, 1874b, p. 21 L— Gorham,
1894, p. 186.-Leng, 1911, p. 303.-Korschefsky, 1931, p. 206.
Diagnosis. Length 3.80 to 4.75, width 2.60 to 3.50 mm. Form oval, somewhat
oblong. Pronotum of male and female black with large lateral area reddish yellow.
Elytron black with reddish yellow maculation varying from a form with only an
apical spot to forms with an additional discal spot, or discal spot plus median lateral
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
spot, discal and lateral spot sometimes feebly connected (Fig. 466e-g). Postcoxal line
slightly angulate; male abdominal cusps connected by a ridge triangularly depressed
medially (Fig. 466d). Male genitalia as in Figure 466a-c.
Discussion. Most examples of B. quadrillum can be recognized by the elytral color
pattern because the presence of a single apical spot on the elytron is a character not
shared by other members of the group. Those specimens having additional macula-
tion can still be recognized as B. quadrillum most of the time because the additional
maculation is obscure and tentative in appearance. The type of B. quadrillum is a
unique female (holotype) labeled “(red disc)”/Type 6704(red paper)/5. quadrillum
LeC. Lindh.“.
Type locality. New Braunfels, Texas.
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NORTH AMERICAN COCCINELLIDAE
571
Fig. 467. Brachiacantha stephani.
Type depository. MCZ.
Distribution. Figure 465. TEXAS: Austin; Beeville; Brownsville; Calvert; Dallas;
Del Rio; Edna; Gainesville; Hondo; Kerrville; Lubbock; New Braunfels; Sabinal; San
Antonio; San Marcos.
Brachiacantha stephani, new species
Fig. 467a-e; Map, Fig. 465
Description. Male, length 4.0 mm, width 2.75 mm. Form oblong, slightly oval.
Head black with orange spot on vertex. Pronotum black except anterior margin
narrowly yellow, small lateral area reddish yellow. Elytron black with reddish yellow
apical spot and median band composed of 2 connected spots (Fig. 467e). Ventral
surface black except metepistemum yellow, apex of femur, tibia, and tarsus reddish
yellow. Punctures on head coarse, dense, separated by a diameter or less; pronotal
punctures equal in size to head punctures, separated by one to 3 times a diameter;
punctures on elytron larger than on pronotum, separated by less then to twice a
diameter. Metastemum densely, coarsely punctured, punctures becoming nearly con-
tiguous laterally. Abdominal sterna densely, coarsely punctured, punctures contig-
uous laterally, nearly contiguous medially. Postcoxal line on first abdominal sternum
slightly angulate, area within line polished, impunctate except along anterior margin
of sternum; cusps on third sternum not reduced, separated by more than the width
of a cusp (Fig. 46 7d); sternum 4 weakly concave medially, sternum 5 slightly more
concave than sternum 4, sternum 6 flat, lacking median depression. Genitalia as in
Figure 467a-c.
Female, length 4.5 mm, width 3.0 mm. Similar to holotype except pronotum black
on anterior margin, yellowish red lateral area larger.
Variation. Length 4.0 to 4.80 mm, width 2.75 to 3.20 mm. The median band on
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 468. Brachiacantha barberi.
each elytron may not be partially divided, therefore does not appear to be composed
of 2 connected spots.
Holotype. Male. ARIZONA: Santa Rita Mts., Madera Canyon, 26 Aug. 1970, K.
Stephan Coll (FSCA).
Allotype. Female. Same data as holotype. (FSCA).
Paratypes. Total 5 (fig. 465). ARIZONA: Santa Rita Mts., Madera Cyn., July 21,
1 969; same data except dates Aug. 3, Aug. 1 8, 1 968, and July, 1970. (FSCA) (USNM).
The head of the male is not entirely yellow in B. stephani, a character which will
separate that sex from B. tau and B. subfasciata, the other species of this group
occuring in Arizona. Females of B. stephani and B. tau are not easily separated; in
fact I have not found any character that will accomplish this. This species is named
for Karl Stephan, the collector of part of the type series and an avid Coleopterist.
Brachiacantha barberi, new species
Fig. 468a-e; Map, Fig. 469
Description. Male, length 3.30 mm, width 2.50 mm. Form oval, slightly rounded.
Head yellow. Pronotum black except wide anterior border and large lateral area
reddish yellow. Elytron black with yellow apical spot, median band, and small hu-
meral spot, median band composed of 2 connected spots (Fig. 468e). Ventral surface
black except metepisternum yellow, leg reddish yellow, abdomen yellowish brown
except median Vs of first 2 abdominal sterna black. Punctures on head extremely fine,
barely perceptible; pronotal punctures coarse, dense, separated by a diameter or less;
punctures on elytron equal in size to pronotal punctures, separated by one to 2 times
a diameter. Metastemum coarsely, densely punctured, punctures separated by a
diameter medially, becoming coarser and contiguous laterally. Abdominal sterna
1985
NORTH AMERICAN COCCINELLIDAE
573
Fig. 469. Distribution. Brachiacantha barberi (triangle); B. subfasciata (star); B. rotunda
(dot).
feebly, sparsely punctured medially, punctures becoming coarse, contiguous laterally.
Postcoxal line on first abdominal sternum rounded, area within line smooth, sparsely
punctured; cusps on 3rd sternum extremely reduced, separated by the width of a
cusp (Fig. 46 8d); sternum 4 with cusps nearly equal in size to those on sternum 3;
sternum 5 feebly depressed medially, sternum 6 flat, not depressed medially. Genitalia
as in Figure 468a-c.
Female, length 3.10 mm, width 2.30 mm. Similar to male except clypeus and
lateral margin of frons brown; abdomen mostly yellowish brown except median */3
of first sternum black.
Variation. Length 2.85 to 3.40 mm, width 2.10 to 2.60 mm. One type specimen
has the median band on the elytron nearly straight, not appearing as 2 connected
spots; abdomen is often almost entirely yellowish brown, the median area of the first
sternum brown or dark brown.
Holotype. Male. TEXAS: Corpus Christi, 4/29/96, Marlatt (USNM 101344).
Allotype. Female. TEXAS: Brownsville, 18.24.V.04, HS Barber collector. (USNM).
Paratypes. Total 6 (Fig. 469). TEXAS: Brownsville, 1.VI.04, H.S. Barber collector;
Brownsville, VI. 25-30, J. O. Martin, collector; Esprza (Esperanza) Rch, Brownsville,
VIII.23; Kingsville, C.T. Reed coll.; Macdona, VII-29, J.W. Green, collector. (CAS)
(USNM).
The only species known to occur in south Texas with which B. barberi might be
confused is B. subfasciata, see remarks under that species. This species is named for
H. S. Barber.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Brachiacantha subfasciata Mulsant
Fig. 470a-f; Map, Fig. 469
Brachyacantha subfasciata Mulsant, 1850, p. 527. — Crotch, 1874b, p. 211.— Gor-
ham, 1894, p. 187.-Leng, 191 l,p. 302.-Leng, 1920, p. 212.- Korschefsky, 1931,
p. 207.
Diagnosis. Length 3.50 to 4.00 mm, width 2.60 to 3.0 mm. Form oblong, slightly
oval. Pronotum of male black with narrow anterior margin and narrow area on
anterolateral angle yellow; female pronotum entirely black or with very small yellow
area on anterolateral angle. Elytron usually black except a median orange band present
(Fig. 470e), south Texas specimens with additional apical spot (Fig. 4701). Postcoxal
line slightly angulate; male abdominal cusps widely separated (Fig. 470d). Male
genitalia as in Figure 470a-c.
Discussion. The typical form occurs in Arizona and is recognizable by the dorsal
1985
NORTH AMERICAN COCCINELLIDAE
575
color pattern. The south Texas specimens have an additional apical spot on each
elytron, resembling H. barberi. However, H. barbed is consistently smaller and more
rounded in body form. Two type specimens were found in the Paris Museum, a male
in the general collection, and a female in the Sicard collection. The male labeled
’’Museum Paris/143/Brachyacantha subfasciata Muls., auct. det.“ is here designated
and labeled the lectotype. The female labeled ’’Coll. Mniszech/Mexique“ is desig-
nated a paralectotype.
Type locality. ”Mexique“ (lectotype here designated).
Type depository. PM.
Distribution. Figure 469. ARIZONA: Cochise Co., Douglas; Pima Co., Pantano;
Sta. Catalina Mts., Molino Basin. TEXAS: Brownsville; Crockett Co., Ozona; Del
Rio; Kerrville; San Antonio; Sinton, Welder Wildlife Res.; Uvalde; Webb Co., Lar-
edo.
ursina group
Anterior tibia not noticeably flanged, or if so, then flange not arcuate, widest at
middle or just before tibial excavation (Fig. 458b); abdomen of male without cusps
on 3rd sternum (Fig. 458d), 5th sternum modified; basal lobe of male genitalia
symmetrical, apically truncate (Fig. 471a), sipho with fan-like membranous lobes
(Fig. 472c).
The ursina group contains all of the species in Leng’s (1911) Group 4 except B.
indubitabilis. I also include B. querceti from Leng’s Group 6 because the male genitalia
are of the B. ursina type.
Brachiacantha rotunda, new species
Fig. 471a-d; Map, Fig. 469
Description. Male, length 3.50 mm, width 2.70 mm. Form round. Head yellow
except clypeus yellowish brown. Pronotum black except narrow anterior margin and
apical angle yellow. Elytron black with 5 yellow spots (Fig. 47 Id). Ventral surface
black to dark brown except mouthparts, tibia, and tarsus yellow. Clypeus narrow,
anterior angle rounded. Punctures on head fine, separated by one to 3 times a di-
ameter; pronotal punctures coarser than on head, separated by a diameter or less;
punctures on elytron slightly coarser than on pronotum, separated by less than to
slightly more than a diameter. Metasternum coarsely punctured, punctures sparse
medially, nearly contiguous laterally. Abdominal sterna finely punctured, punctures
sparse medially, contiguous laterally. Postcoxal line on first abdominal sternum
rounded, area within line alutaceous, sparsely, coarsely punctured; median depression
of 4th and 5th sterna feeble. Genitalia as in Figure 471a-c.
Female, length 3.80 mm, width 2.0 mm. Similar to holotype except yellowish
brown area on clypeus enlarged, vertex black, narrow margin beside eye yellowish
brown; pronotum with anterior margin black.
Variation. Length 3.10 to 4.0 mm, width 2.20 to 3.0 mm.
Holotype. Male. VIRGINIA: Massanutten Mt., Sept. 21, 1 94 1 , E. A. Chapin (USNM
101345).
Allotype. Female. VIRGINIA: same data as holotype. (USNM).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Pamtypes. Total 54 (Fig. 469). ONTARIO: Aubrey Island, St. Lawrence Is. Na-
tional Park, Sept. 15, 1976, Oct. 2, 1976, W. Reid; Bell’s Corners, VI-7- 1950, S.D.
Hicks; Hastings, 8 May-01, Evans; Miners Bay, 26-V-1931, G. S. Walley; Ridgeway,
5/30/91, E.P.U. Coll; St. Lawrence Is. National Park, Grenadier L, Centre, 2-9: VII:
1975, Sigler; Thwartway Island, St. Lawrence Is. National Park, Sept. 9, 1976, W.
Reid. QUEBEC: Aylmer, 18-V-1934, W. J. Brown. DISTRICT OF COLUMBIA:
Washington, 5-IV, H. S. Barber. IOWA: Independence, V-15, Wickham; Iowa City,
5-30-16, L. Buchanan. KENTUCKY: Cadiz, May 24, 1954, K. Stephan. MARY-
LAND: Beltsville, VIII-2 1-1958, H. P. Lanchester; S. Mts. near Myersville, Sept. 2,
15; Oakland, V-24-1942, Dieke; Westminster, V-15- 1940, Dieke. MASSACHU-
SETTS: Blue Hills, W. M. Mann; Mt. Tom; Springfield, 14 May, 1919, Geo Dim-
mock; Tyngsboro, VIII-II- 1 4. MISSOURI: Webster Groves, 6- 10-31. NEW JERSEY:
Big Timber Cr., 1 1-19-1900; Towaco, VI- 19-43, A. Nicolay; Lakehurst, 9/1/07, V-
30-25; Midvale, VIII-30-42, A. Nicolay. NEW YORK: Bangall, IX- 17- 13, G. P.
Engelhard!; Buffalo, 6-5-87, E.P.V. coll; Buffalo, 22 July 1933, J. G. Franclemont;
”Cent.“; Ithaca, 12 May 1937, 1938, J. G. Franclemont; Lancaster, 5/20/89, E.P.V.
coll; L. I. Yaphank VIII- 13- 10; West Point, May 30, 1909, Oct. 10, 1909, May 8,
1910, Sept. 15, 1915, W. Robinson. OHIO: Jefferson, R. J. + M. B. Sim; Knox Co.,
IX- 1 8- 1 940. PENNSYLVANIA: Dauphin Co., VI-5-27, J. N. Knull; Mt. Holly Spgs.,
IX- 1 - 1 9 1 8. VIRGINIA: Top Mt. Elliott, Augusta Co., 4473 ft., 20-6-34, H. A. Allard;
Edinburgh, IX- 13- 1945, Dieke; Fava. Co., Belvoir, VI-2- 1940, Dieke; Glencarlyn,
1985
NORTH AMERICAN COCCINELLIDAE
577
Fig. 472. Brachiacantha felina.
IX- 14,19, H. F. Wickham; same as type data; Nelson Co., Aug. 7, 1 9 1 0, W. Robinson.
(CAS) (CNC) (FSCA) (USNM).
In addition to the key characters, the clypeus with rounded anterior angles and
darkened surface are of help in distinguishing B. rotunda from B. ursina with which
it is most likely to be confused. Males of B. ursina nearly always have the head,
including the clypeus, entirely yellow, and the clypeal angles are abrupt in both sexes.
It is surprising that B. rotunda should have remained undetected after both Casey
and Leng worked on Brachiacantha, particularly in view of the very noticeable
difference in body shape. The specific epithet refers to the round body form.
Brachiacantha felina (F.)
Fig. 472a-d; Map, Fig. 473
CoccinellafelinaF., 1775, p. 87. — Fabricius, 1781, p. 106.— Fabricius, 1787, p. 61.—
Fabricius, 1792, p. 290.— Fabricius, 1801, p. 385. — Olivier, 1791, p. 79.
Brachyacantha felina: Mulsant, 1850, p. 1046. — Crotch, 1873, p. 377.— Crotch,
1874b, p. 307. -Leng, 1911, p. 312.-Korschefsky, 1931, p. 205.-Wingo, 1952,
p. 27. -J. Chapin, 1974, p. 45.
Brachiacantha fulvopustulata Melsheimer, 1847, p. 178.
Brachyacantha fulvopustulata: Crotch, 1874b, p. 43. — Leng, 1911, p. 312.
Brachyacantha felina ym. fulvopustulata: Korschefsky, 1931, p. 205.
Diagnosis. Length 2.20 to 3.0 mm, width 1.65 to 2.50 mm. Form round. Pronotum
of male black with anterior margin and anterolateral angle yellow, apex of black area
irregular, slightly emarginate at middle; female pronotum usually black except an-
terolateral angle yellow, often with narrow, yellow, anterior border. Elytron black
with 5 yellow spots (Fig. 472d). Postcoxal line rounded. Male genitalia as in Figure
472a-^.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Discussion. The round form of this species causes it to resemble B. rotunda, but
there is a distinct size differential between these species, as well as a difference in the
male pronotal color patterns as indicated in the key to species. A specimen in the
LeConte collection is labeled ’’Melsh. fulvopustulata/(ragged red paper“, but I do
not consider it a type because it has 5 spots on each elytron. Melsheimer specifically
stated that his type had 4 spots on each elytron.
1985
NORTH AMERICAN COCCINELLIDAE
579
Fig. 474. Brachiacantha decempustulata.
Type locality. Of felina, ”Amer. bor.“ (type not examined); of fulvopustulata, Penn-
sylvania.
Type depository. Of felina, not located; of fulvopustulata, not located.
Distribution. Figure 473. Massachusetts to North Carolina, west to Iowa and Lou-
isiana.
Brachiacantha decempustulata (Melsheimer)
Fig. 474a-d; Map, Fig. 475
Hyperaspis 10-pustulata Melsheimer, 1847, p. 179.
Brachiacantha 10-pustulata: LeConte, 1852, p. 133.
Brachyacantha 10-pustulata: Mulsant, 1856, p. 149.— Crotch, 1873, p. 378.— Crotch,
1874b, p. 2 11. -Wickham, 1894, p. 304.-Casey, 1899, p. 1 17.-Blatchley, 1010,
p. 520. — Korschefsky, 1931, p. 205.
Brachyacantha felina 10-pustulata: Leng, 1911, p. 313.
Brachyacantha ursina var. troglodytes Mulsant, 1850, p. 534. New Synonymy.
Brachyacantha stellata ab. troglodytes: Korschefsky, 1931, p. 207.
Brachyacantha stellata Casey, 1899, p. 1 17.— Blatchley, 1910, p. 520.— Korschefsky,
1931, p. 207.-Belicek, 1976, p. 317.
Brachyacantha ursina stellata: Leng, 1911, p. 310.
Diagnosis. Length 2.0 to 2.50 mm, width 1.60 to 2.0 mm. Description as for felina
except form oval; spots on elytron smaller (Fig. 474d); male genitalia as in Figure
474a-c.
Discussion. In addition to the key characters and reduced size of the elytral spots,
this species also has sparse elytral punctures that are usually separated by more than
the diameter of a puncture, whereas in B. felina and B. rotunda they are usually
separated by a diameter or less. Melsheimer (1847) had a single type specimen of B.
decempustulata which is now in the LeConte collection labeled ”Melsh./10 pustu-
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 475. Distribution. Brachiacantha decempustulata (shaded, disjunct localities dotted).
lata“. A single female type of B. troglodytes labeled ”Amer. bor., Mannerheim“ is
here designated and labeled as the lectotype. There are 2 female types of B. stellata
in the Casey collection. The first of these is here designated and labeled as the
lectotype, the other specimen as a paralectotype.
Type locality. Of decempustulata, Pennsylvania; of troglodytes, ”Amer. bor.“ (lec-
totype here designated); of stellata, Indiana (lectotype here designated).
Type depository. Of decempustulata, MCZ; of troglodytes, DLM; oi stellata, USNM
(35567).
Distribution. Figure 475. New Brunswick and Nova Scotia to Florida, west to
Wisconsin and Louisiana. Disjunct localities; Barnes Co., Baldhill Dam, and Griggs
Co., North Dakota.
Brachiacantha ursina (F.)
Fig. 476a-f; Map, Fig. 477
Coccinella ursina F., 1787, p. 61.— 1792, p. 291. — 1801, p. 386.— Olivier, 1791, p.
80.
Brachyacantha ursina: Mulsant, 1850, p. 532.— Crotch, 1873, p. 378.— Wickham,
1894, p. 304.-Casey, 1899, p. 1 17.-Blatchley, 1910, p. 520.-Leng, 1911, p.
309.-Leng, 1920, p. 212.-Korschefsky, 1931, p. 207.-Wingo, 1952, p. 27.-J.
Chapin, 1974, p. 45. — Belicek, 1976, p. 317 (in part).
Brachyacantha congruens Casey, 1899, p. 1 17. — Blatchley, 1910, p. 520.— Belicek,
1976, p. 317.
Brachyacantha ursina congruens: Leng, 1911, p. 310.
Brachyacantha ursina ab. congruens: Korschefsky, 1931, p. 207.
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NORTH AMERICAN COCCINELLIDAE
581
Diagnosis. Length 3.0 to 4.0 mm, width 2.10 to 2.80 mm. Form elongate oval.
Pronotum of male black except anterior margin and anterolateral angle broadly
yellow, apical margin of black area indented. Elytron usually with 5 yellow spots
(Fig. 476d), specimens from Iowa and Minnesota often with spots partially confluent
(Fig. 476e, !)• Postcoxal line rounded. Male genitalia as in Figure 476a-c.
Discussion. This is the most commonly collected eastern species of Brachiacantha,
but the dorsal color pattern is remarkably uniform. There is a tendency for the elytral
spots to become somewhat confluent in upper midwest specimens, but, if anything,
these specimens are more easily recognizeable than the typical form because no other
species occurring in that region has a similar appearance. The combination of large
size (3.0 to 4.0 mm long) and elongate, oval body form will separate ursina from
other eastern Brachiacantha having the same color pattern. I agree with Belicek
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
(dot).
(1976) who placed congruens Casey as a synonym of ursina. There are 7 types of
''congruens^" in the Casey collection, however, 4 of these are actually felina. I here
designate and label a male in the type series as the lectotype, the remaining 3 examples
of congruens as paralectotypes.
Type locality. Of ursina, ’’America boreali“ (type not examined); of congruens.
Hot Spring, French Broad River, North Carolina, (lectotype here designated).
Type depository. Of ursina, not located; of congruens, USNM (35568).
Distribution. Figure 477. Nova Scotia to South Carolina, west to Manitoba and
Iowa.
Brachiacantha uteella Casey
Fig. 478a-d; Map, Fig. 477
Brachyacantha uteella Casey, 1908, p. 412. — Belicek, 1976, p. 317.
Brachyacantha ursina uteella: Leng, 1911, p. 310. — Leng, 1920, p. 212.
Brachyacantha stellata ab. uteella: Korschefsky, 1931, p. 208.
Brachyacantha uteella sonorana Casey, 1908, p. 413. — Belicek, 1976, p. 317.
Brachyacantha ursina sonorana: Leng, 1911, p. 311. — Leng, 1920, p. 212.
Brachyacantha ursina ab. sonorana: Korschefsky, 1931, p. 207.
Brachyacantha fenyesiT^ng, 191 1, p. 316. — Leng, 1920, p. 212. — Korschefsky, 1931,
p. 205. New Synonymy.
Brachyacantha Nunenmacher, 1912, p. 449. — Leng, 1920, p. 212.— Korschef-
sky, 1931, p. 205. New Synonymy.
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NORTH AMERICAN COCCINELLIDAE
583
Diagnosis. Length 2.60 to 4.60 mm, width 2.0 to 3.40 mm. Form elongate oval.
Pronotum of male black except anterior margin and anterolateral angle broadly
yellow, apical margin of black area indented. Color pattern on elytron variable, typical
form black with 5 yellow spots (Fig. 478d), basal spot often faint, or spots expanded,
partially confluent. Postcoxal line rounded. Male genitalia as in Figure 478a-c.
Discussion. I regard B. uteella as a valid species, widespread and variable in western
North America. Leng (1911) and Belicek (1976) considered uteella a subspecies of
B. ursina. I have not seen intergrade material between the 2, and they are widely
separated geographically, therefore it seems logical to accord each specific rank. The
unique female type (holotype) of sonorana Casey is not separable from examples of
uteella from Arizona and Utah, therefore I regard sonorana to be a junior synonym
of uteella. Both fenyesi Leng and lengi Nunenmacher are apparently inseparable from
uteella, and I also consider them as junior synonyms. Leng (1911) listed 3 Colorado
localities from which he had type specimens. I can locate only one male from any
of these localities labeled ’’Glenwood Spgs., Col./July/Chas. W. Leng collection/
U.S.N.M. paratype 40413 (red paper)“ which I here designate and label as the lec-
totype. Within the known geographic range, the only species with which B. uteella
might be confused is B. albifrons. The darkest color forms of B. albifrons and the
lightest forms of B. uteella may have very similar color patterns, however B. albifrons
is a narrowly oval species with weakly rounded sides and B. uteella is a broader
species with more strongly rounded sides. They are mostly allopatric with some
overlap (Fig. 477).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Type locality. Of uteella, Milford, Utah; of sonorana, Colonia Garcia, Chihuahua,
Mexico; of fenyesi, Glenwood Springs, Colorado (lectotype here designated); of lengi,
Weitchpee, Humboldt Co., California.
Type depository. Of uteella (35569) and sonorana (35570), USNM: of fenyesi,
USNM (40413), oUengi, CAS.
Distribution. Figure 477. BRITISH COLUMBIA: Osoyoos, Richter Pass. ARI-
ZONA: Bright Angel; Coconino Co.; Flagstaff; Huach. Mts.; Pinal Mts.; Tucson;
Williams. CALIFORNIA: Lundy; Trinity Co., Carrville. COLORADO: Boulder;
Colorado Springs; Durango; Estes Park; Glenwood Springs; Gunnison; Muckana-
wago. IDAHO: Murtaugh. MONTANA: Kalispell. NEVADA: Washoe Co., Gerlach.
NEW MEXICO: Santa Fe. OREGON: Corvallis; Crater Lake; Harney Co.; Hun-
tington. SOUTH DAKOTA: Oglala, White River. UTAH: Bicknell; Milford; Pinto;
Roosevelt; Vernal. WASHINGTON: Naches River; Yakima Co., Satus Creek. WY-
OMING: Carbon Co.; Grand Teton Pk.
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NORTH AMERICAN COCCINELLIDAE
585
Brachiacantha arizonica Schaeffer
Fig. 479a-f; Map, Fig. 481
Brachyacantha arizonica Schaeffer, 1908, p. 125. — Leng, 191 l,p. 314.— Leng, 1920,
p. 212. — Korschfsky, 1931, p. 203.
Diagnosis. Length 2.90 to 3.60 mm, width 2.30 to 2.75 mm. Form round. Pronotum
of male with broad anterior margin and anterolateral angle yellow; female pronotum
similar to male except yellow anterior border and anterolateral angle reduced. Elytron
typically black with 4 yellow spots, basal and humeral spot confluent (Fig. 479e);
pattern variable, often with trace of 5th spot on lateral margin, or all spots confluent
(Fig. 479f). Anterior tibia with prominent tooth on outer margin near tarsal insertion,
tooth angled inward (Fig. 479d). Postcoxal line rounded. Male genitalia as in Figure
479a-c.
Discussion. I have seen this species only from Arizona. The round form, confluent
humeral and basal spots, and unique form of the outer anterior tibial tooth char-
acterize B. arizonica. There are 7 type specimens of B. arizonica', I here designate
and label a male as the lectotype, the remaining 6 as paralectotypes.
Type locality. Huachucha Mts., Arizona (lectotype here designated).
Type depository. USNM (42552).
Distribution. Figure 481. ARIZONA: Cochise Co., Chiricahua Mts.; Palmerlee;
Flagstaff; Oak Creek; Globe; Greenlee Co.; Huachucha Mts.; Santa Rita Mts.; Tucson.
Brachiacantha testudo Casey
Fig. 480a-d; Map, Fig. 481
Brachyacantha testudo Casey, 1899, p. 1 18. — Bowditch, 1902, p. 206. — Leng, 1911,
p. 312.-Korschefsky, 1931, p. 207.
Diagnosis. Length 2.25 to 3.30 mm, width 1 .75 to 2.50 mm. Form rounded, slightly
oval. Male pronotum black except apical margin and anterolateral angle yellow;
female pronotum similar to male except anterior margin narrowly yellow. Elytron
black or brown with 5 yellow spots (Fig. 480d), spots sometimes partially confluent.
Postcoxal line slightly angulate. Male genitalia as in Figure 480a-c.
Discussion. The rounded form, 5 spots on each elytron, and extreme south Texas
distribution make correct identification of this species easy. There are 2 female types
of B. testudo in the Casey collection, and I here designate and label one as the lectotype,
the other as a paralectotype.
Type locality. Brownsville, Texas (lectotype here designated).
Type depository. USNM (35571).
Distribution. Figure 481: TEXAS: Brownsville; Cameron Co.; Pt. Isabel. San An-
tonio; Uvalde Co., Uvalde.
Brachiacantha schwarzi, new species
Fig. 482a-d; Map, Fig. 481
Description. Male, length 3.0 mm, width 2.40 mm. Form round. Head yellow.
Pronotum black except anterior margin and anterolateral angle yellow, apical margin
of black area uneven. Elytron black except 2 yellow, fused spots occupying basal Vi,
large apical spot present (Fig. 482d). Ventral surface black except leg yellow, 6th and
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 480. Brachiacantha testudo.
Fig. 481. Distribution. Brachiacantha arizonica (dot); B. testudo (triangle); B. schwarzi
(square); B. querceti (open star); B. floridensis (circled star); B. bollii (star).
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NORTH AMERICAN COCCINELLIDAE
587
Fig. 482. Brachiacantha schwarzi.
7th abdominal sterna reddish brown. Punctures on head fine, separated by less than
a diameter; pronotal punctures coarser than on head, separated by a diameter or less;
punctures on elytron coarser than on pronotum, separated by a diameter or less.
Metastemum coarsely punctured, punctures sparse medially, slightly denser laterally.
Abdominal sterna finely, densely punctured throughout except median area of first
sternum impunctate, polished. Postcoxal line on first abdominal sternum rounded,
area within line somewhat alutaceous, mostly impunctate. Median depression of 4th
and 5th sterna feeble. Genitalia as in Figure 482a-c.
Female, length 2.75 mm, width 2.10 mm. Similar to holotype except head black,
frons yellow; pronotum entirely black except narrow anterolateral angle yellow; ab-
domen entirely black.
Variation. Length 2.30 to 3.20 mm,"width 1 .80 to 2.60 mm. The basal and humeral
spots on each elytron are either distinctly divided at the hind margin, or nearly
eompletely fused.
Holotype. Male. FLORIDA: Gainesville, ll-V-1930 (USNM 101346).
Allotype. Female. FLORIDA: Jacksonville, Collection Ashmead. (USNM). ,
Paratypes. Total 30 (Fig. 481). FLORIDA: Alachua Co., 30-III-54, H. A. Denmark
coll; Alachua Co., 10-III-55, H. V. Weems, Jr. coll; Alachua Co., 4-VIII-1977, H.
Greenbaum; Alachua Co., Devil’s Millhopper, 26-11-26, G. B. Merrill coll; Alachua
Co., Gainesville, Austin Cary Forest, Vic. Hatchett Creek, 8-III-1976, Flight Trap,
G. B. Fairchild; Alachua Co., Gainesville, Austin Cary Forest, 24-IX-1976, 28-VI-
1976, 20-IX-76, 9-11-IX-75, 12-VII-76, G. B. Fairchild, Flight Trap; Alachua Co.,
Gainesville, Doyle Conner Bldg. 1-6-12-1972, H. V. Weems, Jr.; Alachua Co., 9 mi.
N.W. Gainesville, UF Hort Unit, 5R 232, 18-28-VIII-1977, H. N. Greenbaum;
Alachua Co., W. of Gainesville, Pierce’s Homestead, 22-II-4-III-76, Malaise Trap,
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
W. H. Pierce; Gainesville; Gainesville, V-29-64, VIII-8-64, VIII-9-64, R. E. White
coll; Gainesville, 2-1955, L.A. Hetrick; Gainesville, Doyle Conner Bldg., Dodge, 5-
X-72; Gainesville, Doyle Conner Building, 27-VII-73, H. V. Weems, Jr., Malaise
Trap; Gainesville, Doyle Conner Building, 8-IX-1973, H. V. Weems, Jr., Malaise
Trap; Gainesville, Doyle Conner Building, 5-7-1-1974, H. V. Weems, Jr., Malaise
Trap; Green Cove Spgs., 24-III-1952, J. R. Vockeroth; Jacksonville, 30-IV-67, C.
F. Zeiger coll; Lake Kerr, Ocala Nat. For., 23-III-61, R. E. Woodruff; Wakulla Co.,
20-IV-55, F. W. Mead. GEORGIA: Loundes Co., XI-12-62, collr. E. I. Hazard.
(FSCA) (CNC) (USNM).
This species and B. querceti Schwarz resemble each other closely and are the only
North American species of Brachiacantha having the elytral color pattern as in Figure
48 2d. Characters of value in distinguishing these 2 species are: head of B. querceti
yellow (male) or dark brown (female); head of B. schwarzi yellow (male) or black
with yellow frontal area (female); female pronotum entirely black or dark brown in
B. querceti, with anterolateral angle yellow in B. schwarzi; elytral spots orange in B.
querceti, yellow in B. schwarzi; base of elytron with one apparent spot in B. querceti,
2 partially divided spots or one large spot with ragged posterior margin in B. schwarzi;
body oval in B. querceti, round in B. schwarzi. In addition, they appear to be allopatric
within Florida, but this might be an artifact of collecting.
Brachiacantha querceti Schwarz
Fig. 483a-d; Map, Fig. 481
Brachyacantha querceti Schwarz, 1878, p. 362.— Casey, 1899, p. 118. — Leng, 1911,
p. 324.-Leng, 1920, p. 212.-Korschefsky, 1931, p. 207.
Diagnosis. Length 2.20 to 2.80 mm, width 1.65 to 2.25 mm. Form oval, slightly
elongate. Male head entirely yellow; female head entirely brown. Pronotum of male
black, narrowly yellow on anterior margin, anterolateral angle narrowly yellow; fe-
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NORTH AMERICAN COCCINELLIDAE
589
male pronotum entirely black or brown. Elytron as described for B. schwarzi, except
basal spot entire, not partially divided (Fig. 483d), spots orange. Postcoxal line
rounded. Male genitalia as in Figure 483a-c.
Discussion. See remarks under B. schwarzi, n. sp., for comparative statements.
There are 3 types of B. querceti in the USNM collection, all females. I here designate
and label one specimen as the lectotype, the other 2 as paralectotypes.
Type locality. Tampa, Florida (lectotype here designated).
Type depository. USNM 4514.
Distribution. Figure 481. FLORIDA: Estero; Manatee Co., Myakka Head; Polk
Co.; Punta Gorda; St. Petersburg; Tampa; Venice; Vero Beach.
Brachiacantha floridensis Blatchley
Fig. 484a-d; Map, Fig. 48 1
Brachyacantha floridensis Blatchley, 1916, p. 93. — Leng, 1920, p. 212.— Blatchley,
1930, p. 39.— Korschefsky, 1931, p. 205.
Diagnosis. Length 2.50 mm, width 1.80 mm. Form oval, lateral margin strongly
curved. Male head entirely yellow; female head black except frontal area yellow;
male pronotum black except apical V3 yellow, female pronotum black except antero-
lateral angle yellow. Elytron with distinctly separated basal and humeral spots, apical
spots completely confluent (Fig. 484d). Postcoxal line rounded. Male genitalia as in
Figure 484a-c .
Discussion. I have seen only a type and one other specimen of this species. Su-
perficially B. floridensis resembles B. bollii Crotch, but actually the closest relationship
is with B. schwarzi and B. querceti, from which B. floridensis differs in elytral pattern.
Should the elytral pattern prove to be extremely variable, then B. schwarzi and B.
floridensis are probably synonymous. Blatchley (1930) designated a male specimen
as the lectotype, but the specimen in the PU collection is a female.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 485. Brachiacantha bollii.
Type locality. Ocala, Florida
Type depository. PU.
Distribution. Figure 48 1 . FLORIDA: LaCross; Ocala.
Brachiacantha bollii Crotch
Fig. 485a-e; Map, Fig. 481
Brachyacantha bollii Crotch, 1873, p. 379. — Gorham, 1894, p. 188.— Casey, 1899,
p. 118. — Leng, 191 1, p. 314. — Leng, 1920, p. 212. — Korschefsky, 1931, p. 204.—
J. Chapin, 1974, p. 46.
Diagnosis. Length 2.40 to 3.0 mm, width 1.90 to 2.30 mm. Form rounded, slightly
elongate. Head yellow in both sexes except apex of female clypeus slightly darkened;
male pronotum black except anterior V3 yellow, female pronotum black except an-
terolateral angle broadly yellow. Elytron typically with 5 large, round spots (Fig.
48 5d), spots often confluent in basal Vi (Fig. 48 5e). Postcoxal line rounded. Male
genitalia as in Figure 485a-c.
Discussion. This species and B. testudo are similar in appearance, but the elytral
spots are rarely confluent in B. testudo, when they are confluent the apical spots have
the tendency to coalesce rather than the basal spots as in B. bolli. Crotch (1873) had
more than one type specimen, therefore I here designate and label a male in the
LeConte collection labeled ’’Dallas Tex. Boll/male sign/Type 4462(red paper)/B.
bollii Crotch"" as the lectotype, and a female labeled ’’Dallas Tex. Boll“ as a para-
lectotype.
Type locality. Dallas, Texas (lectotype here designated).
Type depository. MCZ.
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NORTH AMERICAN COCCINELLIDAE
591
Distribution. Figure 481. LOUISIANA: Natchitoches; Rapides Parish; Vernon
Parish; Vowell’s Mill. TEXAS: Ardmore; Dallas; Harrison Co.; Kerrville.
Brachiacantha quadripunctata quadripunctata (Melsheimer)
Fig. 486a-e; Map, Fig. 488
Brachiacantha 4-punctata Melsheimer, 1847, p. 178.
Brachyacantha 4-punctata: Crotch, 1873, p. 378. — Blatchley, 1910, p. 521.— Leng,
1911, p. 316.
Brachyacantha quadripunctata: Leng, 1920, p. 212. — Korschefsky, 1931, p. 206.—
Wingo, 1952, p. 27.— J. Chapin, 1974, p. 46 (in part).
Brachiacantha basalts Melsheimer, 1847, p. 177.
Brachyacantha basalts: Crotch, 1873, p. 378.— Casey, 1899, p. 1 18. — Leng, 191 1, p.
317. — Korschefsky, 1931, p. 206.— Wingo, 1952, p.27.
Brachyacantha diversa Mulsant, 1850, p. 538. — Leng, 191 1, p. 316.— Korschefsky,
1931, p. 206.
Brachyacantha confusa Mulsant, 1850, p. 537.— Crotch, 1874b, p. 212. — Leng, 1911,
p. 319. -Wingo, 1952, p. 27.
Brachyacantha 4-punctata confusa: Leng, 1911, p. 319.
Brachyacantha quadripunctata ab. confusa: Korschefsky, 1931, p. 206.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 487. Brachiacantha quadripunctata flavifrons.
Diagnosis. Length 2.50 to 4.0 mm, width 2.0 to 3.20 mm. Form round. Male head
yellow, female head black; male pronotum usually black except narrow apical margin
and anterolateral angle yellow, but often with anterior Vi yellow. Elytron black with
basal and apical spot (female) or with additional humeral spot often confluent with
basal spot (male) (Fig. 486d, e). Postcoxal line angulate. Male genitalia as in Figure
486a-c.
Discussion. The dorsal color pattern, although somewhat variable, is usually suf-
ficient as an identification character. Melsheimer (1847) apparently had a single
female type specimen of B. quadripunctata now in the LeConte collection labeled
’’Melsh. 4-punctata/(ragged red paper)/“. This specimen will be labeled as the ho-
lotype. Exactly the same statements apply to the holotype of B. basalis labeled ’’Melsh.
basalis/(ragged red paper).“ I here designate and label a male in the Dejean collections
labeled ”Amer. bor., LeConte“ as the lectotype of B. diversa, and a female in the
same collection with identical data as the lectotype of B. confusa.
Type locality, of quadripunctata, Pennsylvania; of basalis, Pennsylvania; of diversa
and confusa, ”Amer. bor.“ (lectotypes here designated).
Type depository, of quadripunctata and basalis, MCZ; of diversa and confusa, DLM.
Distribution. Figure 488. Massachusetts and New York to Virginia and Tennessee,
west to Iowa and Kansas.
Brachiacantha quadripunctata flavifrons Mulsant
Fig. 487a-d; Map, Fig. 488
Brachyacantha flavifrons Mulsant, 1850, p. 531. Crotch, 1874b, p. 212.— Casey,
1899, p. 118.
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NORTH AMERICAN COCCINELLIDAE
593
Fig. 488. Distribution. Brachiacantha quadripunctata quadripunctata (shaded); B. quadri-
punctata flavifrons (dot); B. illustris (circled star); B. albifrons (star).
Brachyacantha ursina flavifrons: Crotch, 1873, p. 378.
Brachyacantha quadripunctata flavifrons: Leng, 1920, p. 212.
Brachyacantha 4-punctata flavifrons: Leng, 1911, p. 319.
Brachyacantha quadripunctata 3b. flavifrons: Korschefsky, 1931, p. 206.
Hyperaspis Carolina C3SQy, 1924, p. 164.— Korschefsky, 1931, p. 185.
Brachyacantha Carolina: Dobzhansky, 1941, p. 85.
Description as for B. quadripunctata, except elytron with one marginal spot in
addition to basal and apical spots (Fig. 487d). Male genitalia as in Figure 487a-c.
I agree with Leng’s (1911) placement of B. flavifrons as a subspecies of B. quad-
ripunctata because the color pattern differences exhibit a geographic correlation; B.
flavifrons having a mostly southern distribution and B. quadripunctata a northern
distribution. I here designate and label a male in the Dejean collection labeled ”Amer.
bor., LeConte“ as the lectotype. I agree with Dobzhansky that Carolina is a synonym
of flavifrons. The type of Carolina is a unique female (holotype).
Type locality. Of flavifrons, ’T’Amerique septentrionale“ (lectotype here desig-
nated); of Carolina, Southern Pines, North Carolina.
Type depository. Of flavifrons, DLM; of Carolina, USNM (35211).
Distribution. Figure 488. ALABAMA: Grand Bay; Mobile; Spring Hill. FLORIDA:
Jacksonville; Leon Co., Tallahassee; Liberty Co., Torreya St. Pk.; Mount Pleasant;
Nassau Co., Hilliard. GEORGIA: Beachton; Grady Co.; Jekyll Island. NORTH
CAROLINA: Chadbome; Tryon. SOUTH CAROLINA: Meredith.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Brachiacantha albifwns (Say)
Fig. 489a-f; Map, Fig. 488
Coccinella albifrons Say, 1824, p. 94. — Mulsant, 1850, p. 1049.
Brachiacantha albifrons: LeConte, 1852, p. 132.
Brachiacantha albifrons: Crotch, 1873, p. 378. — Crotch, 1824b, p. 212.— Casey,
1899, p. 119. — Leng, 1911, p. 320. — Leng, 1920, p. 212. — Korschefsky, 1931, p.
203.
Brachyacantha pacifica Casey, 1899, p. 119. — Leng, 1911, p. 321. — Leng, 1920, p.
212. — Korschefsky, 1931, p. 206. New Synonymy.
Diagnosis. Length 3.50 to 4.40 mm, width 2.40 to 3.0 mm. Form slender, elongate.
Head yellow in both sexes, female with clypeal apex brown; male pronotum black
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NORTH AMERICAN COCCINELLIDAE
595
except anterior ‘A yellow, apex of black area sharply emarginate, female pronotum
similar to male except black area larger, not apically emarginate. Elytron typically
yellow with suture and 2 spots brown (Fig. 489d), pattern often partially coalesced
(Fig. 489e, f). Postcoxal line rounded. Male genitalia as in Figure 489a-c.
Discussion. The typical form of B. albifrons is easily recognizeable, but the forms
with coalesced elytral spots resemble examples of B. illustris (see remarks under that
species). The B. pacifica that Casey described from Santa Monica, California, is a
mislabeled specimen of B. albifrons. No species of Brachiacantha occur in coastal
southern California. Therefore, I do not hesitate to synonymize B. pacifica with B.
albifrons.
Type locality. Of albifrons, ’’Taken on the Missouri by Mr. Nuttall“; of pacifica,
Santa Monica, California.
Type depository. Of albifrons, type lost; of pacifica USNM (35573).
Distribution. Figure 488. ALBERTA: Aden; Conrad; Lethbridge; Manyberries;
Medicine Hat; Scandia. MANITOBA: Aweme; Carberry; Roblin; Russell; Virden.
SASKATCHWAN: Beaver Creek; Cut Knife, Attons Lake; Lebret; Mossbank; St.
Victor; Saskatoon. COLORADO: Boulder; Colorado Springs; Rocky Ford. NE-
BRASKA: War Bonnet Canyon. NORTH DAKOTA: Williston.
Brachiacantha illustris Casey
Fig. 490a-f; Map, Fig. 488
Brachyacantha illustris Casey, 1899, p. 1 18. — Bowdith, 1902, p. 206. — Leng, 1911,
p. 320. — Korschefsky, 1931, p. 205.
Diagnosis. Length 3.40 to 4.60 mm, width 2.50 to 3.0 mm. Form elongate, oval.
Pronotum of male black with anterior margin and anterolateral angle broadly yellow,
apical margin of black area deeply indented; female pronotum as described for male
except apical margin black or narrowly yellow. Color pattern on elytron variable,
typical form with all spots connected except discal spot (Fig. 490d), variations as in
Figure 490e, f Postcoxal line rounded. Male genitalia as in Figure 490a-c.
Discussion. This species is most likely to be confused with B. albifrons, and in fact
may be conspecific with it. However, B. albifrons is apparently a Great Plains species
while illustris is a mountain or high altitude species. I have seen no examples of
illustris which have the clear, sharp elytral pattern of typical B. albifrons, and both
males and females of B. albifrons have the yellow, lateral pronotal areas more ex-
panded than the corresponding sex of B. illustris. The type of B. illustris is a unique
female (holotype) in the Casey collection.
Type locality. Colorado, Beaver Brook, 6000 ft.
Type depository. USNM (35572).
Distribution. Figure 488. COLORADO: Mosca. IDAHO: Beaver Canyon. MON-
TANA: Gallatin Valley; Livingston. WYOMING: Yellowstone National Park, Mam-
moth Hot Springs, Roosevelt Lake.
indubitabilis group
Anterior tibia as in the ursina group except spine very slender; abdomen of male
with 5th sternum modified; basal lobe of male genitalia asymmetrical, apex emar-
ginate on side (Fig. 491a), sipho without fan-like membranous lobe (Fig. 491c).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 490. Brachiacantha illustris.
Brachiacantha indubitabilis is the only North American species in this group, and
I consider it unlikely that any neotropical species possesses the extemely characteristic
male genitalia of this species. The external morphology of B. indubitabilis would
place it in the ursina group, but I propose a monotypic group for this species because
of the distinctive male genitalia.
Brachiacantha indubitabilis Crotch
Fig. 491a-d; Map, Fig. 492
Brachyacantha indubitabilis Crotch, 1873, p. 379. — Casey, 1899, p. 120.— Leng,
191 1, p. 315.-Korschefsky, 1931, p. 205.
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NORTH AMERICAN COCCINELLIDAE
597
Fig. 491. Brachiacantha indubitabilis.
Hyperaspis triplicans Casey, 1924, p. 163. — Korschefsky, 1931, p. 198.— Dobzhan-
sky, 1 94 1 , p. 11. New Synonymy.
Hyperaspis triplicans microsticta Casey, 1924, p. 163. — Korschefsky, 1931, p.
192,198. — Dobzhansky, 1941, p. 11. New Synonymy.
Diagnosis. Length 2.50 to 3.20 mm, with 1.80 to 2.60 mm. Form oval. Male head
yellow, female head yellowish brown except frons usually paler yellow; male prono-
tum black except anterior Vs yellow, female pronotum black except lateral 'A yellow.
Elytron black with 3 yellow spots (Fig. 49 Id). Postcoxal line rounded. Male genitalia
as in Figure 49 1 a-c.
Discussion. The presence of only 3 elytral spots in a unique arrangement, 2 median,
one apical, will usually identify this species. In addition, the anterior tibial spine is
usually long and slender, although some examples of B. indubitabilis do not differ
strikingly from B. ursina in this respect. Dobzhansky (1941) considered Hyperaspis
triplicans and H. t. microsticta Casey junior synonyms of Hyperaspis pratensis LeConte.
Examination of the types shows that they are junior synonyms of B. indubitabilis.
Crotch (1873) had more than one type specimen of B. indubitabilis, therefore I here
designate and label a female in the LeConte collection labeled ’’Type 8244(red paper)/
B. indubitabilis Crotch"" as the lectotype.
Type locality. Of indubitabilis, Illinois (lectotype here designated); of triplicans and
microsticta. Southern Pines, N.C.
Type depository. Of indubitabilis, MCZ; of triplicans (35210) and of microsticta
(35212), USNM.
Distribution. Figure 492. Massachusetts and New York to North Carolina, west to
Iowa and Illinois.
lepida group
Anterior tibia as in the ursina group; abdomen of male with only 5th sterna
modified, apex of 5th sternum barely perceptibly depressed; male genitalia with basal
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Fig. 492. Distribution. Brachiacantha indubitabilis.
lobe asymmetrical, apex obliquely truncate (Fig. 493a), sipho without fan-like mem-
branous lobes (Fig. 493b).
Brachiacantha lepida is the only species I include in the group, but there are
probably other neotropical species that ht the criteria outlined here.
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Brachiacantha lepida Mulsant
Fig. 493a-d
Brachyacantha lepidaMulsanX, 1850, p. 523. — Crotch, 1873, p. 378. — Crotch, 1874b,
p. 210.-Gorham, 1894, p. 185.-Leng, 1911, p. 324.-Leng, 1920, p. 212.-
Korschefsky, 1931, p. 205.
Diagnosis. Length 2.50 to 3.60 mm, width 1.70 to 2.40 mm. Form oval. Head
yellow in both sexes; pronotum with black area in basal Vi sharply emarginate. Elytron
yellow with 2 lateral black spots and a discal and apical spot joined with those on
opposite elytron (Fig. 493d). Postcoxal line rounded. Male genitalia as in Figure
493a-c.
Discussion. The elytral color pattern of B. lepida is unique within the genus, making
it easily recognizeable. Crotch (1873) listed it from ”Texas“, but it seems not to have
been found in Texas or anywhere north of Mexico since. I have included it on the
chance that it might still be collected north of Mexico. I have located 3 type specimens
of lepida and here designate and label a male labeled ”Mexico“ as the lectotype. One
paralectotype in the Paris Museum bears no data and one paralectotype in the British
Museum (NH) is labeled ’’Named by Mulsant.“
Type locality. Mexico (lectotype here designated).
Type depository. DLM.
Distribution. Mexico to Costa Rica.
Tribe Cryptognathini
Cryptognathini Gordon, 1971, p. 181.
Oeniini Casey, 1899, p. 74. — Chapin, 1940, p. 263. — Korschefsky, 1931, p.
218.— Gordon, 1971, p. 184. Type-genus preoccupied.
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Scymninae with form round, convex; dorsal surface glabrous. Head directed ven-
trally or slightly posteriorly, usually at least partly concealed behind prostemum.
Antenna short, compact, 8-10 segmented. Maxillary palpus with apical segment ovate
to securiform. Prosternum narrowly or widely produced in front of coxal cavity, not
emarginate; prostemal lobe widely separating coxae. Epipleuron descending exter-
nally, or horizontal, foveate for reception of femoral apices. Metastemum and first
abdominal sternum deeply impressed for reception of femora. Leg with femur and
tibia expanded and modified for reception of tibia and tarsus, anterior leg with
strongly modified tibia; tarsal claw feebly toothed at base. Abdomen with 5 visible
sterna. Postcoxal line incomplete.
This tribe is native to the neotropics, but at least one species has been introduced
into other regions for biocontrol purposes. Gordon (1971) reviewed the tribe at the
generic level and recognized 4 valid genera, one of which occurs in the United States.
Genus Crypto gnat ha Mulsant
Cryptognatha Mulsant, 1850, p. 497.— Crotch, 1874b, p. 206. — Korschefsky, 1931,
p. 218. — Chapin, 1964, p. 231. — Gordon, 1971, p. 183. Type-species; Cryptog-
natha auriculata Mulsant, by subsequent designation of Crotch, 1874b.
Cryptognathini with length 1 . 1 0-3. 1 0 mm. Clypeus with anterior margin upturned,
anterior angle usually sharp (Fig. 494a). Antenna usually 10-segmented, club appar-
ently 4-segmented (Fig. 494b). Apical segment of maxillary palpus securiform (Fig.
494c). Anterior leg with tibia strongly expanded for reception of tarsus, outer edge
sinuate, slightly narrower than femur. Abdomen with postcoxal line incomplete,
extending nearly to lateral margin (Fig. 494d). Male genitalia symmetrical. Female
genitalia with genital plate elongate, triangular (Fig. 494e).
The genus most likely to be confused with Cryptognatha is Delphastus, but Cryp-
tognatha has the elytral epipleura descending externally and the form of the clypeal
apex (Fig. 494a) is broad and truncate medially. The only species of this genus in
North America is C. nodiceps Marshall which was introduced from Trinidad into
Florida for biocontrol of coconut scales in 1936. I have seen 2 recently collected
specimens from Miami, Florida, the area in which it was released in 1936. It may
have existed there since 1936 and 1938, or these specimens may be the result of an
accidental introduction. Members of this genus are scale feeders with Aspidiotus
destructor (Signoret) and Pseudaulacaspis pentagona (Targioni-Tozzetti) recorded as
hosts, but many other scale insects undoubtedly also serve as hosts. Cryptognatha
has not been taxonomically treated as a whole; Chapin (1964) reviewed those species
occurring in Colombia.
Cryptognatha nodiceps Marshall
Fig. 494a-j
Cryptognatha Marshall, 1912, p. 321. — Korschefsky, 1931, p. 219.— Chap-
in, 1965b, p. 249.
Diagnosis. Length 1.20 to 1.66 mm, width 0.90 to 1.35 mm. Form round, convex.
Color pale yellow; pronotum pale reddish yellow, with or without 2 yellowish brown
basal lines on each side of middle at base; elytron with variable dark brown pattern
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NORTH AMERICAN COCCINELLIDAE
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Fig. 494. Cryptognatha nodiceps. a. Head. b. Antenna, c. Maxillary palpus, d. Postcoxal
lines, e. Female genitalia, f-h. Male genitalia, i-j. Habitus and variation.
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(Figs. 494i, j); pro-, meso- and metastema reddish yellow. Postcoxal line incomplete.
Male genitalia as in Figure 494f-h. Female genitalia as in Figure 494e.
Discussion. The host records I have seen for this species have all been Aspidiotus
destructor (Signoret). In addition to the United States, C. nodiceps has been introduced
and established in Fiji and the Mariana Islands. There are 4 types of nodiceps. The
first of these, a male labeled ’’Type H.T. (white disc with orange border)/Trinidad
C. W. Hewer 1 902-207/Cryptognatha nodiceps, Mshl. Type“, I designate and label
as the lectotype. The remaining 3 are designated as paralectotypes.
Type locality. Cedros, Trinidad (lectotype here designated).
Type depository. BMNH.
Distribution. FLORIDA: Miami.
Subfamily Chilocorinae
Chilocorinae Sasaji, 1968, p. 20.— J. Chapin, 1974, p. 48. — Belicek, 1976, p. 293.
Coccinellidae with clypeus expanded laterally. Antenna reduced, 10-segments or
less. Apical segment of maxillary palpus cylindrical with truncate apex; maxillary
cardo expanded or strongly sclerotized. Pronotum strongly descending laterally, deep-
ly concave on anterior margin. Base of elytron distinctly broader than base of prono-
tum. Metastemum impressed for reception of middle femora. Tibia often angulate
externally.
Sasaji (1968) erected this subfamily for 3 tribes, Telsimiini, Platynaspini, and
Chilocorini. Only the Chilocorini occur in the Western Hemisphere.
Tribe Chilocorini Costa
Chilocorini Costa, 1849, p. 9. — Weise, 1885a, p. 4. — Casey, 1899, p. 104.— Leng,
1908, p. 33.-Blatchley, 1910, p. 517.-Leng, 1920, p. 217.-Mader, 1927, p.
23.-Wingo, 1952, p. 24.-Mader, 1955, p. 772.-Chapin, 1965a, p. 234.-J.
Chapin, 1974, p. 48.-Belicek, 1976, p. 293.
Coccinellinae with length 2.0 to 8.0 mm; form oval to nearly circular; dorsal surface
glabrous or pubescent. Antenna short, terminal segments forming fusiform club, base
concealed beneath genal extension of clypeus which is shelflike and partially divides
eye. Prosternal process without carinae. Abdomen usually with 6 visible sterna in
male, 5 in female, sometimes 5 in both sexes, or 6 in both sexes. Tibia simple or
angulate externally; with or without apical spurs. Tarsus cryptotetramerous; claw
simple, or swollen at base, or with basal tooth. Male genitalia symmetrical or asym-
metrical. Female genitalia with long sperm duct, infundibulum present or absent.
There are 5 native North American genera; one introduced genus is established in
California. The tribe is worldwide in distribution, but most species are tropical. The
presence of a strongly expanded gena that partially divides the eye is an excellent
diagnostic character, particularly in the North American fauna. Chapin (1965a) re-
vised the genera of Chilocorini for the world.
Key to genera of Chilocorini
1 . Postcoxal line on first abdominal sternum merging with posterior margin of sternum
(Fig. 528b) Chilocorus Leach
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NORTH AMERICAN COCCINELLIDAE
603
- Postcoxal line on first abdominal sternum not merging with posterior margin of
sternum 2
2(1). Postcoxal line on first abdominal sternum parallel and close to posterior margin of
first sternum (Fig. 527b) Halmus Mulsant
- Postcoxal line on first abdominal sternum recurved apically, complete or not .... 3
3(2). Postcoxal line on first abdominal sternum complete (Fig. 495b) 4
- Postcoxal line on first abdominal sternum incomplete (Fig. 504c) 5
4(3). Tarsal claw without basal tooth (Fig. 495c) Brumoides Chapin
- Tarsal claw with basal tooth (Fig. 511c) . Exochomus Redtenbacher
5(3). Elytron without reflexed margin, with marginal bead; length less than 4.0 mm . . .
Arawana Leng
Elytron with marginal feebly reflexed, with or without marginal bead; length more
than 5.0 mm Mulsant
Brumoides Chapin
Brumoides Chdipin, 1965a, p. 237. — Belicek, 1976, p. 320. Type-species; Coccinella
suturalis Fabricius, by original designation.
Chilocorini with form oval, convex, upper surface glabrous. Antenna 8-segmented,
club 3-segmented with apical segment partly embedded in penultimate (Fig. 495a).
Apical segment of maxillary palpus securiform. Prostemal lobe narrow, truncate
apically. Elytral margin slightly reflexed; epipleuron descending, not foveolate for
reception of femoral apices. Abdomen with 6 visible sterna in male, 5 in female.
Postcoxal line complete (Fig. 495b). Leg slender, femur not inflated; tibial spur
present; tarsal claw slightly thickened at base, without angular basal tooth (Fig. 495c).
Male genitalia symmetrical. Female genitalia with long sperm duct; small infundi-
blulum present (Fig. 496a).
Brumoides is found worldwide but I am able to recognize only 3 species that occur
in the United States and Canada. This is a genus that will bear further study and I
am not satisfied that the arrangement of species and subspecies presented here is
entirely correct. The male genitalia of all species are extremely similar to one another
and the female infundibulum is apparently somewhat variable within each species.
The dorsal color patterns of the species often overlap, or the same pattern will recur
in more than one species, thus color pattern is often not adequate in itself to distin-
guish species. The meagre host data I have seen recorded is as follows: Coccidohystrix
insolitus (Green), Dactylopius confusus (Cockerell), Pseudococcus sp.
Key to species of Brumoides
1. Dorsal surface completely black; form elongate (Fig. 503e) blumi (Nunenmacher)
Dorsal surface usually not entirely black, or if so, then form rounded (Fig. 501b) . 2
2(1). Form nearly round (Fig. 501b); border of elytron definitely explanate, anterolateral
angle pronounced histrio (Fall)
- Form elongate (Fig. 499b); border of elytron feebly explanate, anterolateral angle
not pronounced 3
3(2). Dorsal punctures dense, coarse; eastern United States, New York and New Jersey
west to Wisconsin septentrionis davisi (Leng)
- Dorsal punctures fine, not dense; western United States and Canada 4
4(3). Elytron strongly alutaceous; pronotal angle and propleuron black; Hudson Bay and
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Fig. 495. Brumoides sp. a. Antenna, b. Postcoxal lines, c. Tarsus, d-g. Habitus and variation
of B. septentrionis septentrionis.
Alaska south to Colorado, northern Arizona and California
septentrionis septentrionis (Weise)
Elytron smooth, shiny; pronotal angle and propleuron not black; western Texas to
Colorado and Idaho septentrionis hogei (Gorham)
Brumoides septentrionis septentrionis (Weise)
Fig. 495d-g; Map, Fig. 497
Brumus septentrionis 1885b, p. 230.— Weise, 1904, p. 360. — Horn, 1886, p.
xiv.— Gorham, 1894, p. 177.— Casey, 1908, p. 412. — Korschefsky, 1932, p. 266.
Exochomus (Brumus) septentrionis: Leng, 1908, p. 42.
Brumoides septentrionis: Chapin, 1965a, p. 239. — Belicek, 1976, p. 320.
Exochomus ovoideus CsLsey, 1899, p. 107. — Belicek, 1976, p. 320.
Brumus ovoideus: Casey, 1908, p. 412.
Exochomus {Brumus) septentrionis var. ovoideus: Leng, 1908, p. 42.
Brumoides ovoideus: Belicek, 1976, p. 320.
Exochomus desertorum Casey, 1899, p. 108. — Fall, 1901, p. 231.
Exochomus californicus var. desertorum: Leng, 1920, p. 217.
Brumus desertorum: Casey, 1908, p. 412.
Brumoides desertorum: Belicek, 1976, p. 320.
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NORTH AMERICAN COCCINELLIDAE
605
Exochomus (Brumus) septentrionis var. nevadensis Leng, 1 908, p. 42. New Synonymy.
Brumus hogei nevadensis: Casey, 1908, p. 412.
Brumoides nevadensis: Chapin, 1965a, p. 239.
Diagnosis. Length 2.90 to 3.0 mm, width 1.60 to 2.40 mm. Form oval. The color
pattern of this subspecies is quite variable as illustrated in Figure 495d-g; the dorsal
surface is black except the background color of the elytron which is yellowish brown
to red. Male genitalia as in B. septentrionis davisi.
Discussion. The combination of black pronotal angle and propleuron, alutaceous
elytron and elongate form distinguish B. s. septentrionis from B. s. hogei except where
the two intergrade in Colorado, Nevada and Arizona. B. s. septentrionis is similar
to B. s. davisi except that the latter has coarse, obvious elytral punctation not seen
in septentrionis.
The names listed in the synonymy of this species have been variously treated as
species, subspecies, aberrations, etc., by authors. I regard them as junior synonyms
of B. septentrionis because they simply represent color forms without geographical
correlation. Belicek (1976) listed B. ovoideus as a junior synonym of septentrionis
but erroneously credited Gordon (1974a) as having established that synonymy. I
here designate and label as the lectotype a female of desertorum in the USNM labeled
“Nev./Type 35559/desertorum Casey”. A second female with the same data is des-
ignated and labeled as a paralectotype. The type of ovoideus is a unique female with
a locality label “Ind.”. Casey (1899) listed this locality as “Indiana?”. This specimen
almost certainly did not come from Indiana and I previously (Gordon, 1976b) noted
a similar instance of a species of Scymnus from the Casey collection with an identical
label. In both instances the specimens in question must have been collected from
somewhere in the western United States. A single type specimen of nevadensis Leng
is in the USNM collection. I here designate and label this specimen labeled “C. W.
Leng dedit/Reno, Nev. V11-18/U.S.N.M. Paratype 40403” as the lectotype.
Type locality. Of septentrionis, Hudson Bay; of ovoideus, Indiana?; of desertorum,
Nevada (lectotype here designated); of nevadensis, Reno, Nevada (lectotype here
designated).
Type depository. Of septentrionis, type not examined; of ovoideus and desertorum,
USNM; of nevadensis, USNM.
Distribution. Figure 497. Hudson Bay to northern Arizona, west to Alaska and
northern California.
Brumoides septentrionis hogei (Gorham)
Fig. 496a-d; Map, Fig. 497
Exochomus hogei Gorham, 1894, p. 180.— Casey, 1899, p. 108.
Brumus hogei: Weise, 1904, p. 358.— Casey, 1908, p. 409. — Korschefsky, 1932, p.
265.
Brumoides hogei: Chapin, 1965a, p. 239.
Exochomus {Brumus) orbiculatus Leng, 1908, p. 41. — Korschefsky, 1932, p. 266.
New Synonymy.
Description as for typical B. s. septentrionis with some variation (Fig. 496b-d),
but typical examples of B. s. hogei are only slightly elongate, with dorsal surface
smooth, polished; pronotal angle and propleuron not black; ventral surface nearly
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 496. Brumoides septentrionis hogei.
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NORTH AMERICAN COCCINELLIDAE
607
Fig. 497. Distribution. Brumoides septentrionis septentrionis (shaded, northern); B. s. hogei
(cross hatch, southern); B. s davisi (shaded, eastern).
always yellowish brown to brownish yellow. Male genitalia as in B. septentrionis
davisi. Female genitalia as in Figure 496a.
The typical form occurs from Mexico and western Texas to Arizona. See discussion
under B. s. septentrionis. I consider E. {B.) orbiculatus Leng a color variant of B. s.
hogei and the name to be a junior synonym of that species. I here designate and label
a female in the USNM collection labeled “Tucson, Ariz. July 13-15, 2300-2500 ft.,
Wickham/C. W. Leng dedit/U.S.N.M. Paratype 40404” the lectotype.
Type locality. Of hogei, Mexico, Villa Lerdo in Durango; of orbiculatus, Tucson,
Arizona (lectotype here designated).
Type depository. Of hogei, BMNH; of orbiculatus, USNM.
Distribution. Figure 497. Colorado to west Texas, west to Idaho and Arizona.
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Brumoides septentrionis davisi (Leng)
Fig. 498a-c, 499a, b; Map, Fig. 497
Exochomus {Brumus) septentrionis var. davisi Leng, 1908, p. 42.— Casey, 1908, p.
412.-Korschefsky, 1932, p. 265.
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609
Fig. 499. Brumoides septentrionis davisi (female genitalia and habitus).
Brumus davisi: Wingo, 1952, p. 47.
Brumoides davisi: Chapin, 1965a, p. 239.
Description as for typical B. s. septentrionis except dorsal color pattern bolder with
large, sometimes confluent, black areas (Fig. 499b); punctation on elytron coarser
and denser than in B. s. septentrionis. Male genitalia as in Figure 498a-c. Female
genitalia as in Figure 499a.
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Fig. 500. Brumoides histrio (male genitalia).
Type locality. Washington, D.C.
Type depository. USNM.
Distribution. Figure 497. Southeastern Canada to Virginia, west to Minnesota.
Brumoides histrio (Fall)
Fig. 500a-c, 501a-d; Map, Fig. 502
Exochomus histrio Fall, 1901, p. 230.
Exochomus (Brumus) histrio: Leng, 1908, p. 43.
Brumus histrio: Casey, 1908, p. 412.
Brumoides histrio: Chapin, 1965a, p. 239.
Exochomus parvicollis Casey, 1908, p. 41 1. — Belicek, 1976, p. 320.
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611
Exochomus (Brumus) parvicollis: Leng, 1920, p. 217.
Brumus septentrionis ab. parvicollis Korschefsky, 1932, p. 261 .
Brumoides parvicollis: Chapin, 1965a, p. 239. — Belicek, 1976, p. 320 (as synonym
of B. septentrionis).
Diagnosis. Length 2.75 to 4.0 mm, width 2.20 to 3.0 mm. Form rounded, slightly
oval. Color pattern similar to those of septentrionis-, typical pattern identical to
Exochomus californicus, another pattern same as B. hogei (Fig. 501b-d). Male gen-
italia as in Figure 500a-c. Female genitalia as in Figure 501a.
Discussion. I am unable to separate B. histrio from B. parvicollis and so regard the
latter name as a junior synonym as did Belicek (1976). The color pattern of B.
parvicollis is the same as that of typical B. histrio and the genitalia are identical.
Fall stated that he had 4 types of E. histrio from near Pomona, California. Two
of these remain in his collection, and I here designate a male labeled “Pom Cal Mts
6/25/95/M. C.Z. Type 24550 (red paper) as the lectotype. The other specimen is
labeled ’Torn Cal Mts 5/30/96/“ and I designate it a paralectotype. A third type in
the USNM labeled ’Torn Cal Mts 1 .6.92/3083/HC Fall Cotype No. 6678/Exochomus
histrio Fall“ is designated and labeled a paralectotype.
There are 2 females and a male in the type series of B. parvicollis, all labeled ”St.
George, Utah, July, Wickham/USNM 35566“; one female is here designated and
labeled the lectotype, and the other 2 specimens as paralectotypes.
Type locality. Of histrio, Pomona, California (lectotype here designated); of parv-
icollis, St. George, Utah (lectotype here designated).
Type depository. Of histrio, MCZ; of parvicollis, USNM.
Distribution. Figure 502. Utah to Arizona and southern California.
Brumoides blumi (Nunenmacher)
Fig. 503a-e; Map, Fig. 502
Brumus blumi Nunenmacher, 1934b, p. 113.
Brumoides blumi: Gordon, 1974a, p. 4.
Diagnosis. Length 2.60 to 3.30 mm, width 1.90 to 2.75 mm. Form elongate (Fig.
503e). Dorsum entirely black; venter black except antenna and mouthparts yellowish
brown. Male genitalia as in Figure 503a-c. Female genitalia as in Figure 503d.
Discussion. The species B. blumi most nearly resembles is Exochomus townsendi
(see Gordon, 1974a).
Type locality. Moraga, Contra Costa Co., California.
Type depository. CAS.
Distribution. Figure 502. CALIFORNIA: Alameda Co., Oakland Hills; Contra
Costa Co.; Berkeley; Gilroy Hot Springs; Paraiso Springs; Santa Clara Co.; Alum
Rock Canyon; Santa Cruz Co.
Axion Mulsant
Exochomus {Axion) Mulsant, 1850, p. 477.
Axion: Crotch, 1874b, p. 191. -Gorham, 1892, p. 176. -Casey, 1899, p. 105.-
Leng, 1908, p. 34.-Leng, 1920, p. 25. -Chapin, 1965a, p. 242. -J. Chapin, 1974,
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 501. Brumoides histrio (female genitalia, habitus and variation).
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613
p. 49. Type-species; Coccinella tripustulata Degeer, by subsequent designation of
Crotch, 1873.
Chilocorini with form subcircular, size large, convex, dorsal surface glabrous. An-
tenna 10-segmented, club 3-segmented with apical segment deeply embedded in
penultimate (Fig. 504a). Apical segment of maxillary palpus wider toward apex, apex
strongly oblique. Prosternal lobe narrow, truncate at apex. Pronotum without fine
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Fig. 503. Brumoides blumi.
marginal line across base. Elytron with lateral margin slightly reflexed; epipleuron
with small foveae for reception of femoral apices. Abdomen with 6 visible sterna in
male, 5 in female, male 5th sternum broadly emarginate at apex. Postcoxal line
incomplete (Fig. 504c). Femur not enlarged; tibia slender; tarsal claw with strong,
quadrate, plate-like basal tooth (Fig. 504b). Male genitalia with basal lobe long,
slender, asymmetrical in apical third. Female genitalia with thick portion of sperm
duct longer than thin portion; infundibulum present, Y-shaped (Fig. 505d).
Several specific names have been proposed within this genus by several authors,
but I am able to recognize only 2 species that occur north of Mexico. As in some
other genera in this tribe, the male and female genitalia are not distinctive for each
species. Axion is apparently restricted in distribution to Mexico and the United States.
Available host data indicates that members of this genus are scale predators. Specific
host records are as follows; Kennes puhescens Bogue, Quadraspidiotus perniciosus
(Comstock).
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615
Fig. 504. Axion sp. a. Antenna, b. Tarsus, c. Postcoxal lines.
Key to species of Axion
1 . Lateral margin of elytron thickened into a distinct border; elytra nearly always with a
pale spot on suture (Fig. 505e) tripustulatum (Degeer)
- Lateral margin of elytron not thickened; elytra without pale sutural spot (Fig. 507d)
plagiatum (Olivier)
Axion tripustulatum (Degeer)
Fig. 505a-e; Map, Fig. 506
Coccinella 3-pustulata Degeer, 1775, p. 393.
Exochomus (Axion) tripustulatus: Mulsant, 1850, p. 478.
Exochomus tripustulatus Crotch, 1873, p. 376.
Axion tripustulatus: Crotch, 1874b, p. 191.
Axion tripustulatum: Casey, 1899, p. 106. — Leng, 1908, p. 34. — Korschefsky, 1932,
p. 248.-Wingo, 1952, p. 47. -J. Chapin, 1974, p. 49.
Coccinella verrucatus Melsheimer, 1847, p. 180.
Axion verrucatus: Crotch, 1874b, p. 191.
Axion tripustulatum var. verrucatus: Leng, 1920, p. 217.
Axion incompletus Nunenmacher, 1911, p. 7 1. — Korschefsky, 1932, p. 248. New
Synonymy.
Diagnosis. Length 5.0 to 7.0 mm, width 4.0 to 6.25 mm. Form rounded, slightly
elongate. Color black except narrow pale area at anterolateral angle of pronotum,
and elytron with red or yellow subhumeral spot and small, elongate sutural area
behind middle (Fig. 505e). Lateral margin of elytron distinctly thickened into bead.
Male genitalia as in Figure 505a-c. Female genitalia as in Figure 505d.
Discussion. The color pattern is very constant in this species but occasionally the
pale sutural spot is absent, and it was one of these specimens that Nunenmacher
(1911) named A. incompletus. I consider this name a junior synonym of A. tripus-
tulatum. Axion incompletus was described from a holotype labeled ’’Lincoln Park
Beach/Chicago Ill./Coll. by Wolcott/ /F. Knab/ Axion incompletus Nun.“.
The color pattern and presence of an elytral ridge distinguish A. tripustulatum from
plagiatum, but the 2 species are also nearly allopatric (Fig. 506). Axion tripustulatum
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Fig. 505. Axion tripustulatum.
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617
Fig. 506. Distribution. Axion tripustulatum (cross hatch, western); A. plagiatum (shaded,
eastern).
is primarily an eastern species occurring as far west as Colorado and Texas, and A.
plagiatum is a western species occurring from Louisiana and Texas west to the Pacific
Coast.
Type locality. Of tripustulatum, ”Pensylvania“; of verrucatus, Pennsylvania; of
incompletus, Lincoln Park Beach, Chicago, Illinois.
Type depository. Of tripustulatum, type not examined; of verrucatus, type not
located; of incompletus, CAS.
Distribution. Figure 506. New York to Florida, west to Colorado and Texas.
Axion plagiatum (Olivier)
Fig. 507a-d; Map, Fig. 506
Coccinella plagiata Olivier, 1808, p. 1044.
Exochomus {Axion) plagiatus: Mulsant, 1850, p. All.
Axion plagiatus: Crotch, 1874b, p. 191.
Axion plagiatum: Casey, 1899, p. 106. — Leng, 1908, p. 34. — Korschefsky, 1932, p.
248.— Wingo, 1952, p. 47. — Hatch, 1961, p. 163.— J. Chapin, 1974, p. 50.
Exochomus {Axion) pilatii Mulsant, 1850, p. 478. New Synonymy.
Axion pilatii: Crotch, 1874b, p. 191.
Axion pilatei: Casey, 1899, p. 106. — Leng, 1908, p. 34. — Korschefsky, 1932, p. 248.
Exochomus texanus LeConte, 1858, p. 88.
Axion texanus: Crotch, 1874b, p. 191.— Casey, 1899, p. 166. — Casey, 1908, p. 409.
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Fig. 507. Axion plagiatum.
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NORTH AMERICAN COCCINELLIDAE
619
Axion plagiatum var. texanum: Leng, 1908, p. 36. — Korschefsky, 1932, p. 248.
Chilocorus pleuralis LeConte, 1859a, p. 90.— Crotch, 1874b, p. 185.
Axion pleuralis: Casey, 1899, p. 106.— Casey, 1908, p. 409.
Axion plagiatum var. pleurale: Leng, 1908, p. 36. — Korschefsky, 1932, p. 248.
Axion alutaceum Casey, 1899, p. 106. — Korschefsky, 1932, p. 248.
Axion plagiatum var. alutaceum: Leng, 1908, p. 36.
Description as for plagiatum except elytron lacking sutural spot (Fig. 507). Male
genitalia as in Figure 507a-c.
The size of the pale elytral areas varies as does the body form. At first glance it
seems possible to segregate 2 or more species from what I here consider to be yl.
plagiatum. The principal characteristic involved in giving some specimens a different
appearance is the degree to which the outer epipleural margin descends. In some
specimens, principally from California, this margin does not descend vertically as in
most examples, but descends somewhat obliquely. This gives the specimen a dorso-
ventrally flattened appearance, however, when a large number of specimens from
several localities are available for study, all degrees of epipleural explanation between
the 2 extremes exist. The pale elytral spots are also extremely variable in size and
shape, but I cannot detect any consistent pattern of variation. In the absence of
genitalic differences or any consistent external morphological characters, I regard A.
plagiatum as a somewhat polymorphic species. One form of A. plagiatum that is
apparently restricted to the San Francisco Bay area of California is fairly distinctive
in appearance. The elytral spots are small and light yellow rather than red. This is
the only readily recognizeable variation of A. plagiatum that I have seen. LeConte
specifically stated that he had one specimen of E. texanus when he described it,
therefore the male in his collection labeled ’’(red disc)/male sign/Type 6693(red
paper)/£’. pilatei Muls. texanus Lee. pleuralis Lec.“ must be considered the holotype.
It is not clear how many specimens LeConte had when he described C. pleuralis,
therefore the female in his collection labeled ’’(gold disc)/Type 6694/C. pleuralis
Rathv. Lee. is designated and labeled the lectotype. The type of alutaceus is a unique
female (holotype) in the Casey collection.
Type locality. Of plagiatum, “les iles de la mer des Indes” (West Indies); of pilatei,
not stated; of texanum, Texas; of pleuralis, California (lectotype here designated); of
alutaceum. Las Vegas, New Mexico.
Type depository. Of plagiatum, type not examined; of pilatei, type not examined;
of texanum and pleuralis, MCZ; of alutaceum (35554) USNM.
Distribution. Figure 506. Lousiana to Oregon and southern California.
Arawana Leng
Exochomus (Arawana) Leng, 1908, p. 34, 38. — Casey, 1908, p. 409.— Korschefsky,
1932, p. 245.
Arawana: Leng, 1920, p. 217.— Chapin, 1965a, p. 245. Type-species; Exochomus
arizonicus Casey, by original designation of Leng, 1908.
Chilocorini with form broadly oval, strongly convex, dorsal surface glabrous. An-
tenna 10-segmented, club 3-segmented with apical segment embedded in apex of
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 508. Arawana scapularis. a. Antenna, b. Postcoxal lines, c. Front tibia and tarsus.
penultimate (Fig. 508a). Apical segment of maxillary palpus strongly securiform,
apex oblique. Prostemal lobe broad, apex truncate. Elytral margins not reflexed,
finely beaded; epipleuron foveolate for reception of femoral apices. Abdomen with
5 visible sterna in both sexes. Postcoxal line nearly complete (Fig. 508b). Anterior
tibia with outer margin expanded into a thin keel (Fig. 508c); middle and hind tibiae
with apical spurs; tarsal claw strong, abruptly bent, with triangular basal tooth (Fig.
508c). Male genitalia with basal lobe long, lanceolate; paramere with finger-like
process at apex. Female genitalia with thick portion of sperm duct longer than thin
portion; infundibulum present, Y-shaped (Fig. 509d).
Chapin (1965a) placed 3 species in this genus, one from Central America, one from
Cuba, and the type-species, which is known only from Arizona.
I have not seen any host data for members of this genus, but they are most likely
to be predators of scale insects.
Arawana arizonica (Casey)
Fig. 509a-e; Map, Fig. 510
E xochomus arizonicus CdiSQy, 1899, p. 107.
Exochomus {Arawana) arizonica: Leng, 1908, p. 38.— Casey, 1908, p. 409.— Kor-
schefsky, 1932, p. 245.
Arawana arizonica: Leng, 1920, p. 217.— Chapin, 1965a, p. 245.
Diagnosis. Length 3.25 to 3.50 mm, width 3.0 to 3.20 mm. Form nearly round.
The generic characters separate A. arizonica from other North American chilocorines.
The dorsal color pattern is basically black or piceous with an elongate red spot on
each elytron (Fig. 509e). Male genitalia as in Figure 509a-c. Female genitalia as in
Figure 509d.
Discussion. The presence of a keel on the anterior tibia and the finger-like process
on the paramere are characteristics not found elsewhere in this subfamily. There are
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621
Fig. 509. Arawana scapularis.
2 type specimens of E. arizonica in the Casey collection, labeled “Ari.”. I here
designate and label one as the lectotype and one as a paralectotype.
Type locality. Arizona (lectotype here designated).
Type depository. USNM (35555).
Distribution. Figure 510. ARIZONA: Catalina Springs; Santa Rita Mts.
Exochomus Redtenbacher
Redtenbacher, 1843, p. 11. — Mulsant, 1850, p. 476.— Crotch, 1873, p.
376. -Crotch, 1874b, p. 192.-Casey, 1899, p. I06.-Barovsky, 1922, p. 293.-
Korschefsky, 1932, p. 25. — Wingo, 1952, p. 25.— Chapin, 1965a, p. 247.— J. Chap-
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 510. Distribution. Arawana arizonica (dot); Exochomus marginipennis (shaded).
in, 1974, p. 51. — Belicek, 1976, p. 319. Type-species; Coccinella 4-pustulata L.,
by subsequent designation of Korschefsky, 1932.
Exochomus {Exochomus): Leng, 1908, p. 39.
Chilocorini with form broadly oval to almost round; upper surface glabrous or
pubescent. Antenna 10-segmented; last 3 segments forming a slender fusiform club,
10th segment embedded in 9th (Fig. 511a). Terminal segment of maxillary palpus
subsecuriform, apex strongly oblique. Prosternal lobe narrow, truncate at apex, an-
terior coxae nearly contiguous. Pronotum hnely margined across base, lateral margin
slightly reflexed. Elytral margin strongly beaded, epipleuron not foveolate for recep-
tion of femoral apices. Abdomen with 6 visible sterna in male, 5 in female. Postcoxal
line complete or nearly so (Fig. 511b). Leg with robust femora, tibia slender, tarsal
claw with subquadrate basal tooth (Fig. 511c). Male genitalia with basal lobe asym-
metrical. Female genitalia with long sperm duct; infundibulum present (Fig. 5 1 2d).
Exochomus is distributed worldwide with 10 representatives in the United States
and Canada, 3 of which have been introduced as biocontrol agents. Three of the
native species are readily recognizeable and seem to represent clearly defined taxa.
Another species, E. subrotundus, is also not likely to be confused with anything else
but does bear some resemblance to E. marginipennis and related taxa. Exochomus
marginipennis and E. childreni contain a complex assemblage of forms and I am not
convinced that the classification presented herein is entirely accurate. Previous au-
thors have shuffled species and subspecies in various ways, but the arrangement
presented here seems the most logical at the present time. Specific host data records
are as follows. Aphids; Aphis pomi Degeer, Eriosoma lanigerum (Hausmann), Tox-
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NORTH AMERICAN COCCINELLIDAE
623
a
b
Fig. 511. Exochomus sp. a. Antenna, b. Postcoxal lines.
optera aurantii (Boyer de Fonscolombe). Scales; Aonidiella aurantii (Maskell), Aon-
idiella taxus Leonard!, Asterolecanium cojfeae Newstead, Asterolecanium miliaris
(Boisduval), Asterolecanium bambusae (Boisduval), Ceroplastes rusci (L.), Cero-
plastes sinensis (Del Guercio), Chionaspis furfura (Fitch), Chionaspis minor Maskell,
Chionaspis salicis (L.), Chrysomphalus dictyospermi (Morgan), Chrysomphalus aon-
idum (L.), Coccus viridis (Green), Cryptococcus fagisuga Lindinger, Dactylopius opun-
tiae (Cockerell), Eulecanium tiliae (L.), Filippia oleae (Costa), Hcmiberlcsia lataniae
(Signoret), Lepidosaphes bcckii (Newman), Lepidosaphes gloverii (Packard), Parthen-
olccanium corni (Bouche), Parlatoria camclliae Comstock, Parlatoria oleae (Colvee),
Pinnaspis buxi (Bouche), Planococcus citri (Risso), Planococcus lilacinus (Cockerell),
Pollinia pollini (Costa), Pulvinaria floccifera (Westwood), Pseudaulacaspis pcntagona
(Targioni-Tozzeti), Pseudoparlatoria ostreata Cockerell, Quadraspidiotus marani
Zahradnik, Quadraspidiotus ostreaeformis (Curtis), Saissetia oleae {0\W\qv), Ischnas-
pis longirostris (Signoret), Situlaspis yuccae (Cockerell), Sphaerolecanium prunastri
(Boyer de Fonscolombe), Toumeyella liriodendri (Gmelin), Unaspis citri (Comstock),
Unaspis yanonensis (Kuwana).
Key to species of Exochomus
1 . Dorsal surface including pronotum completely black 2
- Dorsal surface not completely black 3
2(1). Form rounded (Fig. 518e); lateral margin of elytron strongly explanate
aethiops (Bland)
Form elongate (Fig. 520e); lateral margin of elytron weakly explanate
townsendi Casey
3(1). Elytron metallic green, pubescent; California rnetallicus Korschefsky
- Elytron not metallic green, not pubescent; California and elsewhere 4
4(3). Elytron entirely black; pronotum black with large, yellow spot on anterolateral
angle (Fig. 525e) flavipes {Thxmhcr^
- Elytron never entirely black; pronotal color pattern variable 5
5(4). Elytron black with humeral area and spot on apical V2 of elytron yellow or orange
(Fig. 522b)
Elytral color pattern not as above
6
7
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93( 1 )
6(5). Pronotum entirely black; elytron with spot near apex (Fig. 522b) . . californicus Casey
- Pronotum with marginal bead extremely narrowly yellow; elytron with spot just
behind middle (Fig. 523e) quadripustulatus (L.)
7(5). Elytral suture without median pale area (Fig. 517e); west Texas to Utah and Cal-
ifornia subrotundus Casey
- Elytral suture with pale median area (Fig. 516d, e); California or eastern North
America west to central Texas 8
8(7). Known only from California fasciatus Casey
- Known only from eastern North America to central Texas 9
9(8). Form elongate; elytral punctation coarse, distinct; dorsal color patterns as in Figure
512e-h marginipennis (LeConte)
- Form rounded; elytral punctation fine, nearly obsolete, dorsal color patterns as in
Figures 513a, b; 514a, b 10
10(9). Male with pronotal margin and leg yellow; Louisiana, southern Texas
chUdreni guexi LeConte
Male with pronotal margin and leg dark as in female; Florida
childreni chUdreni Mulsant
Exochomus marginipennis (LeConte)
Fig. 512a-h; Map, Fig. 510
Coccinella marginipennis LeConte, 1824, p. 173.
Exochomus marginipennis: Mulsant, 1850, p. 485.— Gorham, 1894, p. 177.— Crotch,
1873, p. 377. — Casey, 1899, p. 108. — Korschefsky, 1932, p. 263. — Wingo, 1952,
p. 47. -J. Chapin, 1974, p. 51.
Coccinella praetextatus Melsheimer, 1847, p. 180.
Exochomus praetextatus: Mulsant, 1856, p. 149.— Crotch, 1874b, p. 193.
Exochomus latiusculus deflectens Casey, 1908, p. 410. — Korschefsky, 1932, p. 263.
New Synonymy.
Diagnosis. Length 2.50 to 3.60 mm, width 2.0 to 2.70 mm. Form oval, slightly
flattened dorsoventrally. Male with anterolateral angle of pronotum and leg yellow,
female with anterolateral angle of pronotum black or slightly pale, leg dark, concol-
orous with ventral surface; elytron typically reddish yellow with black maculae (Fig.
5 1 2g), variable (Fig. 5 1 2e, f, h). Elytral punctation coarse, dense; epipleuron nearly
flat to obliquely or vertically descending externally. Male genitalia as in Figure 5 1 2a-
c. Female genitalia as in Figure 5 1 2d.
Discussion. There are several color forms of what I consider to be E. marginipennis
(Fig. 5 1 2e-h). The dark colored specimens are mostly restricted to the eastern and
southeastern United States. Some specimens from northern Florida and Louisiana
have the pale childreni pattern (Fig. 512h), and all specimens from Texas possess
that pattern. These pale specimens also exhibit strong sexual dimorphism in that the
male has large lateral and sometimes anterior pronotal areas and all legs yellow. The
legs in the female are always dark and the pale lateral areas on the pronotum are
usually reduced to just the anterolateral angle. In addition to changes in color pattern,
a gradual change in body shape can be observed in a dine from New York to Florida
and Alabama. This is a change in the form of the epipleuron from the obliquely
descending type (New York) to a more strongly descending, almost vertical type
(Florida). This also causes the body to appear more slender and elongate in dorsal
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625
Fig. 512. Exochomus marginipennis.
view which, along with the coarse elytral punctation, distinguishes pale specimens
of E. marginipennis from typical specimens of E. childreni. Because of the lack of
genitalic differences I at first considered E. childreni as either a synonym or possible
subspecies of E. marginipennis. However, I have examined specimens of both phe-
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 513. Exochomus childreni childreni.
notypes collected together at Enterprise, Osceola, and Crescent City, Florida, which
forces me to regard them as distinct entities. Exochomus childreni does not exhibit
the secondary sexual dimorphism of E. marginipennis and is distinctly convex and
round in form with the elytral punctures very fine, sparse, often nearly invisible. In
addition, the elytral epipleuron is strongly descending externally in E. childreni, more
so than in most specimens of E. marginipennis. The western form of E. margini-
pennis, which possesses the color pattern of E. childreni, is sometimes difficult to
separate from that species, but if examples of both are present the distinction is
usually apparent. Exochomus deflectens Casey is based on a single female from
Missouri and cannot be maintained even as a subspecies. I here consider it a junior
synonym of E. marginipennis. There are several specimens in the LeConte collection
under the name C. marginipennis, but only 2 bear the orange disc indicative of the
southern states. It is not apparent from the original description how many type
specimens LeConte had, therefore I here designate and label a female labeled “(orange
disc)/E. marginipennis (Lee.) praetextus Mels” the lectotype. The type or types of C.
praetextatus Melsheimer are apparently no longer in existence, but there is no doubt
that this species is E. marginipennis.
Type locality. Of marginipennis, Georgia (lectotype here designated); of praetex-
tatus, Pennsylvania; of deflectens, Missouri.
Type depository. Of marginipennis, MCZ; of praetextatus, not located; of deflectens,
USNM (35565).
Distribution. Figure 510. New York to Florida, west to Kansas and eastern Texas.
Exochomus childreni childreni Mulsant
Fig. 513a, b; Map, Fig. 515
Exochomus childreni MulsanX, 1850, p. 1035.— Crotch, 1873, p. 377.— Crotch, 1874b,
p. 193. -Casey, 1899, p. 108. -Casey, 1908, p. 410.
Exochomus marginipennis var. childreni: Leng, 1908, p. 39. — Leng, 1920, p. 217.
Diagnosis. Length 2.80 to 3.60 mm, width 2.30 to 3.0 mm. Form rounded, slightly
oval, strongly convex. Pronotum and head black; elytron yellowish red except black
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627
Fig. 514. Exochomus childreni guexi.
spot subapically (Fig. 5 1 3a), variable (Fig. 5 1 3b). Elytral punctures fine, sparse, nearly
obsolete; epipleuron abruptly descending externally. Male and female genitalia not
separable from those illustrated for marginipennis.
Discussion. E. c. childreni appears to be restricted to Florida where the normal
color form cannot be confused with that of any other coccinellid. The 4-spotted or
4-banded forms however, resemble E. marginipennis which also occurs in Florida.
See discussion under marginipennis. One “Cotype” exists in the BM collection,
another in the Oxford University collection, not examined (R. D. Pope, pers. comm.).
Type locality. Florida.
Type depository. Lectotype not designated.
Distribution. Figure 515. FLORIDA: Apopka; Biscayne; Clay Co.; Crescent City;
Dundee; Dunedin; Enterprise; Estero; Ft. Myers; Gainesville; Haulover; Hudson;
Interlachen; Key West; Lake Co.; Miami; Orange Co.; Orlando; Osceola; Pinellas;
St. Petersburg; Tampa.
Exochomus childreni guexi LeConte, new combination
Fig. 514a, b; Map, Fig. 515
Exochomus guexi EQConiQ, 1852, p. 132. — Crotch, 1874b, p. 193.
Exochomus marginipennis var. guexi: Leng, 1920, p. 217.
Exochomus latiusculus Casey, 1899, p. 108.— Casey, 1908, p. 410.— Korschefsky,
1932, p. 264.
Exochomus marginipennis var. latiusculus: Leng, 1908, p. 40. — Leng, 1920, p. 217.
This subspecies exhibits the major color patterns of E. c. childreni (Fig. 514a, b),
and also the same secondary sexual dimorphism described forF". marginipennis. The
genitalia are not separable from those illustrated for E. marginipennis.
In spite of the lack of specimens from the area between Florida and Louisiana, I
consider E. guexi a subspecies of E. childreni rather than a distinct species. The
apparent difference between E. c. guexi and E. c. childreni is the striking sexual
dimorphism exhibited by E. c. guexi, in other morphological respects they appear
to be extremely similar. There are 5 type specimens of E. latiusculus, all labeled
“Brownsville, Texas, Wickham”. One female is here designated and labeled the
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 515. Distribution. Exochomus childreni childreni (dot); E. c. guexi (shaded, Texas); E.
fasciatus (shaded, California); E. subrotundus (open circle).
lectotype, the other 4 specimens are designated and labeled as paralectotypes. Ex-
ochomus guexi has been considered a synonym of E. childreni by previous authors
but I believe a subspecific ranking is justified. There are 4 specimens of this species
in the LeConte collection, all of which I consider types. One pin bears 2 males labeled
“(orange disc)/Type 6695 (red paper)/E. guexi LeC.”. I here designate the first male
marked with a red dot on the point as the lectotype and the other male as a para-
lectotype. The second pin bears a male and a female, both of which I designate as
paralectotypes. LeConte specifically stated that these specimens were from Louisiana.
Type locality. Of guexi, Louisiana (lectotype here designated); of latiusculus,
Brownsville, Texas (lectotype here designated).
Type depository. Of guexi, MCZ; of latiusculus, USNM (35564).
Distribution. Figure 515. Louisiana. Southern Texas.
Exochomus fasciatus Casey
Fig. 5 1 6a-e; Map, Fig. 5 1 5
Exochomus fasciatus Casey, 1899, p. 108. — Leng, 1908, p. 40.— Casey, 1908, p. 412.
Exochomus marginipennis ah. fasciatus: Weise, 1904, p. 359. — Leng, 1920, p. 217.—
Korschefsky, 1932, p. 264.
Diagnosis. Length 2.60 to 3.70 mm, width 2.20 to 2.80 mm. Form distinctly
elongate. Color pattern similar to that of E. marginipennis (Fig. 5 1 6d), or reduced
to that of typical E. childreni (Fig. 5 1 6e). Dorsal surface smooth, polished, punctures
nearly invisible; epipleuron weakly descending externally. Male genitalia as in Figure
516a-c.
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629
Fig. 516. Exochomus fasciatus.
Discussion. Exochomus fasciatus does not exhibit the secondary sexual dimor-
phism described for E. marginipennis and E. c. guexi\ both male and female have
dark legs and uniformly dark pronota. The elongate body form and highly polished
dorsal surface characterize fasciatus and serve to separate it from E. marginipennis
and E. c. guexi. I have not seen specimens of E. fasciatus from anywhere other than
California, nor have I seen examples of E. marginipennis or E. c. guexi from the
region between California and Texas.
There are 6 type specimens of fasciatus in the Casey collection. A male labeled
“Cal” is here designated and labeled lectotype. The remaining 5 specimens variously
labeled “Cal”, “Pasadena, Cal, Fall”, “Apr., Los Angeles Co. Cal”, are designated
and labeled as paralectotypes.
Type locality. California, San Diego (lectotype here designated).
Type depository. USNM (35562).
Distribution. Figure 515. Southern California.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Exochomus subrotundus Casey
Fig. 517a-g; Map, Fig. 515
Exochomus subrotundus Casey, 1899, p. 108. — Casey, 1908, p. 412. — Leng, 1920,
p. 2 1 7. — Korschefsky, 1932, p. 264.
Exochomus fasciatus var. subrotundus: Leng, 1908, p. 40.
Diagnosis. Length 2.70 to 3.40 mm, width 2.30 to 2.80 mm. Form round, convex.
Head and pronotum black, sometimes clypeus and anterolateral pronotal angle yel-
low; elytron dark brown or black with lateral border and basal spot beside suture
yellow or orange (Fig. 5171), or with both basal and apical areas yellow (Fig. 517c,
g); ventral surface brown or black except epipleuron yellow or orange. Dorsal surface
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631
smooth, polished; epipleuron abruptly descending externally. Male genitalia as in
Figure 517a-c. Female genitalia as in Figure 517d.
Discussion. Except for the entirely black species of Exochomus, E. subrotundus is
the most easily recognized member of this genus because the body shape is like that
of Brumoides hogei or B. histho. The distribution of E. subrotundus apparently does
not overlap that of either E. fasciatus or E. marginipennis, but does overlap that of
E. aethiops.
The type of E. subrotundus is a unique (holotype) male in the Casey collection.
Type locality. Texas, El Paso.
Type depository. USNM (35563).
Distribution. Figure 5 1 5. ARIZONA: Coconino Co., Page; Duncan; Grand Canyon;
Huachucha Mts.; Peach Springs; Tempe; Tucson. CALIFORNIA: Claremont; In-
dependence; Inyo Co., Inyo Mts.; Inyo Co., Saline Valley; Joshua Tree; Kem Co.,
Hobo Hot Springs; Kern Co., Tehachapi Pass; Mohave; Palmdale; Riverside Co.,
Black Hill; San Bernardino Co., Keys Ranch; San Diego Co., Warner Hot Spring;
Vidal. TEXAS: Chisos Mts.; Terrell Co., Langtry. UTAH: Chad’s Ranch.
Exochomus aethiops (Bland)
Fig. 518a-e; Map, Fig. 519
Coccinella aethiops Bland, 1864, p. 72.
Exochomus marginipennis var. aethiops: Crotch, 1873, p. 377.
Exochomus aethiops: CdiSQy, 1899, p. 109. — Chapin, 1965a, p. 249. — Gordon, 1974a,
p. 2.-Belicek, 1976, p. 320.
Exochomus (Brumus) aethiops: Leng, 1908, p. 41.
Brumus aethiops: Korschefsky, 1932, p. 265. — Hatch, 1961, p. 163.
Exochomus mormonicus Casey, 1908, p. 41 1.— Gordon, 1974a, p. 2.
Exochomus {Brumus) aethiops ab. mormonicus: Leng, 1920, p. 217.
Brumus aethiops dih. mormonicus: Korschefsky, 1932, p. 265.
Diagnosis. Length 3.0 to 4.20 mm, width 2.50 to 3.50 mm. Form rounded, convex
(Fig. 5 1 8e). Color entirely black except antenna and mouthparts yellowish brown.
Dorsal surface smooth, polished, punctures nearly invisible; epipleuron moderately
descending externally, obliquely inclined. Male genitalia as in Figure 5 1 8a-c. Female
genitalia as in Figure 5 1 8d.
Discussion. The rounded, very convex form, and strongly explanate lateral margin
of the elytron are characters which will distinguish E. aethiops from E. townsendi
which it most closely resembles. See Gordon (1974a) for further comments.
Type locality. Of aethiops. Rocky Mts., Colorado Territory; of mormonicus, Mar-
ysvale, Utah (lectotype designated by Gordon, 1974a).
Type depository. Of aethiops, PAS; of mormonicus, USNM (35556).
Distribution. Figure 519. South Dakota to New Mexico, west to Alberta, and
southern California.
Exochomus townsendi Casey
Fig. 520a-e; Map, Fig. 519
Exochomus townsendi Casey, 1908, p. 41 1. — Korschefsky, 1932, p. 264.— Black-
welder, 1945, p. 451. — Gordon, 1974a, p. 2.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 518. Exochomus aethiops.
Diagnosis. Length 2.80 to 3.30 mm, width 2.0 to 2.70 mm. Form oval, somewhat
elongate, not strongly convex (Fig. 520e). Color as described for aethiops. Dorsal
surface somewhat dull, punctures fine, barely visible; epipleuron feebly descending,
obliquely inclined. Male genitalia as in Figure 520a-c. Female genitalia as in Figure
520d.
Discussion. In addition to the comments under E. aethiops, E. townsendi differs
from that species in having the epipleuron less strongly descending, lateral margin
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633
of elytron less strongly explanate, and the sides of the pronotum and elytra appear
continuous in dorsal view, discontinuous in E. aethiops.
Type locality. Colonia Garcia, Chihuahua, Mexico (lectotype designated by Gor-
don, 1974a).
Type depository. USNM (35557).
Distribution. Figure 519. COLORADO: Buena Vista; Estes Park; Garland; Gun-
nison.
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Fig. 520. Exochomus townsendi.
Exochomus californicus Casey
Fig. 521a-c, 522a-c; Map, Fig. 524
Exochomus californicus CdiSQy, 1899, p. 107.— Casey, 1908, p. 410. — Leng, 1908, p.
40. — Hatch, 1961, p. 163. — Chapin, 1965a, p. 249.
Exochomus marginipennis var. californicus: Weise, 1904, p. 359. — Leng, 1920, p.
217.-Korschefsky, 1932, p. 263.
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NORTH AMERICAN COCCINELLIDAE
635
Diagnosis. Length 3.50 to 4.0 mm, width 2.90 to 3.20 mm. Form oval, not strongly
convex. Head and pronotum black; elytron black or brown with yellow or orange
rectangular humeral area and subapical spot (Fig. 522b, c), humeral spot sometimes
extended to apical Vs of elytron. Dorsal surface smooth, polished, punctures barely
visible; epipleuron feebly descending externally, obliquely inclined. Male genitalia
as in Figure 521a-c. Female genitalia as in Figure 522a.
Discussion. The color pattern of this species is quite popular among members of
the Chilocorini. Both Brumoides histrio and B. hogei exhibit the same pattern and
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol 93(1)
Fig. 522. Exochomus californicus (female genitalia, habitus).
care must be taken to examine the generic characters to insure correct generic place-
ment.
There are 5 type specimens of E. californicus in the Casey collection. The localities,
all in California, represented are: “Cal” (Hoopa Valley, Trinity River, Humboldt
Co.); Siskiyou Co.; Alameda Co. I here designate and so label a male from Alameda
Co. as the lectotype. The other 4 specimens are designated and labeled as paralec-
totypes.
Type locality. Alameda Co., California (lectotype here designated).
Type depository. USNM (35558).
Distribution. Figure 524. Washington to Nevada and northern California (Montana
specimen possibly mislabeled).
Exochomus quadripustulatus (L.)
Fig. 523a-e; Map, Fig. 524
Coccinella 4-pustulata L., 1758, p. 367.
Exochomus 4-pustulatus: Redtenbacher, 1843, p. 11. — Weise, 1879, p. 132.
Exochomus quadripustulatus: Mulsant, 1846, p. 172. — Mulsant, 1850, p. 485.—
Crotch, 1874b, p. 192. — Chapin, 1965a, p. 249.— Clausen et al., 1978, pp. 50, 51,
101 (for complete synonymy see Korschefsky, 1932).
Exochomus quatuorpustulatus: Korschefsky, 1932, p. 256.
Diagnosis. Length 3.60 to 4.80 mm, width 2.85 to 4.0 mm. Form oval, lateral
margin of elytron strongly beaded and somewhat explanate. Head and pronotum
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NORTH AMERICAN COCCINELLIDAE
637
Fig. 523. Exochomus quadripustulatus.
black except anterior margin and angle of pronotum narrowly yellow; elytron with
comma-shaped yellow or orange humeral spot (spot sometimes rectangular) and spot
on suture just behind middle (Fig. 523e); ventral surface black except inner ‘/2 of
epipleuron in basal Vi and broad border around abdomen reddish yellow. Dorsal
surface smooth, punctures fine, distinct; epipleuron moderately descending, obliquely
inclined. Male genitalia as in Figure 523a-c. Female genitalia as in Figure 523d.
Discussion. The similarities in color pattern and shape between this introduced
palearctic species and the native E. californicus are striking. The distributions barely
overlap however, and the key characters are sufficient for recognition. I have seen
specimens of E. quadripustulatus from only a few localities, but K. Hagen (pers.
comm.) informs me that the known distribution is from the San Francisco Bay area
south to Monterey.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 524. Distribution. Exochomus californicus (shaded, disjunct locality circled star); E.
quadripustulatus (cross hatch); E. metallicus (dot); Halmus chalybeus (star).
Type locality. “Europa”.
Type depository. Type not examined.
Distribution. Figure 524. CALIFORNIA: San Francisco Bay area south to Monterey
(coastal). Specimens examined: Hayward; San Mateo Co., Stanford; Santa Clara Co.,
Alum Rock Park, Stevens Creek Area, San Jose.
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NORTH AMERICAN COCCINELLIDAE
639
Exochomus flavipes (Thunberg)
Fig. 525a-e
Coccinella flavipes Thunberg, 1781, p. 21.
Exochomus flavipes: Crotch, 1874b, p. 192.— Sicard, 1909, p. 99.— Reitter, 1911, p.
135.-Korschefsky, 1932, p. 254.-Chapin, 1965a, p. 249. -Bartlett, et al., 1978,
pp. 69,81, 157, 167, 371.— Tassan et al., 1982, p. 1 6 (for complete synonymy see
Korschefsky, 1932).
Diagnosis. Length 3.0 to 4.0 mm, width 2.50 to 3.70 mm. Form oval, convex.
Color black (Fig. 525e) except lateral Vs of pronotum, propleuron, leg, and most of
abdomen yellow. Dorsal surface smooth, punctation fine, distinct; epipleuron de-
scending externally, nearly vertical. Male genitalia as in Figure 525a-c. Female gen-
italia as in Figure 525d.
Discussion. The entirely black dorsal surface with broadly yellow lateral margins
on the pronotum are distinctive. This is another introduced species naturally occur-
ring in Africa and the Palearctic Region which is known to be established only in
the San Francisco Bay area (K. Hagen, pers. comm.).
Type locality. Cape of Good Hope, South Africa.
Type depository. Type not examined.
Distribution. CALIFORNIA: San Francisco Bay area.
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Fig. 526. Exochomus metallicus.
Exochomus metallicus Korschefsky
Fig. 526a-e; Map, Fig. 524
Exochomus metallicus Korschefsky, 1935, p. 60.— Clausen et al., 1978, p. 151.
Diagnosis. Length 3.0 to 4.50 mm, width 2.10 to 3.50 mm. Form oval, somewhat
elongate, not strongly convex. Dorsal surface metallic green except lateral ‘A to
of pronotum yellow (Fig. 526e); ventral surface black except mouthparts, prostemum,
leg and abdomen yellow. Dorsal surface pubescent, coarsely, densely punctured;
epipleuron not strongly descending, obliquely inclined. Male genitalia as in Figure
526a-c. Female genitalia as in Figure 526d.
Discussion. This striking species resembles only one other imported species in the
North American fauna, Halmus chalybeus, which is not pubescent. Exochomus me-
tallicus is known to be established in the Santa Barbara, California, area (K. Hagen,
pers. comm.) and I have seen specimens from Santa Barbara and Oxnard.
Type locality. Abyssinia, between Addis Alam and Jem Jem, 7,000-8,000 ft.
Type depository. BMNH.
Distribution. Figure 524. CALIFORNIA: Oxnard; Santa Barbara.
Genus Halmus Mulsant
Orcus {Halmus) MuXsdinX, 1850, p. 471. — Crotch, 1874b, p. 188. — Korschefsky, 1932,
p. 249.
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NORTH AMERICAN COCCINELLIDAE
641
Orcus: Leng, 1920, p. 217.
Halmus: Weise, 1923, p. 134. — Chapin, 1965a, p. 257. Type-species; Coccinella
chalybea Boisduval, by monotypy.
Chilocorini with form nearly circular, convex, upper surface glabrous. Antenna 7-
segmented, first segment bent, apically produced in conical lobe (Fig. 527a). Apical
segment of maxillary palpus oval, apical margin oblique. Prostemal lobe broad,
truncate apically. Elytron with reflexed lateral margin; epipleuron not foveolate for
reception of femoral apices. Abdomen with 6 visible sterna in both sexes. Postcoxal
line incomplete, of Nephus type (Fig. 527b). Leg slender, femur not inflated; tibial
spur absent; tarsal claw with large basal tooth (Fig. 527c). Male genitalia symmetrical.
Female genitalia with long sperm duct; small infundibulum present (Fig. 527g).
One species, Halmus chalybeus (Boisduval), has been introduced into California
from Australia and is established (K. Hagen, pers. comm.). This species is unique
among North American Chilocorini because the dorsum is a brilliant blue or green
and lacks pubescence. Members of this genus are scale predators with specific host
records as follows. Anoidiella aurantii (Maskell), Quadraspidiotus perniciosus (Com-
stock), Pseudococcus calceolariae (Maskell), Saissetia oleae (Olivier).
Halmus chalybeus (Boisduval)
Fig. 527a-h; Map, Fig. 524
Coccinella chalybea Boisduval, 1835, p. 595.
Orcus {Halmus) chalybeus: Mulsant, 1850, p. 471.— Crotch, 1874b, p. 188.
Orcus chalybeus: Timberlake, 1920a, p. 145.
Halmus chalybeus: Weise, 1923, p. 134.— Chapin, 1965a, p. 257.
Diagnosis. Length 3.0 to 4.25 mm, width 2.85 to 3.90 mm. Form rounded, convex.
Color entirely green or blue except male head and lateral margin of pronotum yellow
(Fig. 527h); mouthparts, mesepimeron, and abdomen yellow. Male genitalia as in
Figure 527d-e. Female genitalia as in Figure 527g.
Type locality. “Nouvelle Hollande”.
Type depository. DLM.
Distribution. Figure 524. CALIFORNIA: Costa Mesa; Los Angeles; Santa Barbara.
Genus Chilocorus Leach
C hi locorus heach, 1815b, in Brewster, p. 116. — Redtenbacher, 1843, p. 11.— Mulant,
1850, p. 452. — Crotch, 1873, p. 376. — Crotch, 1874b, p. 183. — Gorham, 1892, p.
175.-Korschefsky, 1932, p. 237.-Wingo, 1952, p. 25.-Belicek, 1976, p. 318.-
Chapin, 1965a, p. 263.— J. Chapin, 1974, p. 50. Type-species; Coccinella cacti L.,
by monotypy.
Chilocorini with form broadly oval, convex, dorsal surface glabrous. Antenna 8-
segmented, club 4-segmented, fusiform (Fig. 528a). Apical segment of maxillary
palpus with lateral margins nearly parallel, apical margin strongly oblique. Prostemal
lobe flat, wide. Elytral margin not reflexed, finely beaded; epipleuron descending
externally, shallowly foveolate for reception of femoral apices. Abdomen with 6
visible sterna in male, 5 in female. Postcoxal line incomplete (Fig. 528b), merging
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 527. Halmus chalybeus. a. Antenna, b. Postcoxal lines, c. Tarsus, d-f. Male genitalia,
g. Female genitalia, h. Habitus.
with posterior margin of abdominal sternum. Leg with stout femora; tibia with
external, triangular tooth at basal third; tarsal claw with small, quadrate tooth at
base. Male genitalia with basal lobe slightly asymmetrical; trabes slender, longer than
phallobase; sipho stout, twisted near apex. Female genitalia with spermathecal capsule
large, without differentiation into nodulus and ramus, cornu short, bent, with falci-
form appendix at apex; infundibulum absent.
This is a large genus with approximately 70 species distributed worldwide. Nine
of these, including 2 species (C. kuwanae Silvestri and C. bipustulatus L.) imported
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NORTH AMERICAN COCCINELLIDAE
643
Fig. 528.
Chilocorus sp. a. Antenna, b. Postcoxal lines.
for biocontrol purposes, occur north of Mexico. Identification of the imported species
does not present a problem because both are distinctive; however, the native species
are a different matter. They are amazingly similar in appearance, and some are
apparently recognizable only through karyological studies. Smith (1959) recognized
2 such species and it is quite likely that still others occur in populations as yet untested.
Drea (1956) did a biological analysis of the California species of Chilocorini, and
was able to differentiate the species of Chilocorus that occur there. I use the conclu-
sions of Drea and Smith in this taxonomic treatment because I have not been able
to find any additional morphological criteria to aid in distinguishing the species. As
a result, it is only possible to approximate the distribution of the various species,
the degree of sympatry or allopatry cannot be determined except in a rather hazy
fashion. There are a number of specific host records available for members of Chil-
ocorus-, most are scale insects, but some species at least accept aphids as prey although
aphids may not be the preferred food. These host records are listed below. Adelgid;
Adelges picea (Ratzeburg). Aphids; Acyrthosiphon solani (Kaltenbach), Anoecia corni
(F.), Aphis cytisorum Hartig, Aphis donacis Passerini, Chromaphis juglandicola (Kal-
tenbach), Eriosoma lanigerum (Hausmann), Macrosiphum avenae (F.), Mone/lia cal-
ifornica Essig, Monellia caryae (Monell), Monellia caryella (Fitch), Myzus malisuctus
Matsumura, Phorodon humuli (Schrank), Rhopalomyzus lonicerae (Siebold), Rho-
palosiphum padi (L.), Schizaphis graminum (Rondani), Schizaphis piricola (Matsu-
mura), Toxoptera citricidus (Kirkaldy). Scales; Africaspis chionaspiformis (Newstead),
Antonina bambusae (Masked), Aonidia lauri (Bouche), Aonidiella aurantii (Masked),
Aonidiella citrina (Coquidett), Aonidiella taxus Leonard!, Aonidomytilus albus (Cock-
ered), Aspidiotus nerii Bouche, Asterolecanium coffeae Newstead, Asterolecanium
phoenicis Rao, Asterolecanium pustulans (Cockered), Aulacaspis difficilis (Cockered),
Aulacaspis rosae (Bouche), Aulacaspis tubercularis (Newstead), Ceroplastes destructor
Newstead, Ceroplastes Jloridensis Comstock, Ceroplastes japonicus Green, Cero-
plastes rubens Masked, Ceroplastes rusci (L.), Ceroplastes sinensis Del Guercio, Cer-
oplastes zonatus Newstead, Chionaspis salicis (L.), Chrysomphalus aonidum (L.),
Chrysomphalus dictyospermi (Morgan), Coccus africanus (Newstead), Coccus cole-
mani Kannan, Coccus hesperidum L., Coccus longulus (Douglas), Coccus viridis
(Green), Cryptes baccatus (Masked), Cryptococcus fagisuga Lindinger, Drosicha cor-
pulenta (Kuwana), Dysmicoccus brevipes (Cockered), Duplachionaspis saccharifolii
(Zehntner), Ehrhornia cupressi (Ehrhorn), Ericerus pela Chavannes, Eriococcus cas-
uarinae (Masked), Eriococcus coriaceus Masked, Eriococcus ironsidei Williams, Er-
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
iococcus leptospermi Maskell, Eulecanium kunoensis Kuwana, Eulecanium tiliae{L.),
Eilippia oleae Costa, Eiorinia theae (Green), Gossyparia casuarinae Maskell, Gos-
syparia spuria (Modeer), Hemiberlesia lataniae (Signoret), Hemiberlesia rapax (Com-
stock), Icerya purchasi Maskell, Inchoaspis dentilobis (Newstead), Inglisia conchi-
formis Newstead, Ischnaspis longirostris (Signoret), Kermes ilicis (L.), Kermes miyasakii
Kuwana, Kermes nakagawae Kuwana, Lepidosaphes afganensis Borchsenius, Lepi-
dosaphes beckii (Newman), Lepidosaphes conchiformis (Gmelin), Lepidosaphes glov-
erii (Packard), Lepidosaphes olivina Leonard), Lepidosaphes ulmi (L.), Leucaspis sp.,
Lineaspis striata (Newstead), Mesolecanium nigrofasciatum (Pregande), Miscan-
thaspis tegalensis (Zehntner), Monophlebulus sp., Nelaspis humilis (Brain), Nipae-
coccus aurilanatus (Maskell), Nipaecoccus filamentosus (Cockerell), Nipaecoccus ni-
pae (Maskell), Paralecanium frenchii (Maskell), Parlatoria blanchardi (Targioni-
Tozzetti), Parlatoria oleae (Colvee), Parlatoria pergandii Comstock, Parlatoria pro-
teus (Curtis), Parlatoria ziziphi Lucas, Parthenolecanium corni Bouche, Partheno-
lecanium persicae (F.), Parthenolecanium quercifex (Fitch), Phenacaspis grandilobis
(Maskell), Phenacoccus solani Ferris, Pinnaspis strachani (Cooley), Planococcus citri
(Risso), Planococcus kenyae (LePelley), Planococcus lilacinus (Cockerell), Pollinia
pollini (Costa), Protopulvinaria mangiferae (Green), Pseudococcus longispinus (Tar-
gioni-Tozzetti), Pseudaonidia duplex (Cockerell), Pseudaonidia paeoniae (Cockerell),
Pseudoparlatoria ostreata (Cockerell), Pulvinaria aurantii (Cockerell), Pulvinaria
maxima Green, Pulvinaria okitsuensis Kuwana, Pulvinaria psidii Maskell, Quadras-
pidiotus ostreaeformis (Curtis), Saissetia cojfeae (Walker), Saissetia oleae (Olivier),
Unaspis yanonensis (Kuwana).
Key to species of Chilocorus
1 . Elytron brown or reddish brown with 3 small discal spots in transverse row, usually
partially fused (Fig. 537d) bipustulatus (L.)
Elytron black or brown with one yellow or red spot (Fig. 536d) 2
2(1). Spot on elytron median or slightly behind middle (Fig. 536d); form orbicular ....
kuwanae Silvestri
Spot on elytron anterior to middle (Fig. 53 Id); form elongate or orbicular 3
3(2). Venter mostly yellow or red, only prostemum black 4
- Venter mostly black, only abdomen red or yellow 5
4(3). Form orbicular, strongly convex; spot on elytron small (Fig. 53 Id); mid Atlantic
states tumidus Leng
- Form somewhat elongate, not strongly convex; spot on elytron large (Fig. 529e);
extreme southern United States cacti (L.)
5(3). Species known only from California 6
Species not occurring in California 7
6(5). Paramere with setae long, approximately 103 to 159 setae per paramere, setae in
inner margin extending into basal V2 (Fig. 532a) orbus Casey
- Paramere with setae short, approximately 47 to 88 setae per paramere, setae on
inner margin confined to apical ‘/2 fraternus LeConte
7(5). Species occurring from the Atlantic coast to the Rocky Mts. in Alberta, and the
Sierra Nevada in the United States; karyotype 2n = 22 + s stigma (Say)
- Species occurring from southern Alberta and Saskatchewan to British Columbia . 8
8(7). Karyotype 2n = 14 hexacyclus Smith
Karyotype 2n = 20 tricyclus Smith
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NORTH AMERICAN COCCINELLIDAE
645
Fig. 529. Chilocorus cacti.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 530. Distribution. Chilocorus cacti (shaded, southern); C tumidus (star); C. orbus (shad-
ed, west coast).
Chilocorus cacti (L.)
Fig. 529a-e; Map, Fig. 530
Coccinella cacti L., 1767, p. 584. — Say, 1835, p. 202.
Chilocorus cacti: Mulsant, 1850, p. 459. — Crotch, 1873, p. 376.— Crotch, 1874b, p.
184.-Gorham, 1892, p. 175.-Leng, 1908, p. 38. -Casey, 1908, p. 408. -Kor-
schefsky, 1932, p. 245.
Chilocorus confusor Casey, 1899, p. 105. — Casey, 1908, p. 408.
Chilocorus cacti var. confusor: Leng, 1908, p. 38. — Leng, 1920, p. 217.— Korschefsky,
1932, p. 246.
Chilocorus cacti confusor: Drea, 1956, p. 76.
Diagnosis. Length 4.0 to 6.20 mm, width 3.60 to 5.20 mm. Form oval, convex.
Color black except large transverse spot on elytron (Fig. 529e) and mesosternum,
metasternum, and abdomen yellow or red. Dorsal surface smooth, polished, punc-
tures fine, distinct. Male genitalia as in Figure 529a-c. Female genitalia as in Figure
529d.
Discussion. The red or yellow ventral surface (except prosternum) will separate C.
cacti from C. fraternus and allies which have the mesosternum and metastemum
black. This species occurs primarily in northern South America, Central America,
Mexico, and the Caribbean islands, with the northern range limit in the southern
United States. There are 2 type specimens of C. confusor in the Casey collection, I
here designate and label a male as the lectotype, and the other specimen as a para-
lectotype.
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NORTH AMERICAN COCCINELLIDAE
647
Fig. 531. Chilocorus tumidus.
Type locality. Of cacti, “America”, of confusor, San Diego, California (lectotype
here designated).
Type depository. Of cacti, type not examined; of confusor, USNM (35552).
Distribution. Figure 530. Southern Arizona, California, Florida, and Texas.
Chilocorus tumidus Leng
Fig. 531a-d; Map, Fig. 530
Chilocorus tumidus Leng, 1908, p. 37. — Korschefsky, 1932, p. 247.
Diagnosis. Length 4.20 to 5.75 mm, width 3.75 to 5.0 mm. Form orbicular, ex-
tremely convex, not tapered posteriorly. Color dark brown to black except elytron
with transverse spot (Fig. 53 Id) and mesosternum, metasternum, and abdomen
yellow or red. Dorsal surface smooth, polished, punctures fine, distinct. Male genitalia
as in Figure 531a-c.
Discussion. This species resembles C. cacti which does not occur in the same
geographic region. In C. tumidus, the elytral spots are much smaller and the form is
distinctly rounded and strongly convex, while C. cacti is slightly elongate and not as
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strongly convex. There are 2 female type specimens of C. Xumidus in the USNM
collection, one of which I designate and label the lectotype and the other as a para-
lectotype.
Type locality. Fortress Monroe, Virginia (lectotype here designated).
Type depository. USNM (40901).
Distribution. Figure 530. MARYLAND: 2 mi. N. of Priest Bridge. VIRGINIA:
type locality.
Chilocorus orbus Casey
Fig. 532a-d; Map, Fig. 530
Chilocorus orbus Casey, 1899, p. 105.— Casey, 1908, p. 408. — Drea, 1956, p. 58.—
Smith, 1959, p. 445. -Hatch, 1961, p. 163.
Chilocorus bivulnerus var. orbus: Leng, 1908, p. 37. — Korschefsky, 1932, p. 247.
Chiloocorus stigma (Say): Korschefsky, 1932, p. 246 (in part).
Diagnosis. Length 4.0 to 5.10 mm, width 3.30 to 4.50 mm. Form oval, slightly
tapered posteriorly, convex. Color black except spot on elytron (Fig. 532d) and
abdomen yellow or red. Dorsal surface smooth, polished, punctures fine, distinct.
Male genitalia as in Figure 532a-c.
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NORTH AMERICAN COCCINELLIDAE
649
Fig. 533. Chilocorus fraternus.
Discussion. Male genitalia must be examined to separate C. orbus and C. fraternus
(see key to species). The California species of Chilocorus, except for cacti, have been
identified as C. stigma Say or C. bivulnerus Mulsant since Leng’s publication in 1908.
Drea (1956) and Smith (1959) have demonstrated that both C. orbus and C. fraternus
are valid species and that C. stigma does not occur in California. There are 3 type
specimens of C. orbus in the Casey collection; I here designate and label a male the
lectotype, the other 2 specimens as paralectotypes.
Type locality. California (lectotype here designated).
Type depository. USNM (35553).
Distribution. Figure 530. Western Washington and Oregon to southern California.
Chilocorus fraternus LeConte
Fig. 533a-c
Chilocorus fraternus LeConte, 1860, (1857), p. 70.— Casey, 1899, p. 105.— Casey,
1908, p. 408.-Drea, 1956, p. 70.-Smith, 1959, p. 445. -Hatch, 1961, p. 162.-
Smith, 1965, p. 1614.
Chilocorus bivulnerus var. fraternus: Crotch, 1873, p. 376. — Leng, 1920, p. 217.
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Fig. 534.
Chilocorus stigma.
Chilocorus bivulnerus Mulsant: Leng, 1908, p. 37 (in part).
Chilocorus stigma (Say): Korschefsky, 1932, p. 246 (in part).
Diagnosis. Length 3.40 to 5.10 mm, width 3.0 to 4.30 mm. Description as for C.
orbus. Male genitalia as in Figure 533a-c.
Discussion. The number and distribution of the setae on the parameres are diag-
nostic characters for this species (see key to species and comments under orbus). The
actual distribution of C. fraternus is not known. Drea (1956) stated that “It appears
that this species is found chiefly in the Central Valley of the state” (California). Hatch
(1961) reported it as “common” in the Pacific Northwest. Since Hatch did not
examine the male genitalia this may or may not be the case. Until the genitalia of
all available specimens are examined, and until karyotype studies are carried out to
determine the distribution of C. tricyclus, the actual distribution of any of these
species cannot be accurately stated.
Type locality. Sacramento, California.
Type depository. MCZ.
Distribution, (see comments above) CALIFORNIA: Central Valley. PACIFIC
NORTHWEST: ?
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NORTH AMERICAN COCCINELLIDAE
651
Chilocorus stigma (Say)
Fig. 534a-e; Map, Fig. 535
Coccinella stigma Say, 1835, p. 202.
Chilocorus stigma: Melsheimer, 1853, p. 130.— Mulsant, 1856, p. 148.— Korschef-
sky, 1932, p. 246.-Smith, 1959, p. 445. -J. Chapin, 1974, p. 50.-Belicek, 1976,
p. 319.
Chilocorus bivulnerusMu\s3.nX, 1850, p. 460. — Mulsant, 1856, p. 148. — Crotch, 1873,
p. 376. -Crotch, 1874b, p. 185. -Casey, 1899, p. 105.-Leng, 1908, p. 37.-
Wingo, 1952, p. 47.
Diagnosis. Length 3.75 to 5.0 mm, width 3.0 t 4.25 mm. Form oval, tapered
slightly posteriorly, convex. Color black except spot on elytron (Fig. 534e) and ab-
domen yellow or red. Dorsal surface smooth, punctures larger, denser than in C.
orbus. Male genitalia as in Figure 534a-d. Karyotype: 2n = 22 + s. Polymorphic:
males, 2n = 19-25; females, 2n = 20-26.
Discussion. Two species, cacti and tumidus, occur in parts of the eastern range of
C. stigma. Both of these species have yellow or red meso- and metasterna, in C.
stigma these areas are black or at least brown. Chilocorus stigma apparently does
not occur west of the Rocky Mountains in Alberta or the Sierra Nevada in the United
States. In Alberta it is sympatric with C hexacyclus from which C stigma can be
separated by karyotype.
Type locality. Not stated.
Type depository. DLM (not examined).
Distribution. Figure 535. Nova Scotia to Florida, west to Alberta and Arizona.
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Chilocorus hexacyclus Smith
Chilocorus hexacyclus Smith, 1959, p. 446. — Belicek, 1976, p. 319.
Morphologically not separable from C. stigma. Karyotype: 2n = 14. Meiotic for-
mula—6 ring II + 1; neo-XY (male): XX (female) II.
Type locality. Conquest, Saskatchewan (subsequently established by Belicek, 1976).
Type depository. CNC.
Distribution. ALBERTA: “eastern foothills of the Rocky Mountains”. SAS-
KATCHEWAN: type locality.
Chilocorus tricyclus Smith
Chilocorus tricyclus Smith, 1959, p.446. — Hatch, 1961, p. 163. — Belicek, 1976, p.
319.
Morphologically not separable from stigma. Karyotype: 2n = 20. Meiotic for-
mula-3 ring II + 6 non-ring II; II + 1 neo-XY (male): XX (female) II.
Type locality. Grand Forks, British Columbia (subsequently established by Belicek,
1976).
Type depository. CNC.
Distribution. BRITISH COLUMBIA: “interior of British Columbia”. WASHING-
TON: Seattle.
Chilocorus kuwanae Silvestri
Fig. 536a^
Chilocorus kuwanae Silvestri, 1909, p. 126. — Korschefsky, 1932, p. 240.— Mader,
1955, p. 781.-Kamiya, 1959, p. lOl.-Sasaji, 1971a, p. 226.
Chilocorus similis: Lewis, 1896, p. 31 (not Rossi, 1790) (misidentification).—
Essig, 1931, p. 289.— Clausen, 1956, p. 2. — Smith, 1965, p. 1614. — Clausen et al.,
1978, pp. 81, 121, 123, 125.
Chilocorus similis y?lt. japonicus Sicard, 1907, p. 21 1.— Sasaji, 1971a, p. 226.
Chilocorus renipustulatus: Lewis, 1873, p. 56 (not renipustulatus Scriba, 1792).—
Sasaji, 1971a, p. 226.
Diagnosis. Length 3.0 to 4.75 mm, width 2.90 to 4.50 mm. Form orbicular, lateral
margin of elytron broadly explanate. Color black except spot on elytron at or behind
middle (Fig. 536d) and abdomen yellow or red. Dorsal surface smooth, polished,
punctures fine, distinct. Male genitalia as in Figure 536a-c.
Discussion. This species, imported from Japan, is very similar in appearance to
the native species of Chilocorus (fraternus, orbus, etc.), but the spots on the elytra
are located at the middle or slightly behind the middle of each elytron, and the lateral
margin of the elytron is more strongly flared. The name applied to the species in-
troduced into California as C. similis Rossi has been the subject of debate for some
years. Chilocorus kuwanae is the name in current useage by Japanese scientists, and
is the correct name. Type specimens of C. similis have been examined and compared
with examples of C. kuwanae. The head of C. similis is somewhat shiny with distinctly
separated punctures; the pronotum has the punctures coarse and quite close together;
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NORTH AMERICAN COCCINELLIDAE
653
Fig. 536. Chilocorus kuwanae.
and the lateral margin of the elytron feebly explanate. The head of C. kuwanae is
dull, strongly alutaceous with punctures extremely close together; the pronotumm
has the punctures fine and widely separated; and the lateral margin of the elytron
widely explanate. The type series consists of 2 female specimens, one of which, labeled
“28030/Typus(red paper)/ Affinis R Etrur Rossi/Zool. Mus. Berlin“ I designate and
label as the lectotype. The other specimen, labeled ’’affinis Ross Etr/Etrur Rossi Hist.
Coll. No. 28030“, is designated as a paralectotype.
Type locality. ’’Cine e Giappone“.
Type depository. MNHUB.
Distribution. CALIFORNIA: Santa Barbara Co. (K. Hagen, pers. comm.).
Chilocorus bipustulatus (L.)
Fig. 537a-d
Coccinella bipustulata L., 1758, p. 367.
Chilocorus bipustulatus: Mulsant, 1846, p. 170.— Crotch, 1874b, p. 185.— Smith,
1915, p. 523.-Essig, 1931, p. 291. -Clausen et. al., 1978, pp. 69, 80, 84, 85, 100,
101, 113, 115, 116, 121.
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Fig. 537. Chilocorus bipustulatus.
Diagnosis. Length 3.0 to 4.50 mm, width 3.10 to 4.0 mm. Form oval, tapered
posteriorly, moderately convex. Color light to dark brown dorsally, elytron with
narrow, irregular band of 3 partially connected spots on disc, external spot often free
(Fig. 537d), venter light brown to black except prostemum, mesosternum, and meta-
stemum usually black. Dorsal surface smooth, polished, punctures fine, distinct. Male
genitalia as in Figure 537a-c.
Discussion. The dorsal color pattern is diagnostic, being unlike that of any other
imported or native species. This species has been imported from Europe and is now
established in the San Joaquin Valley of California (pers. comm. K. Hagen). Huffaker
and Doutt (1965) state that C. bipustulatus is established in 3 California counties;
Fresno, Madera, and Merced.
Type locality. ”Europa“.
Type depository. Type not examined.
Distribution. CALIFORNIA: San Joaquin Valley.
Subfamily Coccidulinae
Coccidulinae Sasaji, 1968, p. 22.— J. Chapin, 1974, p. 52. — Belicek, 1976, p. 293.
Trichosomides Mulsant, 1846, p. 27. — Mulsant, 1850, p. 696 (in part).
Coccinellidae with dorsum weakly or moderately convex, pubescent. Head capsule
normal; apex truncate; clypeus expanded or not; compound eye sometimes coarsely
faceted; apical segment of maxillary palpus usually strongly divergent apically and
securiform. Antenna 8 to 1 1 -segmented. Meso- and metasternum narrowly articu-
lated. Epipleuron usually broad and entire without distinct foveae (except Azyini).
Female genital plates very elongate.
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NORTH AMERICAN COCCINELLIDAE
655
Sasaji (1968) included 4 tribes in this subfamily, all of which are represented in
North America except the tribe Lithophilini. He did not mention the tribe Azyini
which I here include in this subfamily. The host preferences of the New World
members of the Coccidulinae are not well known, but they are apparently scale
predators for the most part.
Key to tribes of Coccidulinae
1 . Abdomen with 5 visible sterna 2
Abdomen with 6 visible sterna 3
2( 1 ). Epipleuron with deep foveae for reception of femoral apices of middle and hind legs;
eye undivided Azyini
Epipleuron not foveate; eye almost completely divided (Fig. 550a) Exoplectrini
3(1). Antenna 8-segmented, weakly clubbed, short (Fig. 546a); tarsus trimerous; apex of
clypeus thickened, narrower than labrum (Fig. 547a) Noviini
- Antenna 10 or 1 1 -segmented, strongly clubbed, long (Fig. 538a); tarsus cryptotetra-
merous; apex of clypeus not thickened, wider than labrum Coccidulini
Tribe Coccidulini
Coccidulini Costa, 1849, p. 9, 104.— Casey, 1899, p. 74, 162. — Korschefsky, 1931,
p. 80.-Sasaji, 1968, p. 23.-J. Chapin, 1974, p. 52.-Belicek, 1976, p. 293.
Rhizobiares Mulsant, 1846, p. 261. — Mulsant, 1850, p. 938.
Cocciduliens Mulsant, 1846, p. 266. — Mulsant, 1850, p. 1007.
Coccidulides Crotch, 1873, p. 363.
Rhizobiides Crotch, 1874, p. 288.
Rhizobiini Weise, 1885a, p. 6.— Casey, 1899, p. 161.
Rhizobiinae Della Beffa, 1912, p. 167.
Coccidulina Jacobson, 1916, p. 969.
Coccidulinae of widely varying size, length ranging from 2.0 to 7.50 mm; form
elongate, slender, or oval. Gena not extending onto eye; eye finely or coarsely faceted.
Antenna inserted laterally beside eye, insertion exposed, 1 0 or 11 -segmented, very
elongate with loose 3-segmented club. Prostemum with intercoxal process narrow,
bicarinate. Leg slender, not angulate or dentate. Tarsus cryptotetramerous. Abdomen
with 6 visible sterna.
Two genera represent this tribe in America north of Mexico. One genus, Coccidula,
is European with one American representative; the other, Rhyzobius, is represented
by 2 species introduced from Australia as biocontrol agents. The long, slender an-
tenna, slender, unarmed legs and undivided eye distinguish this tribe from the Noviini
and Exoplectrini to which it is most closely related. There are many Old World genera
in the Coccidulini and it is probable that this tribe will be further divided when these
genera have been completely studied.
Key to genera of Coccidulini
1. Eye extremely coarsely faceted; dorsal pubescence uniform, decumbent
Coccidula Kugelann
- Eye moderately coarsely faceted; dorsal pubescence composed of mostly decumbent
hairs with some long, erect hairs scattered throughout Rhyzobius Stephens
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 538. Coccidula lepida. a. Antenna, b. Maxillary palpus, c. Prostemum. d. Postcoxal
lines.
Genus Coccidula Kugelann
Coccidula Kugelann, 1798, p. 421. — Mulsant, 1850, p. 1007. — LeConte, 1852, p.
130. — Crotch, 1874b, p. 300. — Casey, 1899, p. 162.— Korschefsky, 1931, p. 81.—
Wingo, 1952, p. 22. — Belicek, 1976, p. 321. Type-species; Chrysomela scutellata
Herbst, 1783, by subsequent designation of Crotch, 1874b.
Strongylus Panzer, 1 8 1 3, p. 114.
Cacidula Curtis, 1827, p. 114.
Cacicula Stephens, 1828, p. 319.
Coccidulini with form extremely elongate, slender (Fig. 5391); dorsal pubescence
short, decumbent. Antenna long, slender, 1 1 -segmented, club serrate, 3-segmented
(Fig. 538a). Head mostly exposed; eye coarsely faceted; apical segment of maxillary
palpus securiform (Fig. 538b). Prostemum with carinae joined apically (Fig. 538c).
Epipleuron narrow, not descending externally. Tarsal claw with feeble tooth. Po-
stcoxal line complete, as in Pullus (Fig. 538d). Male genitalia symmetrical; female
genitalia lacking infundibulum (Fig. 539e).
The key characters will separate Coccidula from Rhyzobius\ in addition, Coccidula
has the prostemal carinae joined apically, the body is more slender and elongate and
the head mostly exposed. Most species of this genus are European with one, lepida
LeConte, occurring in North America. Members of this genus are said to be scale
predators, but I have not seen any host data to prove or disprove this statement.
There has been no modern taxonomic treatment of Coccidula.
Coccidula lepida LeConte
Fig. 539a-g; Map, Fig. 540
Coccidula lepida LeConte, 1852, p. 132.— Crotch, 1874b, p. 301. — Horn, 1895, p.
113.— Casey, 1899, p. 162. — Korschefsky, 1931, p. 82. — Dodge, 1938, p. 222.—
Wingo, 1952, p. 45.
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NORTH AMERICAN COCCINELLIDAE
657
Fig. 539. Coccidula lepida.
Coccidula lepida var. suturalis Weise, 1895, p. 132.— Casey, 1899, p. 163.— Kor-
schefsky, 1931, p. 82.— Dodge, 1938, p. 221.
Coccidula occidentalis Horn, 1895, p. 114.— Weise, 1898c, p. 238.— Frost, 1920, p.
231.-Korschefsky, 1931, p. 82.-Dodge, 1938, p. 222.-Belicek, 1976, p. 321.
New Synonymy.
Coccidula suturalis Reiner, 1897, p. 127 (not suturalis Weise, 1895).
Coccidula suturalis: Leng, 1920, p. 215.— Dodge, 1938, p. 222.— Wingo, 1952, p.
22.
Coccidula reitteri Dodge, 1938, p. 222 (unnecessary replacement name for suturalis
Reitter, 1897).
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Diagnosis. Length 2.75 to 3.45 mm, width 1.50 to 1.85 mm. Form extremely
elongate, parallel sided. Color yellow; head, pro-, meso- and metasterna, and basal
2 abdominal sterna black; epipleuron piceous; elytron yellow with black maculation
as in Figure 539f, color pattern variable (Fig. 539g). Male genitalia as in Figure 539a-
d. Female genitalia as in Figure 539e.
Discussion. The generic characteristics are sufficient to distinguish this species
because it does not closely resemble any other North American species. This species
is most similar to the European C scutellata (Herbst), but the male genitalia are
quite different in each species. Coccidula occidentalis Horn {suturalis Weise) is a
junior synonym of lepida. The male genitalia are identical in the nominate forms
and the dorsal color patterns are almost identical except that the apical spots are
connected to the base along the suture in C. occidentalis (Fig. 539f). LeConte stated
that he had one type specimen of C. lepida. That specimen, a female, labeled “(white
disc)/4641/Type 6748 (red pdLpQv)/ coccidula lepida LeC.” must be considered the
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NORTH AMERICAN COCCINELLIDAE
659
Fig. 541. Rhyzobius sp. a. Antenna, b. Maxillary palpus, c. Prostemum. d. Tarsus, e. Po-
stcoxal lines.
holotype. Horn (1895) had several specimens when he described C. occidentalism 6
of which are now in the Horn collection. The first of these, a female labeled “Van./
Lectotype 32 1 7/C. occidentalis Horn” is here designated and labeled as the lectotype,
the remaining 5 are designated as paralectotypes.
Type locality. Of lepida, Vermont; of occidentalis, Vancouver, B.C. (lectotype here
designated).
Type depository. Of lepida and occidentalis, MCZ.
Distribution. Figure 540. Quebec to New Jersey, west to Alaska and Colorado.
Genus Rhyzobius Stephens
Rhyzobius Stephens, 1829, p. 239. — Stephens, 1832, p. 396. — Pope, 1981, p. 22.
Type-species; Nitidula litura F., 1787, by monotypy.
R/z/z(9/7/w5 Stephens, 1832, p. 373 (error). — Leng, 1920, p. 214. — Korschefsky, 1931,
p. 88.
Rhizobius Agassiz, 1846, p. 325 (unjustified emendation).
Casey, 1899, p. 161. — Leng, 1920, p. 214.— Chapin, 1974, p. 52.— Pope,
1981, p. 22. Type-species; Scymnus lophanthae Blaisdell, 1892, by monotypy.
Rhizobiellus Oke, 1 95 1 , p. 2 1 (unnecessary replacement name for Rhizobius Agassiz,
1846 (not Burmeister, 1835).
Coccidulini with form elongate or oval; dorsal pubescence composed of dense,
decumbent hairs with sparse, erect hairs intermixed. Antenna long, slender, 1 1 -
segmented, club serrate (Fig. 541a). Head partly concealed beneath pronotum; eye
moderately coarsely faceted; apical segment of maxillary palpus securiform (Fig.
541b). Prostemum with carinae widely separated, usually not joined apically (Fig.
541c). Epipleuron narrow, not descending externally. Tarsus cryptotetramerous; tar-
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
sal claw not toothed, slightly angulate at base (Fig. 54 Id), or appendiculate at least
on hind leg. Postcoxal line on first abdominal sternum complete, as in Pullus (Fig.
541e).
The partly concealed head and dual type of pubescence distinguish Rhyzobius from
the genera of North American Scymnini which it superficially resembles. This is an
Old World genus with 2 species having been introduced and established in North
America for biocontrol purposes. These species have been considered as belonging
in separate genera, Lindorus Casey and Rhyzobius, but Pope (1981) synonymized
Lindorus with Rhyzobius. Species of Rhyzobius (at least the Australian species) are
apparently scale feeders on such species as Aonidiella aurantii (Masked), Chrysom-
phalus dictyospermi (Morgan), Coccus hesperidum L., Fiorinia theae Green, Piano-
coccus citri (Risso), Pseudaulacaspis pentagona (Targioni-Tozzetti), Pseudococcus
acaciae (Masked), Pseudococcus calceolariae (Masked), Pseudococcus maritimus
(Ehrhorn), Quadraspidiotus perniciosus (Comstock), Saissetia oleae (Olivier). The
European Rhyzobius litura (F.) has been recorded as feeding on the aphids Dactynotus
cirsii (L.), Dactynotus jaceae (L.), Macrosiphum avenae (F.).
Key to species of Rhyzobius
1. Ventral surface and leg black except abdomen red forestieri (Mulsant)
- Ventral surface including leg red or yellow lophanthae (Blaisdell)
Rhyzobius lophanthae (Blaisdell)
Fig. 542a-e; Map, Fig. 543
Scymnus lophanthae 1892, p. 51. — Riley, 1892, p. 127.
Rhizobius lophanthae: Horn, 1895, p. 112. — Essig, 1911, p. 518. — Weise, 1923, p.
149.
Lindorus lophanthae: Casey, 1899, p. 162. — Korschefsky, 1931, p. 86.— Clausen,
1956b, p. 109.— J. Chapin, 1974, p. 53.
Lindorus lophantae: Leng, 1920, p. 214 (misspelling).
Rhizobius toowoombae Blackburn, 1892, p. 254. — Horn, 1895, p. 112.
Rhyzobius lophanthae: Pope, 1981, p. 22.
Diagnosis. Length 1.70 to 2.85 mm, width 1.35 to 2.0 mm. Form elongate, oval
(Fig. 542e). Color yellowish brown; pronotum light reddish brown, elytron dark
reddish brown with faint, green metallic tint. Male genitalia as in Figure 542a-c.
Female genitalia as in Figure 542d.
Discussion. This species was introduced into California from Australia in 1892 for
control of the black scale. It has been highly successful against a variety of scales,
not only in California, but in other areas of the United States as indicated in Figure
543. Blaisdell stated that he had many specimens of this species and one, a female
labeled “Coronado, Cal., XI-3-90/F.E. Blaisdell collector/female sign/Blaisdell col-
lection/Allotype lophanthe Blais, (red paper)”, is designated as the lectotype. Eight
other specimens, all bearing the same locality data, are designated and labeled as
paralectotypes.
Type locality. Coronado, California (lectotype here designated).
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NORTH AMERICAN COCCINELLIDAE
661
Fig. 542. Rhyzobius lophanthae.
Type depository. CAS.
Distribution. Figure 543. Maryland to Florida, west to California.
Rhyzobius forest ieri (Mulsant)
Fig. 544a-e, 545; Map, Fig. 543
Platyomus forestieri Mulsant, 1853, p. 158.
Scymnodes forestieri: Korschefsky, 1931, p. 85.
Scymnus circularis Sharp, 1889, p. 365. — Pope, 1981, p. 26.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 543. Distribution. Rhyzobius lophanthae (shaded, southern); R. forestieri (cross hatch).
Rhizobius ventralis: sensu auct.
Lindorus ventralis: Timberlake, 1927, p. 532.
Rhyzobius forestieri: Pope, 1981, p. 26.
Diagnosis. Length 2.60 to 3.70 mm, width 1.90 to 2.30 mm. Form oval, lateral
border of pronotum and elytron discontinuous (Fig. 544e). Color black except mouth-
parts and abdomen yellow or reddish. Male genitalia as in Figure 544a-c. Female
genitalia as in Figure 555.
Discussion. Rhyzobius forestieri was introduced into California in 1892 under the
name R. ventralis Erichson for control of lecaniine scales (Coccidae). It was released
at Santa Barbara and San Jose and is now established in coastal California. Unlike
R. lophanthae, it apparently has not spread to other parts of the United States.
Type locality. Australia.
Type depository. Of forestieri, PM; of circularis, BMNH.
Distribution. Figure 543. Coastal California.
Tribe Noviini
Noviini Gangelbauer, 1899, p. 954. — Leng, 1920, p. 214. — Mader, 1924, p. 7.—
Korschefsky, 1931, p. 96.— Sasaji, 1968,p. 26.— Gordon, 1972a, p. 23.— J. Chapin,
1974, p. 53.-Belicek, 1976, p. 293.
Coccidulinae with form broad, somewhat oblong; dorsal surface pubescent. Head
directed ventrally, not deeply inserted in pronotum; clypeus thick with labrum on
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NORTH AMERICAN COCCINELLIDAE
663
Fig. 544. Rhyzobius forestieri.
lower plane than clypeus; gena not extending onto eye. Antennal insertion concealed;
antenna 8-segmented with club weakly 3-segmented, basal 2 segments large. Apical
segment of maxillary palpus large, strongly securiform. Epipleuron obliquely inclined
with external margin lower than internal margin, not foveate for reception of legs.
Prostemum raised medially, protuberant, narrowly separating anterior coxae. An-
terior femur deeply emarginate for reception of tibia, middle and hind femora less
so; tibia on all legs compressed laterally, with weak external angulation at basal ‘/a;
tarsus trimerous. Abdomen with 6 visible sterna; postcoxal line on first sternum
complete or nearly so; apex of 6th sternum in male emarginate medially. Male
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
genitalia symmetrical. Female genitalia with spermathecal capsule short, stout, lack-
ing infundibulum.
There are 2 genera of this tribe in the New World, one (Anovia) native and one
(Rodolia) introduced for biocontrol purposes. The 6-segmented abdomen, protuber-
ant prostemal process, and unequal planes of the clypeus and labrum distinguish the
Noviini. The New World members of the tribe were treated by Gordon (1972a).
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NORTH AMERICAN COCCINELLIDAE
665
Fig. 546. Rodolia cardinalis. a. Antenna, b. Maxillary palpus, c. Postcoxal lines.
Key to genera of Noviini
1. Postcoxal line complete (Fig. 546c); labrum flat, slightly emarginate anteriorly (Fig.
547a) Rodolia Mulsant
- Postcoxal line incomplete (Fig. 549e); labrum convex, apical margin broadly, feebly
concave Anovia Casey
Genus Rodolia Mulsant
RodoliaM.\x\s?inX, 1850, p. 902.— Crotch, 1874b, p. 280. — Korschefsky, 1931,p. 98.—
Gordon, 1972a, p. 25.— J. Chapin, 1974, p. 53. Type-species; Rodolia ruficollis
Mulsant, by subsequent designation of Crotch, 1874b.
Rodolia {Macronovius)V^Q\SQ, 1885a, p. 63.— Weise, 1895b, p. 149. — Sicard, 1907b,
p. 68.
Noviini with antenna as in Figure 546a. Labrum flat (Fig. 547) or concave, anterior
margin usually feebly emarginate. Maxillary palpus as in Figure 546b. Prosternal
protuberance margined apically, densely pubescent. Abdomen with postcoxal line
on first sternum complete (Fig. 546c); 6th sternum of male with apical emargination
strong.
See Gordon (1972a) for a more detailed discussion of this genus in the New World.
The tribe is composed of a group of very closely related genera and it is difficult to
separate them satisfactorily except in the larval stage. The postcoxal line is definitely
complete in Rodolia, narrowly incomplete in Anovia. Rodolia cardinalis, the only
species of Rodolia occurring in the United States, was introduced into California
from Australia in 1888 for control of the cottony cushion scale, Icerya purchasi
Maskell. It has since been introduced and become established in many parts of the
world. Rodolia koebelei (Coquillett) was also introduced into California from Aus-
tralia in 1891, but does not exist there now, although it was thought to have been
established for some time.
Species of Rodolia prey primarily on scales of the genus Icerya. Recorded hosts
include Icerya purchasi Maskell, Icerya seychellarum (Westwood), Pseudococcus sp.,
Pseudaulacaspis pentagona (Targioni-Tozzetti).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 547. Rodolia cardinalis.
Rodolia cardinalis (Mulsant)
Fig. 547a-g; Map, Fig. 548
Vedalia cardinalis Mulsant, 1850, p. 906.
Rodolia cardinalis: Weise, 1905, p. 220.— J. Chapin, 1974, p. 54. (for detailed syn-
onymy see Gordon, 1972a, p. 25).
Diagnosis. Length 2.65 to 4. 1 8 mm, width 2.00 to 3.33 mm. Form elongate, elytron
nearly parallel sided, widest at middle. Color red; basal area of pronotum and head
black; meso- and metasternum, femur and median area of basal 2 abdominal sterna
piceous; elytron with black maculation (Fig. 547g). Male genitalia as in Figure 547b-
e. Female genitalia as in Figure 547f
Discussion. See Gordon (1972a) for a discussion of this species. In addition to the
characters used in the generic key, R. cardinalis can usually be distinguished from
A. virginalis by body form. Anovia virginalis is definitely widest just posterior to the
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NORTH AMERICAN COCCINELLIDAE
667
Fig. 548. Distribution. Rodolia cardinalis (shaded, disjunct localities circled star); Anovia
virginalis (star); Exoplectra schaefferi (open star); Azya orbigera (dot).
humeral callus; R. cardinalis is widest at the middle of the elytra. This species is
definitely established in California and Florida. The data listed is from actual spec-
imens examined.
Type locality. New Holland.
Type depository. Oxford University.
Distribution. Figure 548. CALIFORNIA: San Francisco Bay area to San Diego.
LOUISIANA: Baton Rouge. SOUTH CAROLINA: Charleston. TEXAS: Browns-
ville.
Genus Anovia Casey
Anovia Casey, 1908, p. 408. — Leng, 1920, p. 214.-Korschefsky, 1931, p. 96.— Gor-
don, 1972a, p. 26. Type-species; Scymnus virginalis Wickham, by monotypy.
Noviini with description as for Rodolia except labrum convex, apical margin
broadly, feebly concave; prostemal protuberance not margined apically, pubescence
sparse; abdomen with postcoxal line on first sternum narrowly incomplete (Fig. 549e);
6th abdominal sternum of male with apical emargination feeble.
Anovia is the only genus of the Noviini native to the New World, and it closely
resembles Rodolia (see discussion under Rodolia, and Gordon, 1972a, pp. 27, 28).
There are 6 described species in this genus, only one of which, A. virginalis, occurs
north of Mexico. The remainder are neotropical. Scale species recorded as hosts for
members of Anovia are Steatococcus plucheae (Cockerell), leery a purchasi Maskell,
Icerya rileyi Cockerell, and Icerya montserratensis Riley and Howard.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Anovia virginalis (Wickham)
Fig. 549a-g; Map, Fig. 548
Scymnus virginalis Wickham, 1905, p. 166.
Anovia virginalis: Casey, 1908, p. 408. — Korschefsky, 1931, p. 96.— Gordon, 1972a,
p. 27.
Diagnosis. Length 2.43 to 3.05 mm, width 2.00 to 2.44 mm. Form elongate, oval,
widest anterior to middle of elytron. Color red; pronotum except anterior angle, head,
and basal portion of femur piceous; elytron typically with a median red spot and
subhumeral red area (Fig. 5491), variation in pattern shown in Figure 549g. Male
genitalia as in Figure 549a-c. Female genitalia as in Figure 549d.
Discussion. There are 5 cotypes of virginalis in the USNM collection. One of these,
a male, is here designated as the lectotype and so labeled. The remaining 4 types are
designated as paralectotypes.
Type locality. Chad’s Ranch, Utah (lectotype here designated).
Type depository. USNM (50212).
Distribution. Figure 548. ARIZONA: Benson; Capitan Mt.; Cottonwood; Mojave
Co., Hualapai Mts.; Sabino Canyon foothills; Santa Rita Range Exp. Sta.; Tombstone;
Tucson. NEW MEXICO: Mesilla Valley. TEXAS: El Paso; Finlay; Presidio; Rio
Grande City. UTAH: Leeds.
Tribe Exoplectrini
Exoplectrini Casey, 1908, p. 407. — Korschefsky, 1932, p. 225. — Blackwelder, 1945,
p. 450. — Sasaji, 1968, p. 26.
Chnoodiaires Mulsant, 1850, p. 907.
Exoplectrae Crotch, 1874b, p. 280.
Exoplectrides Gorham, 1895, p. 211.
Exoplectrinae Weise, 1904, p. 362.
Coccidulinae of widely varying size, length ranging from 2.0 to 8.0 mm. Head
deeply inserted in pronotum; clypeus extending well beyond antennal insertion, apex
broadly emarginate (Fig. 550a); gena extending onto eye, nearly completely dividing
eye. Antenna inserted under clypeal margin anterior to eye, 1 1 -segmented, club large,
asymmetrical, 3-segmented, basal segment extremely large, laterally compressed (Fig.
550b). Apical segment of maxillary palpus large, strongly securiform (Fig. 550c).
Mandible with 2 strong apical teeth and a large basal tooth. Epipleuron broad,
obliquely inclined, not fovea te for reception of leg. Prostemum flat, simple, narrowly
separating coxae. Leg variable, with or without external angulation at base of tibia.
Tarsus cryptotetramerous; tarsal claw strongly bifid (Fig. 550d). Abdomen with 5
visible sterna. Postcoxal line on first abdominal sternum complete or incomplete
(Fig. 550e, 0.
Exoplectra is the only representative of this tribe occurring north of Mexico. There
are 4 other genera as well as Exoplectra represented in the Neotropical region and 5
genera known from the Old World. The New World genera are closely similar to
each other with the essential characteristics of the head and antennae virtually iden-
tical. The partially divided eyes, pubescent dorsum, large basal segment of the an-
tenna, and partially concealed head characterize this tribe.
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Fig. 549. Anovia virginalis.
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Fig. 550. Exoplectra sp. a. Head. b. Antenna, c. Maxillary palpus, d. Front tibia.
Genus Exoplectra
Exoplectra Chevrolat, 1837, p. 461. — Mulsant, 1850, p. 916. — Crotch, 1874b, p.
284. — Korschefsky, 1932, p. 227. — Blackwelder, 1945, p. 450. Type-species; Coc-
cinella coccinea Fabricius, 1801, by subsequent designation of Korschefsky, 1932.
Exoplectrini with tibia angulate or almost dentate externally at base (Fig. 550d).
Postcoxal line incomplete. Male genitalia simple, symmetrical. Female genitalia lack-
ing infundibulum, accessory gland present.
Exoplectra schaefferi, new species, is the only representative of this genus occurring
north of Mexico. The tribal characteristics cause it to be easily recognized as there
are no similar appearing coccinellids in this region. This species has been known as
E. subaenescens Gorham since Schaeffer (1905) recorded it from Arizona. I have
examined the type series of subaenescens in the BMNH and find that the Arizona
species of Exoplectra is not subaenescens nor any presently described species of that
genus. I have not seen any host data for members of this genus except one record in
Schilder and Schilder (1928) of a species feeding on ""Aleurodicus cocois."' I seriously
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NORTH AMERICAN COCCINELLIDAE
671
doubt that members of this genus (or tribe) feed on whiteflies because of the robust
mandibular structure and lack of confirming evidence. The absence of host data and
the presence of large, strong teeth on the mandible causes me to suspect that they
may be plant feeders rather than predators. I have collected a series of an undescribed
species of Exoplectra in Peru, apparently making feeding marks on bamboo leaves,
but the act of feeding could not be positively established. There has been no modern
taxonomic treatment of members of this genus.
Exoplectra schaefferi, new species
Fig. 551a-d, 552a, b; Map, Fig. 548
Description. Male, length 4.20 mm, greatest width 3.29 mm. Form elongate, oval,
feebly convex (Fig. 552b). Color yellowish red; elytron dark brown with brassy green
tint; head and median Va, of pronotum black; meso- and metasternum dark brown,
epipleuron reddish brown. Dorsum densely pubescent with grayish white, decumbent
hairs. Punctures on head moderately coarse, separated by a diameter. Punctures on
pronotum fine, separated by one to 3 times a diameter. Elytral punctation very coarse,
dense, punctures separated by less than a diameter, nearly contiguous. Punctures on
ventral surface very fine, sparse, widely separated. Leg with external tibial angulation
feeble, most pronounced on anterior tibia. Postcoxal line incomplete, of the Pullus
type. Genitalia as in Figure 551a-d.
Female, similar to holotype except length 4.0 mm, width 3.20 mm; color pale
yellow; postcoxal line of the Diomus type; genitalia as in Figure 552a.
Variation. Length 2.75 to 4.20 mm, width 2.25 to 3.29 mm. The normally yellowish
red color is a paler yellow in some specimens.
Holotype. Male. ARIZONA: Huachucha Mts., VII- 15, from Ch. Schaeffer, Exo-
plectra subaenescens Gorh. (USNM 101347).
Allotype. Female. ARIZONA: Palmerly, Cochise Co., VIII- 17. (USNM).
Paratypes. Total 4 (Fig. 548). ARIZONA: Huachucha Mts.; Palmerly, Cochise Co.,
VI. (USNM).
The specimens described here are the same specimens that Schaeffer identified as
E. subaenescens. Exoplectra schaefferi is very similar to subaenescens in external
appearance, but the male genitalia are quite different. In addition, the elytral punctures
of E. subaenescens are much finer than those of E. schaefferi, and the areas that are
normally yellowish red in E. schaefferi are always yellow in E. subaenescens. The
type locality of E. subaenescens is Ventanas, in Durango, Mexico, and I have not
seen it from any other locality. This species is named for Charles Schaeffer who
collected and described several coccinellids from the southwestern United States.
Tribe Azyini
Azyini Schilder and Schilder, 1928, p. 217. — Korschefsky, 1932, p. 230.— Balduf,
1935, p. 152. — Blackwelder, 1945, p. 451.— Gordon, 1980, p. 153.
Azyaires Mulsant, 1850, p. 927.
Azyae Crotch, 1874b, p. 279.
Coccidulinae with form compact; dorsal surface black, pubescent, without color
pattern except many species with spots formed from brown and white pubescence.
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Fig. 551. Exoplectra schaejferi (male genitalia).
Head deeply inserted under pronotum; apex of clypeus emarginate medially; gena
only slightly extending onto eye; antennal insertion mostly exposed (Fig. 553a).
Antenna 1 1 -segmented, club 3-segmented, asymmetrical, each club segment with
small papilla on outer angle (Fig. 553b). Apical segment of maxillary palpus barrel-
shaped (Fig. 553c). Pronotum with anterolateral angle thickened, obtuse (Fig. 553a).
Epipleuron deeply notched for reception of femur. Leg broad, flat; anterior tibia with
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673
Fig. 552. Exoplectra schaefferi (female genitalia and habitus).
dually rounded external border (Fig. 553d). Tarsus cryptotetramerous (Fig. 553d);
claw with tooth near apex (Fig. 553e). Abdomen with 5 visible sterna; postcoxal line
of first sternum incomplete (Fig. 554b). Female genitalia without infundibulum,
spermatheca without development of ramus or nodulus.
Members of this tribe most nearly resemble members of the Exoplectrini and
Noviini. Members of the Noviini have simple legs without the armature found in
the Azyini. Some members of the Exoplectrini have legs similar to those found in
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b
e
Fig. 553. Azya sp. a. Head. b. Antenna, c. Maxillary palpus, d. Front leg.
the Azyini, but all members of the Exoplectrini have a broad, flat, basal antennal
segment and the clypeal structure is quite different. In addition, characters diagnostic
for the Azyini are the obtuse anterolateral angle of the pronotum and the papillae
on the antennal club segments.
Key to Genera of Azyini
1. Prostemum with intercoxal process elevated, narrow (Fig. 554a); apex of male sipho
with ventral flap before apex (Fig. 555d) Azya Mulsant
- Prosternum with intercoxal process flat, not elevated (Fig. 556a); apex of male sipho
bifid (Fig. 557d) Pseudoazya Gordon
Genus Azya Mulsant
Azya Mulsant, 1850, p. 928. — Crotch, 1874b, p. 279. — Schilder and Schilder, 1928,
p. 245. — Korschefsky, 1932, p. 230. — Blackwelder, 1945, p. 451.— Chapin, 1965b,
p. 246. — Woodruff and Sailer, 1977, p. 1. — Gordon, 1980, p. 155. Type-species;
Azya luteipes Mulsant, by subsequent designation of Crotch, 1874b.
Azyini with length ranging from 2.90 to 4.40 mm. Form oval. Dorsal surface black
except male head yellow, often with metallic lustre; clothed with dense, appressed
pubescence, pubescence short or long, usually white with spot on elytron composed
of brown hairs. Venter usually black or piceous except leg and abdomen yellow.
Prosternum lobed anteriorly, partially concealing mouthparts, deeply excavated at
side for reception of antenna, intercoxal process elevated, narrow, bicarinate or
medially ridged (Fig. 554a). Male genitalia with paramere slender; apex of sipho
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675
Fig. 554. Azya sp. a. Prostemum. b. Abdomen, c. Female genitalia.
slender, with ventral flap before apex. Female genitalia with apex of spermathecal
capsule broader than base; genital plate extremely elongate, triangular (Fig. 554c).
This is a distinctive genus in the North American fauna. The antennal club with
papillae, strong armature of the legs, and deeply foveate epipleuron render it easily
recognizeable. All species of Azya are neotropical but one species, A. trinitatis Mar-
shall (now in Pseudozya), was introduced into Florida from Trinidad in 1938 for
control of the coconut scale {Aspidiotus destructor Signoret). This species is probably
not established now, but a survey taken in 1939 showed that it was established at
that time. A second species, A. orbigera orbigera Mulsant, has recently (1976) been
collected in the vicinity of Miami. Specific host records for this scale feeding genus
are as follows; Asterolecanium bambusae (Boisduval), Asterolecanium miliaris (Bois-
duval), Aulacaspis tubercularis (Newstead), Coccus viridis (Green), Dysrnicoccus brev-
ipes (Cockerell), Ferrisia virgata (Cockerell), Lecanium sp., Lecanium viride Green,
Parasaissetia nigra (Nietner), Pseudococcus sp., Saissetia coffeae (Walker), Saissetia
oleae (Olivier), Selenaspidius sp. Azya was revised by Gordon (1980), see that pub-
lication for detailed discussion.
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Fig. 555. Azya orbigera.
Azya orbigera orbigera Mulsant
Fig. 555a-e; Map, Fig. 548
Azya orbigera Mulsant, 1850, p. 930.— Crotch, 1874b, p. 279.— Gorham, 1895, p.
21 l.-Korschefsky, 1932, p. 230.-Blackwelder, 1945, p. 45 1. -Wolcott, 1950, p.
310. — Chapin, 1965b, p. 247. — Leeper, 1976, p. 286.
Azya orbigera orbigera: Gordon, 1980, p. 157.
Azya luteipes: Woodruff and Sailer, 1977, p. 1 (not luteipes Mulsant, 1850).
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NORTH AMERICAN COCCINELLIDAE
677
Diagnosis. Length 2.90 to 4.25 mm, width 2.50 to 3.45 mm. Form oval. Head
yellow; dorsum greenish black, each elytron with round discal spot (Fig. 555e); venter
black except leg and abdomen yellow. Male genitalia as in Figure 555a-d.
Discussion. This is the only species of Azya that is known to be established in the
United States (see Gordon, 1980), apparently the result of an accidental introduction.
I originally identified this species as A. luteipes Mulsant, but subsequent examination
of the type specimens proved this identification to be incorrect.
Type locality. Colombia (lectotype designated by Gordon, 1980).
Type depository. UCCC.
Distribution. Figure 548. FLORIDA: Boca Raton; Dania; Davie; Ft. Lauderdale;
Hollywood; Miami; North Miami; Pompano Beach; West Palm Beach.
Genus Pseudoazya Gordon
Pseudoazya Gordon, 1980, p. 192. Type-species; Azya trinitatis Marshall, by original
designation.
Description as for Azya except length range from 2.20 to 2.65 mm; form nearly
round (Fig. 557e); male head black or mostly black; dorsal pubescence appressed or
erect; prosternum short, flat, intercoxal process not elevated or ridged (Fig. 556a).
Male genitalia with paramere paddle-shaped, apex of sipho bifid; female spermathecal
capsule with apex narrower than base, genital plate somewhat elongate, shorter than
in Azya (Fig. 556b).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 557. Pseudoazya trinitatis.
Pseudoazya trinitatis (Marshall)
Fig. 557a-e
Azya trinitatis Marshall, 1912, p. 320. — Korschefsky, 1932, p. 231.— Taylor, 1935,
p. 70. — Dohanian, 1937, p. 246. — Clausen, 1939,, p. 340.— Clausen, 1940, p.
573. — Blackwelder, 1945, p. 451. — Wolcott, 1950, p. 3 10.— Cochereau, 1969, p.
57. -Woodruff and Sailer, 1977, pp. 1-2.
Pseudazya trinitatis: Gordon, 1980, p. 194.
Diagnosis. Length 2.35 to 2.65 mm, width 2.0 to 2.38 mm. Form nearly round,
subdepressed (Fig. 557e). Head greenish black except apex of clypeus yellow; prono-
tum greenish black; elytron bluish black; venter black except leg and abdomen yellow.
Male genitalia as in Figure 557a-d.
Discussion. This species was released in the Miami, Florida, area in 1938. Speci-
mens were taken in 1939, but there is no evidence to suggest that A. trinitatis survived
in Florida after 1939.
Type locality. Cedros, Trinidad (lectotype designated by Gordon, 1980).
Type depository. BMNH.
Distribution. See discussion above.
Subfamily Coccinellinae
Coccinellinae Ganglbauer, 1899, pp. 954, 986. — Della Beffa, 1912, p. 167.— Mader,
1924, p. 6. — Korschefsky, 1931, p. 79. — Wingo, 1952, p. 16. — Sasaji, 1968, p.
21.— J. Chapin, 1974, p. 54.
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NORTH AMERICAN COCCINELLIDAE
679
Aphidiphages LaPorte, 1840, p. 523. — Chevrolat, 1849, p. 43.
Gymnosomides Mulsant, 1846, p. 27. — Mulsant, 1850, p. 2 (in part).
Coccinelliti Costa, 1849, p. 9.
Coccinellidae Crotch, 1873, p. 363.— Crotch, 1874b, p. 53.
Coccinellides Aphidiphages Chapuis, 1876, p. 166.
Coccinellidae Aphidiphages Weise, 1885a, p. 4.
Coccinellidae with dorsal surface glabrous; size medium to large. Mandible with
single basal tooth, apex bifid or with several teeth arranged in a row. Apical segment
of maxillary palpus securiform. Mentum narrowly articulated with submentum, ex-
panded apically. Antenna long, 1 1 -segmented, inserted more or less dorsally. Pro-
sternum T-shaped. Mesosternum narrowly articulated with metasternum. Mesepi-
meron triangular, posterior margin feebly bent. Each femur elongate, not flattened;
each leg simple, free. Tarsus cryptotetramerous. Female genital plate with stylus near
inner angle of plate, inner margin lacking distinct emargination. Male genitalia with
sipho long, usually strongly curved with well developed capsule.
This subfamily contains the species referred to as “ladybeetles” in the classic sense.
These are the commonly collected “aphid” predators that are often red with black
spots. I follow Sasaji (1968) in attributing only 2 tribes (Coccinellini and Psylloborini)
to the subfamily, both of which are represented north of Mexico.
Key to tribes of Coccinellinae
1. Gena extending onto eye (Fig. 558b); anterolateral angle of clypeus usually produced
forward; mandible bifid at apex Coccinellini
- Gena not extending onto eye (Fig. 558a); anterolateral angle of clypeus not produced,
apex truncate; mandible multidentate at apex Psylloborini
Tribe Coccinellini
Coccinellini Weise, 1885a, p. 7.— Casey, 1899, pp. 73, 82. — Blatchley, 1910, p. 512.—
Leng, 1920, p. 215. — Korschefsky, 1932, p. 310. — Wingo, 1952, p. 23. — Watson,
1956, p. 43 (in part). — Sasaji, 1968, p. 21.— J. Chapin, 1974, p. 55. — Belicek, 1976,
p. 295.
Adoniates Mulsant, 1846, p. 35. — Mulsant, 1850, p. 36.
Coccinellaires Mulsant, 1846, p. 29. — Mulsant, 1850, p. 35.
Coccinellates Mulsant, 1846, p. 35. — Mulsant, 1850, p. 74.
Coccinelliens Mulsant, 1846, p. 28. — Mulsant, 1850, p. 2.
Halyziaires Mulsant, 1846, p. 29. — Mulsant, 1850, p. 131.
Halyziates Mulsant, 1846, p. 147. — Mulsant, 1850, p. 162.
Hippodamiaires Mulsant, 1846, p. 30. — Mulsant, 1850, p. 5.
Micraspaires Mulsant, 1846, p. 162. — Mulsant, 1850, p. 212.
Mysiates Mulsant, 1846, p. 125. — Mulsant, 1850, p. 132.
Aliziarii Costa, 1849, p. 11.
Hippodamiini Costa, 1849, p. 10.— Weise, 1885a, p. 6.— Casey, 1899, pp. 73, 75.—
Blatchley, 1910, p. 509.— Wingo, 1952, p. 22. — Watson, 1956, p. 44. — Brown and
de Ruette, 1962, p. 643. — Sasaji, 1968, p. 21.
Micraspidarii Costa, 1849, p. 11.
Cariaires Mulsant, 1850, p. 228.
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Fig. 558. a. Psyllobora sp. head. b. Myzia sp. head.
Alesiaires Mulsant, 1850, p. 343.
Coelophoraires Mulsant, 1850, p. 374.
Cydoniaires Mulsant, 1850, p. 429.
Coccinellina Thomson, 1866, p. 332.
Coccinellides Thomson, 1866, p. 332. — Crotch, 1874, p. 91. — Gorham, 1891, p. 153.
Coccinellidae Berg, 1874, p. 288.
Hippodamiidae Berg, 1874, p. 287.
Tytthaspides Crotch, 1874b, p. 181.
Cariites Chapuis, 1876, p. 166.
Coccinellites Chapuis, 1876, p. 171.
Hippodamiites Chapuis, 1876, p. 167.
Synonychini Weise, 1885a, p. 7. — Mader, 1927, p. 2 1 . — Korschefsky, 1932, p. 268.
Halyziides Gorham, 1892, p. 161.
Synonychinae Della Beffa, 1912, p. 167.
Anisostictini Jacobson, 1916, p. 969. — Watson, 1956, p. 44.
Coccinellina Jacobson, 1916, p. 969. — Dobzhansky, 1926c, p. 1560.
Synonychina Jacobson, 1916, p. 969. — Dobzhansky, 1926c, p. 1574.
Hippodamiina Dobzhansky, 1926c, p. 200.
Coccinellinae with body length usually 3.0 mm or more. Head with gena extending
onto eye (Fig. 558b); eye usually finely faceted; mandible bifid at apex; clypeus slightly
narrower than frons, apex usually emarginate with anterolateral angle produced (Fig.
558b) (except in CeratomegiUa, Paranaemia, and some species of Hippodamia).
Anterior border of pronotum deeply excavate around head.
Twenty genera represent this tribe north of Mexico. Several of these genera are
holarctic, and representatives of some are neotropical. The tribe is extremely wide-
spread worldwide and the generic divisions are not as sharply defined as in most
other coccinellid tribes. The key characters plus the deeply excavated pronotum and
finely faceted eyes distinguish the Coccinellini from the Psylloborini to which it most
closely related. The Coccinellini as treated here have not been taxonomically con-
sidered as a whole, but Brown and de Ruette (1962) discussed the genera formerly
placed in the tribe Hippodamiini, and lablokoff-Khnzorian (1982) revised the tribe
for the Palearctic and Oriental Regions.
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681
1.
2(1).
3(2).
4(2).
5(1).
6(5).
7(6).
8(7).
9(6).
10(9).
11(10).
12(11).
13(12).
14(13).
15(14).
16(15).
Key to genera of Coccinellini
Tarsal claw not toothed or cleft, simply widened basally (Fig. 567a) 2
Tarsal claw toothed or apically cleft (Fig. 587i, 614c) 5
Pronotal base with fine, entire marginal bead 3
Pronotal base not margined 4
Metastemum with postcoxal line; elytron with large black spots (Fig. 570g) ....
Naemia Mulsant
Metastemum without postcoxal line; elytron vittate (Fig. 567g) . Paranaemia Casey
Apex of middle and hind tibia each with 2 spurs; elytron vittate (Fig. 5651);
epipleuron sloping downward internally Macronaemia Casey
Apex of middle and hind tibia each with a single spur; elytron spotted or very
irregularly vittate (Fig. 5601); epipleuron horizontal Anisosticta Dejean
Each tarsal claw cleft near apical y^ (Fig. 587i); form slender, legs distinctly visible
beyond body in dorsal view Hippodamia Dejean
Each tarsal claw with subquadrate basal tooth (Fig. 614c); or if tooth median,
then form rounded, legs barely visible beyond body in dorsal view (genus Myzid)
6
7
9
Pronotal base with marginal bead
Pronotal base without marginal bead
Metastemum and first abdominal sternum with distinct postcoxal line (fig. 1) . . 8
Metastemum and first abdominal sternum without postcoxal line
Coleomegilla Timberlake
Dorsal color mostly yellow, occasionally with some obscure dark markings (Fig.
680a, b) Aphidecta Weise
Dorsal color pattern red and black (Fig. 579f, g) Ceratomegilla Crotch
Length 7.40 to 10.0 mm; form rounded, highly convex; elytron reddish yellow
with 7 discrete black spots (Fig. 676a); one introduced species established in
Florida Harmonia Mulsant
Length usually less than 7.50 mm (except Anatis)\ other statements not as above
10
Prostemum strongly convex medially, protuberant at apex (Fig. 614b); mesoster-
num deeply emarginate for reception of sternal process Anatis Mulsant
Prostemum normally rounded, not protuberant at apex; mesostemum tmncate
or weakly emarginate for reception of prostemal process 11
Postcoxal line on first abdominal sternum complete, of the Pullus type (Fig. 637a)
Adalia Mulsant
Postcoxal line on first abdominal sternum incomplete, of the Diomus or Nephus
type (Figs. 634b, 682a) 12
Elytron yellow with black sutural margin and 4 black spots (Fig. 672g), spots
often somewhat coalesced; Oriental genus, one species possibly established in
Rorida Coelophora Mulsant
Elytron with color pattern not resembling that of Coelophora', North American
or European genera 13
Apex of each middle and hind tibia without spurs Mulsantina Weise
Apex of each middle and hind tibia with 2 spurs (Fig. 626a) 14
Tarsal claw with median tooth (Fig. 626a) Myzia LeConte
Tarsal claw with subquadrate basal tooth (Fig. 664b) 15
Pronotal surface polished, shiny, not alutaceous between punctures Calvia Mulsant
Pronotal surface alutaceous, often dull, not polished 16
Pronotum black with a large, subtrapezoidal or triangular white spot on each
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Fig. 559. Anisosticta sp. a. Postcoxal lines, b. Hind tibia, c. Tarsus.
anterolateral angle (Fig. 643g) (apical margin of pronotum sometimes narrowly
pale); epipleuron nearly horizontal, not descending externally Coccinella L.
- Pronotum not as above; epipleuron horizontal or descending externally 17
17(16). Postcoxal line on first abdominal sternum with oblique dividing line 18
- Postcoxal line on first abdominal sternum without oblique dividing line 19
18(17). Epipleuron strongly, abruptly descending externally; punctures on elytron fine,
nearly invisible Olla Casey
- Epipleuron nearly horizontal, not abruptly descending externally; punctures on
elytron coarse, dense Neoharrnonia Crotch
1 9( 1 7). Apex of mesosternum notched for reception of prostemal process European genus
(one species) presently established only in southeastern Canada Propylaea Mulsant
- Apex of mesosternum truncate; occurring over most of North America from
southern Canada to Mexico Cycloneda Crotch
Genus Anisosticta Dejean
Anisosticta Dejean, 1837, p. 456. — Mulsant, 1850, p. 36. — Mulsant, 1866, p. 25.—
LeConte, 1852, p. I30.-Crotch, 1873, p. 369.-Crotch, 1874b, p. 93. -Wickham,
1894,p. 299.-Casey, 1899, p. 75.-Leng, 1903a, p. 36.-Blatchley, 1910,p.510.-
Korschefsky, 1932, p. 367.— Timberlake, 1943, p. 45. — Bielawski, 1958, p. 91.—
Wingo, 1952, p. 22. — Brown and de Ruette, 1962, p. 644. — Belicek, 1976, p. 351.—
lablokoff-Khnzorian, 1982, p. 113. Type-species; Coccinella novemdecimpunctata
L., by subsequent designation of Crotch, 1874b.
Coccinellini with length 2.50 to 4.0 mm. Form elongate, dorsoventrally flattened,
femur visible beyond lateral margin of elytron. Dorsal color yellow with head black
basally or almost entirely; pronotum and elytron with brown or black maculae (Fig.
560f). Apex of clypeal margin truncate, anterolateral angle produced forward. Lateral
margin of elytron broadly reflexed; epipleuron nearly flat. Intercoxal process of pro-
sternum narrow with broad lateral ridge. Apical margin of mesosternum truncate,
ridged. Apex of middle and hind tibiae each with a single spur (Fig. 559h). Tarsal
claw widened basally, not toothed (Fig. 559c). Postcoxal line nearly complete, but
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NORTH AMERICAN COCCINELLIDAE
683
not entirely so (Fig. 559a). Male genitalia symmetrical. Female genitalia lacking
infundibulum; coxal plate elongate, with strong apical stylus (Fig. 560e).
The simple tarsal claw, single tibial spur, and the dorsal color pattern characterize
members of this genus. The only other genus with remotely similar habitus is Naemia
Mulsant which has two tibial spurs and the head entirely black. Anisosticta is a
holarctic genus with 7 names currently recognized as valid. Two of these species
occur in North America. The genus was revised by Bielawski (1958) and further
treated by Brown and de Ruette (1962). The name Anisosticta has incorrectly been
attributed to Duponchel (see comments by Brown and de Ruette, 1962) and more
recently to Dejean (Belicek, 1976). Members of Anisosticta are said to be aphid
predators, but I have not seen any specihc host data.
Key to species of Anisosticta
1. Abdomen entirely black; mesepimeron darkened or black borealis Timberlake
- Abdomen with lateral margin pale; mesepimeron pale bitriangularis (Say)
Anisosticta bitriangularis (Say)
Fig. 560a-h; Map, Fig. 561
Coccinella bitriangularis Say, 1824, p. 269.
Anisosticta bitriangularis: Casey, 1899, p. 76.— Timberlake, 1943, p. 45.— Wingo,
1952, p. 45. — Bielawski, 1958, p. 101. — Brown and de Ruette, 1962, p. 645.—
Belicek, 1976, p. 352. — lablokoff-Khnzorian, 1982, p. 115.
Anisosticta strigata ab. bitriangularis: Korschefsky, 1932, p. 373.
Coccinella multiguttata Randall, 1838, p. 51. — Mulsant, 1850, p. 35.— LeConte,
1859c, p. 197.
Anisosticta strigata multiguttata: Gemminger and Harold, 1876, p. 3744.
Anisosticta 19-punctata multiguttata: Weise, 1895a, p. 126.
Anisosticta strigata ab. multiguttata: Korschefsky, 1932, p. 373.
Anisosticta strigata: Crotch, 1873, p. 369. — Crotch, 1874b, p. 93.— Leng, 1903a, p.
37. -Wickham, 1894, p. 299.-Blatchley, 1910, p. 510.
Anisosticta novemdecimpunctata irregularis Weise, 1879, p. 94— Brown and de Ruette,
1962, p. 645.
Diagnosis. Length 3.0 to 4.0 mm, width 1.90 to 2.40 mm. Dorsal color pattern as
in Figure 560f-h. Lateral margin of abdomen and meso- and metepimeron pale.
Male genitalia as in Figure 560a-d. Female genitalia as in Figure 560e.
Discussion. Specimens from the southern portion of the range have the elytral and
pronotal spots free. Northern specimens tend to become more heavily maculate until
the most northern specimens have both elytral and pronotal spots fused into almost
regular vittae (Fig. 560f). See Bielawski (1958) for a detailed discussion of the syn-
onymy of A. bitriangularis. This species was considered a synonym of the European
A. strigata by almost all authors until Casey (1899) recognized it as a valid species.
Examination of the genitalia by Bielawski (1958) and Brown and de Ruette (1962)
confirm Casey’s decision.
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Fig. 560. Anisosticta bitriangularis.
Type locality. Of bitriangularis, “Northwest Territory”; of multiguttata, Cam-
bridge, Massachusetts; of irregularis, Oregon.
Type depository. Of bitriangularis, type lost; of multiguttata, not located; of irre-
gularis, not examined.
Distribution. Figure 561. Labrador to New Jersey, west to Alaska, California, and
British Columbia.
Anisosticta borealis Timberlake
Fig. 562a-g; Map, Fig. 563
Anisosticta borealis 1943, p. 45. — Bielawski, 1958, p. 108.— Brown and
de Ruette, 1962, p. 645. — Belicek, 1976, p. 352. — lablokoff-Khnzorian, 1982, p.
115.
Description as for bitriangularis except head mostly black; dorsal maculation heavy
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(Fig. 562f, g); lateral margin of abdomen and meso- and metepimeron black or at
least partially darkened. Male genitalia as in Figure 562a-d. Female genitalia as in
Figure 562e.
Bielawski (1958) and lablokofF-Khnzorian (1982) regard A. borealis as a junior
synonym of A. strigata (Thunberg). Brown and de Ruette (1962) maintained A.
borealis as a valid species. Anisosticta borealis is probably deserving of at least
subspecific standing because the differences in size and dorsal coloration between it
and A. strigata are at least as significant as those found in similar situations in the
genera Coccinella and Hippodamia, and differences in distribution of the same nature
have been accepted also.
Type locality. Nulato, Alaska.
Type depository. USNM.
Distribution. Figure 563. Manitoba to Alaska.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 562. Anisosticta borealis.
Genus Macronaemia Casey
Macronaemia Casey, 1899, pp. 75, 76. — Leng, 1903a, p. 37. — Casey, 1908, p. 394.—
Timberlake, 1943, p. 45. — Wingo, 1952, p. 22. — Brown and de Ruette, 1962, p.
646. — Belicek, 1976, p. 349. — lablokoff-Khnzorian, 1982, p. 120. Type-species;
Coccinella episcopalis Kirby, by monotypy.
Micronaemia Weise, 1905, p. 218. — Casey, 1908, p. 394.
Coccinellini with length 3.25 to 4.0 mm; form extremely elongate, parallel sided,
dorsoventrally flattened, femur visible beyond lateral margin of elytron. Dorsal color
pattern yellow with black vittae (Fig. 5651)- Anterior margin of clypeus nearly trun-
cate, anterolateral angle produced forward. Lateral margin of elytron narrowly, abruptly
explanate; epipleuron sloping downward internally, entirely visible when viewed
laterally. Intercoxal process of prosternum narrow, strongly convex, lacking carinae
but with fine lateral ridge. Apical margin of mesostemum narrowly produced forward
between anterior coxae. Apex of middle and hind tibiae each with 2 spurs. Tarsal
claw widened basally, not toothed (Fig. 564a). Postcoxal line narrowly incomplete
as in Anisosticta (Fig. 564b). Male genitalia symmetrical. Female genitalia lacking
infundibulum; coxal plate elongate, with strong apical stylus (Fig. 565e).
Paranaemia and Macronaemia are the only genera of North American Coccinellini
with a distinctly vittate dorsum. Macronaemia is much smaller than Paranaemia,
and the pronotal base is not margined. Macronaemia is monobasic in North America
but 2 other species have been described from China.
Macronaemia episcopalis is said to be an aphid predator, but I have not seen any
specific host data.
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Macronaemia episcopalis (Kirby)
Fig. 565a-f; Map, Fig. 566
Coccinella episcopalis Kirby, 1837, pp. 228.
Naemia episcopalis: Mulsant, 1850, p. 34. — Mulsant, 1866, p. 24. — Crotch, 1874b,
p. 93.-Wickham, 1894,, p. 300.
Anisosticta episcopalis: Crotch, 1873, p. 369. — Leng, 1903a, p. 37.
Macronaemia episcopalis: Casey, 1899, p. 76.— Timberlake, 1943, p. 10.— Wingo,
1952, p. 45. — Brown and de Ruette, 1962, p. 646. — Belicek, 1976, p. 349. — la-
blokoff-Khnzorian, 1982, p. 121.
Micronaemia episcopalis: Weise, 1905, p. 217.
Diagnosis. Length 3.25 to 4.0 mm, width 1.60 to 2.0 mm. Color yellow with head
black except frons and clypeus yellow; 3 black vittae on elytron, sutural vitta narrow;
pronotum with 3 black spots on each side, spots often confluent (Fig. 5651); ventral
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Fig. 564. Macronaemia episcopalis. a. Tarsus, b. Postcoxal lines.
surface except leg mostly black except meso- and metepimeron, lateral and apical
margins of abdomen yellow; male with prostemum, anterior coxa, and median area
of mesosternum yellow. Male genitalia as in Figure 565a-d. Female genitalia as in
Figure 565e.
Type locality. “Journey from New York to Cumberland-house”.
Type depository. BMNH (not examined).
Distribution. Figure 566. Ontario to New York, west to Yukon Territory and
northern California.
Fig. 565. Macronaemia episcopalis.
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Genus Paranaemia Casey
Paranaemia Cdisey, 1899, p. 76. — Leng, 1903a, p. 38. — Dobzhansky, 1926b, p. 201.—
Timberlake, 1943, p. 45. — Hatch, 1961, p. 165. — Brown and de Ruette, 1962, p.
644. Type-species; Hippodamia vittigera Mannerheim, by original designation.
Cemtomegilla {Paranaemia): Leng, 1920, p. 21 5.— Korschefsky, 1932, p. 312.
Coccinellini with length 4.50 to 6.60 mm. Form elongate, dorsoventrally flattened,
femur visible beyond lateral margin of elytron. Dorsal color yellow with black vittae
(Fig. 567g). Apex of clypeus truncate, anterolateral angle produced forward. Base of
pronotum finely margined. Lateral margin of elytron feebly reflexed; epipleuron
nearly flat, sloping downward slightly internally. Intercoxal process of prosternum
narrow, feebly convex, lacking carinae but with fine lateral ridge. Apical margin of
mesostemum weakly emarginate, ridged. Metasternum lacking postcoxal line. Apex
of middle and hind tibiae each with 2 spurs. Tarsal claw widened basally, not toothed
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Fig. 567. Paranaemia vittigera.
(Fig. 567a). Postcoxal line on abdomen lacking (Fig. 567b). Male genitalia symmet-
rical. Female genitalia with small infundibulum; coxal plate elongate, stylus distinct
(Fig. 568).
Paranaemia may be confused with some vittate specimens of Hippodamia, but
the pronotal base is margined and the claws are not apically cleft in Paranaemia (see
comparative remarks under Macronaemia). Paranaemia is monobasic. Paranaemia
vittigera is said to be an aphid predator, but I have not seen any specific host data.
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691
Paranaemia vittigera (Mannerheim)
Fig. 567a-g, 568; Map, Fig. 569
Hippodamia vittigera Mannerheim, 1843, p. 312.
Coccinella {Hippodamia) vittigera: Guerin, 1844, p. 322.
Naemia vittigera: Mulsant, 1850, p. 33. — Mulsant, 1866, p. 23. — Crotch, 1874b, p.
93.-Chapuis, 1876, p. 171. -Gorham, 1891, p. 153.
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Megilla vittigera: Crotch, 1873, p. 364.
Paranaemia vittigera: Casey, 1899, p. 76. — Leng, 1903a, p. 38.— Timberlake, 1943,
p. 9. -Hatch, 1961, p. 166.
Ceratomegilla {Paranaemia) vittigera: Korschefsky, 1932, p. 315.
Paranemia similis Casey, 1899, p. 76. — Leng, 1903a, p. 39.— Timberlake, 1943,
p. 9.
Ceratomegilla {Paranaemia) vittigera ab. similis: Korschefsky, 1932, p. 315.
Diagnosis. Length 4.50 to 6.60 mm, width 2.90 to 3.40 mm. Color yellow with
head black except narrow yellow area on frons; 3 black vittae on elytron, sutural
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693
vitta wide; pronotum with large, black lateral spot (Fig. 567g); ventral surface in-
cluding legs black except prostemum yellow. Male genitalia as in Figure 567c-f.
Female genitalia as in Figure 568.
Discussion. There are 6 types of vittigera in Helsinki. One of these, a male labeled
“Cygnaeus/Califbor./coll. Mannerh.”, I here designate and label as the lectotype,
the remainder as paralectotypes. There are 5 types of P. similis in the Casey collection;
I here designate and label the first of these, a female as the lectotype and the remainder
as paralectotypes.
Type locality. Of vittigera, California (lectotype here designated); of similis, Arizona
(lectotype here designated).
Type depository. Of vittigera, UMZH; oi similis, USNM (35491).
Distribution. Figure 569. Alberta to west Texas, west to British Colombia and
California.
Genus Naemia Mulsant
Naemia Mulsant, 1850, p. 30. — Mulsant, 1866, p. 21.— Crotch, 1874b, p. 92.—
Chapuis, 1876, p. 170.— Gorham, 1891, p. 152.— Wickham, 1894, p. 300. — Casey,
1899, p. 76. — Dobzhansky, 1926b, p. 201. — Mader, 1929, p. 90. — Korschefsky,
1932, p. 317.— Timberlake, 1943, pp. 9, 45. — BrownanddeRuette, 1962, p. 644.—
J. Chapin, 1974, p. 56. — lablokoff-Khnzorian, 1982, p. 126. Type-species; Coc-
cinella seriata Melsheimer, by subsequent designation of Crotch, 1874.
Coccinellini with length 4.0 to 6.70 mm. Form elongate, somewhat flattened, femur
visible beyond lateral margin of elytron. Dorsal color yellow with strong, variable,
black maculae. Apex of clypeal margin broadly emarginate, anterolateral angle pro-
duced forward. Base of pronotum finely margined. Lateral margin of elytron broadly,
feebly reflexed; epipleuron nearly flat, sloping downward slightly internally. Intercoxal
process of prostemum narrow, feebly convex, lacking carinae but with fine lateral
ridge. Metastemum with postcoxal line. Middle and hind tibia each with 2 spurs.
Tarsal claw widened basally, not toothed (Fig. 570a). Postcoxal line on abdomen
lacking. Male genitalia symmetrical. Female genitalia with small infundibulum; coxal
plate elongate, stylus distinct (Fig. 5701).
Naemia is easily confused with Coleomegilla because the dorsal color patterns are
very similar, but the tarsal claw is toothed and the metasternum lacks postcoxal lines
in Ceratomegilla. Naemia is an American genus ranging from southern New England
and southwestern United States to the Antilles and Central America. The distribution
is mainly coastal or insular, possibly because of a high humidity requirement. There
have been 3 names proposed within the genus, and I consider one of these a synonym
and 2 valid subspecies. Members of Naemia are said to be aphid predators, but I
have not seen any specific host data.
Key to subspecies of Naemia seriata (Melsheimer)
1. Head usually entirely black; surface of elytron not strongly alutaceous; eastern and
southern United States, Antilles seriata seriata Melsheimer
- Head with clypeus and triangular median area on frons pale; surface of elytron strongly
alutaceous; southern California seriata litigiosa Mulsant
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Fig. 570. Naemia serial a seriata.
Naemia seriata seriata (Melsheimer)
Fig. 570a-g; Map, Fig. 571
Coccinella seriata Melsheimer, 1847, p. 177.
Anisosticta seriata: LeConte, 1852, p. 130. — Crotch, 1873, p. 369.— Leng, 1903a,
p. 37.
Naemia seriata: Mulsant, 1866, p. 21.— Crotch, 1874b, p. 92.— Gorham, 1891, p.
152. — Casey, 1899, p. 76. — Korschefsky, 1932, p. 3 1 7.— Timberlake, 1943, p. 9.
Naemia seriata seriata: Timberlake, 1943, p. 46.
Megilla fuscilabris decepta Blatchley, 1914, p. 64. New Synonymy.
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695
Ceratomegilla fuscilabris a. decepta: Leng, 1920, p. 215.
Ceratomegilla maculata ab. decepta: Korschefsky, 1932, p. 313.
Naemia seriata decepta: Timberlake, 1943, pp. 9, 46.— J. Chapin, 1976, p. 56.
Diagnosis. Length 4.0 to 6.70 mm, width 2.50 to 3. 10 mm. Head black; pronotum
typically yellow with an irregular, black, central macula (Fig. 570g), but many south-
ern specimens have macula broken into ill-dehned spots; elytron yellow with 6 black
spots more or less fused in northern specimens (typical form). Ventral surface in-
cluding leg black except prostemum and lateral abdominal margin yellow. Male
genitalia as in Figure 570b-e. Female genitalia as in Figure 570f
Discussion. Naemia decepta is a color form of N. seriata occurring mainly in Florida
and along the Gulf Coast, also in the Antilles. The N. decepta color type occurs in
occasional specimens as far north as Maryland; there are no genitalic differences
between N. decepta and N. seriata and numerous specimens cannot be assigned to
either N. seriata or N. decepta\ therefore I consider decepta a junior synonym.
Type locality. Of seriata, Pennsylvania; of decepta, Ormond, Florida.
Type depository. Of seriata, not known; of decepta, PU.
Distribution. Figure 571. Atlantic and Gulf coasts, Rhode Island to south Texas.
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Fig. 572. Naemia seriata litigiosa.
Naemia seriata litigiosa Mulsant
Fig. 572a-c; Map, Fig. 571
Naemia litigiosa Mulsant, 1850, p. 31. — Mulsant, 1853, p. 22. — Crotch, 1874b, p.
92.
Anisosticta litigiosa: Crotch, 1873, p. 369. — Leng, 1903a, p. 37.
Naemia seriata litigiosa: Leng, 1920, p. 215. — Korschefsky, 1932, p. 317.
Description as for seriata, s. str., except clypeus and triangular area on frons pale,
spots on elytron not particularly confluent (Fig. 572c, d); surface of elytron strongly
alutaceous and densely punctured; male genitalia with basal lobe curved in lateral
view, sipho sinuate (Fig. 572a, b).
The southwestern United States populations of seriata are apparently widely dis-
junct, unless there are Mexican populations connecting them with the eastern form.
The California specimens are separable on both male genitalia and external appear-
ance, therefore I elect to apply the name litigiosa Mulsant, long considered a junior
synonym of seriata. The question seems to be whether or not to consider litigiosa a
valid species rather than a subspecies, but I prefer to follow the more conservative
course at present.
Type locality. “FAmerique du nord”.
Type depository. DLM (type not examined).
Distribution. Figure 571. CALIFORNIA: San Diego. NEW MEXICO: La Cuera,
Organ Mts.
Genus Coleomegilla Timberlake
Co/comc’^/7/fl Timberlake, 1920a, p. 139. — Timberlake, 1920b, p. 96.— Timberlake,
1943, pp. 9, 46. — Wingo, 1952, p. 23. — Brown and de Ruette, 1962, p. 646.— J.
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697
Fig. 573. Coleomegilla maculata lengi.
Chapin, 1974, p. 57. Type-species; Coccinella maculata Degeer, by original des-
ignation.
Megilla Mulsant, 1850, p. 24. (not Megilla F., 1805, nor Megilla Erichson, 1849).—
Mulsant, 1866, p. 16. — LeConte, 1852, p. 130.— Crotch, 1873, p. 364. — Crotch,
1874b, p. 92.-Chapuis, 1876, p. 169. -Gorham, 1891, p. 151. -Wickham, 1894,
p. 300.-Casey, 1899, p. 76.-Leng, 1903a, p. 38.-Blatchley, 1910, p. 510.-
Dobzhansky, 1926b, p. 201. — Korschefsky, 1932, p. 312. Type-species; Coccinella
maculata Degeer, by subsequent designation of Crotch, 1873.
Coccinellini with length 4.0 to 8.0 mm. Form elongate, somewhat dorsoventrally
flattened, femur visible beyond lateral margin of elytron. Dorsal color red or yellowish
orange with black maculae (Fig. 573g). Apex of clypeus broadly, feebly emarginate,
anterolateral angle produced forward. Base of pronotum finely margined. Lateral
margin of elytron broadly, feebly reflexed; epipleuron nearly flat, sloping downward
slightly internally. Intercoxal process of prostemum narrow, feebly convex, lacking
carinae but with fine lateral ridge. Apical margin of mesosternum triangularly notched
medially, ridged. Metastemum lacking postcoxal line. Apex of middle and hind tibiae
each with 2 spurs. Tarsal claw with subquadrate basal tooth (Fig. 573a). Postcoxal
line on abdomen lacking. Male genitalia symmetrical. Female genitalia with small
infundibulum (Fig. 573f); coxal plate elongate, stylus distinct.
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Coleomegilla and Naemia are very similar in appearance (see remarks under Nae-
mia) but quite distinct from each other in claw structure. Coleomegilla is an American
genus ranging from southern Canada to Georgia and southern California, south to
Venezuela and Peru. There are currently about 15 names in use within the genus as
species, subspecies, or varieties. Of these names, 3 have been listed as subspecies of
C. maculata from America north of Mexico (Timberlake, 1943). Members of this
genus are usually considered to be primarily aphid predators, but Forbes (1883)
found that more than 50% of the diet of C maculata lengi was composed of pollen
from various plants. The bulk of the animal portion of the diet was composed of
aphids. Specific aphid and adelgid hosts are as follows: Acyrthosiphon dirhodum
(Walker), Acyrthosiphon pisum (Harris), Aphis gossypii Glover, Aphis rumicis L.,
Brevicoryne brassicae (L.), Hyadaphis erysimi (Kaltenbach), Macrosiphum avenae
(F.), Macrosiphum euphorbiae (Thomas), Nearctaphis crataegifoliae (Fitch), Pem-
phigus bursarius (L.), Pineus strobi (Hartig).
Key to subspecies of Coleomegilla maculata (Degeer)
1. Pronotal spots large, triangular; median elytral spot large, oval, spot on apical de-
clivity touching sutural margin (Fig. 573g) 2
Pronotal spots small, reduced, oval or curvilinear; median elytral spot reduced or
divided into a small lateral spot and a large median spot; spot at apical declivity not
touching sutural margin (Fig. 576f) maculata fuscilabris (Mulsant)
2(1). Basal lobe of male genitalia with apex feebly notched (Fig. 573g)
maculata lengi Timberlake
Basal lobe of male genitalia with apex distinctly notched (Fig. 575a)
maculata strenua (Casey)
Coleomegilla maculata lengi Timberlake
Fig. 573a-h; Map, Fig. 574
Chrysomela 10- maculata F., 1781, p. 98.
Megilla maculata: Mulsant, 1850, p. 28. — Mulsant, 1866, p. 20. — Crotch, 1873, p.
364.-Crotch, 1874b, p. 92.-Gorham, 1891, p. 1 5 1. -Wickham, 1894, p. 300.-
Leng, 1903a, p. 38. — Blatchley, 1910, p. 510.
Hippodamia maculata: LeConte, 1852, p. 131. — Forbes, 1883, p. 51.
Ceratomegilla maculata: Leng, 1920, p. 215. — Korschefsky, 1932, p. 312.
Coleomegilla maculata Timberlake, 1943, p. 9. — Wingo, 1952, p. 45.— Brown
and de Ruette, 1962, p. 646.— J. Chapin, 1974, p. 57.
Megilla fuscilabris: (of authors, not Mulsant, 1866).— Casey, 1899, p. 76.— Leng,
1903a, p. 38. -Blatchley, 1914, p. 64.
Diagnosis. Length 4.20 to 6.60 mm, width 2.80 to 3.80 mm. Head black with
triangular pale area on frons; pronotum yellow with triangular black macula on each
side; elytron pink to red with 6 black maculae (Fig. 573g, h). Ventral surface including
legs black except prostemum and lateral abdominal margin yellow. Male genitalia
as in Figure 573b-e. Female genitalia as in Figure 573f
Discussion. Prior to Timberlake (1943), the names and combinations thereof re-
lating to C. maculata were greatly confused in the literature. Thanks to his work they
can now be sorted out in a reasonable fashion although at least a few of the names
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699
Fig. 574. Distribution. Coleomegilla maculata lengi (shaded, eastern); C. m. strenua (shad-
ed, southwest).
currently in use will surely prove to be synonyms. As pointed out by Timberlake,
the name C. fuscilabris (Mulsant) was incorrectly applied to this subspecies by most
authors subsequent to Mulsant’s original description. Timberlake (1943) said that
subspecies lengi was generally distributed east of the Rocky Mountains, referring
western specimens to the subspecies strenua. I follow this course here, but with some
reservations about the necessity of maintaining 2 names. If it becomes apparent that
strenua and lengi are synonymous, strenua will have priority.
Type locality. Columbus, Ohio.
Type depository. Type not examined.
Distribution. Figure 574. Ontario to Georgia, west to Minnesota and Texas.
Coleomegilla maculata strenua (Casey)
Fig. 575a-d; Map, Fig. 574
Megilla strenua Casey, 1899, p. 76.
Megilla maculata var. strenua: Leng, 1903a, p. 38.
Ceratomegilla maculata ab. strenua: Korschefsky, 1932,p. 313.
Coleomegilla maculata strenua: Timberlake, 1943, p. 46.
Diagnosis. Length 6.40 to 7.0 mm, width 3.40 to 4.30 mm. Description as for
lengi (Fig. 575f) except male genitalia as in Figure 575a-d. Female genitalia as in
Figure 575e.
Discussion. This subspecies is noticeably larger than C. m. lengi on the average,
but the only other difference I can find is in the shape of the apex of the basal lobe
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Fig. 575. Coleomegilla maculata strenua.
of the male genitalia. There are 5 types of C. m. strenua in the Casey collection, the
first of which (female) I here designated and label as the lectotype, the remainder as
paralectotypes.
Type locality. Brownsville, Texas (lectotype here designated).
Type depository. USNM (35493).
Distribution. Figure 574. Texas to southern California.
Coleomegilla maculata fuscilabris (Mulsant)
Fig. 576a-f; Map, Fig. 577
Naemia fuscilabris Mulsant, 1866, p. 22.
Megilla fuscilabris: Crotch, 1873, p. 364 (in part); Gorham, 1891, p. 152.
Ceratomegilla maculata ?ih. fuscilabris: Korschefsky, 1932, p. 313.
Coleomegilla maculata fuscilabris: Timberlake, 1943, p. 46.— J. Chapin, 1974, p.
58.
Megilla maculata var. floridana Leng, 1903a, p. 38. — Blatchley, 1914, p. 64.— Tim-
berlake, 1943, p. 9.
Ceratomegilla maculata ab. floridana: Leng, 1920, p. 2 1 5. — Korschefsky, 1932, p.
313.
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701
Diagnosis. Length 4.0 to 5.75 mm, width 2.30 to 3.50 mm. Description as for
lengi except pronotum with black spots reduced, oval or curvilinear; median spot
on elytron reduced, usually divided into larger median spot and small sublateral spot
(Fig. 5761), lateral spot often lacking. Male genitalia as in Figure 576a-d. Female
genitalia as in Figure 576e.
Discussion. The dorsal background color in C. m. fuscilabris is usually paler yellow
than in the other two subspecies, and is almost never pink or reddish as is often the
case with C. m. lengi and C. m. strenua.
Type locality. Of fuscilabris. New Orleans, Louisiana; of floridana, Florida and
Louisiana.
Type depository. Of fuscilabris, type not examined; of floridana, type not examined.
Distribution. Figure 577. South Carolina to Florida, west to Louisiana (coastal
localities); disjunct locality— Shaw Pond, Washington, D.C.
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Fig. 577. Distribution. Coleomegilla maculata fuscilabris (shaded, disjunct locality dotted).
Genus Ceratomegilla Crotch
Cemtomegilla CroXch, 1873, p. 365. — Crotch, 1874b, p. IX. — Casey, 1899, p. 75.—
Leng, 1920, p. 215.-Korschefsky, 1932, p. 312.-Scott, 1933, p. 126. -Timber-
lake, 1943, p. 45. — Brown and de Ruette, 1962, p. 646. — Belicek, 1976, p. 338.
Type-species; Ceratomegilla ulkei Crotch, by monotypy.
Spiladelpha Semenov-Tian-Shanskij and Dobzhansky, 1923, p. 99. — Mader, 1929,
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703
c
Fig. 578. Ceratomegilla ulkei. a. Antenna, b. Tarsus, c. Postcoxal lines.
p. 87.— Brown and de Ruette, 1962, p. 646. Type-species; Spiladelpha barovskii
Semenov-Tian-Shanskij and Dobzhansky, by monotypy.
Hippodamia {Ceratomegilla) lablokolf-Khnzorian, 1982, p. 327.
Coccinellini with length 3.70 to 4.70 mm. Form elongate, slender, apex of elytron
acute or subacute, not rounded. Apex of clypeus truncate; anterolateral angle without
projection. Third antennal segment in male strongly triangular, wider than second
or fourth segment, anterior apical angle ciliate (Fig. 578a). Base of pronotum finely
margined. Lateral margin of elytron not reflexed; epipleuron sloping downward in-
ternally. Intercoxal process of prosternum narrow, feebly convex, lacking carinae but
with fine lateral ridge. Apical margin of mesosternum truncate, ridged. Metastemum
with postcoxal line, line usually reaching lateral margin. Apex of middle and hind
tibiae each with 2 spurs. Anterior tarsus of male with basal 2 segments strongly
dilated; of middle tarsus almost as strongly dilated. Tarsal claw with basal tooth,
tooth not as wide as long, apex acute (Fig. 578b). Postcoxal line on abdomen distinct,
complete, of Pullus type (Fig. 578c). Male genitalia symmetrical. Female genitalia
with small infundibulum; coxal plate somewhat elongate, stylus distinct (Fig. 579e).
Ceratomegilla has been confused with other genera over the years, mainly because
of its rarity in collections. Crotch (1873) included one species, C. ulkei, known from
arctic and subarctic North America in the genus. Brown and de Ruette (1962) syn-
onymized Spiladelpha with Ceratomegilla, Spiladelpha contains 3 species from Si-
beria, Tibet, and Russian Turkestan. Belicek (1976) placed Ceratomegilla as a junior
synonym of Hippodamia, but this placement cannot be maintained because the
characters distinguishing Ceratomegilla are at least as definitive as those of any other
genus in the Coccinellini. lablokoff-Khnzorian treated Ceratomegilla as a subgenus
of Hippodamia. Some species of Hippodamia have the male tarsal segments strongly
dilated as in Ceratomegilla, but in Hippodamia the pronotal base is not margined,
and each tarsal claw is cleft at the middle. Members of Ceratomegilla are probably
aphid predators, but no specific host data is known.
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Fig. 579. Ceratomegilla ulkei.
Cemtomegilla ulkei Crotch
Fig. 579a-g; Map, Fig. 580
Cemtomegilla ulkei Crotch, 1873, p. 365. — Crotch, 1874b, p. IX.— Wickham, 1894,
p. 305. -Casey, 1899, p. 75.-Leng, 1903a, p. 39.-Korschefsky, 1932, p. 315.-
Scott, 1933, p. 136. — Brown and de Ruette, 1962, p. 647.
Hippodamia ulkei: Belicek, 1976, p. 340.
Spiladelphia barovskii Semenov and Dobzhansky, 1923, p. 99. — lablokoff- Khnzo-
rian, 1982, p. 328.
Hippodamia parva Watson, 1954, p. 45. — Belicek, 1976, p. 340.
Cemtomegilla parva: Brown and de Ruette, 1962, p. 647.
Hippodamia {Ceratomegilla) ulkei: lablokoff-Khnzorian, 1982, p. 327.
Diagnosis. Length 3.70 to 4.70 mm. Head black with 2 yellow spots; pronotum
black with yellow lateral margin; elytron typically black with pale margins, but pattern
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705
variable (Fig. 579f, g). Ventral surface including legs black except abdomen reddish
brown. Male genitalia as in Figure 579a-d. Female genitalia as in Figure 579e.
Discussion. Brown and de Ruette (1962) indicated the similarity of C. ulkei and
H. parva, and Belicek (1976) synonymized the two names. The only species with
which C. ulkei is likely to be confused is Hippodamia arctica (Schneider), but the
generic characters will separate them. Crotch apparently had only one type specimen
(holotype) when he described this species.
Type locality. Of ulkei, “Hudson’s Bay”, of parva. Cape Henrietta Maria, Ontario.
Type depository. Of ulkei, type not located; of parva, CNC.
Distribution. Figure 580. ALASKA: Cape Thompson, Ogotonuk Cr.; Rampart
House, 60-75 mi. North; Sheenjek R.; Umiat. BRITISH COLUMBIA: Summit Lake,
mi 379 and 392, Alaska Hwy. NORTH WEST TERRITORIES: Coppermine; Fort
McPherson; Muskox Lake; Reindeer Depot; Tuktoyaktuk; Tununuk. ONTARIO:
Cape Henrietta Maria. YUKON TERRITORY: Selkirk.
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Genus Hippodamia Dejean
Hippodamia Dejean, 1837, p. 456. — Mulsant, 1846, p. 30. — Mulsant, 1850, p. 10.—
Mulsant, 1866, p. 8. — LeConte, 1852, p. 130.— Crotch, 1873, p. 365.— Crotch,
1874b, p. 94. — Chapuis, 1876, p. 168. — Gorham, 1891, p. 106.— Wickham, 1894,
p. 298.— Casey, 1899, p. 77. — Leng, 1903a, p. 36. — Blatchley, 1910, p. 511.—
Timberlake, 1919, p. 162. — Mader, 1926, p. 1 7. — Korschefsky, 1932, p. 318.—
Timberlake, 1943, p. 45.— Chapin, 1946, p. 2. — Wingo, 1952, p. 22. — Brown and
de Ruette, 1962, p. 648.— J. Chapin, 1974, p. 58. — Belicek, 1976, p. 338. Type-
species; Cocci nella tredecimpunctata L., by subsequent designation of Crotch, 1873.
Hippodamia {Hippodamia): lablokoff-Khnzorian, 1982, p. 308.
Hemisphaerica Hope, 1840, p. 1 57. — Mulsant, 1850, p. 16. — Korschefsky, 1932, p.
439. — Belicek, 1976, p. 338. Type-species; Coccinella quinquesignata BCirby, by
monotypy.
AdoniaMu\s2inX, 1846, p. 39. — Mulsant, 1850, p. 36. — Mulsant, 1866, p. 27.— Crotch,
1873, p. 368. — Crotch, 1874b, p. 94. — Mader, 1926, p. 18. — Korschefsky, 1932,
p. 345. — Timberlake, 1943, p. 45. — Brown and de Ruette, 1962, p. 648. — Belicek,
1976, p. 338. Type-species; Coccinella mutabilis Scriba, a synonym of Adonia
variegata (Goeze), by monotypy.
Hippodamia {Adonia): lablokolf-Khnzorian, 1982, p. 308.
Hippodamia {Parippidamia) lablokoff-Khnzorian, 1979, p. 5 1 . — lablokoff- Khnzo-
rian, 1982, p. 308. Type-species, Coccinella arctica Schneider, by original desig-
nation.
Coccinellini with length 3.0 to 8.0 mm. Form elongate, oval, femur visible beyond
lateral margin of elytron. Dorsal color usually red with black maculae. Apex of clypeus
truncate or feebly concave; anterolateral angle without projection or with slight for-
ward projection. Base of pronotum not margined. Lateral margin of elytron feebly
reflexed; epipleuron nearly flat, sloping downward slightly internally. Intercoxal pro-
cess of prosternum narrow, usually feebly convex, lacking carinae but with fine lateral
ridge. Mesosternum protuberant medially; with apical fossa for reception of pros-
ternal process. Metasternum with or without postcoxal line. Apex of middle and hind
tibiae each with 2 spurs. Each front and middle tarsus of male dilated in some species,
unmodified in others. Tarsal claw cleft (Fig. 587i). Postcoxal line on abdomen present,
or absent. Male genitalia symmetrical. Female genitalia with large infundibulum;
coxal plate elongate, stylus distinct (Fig. 595e).
Species of Hippodamia are superficially similar to members of several other coc-
cinelline genera, but the cleft tarsal claws are unique to members of Hippodamia.
Hippodamia is mostly nearctic and palearctic in distribution with 9 species either
restricted to the Palearctic Region, or are holarctic. I recognize 25 names as valid in
America north of Mexico, and one species, H. koebelei Timberlake, is known only
from Mexico. The genus was revised by Chapin ( 1 946) and I have essentially followed
his classification except for the elimination of a few subspecific names and the addition
of a key to species. I have attempted to construct a key based on external characters,
but with limited success. Male genitalia or female still must be examined in many
instances because I have not been able to find external characters to distinguish some
species. Chapin (1946) divided the American species into 4 groups based on the
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NORTH AMERICAN COCCINELLIDAE
707
structure of the male genitalia and, in part, the presence or absence of denticles in
the female bursa. Chapin also included illustrations of almost all the dorsal color
variations of species of Hippodamia; and his paper should be consulted for these
because I have included only illustrations of the basic patterns. Brown and de Ruette
(1962) compared the genera Hippodamia and Adonia but did not synonymize the 2
names. Belicek (1976) placed Adonia as a junior synonym of Hippodamia and I
follow this course here. lablokoff-Khnzorian treated Adonia as a subgenus of Hip-
podamia.
Animal food of members of Hippodamia consists of aphids, with specihc host
records as follows; Acyrthosiphon dirhodum (Walker), Acyrthosiphon pisum (Harris),
Aphis forbesi Weed, Aphis gossypii Glover, Aphis nerii Boyer de Fonscolombe, Aphis
pomi Degeer, Aphis rumicis L., Aphis viburni Scopoli, Brevicoryne brassicae (L.),
Chromaphis juglandicola (Kaltenbach), Capitophorus eleagni (del Guercio), Eho-
soma lanigemm (Hausman), Hyadaphis erysimi (Kaltenbach), Macrosiphum avenae
(F.), Macrosiphum euphorbiae (Thomas), Macrosiphum rosae (L.), Monellia caryella
(Fitch), Monelliopsis californica (Essig), Monelliopsis caryae (Monell), Myzus cerasi
(F.), Myzus persicae (Sulzer), Nearctaphis bakeri (Cowen), Nearctaphis crataegifoliae
(Fitch), Periphyllus negundinis (Thomas), Phorodon humuli (Schrank), Rhopalosi-
phum (Fitch), Schizaphis graminum (Rondani).
Key to species of Hippodamia
1 . Mesepimeron black (occasionally bicolored in arctica) 2
- Mesepimeron entirely yellow 4
2(1). Pronotal black area with apical and basal pale indentations (Fig. 594g); form short,
broad arctica (Schneider)
- Pronotal blaek area without pale indentations (Fig. 5831); form elongate, slender
3
3(2). Head dull, strongly alutaceous, densely punctured, with small, oval, median pale
spot; elytral spots confluent (Fig. 5831) americana Crotch
Head shiny, feebly alutaceous, not densely punctured, with transverse pale spot;
elytral spots not confluent or feebly so (Fig. 5861) washingtoni Timberlake
4(1). Elytron almost entirely black except for apical pale spot or band (Fig. 6041), or
maculate but with lateral margin of elytron black (Fig. 604g)
moesta moesta LeConte
- Elytron not black, lateral margin of elytron never black 5
5(4). Elytron dull, strongly alutaceous, bivittate, often with median vitta broken apically
6
- Elytron shiny, not vittate 8
6(5). Pronotum with pale convergent spots (Fig. 6 1 21) 7
- Pronotum without pale convergent spots (Fig. 5851) falcigera Crotch
7(6). Median vitta of elytron complete (Fig. 6121); coastal California
sinuata sinuata Mulsant
- Median vitta of elytron broken apically (Fig. 6 1 2g); not occurring in coastal Cal-
ifornia sinuata crotchi Casey
8(5). Pronotum without convergent pale spots (Fig. 5811); elytron with 7 discrete black
spots; form elongate, slender tredecimpunctata tibialis (Say)
- Species not agreeing with all above statements 9
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9(8). Pronotum with pale median spot at base (Fig. 5891) 10
Pronotum without pale median spot at base (Fig. 596a) 14
10(9). Apex of elytron with sutural margin black (Figs. 589f) 11
- Apex of elytron wih sutural margin never black (Fig. 587f-h) 12
11(10). Male genitalia as in figure 589a-d apicalis Casey
- Male genitalia as in Figure 592a-d expur gata Casey
12(10). Tarsal claw with tooth not closely appressed (Fig. 59 li); Pacific coastal region . 13
- Tarsal claw with tooth closely appressed (Fig. 587i); transcontinental
parenthesis (Say)
1 3( 1 2). Markings on elytron heavy; pronotal markings reduced (Fig. 59 Ig); Pacific North-
west lunatomaculata dobzhanskyi Chapin
- Markings on elytron reduced (Fig. 59 If); pronotum with black area heavy; Cali-
fornia lunatomaculata lunatomaculata Motschulsky
14(9). Elytron immaculate except scutellar spot usually present 15
Elytron with at least a transverse basal band, usually with addditional black
maculae 20
15(14). Pronotum without convergent pale spots quinquesignata ambigua LeConte
Pronotum with convergent pale spots 16
16(15). Apex of basal lobe of male genitalia broadly triangular (Fig. 612a); northern
California sinuata crotchi Casey
- Apex of basal lobe of male genitalia not broadly triangular 17
17(16). Male genitalia with ventral ala well developed, visible in dorsal view (Fig. 606a)
19
- Male genitalia with ventral ala not visible in dorsal view (Fig. 598a) 18
18(17). Male genitalia as in Figure 598a glacialis extensa Mulsant
- Male genitalia as in Figure 595a quinquesignata ambigua LeConte
19(17). Basal lobe of male genitalia slender (Fig. 606a) convergens Guerin
- Basal lobe of male genitalia broad (Fig. 604a) moesta politissima LeConte
20(14). Pronotum with strong convergent pale spots (Fig. 6021); elytron with 6 or more
black maculae, maculae often confluent; Ontario to Missouri, west to Saskatch-
ewan quindecimmaculata Mulsant
- Pronotum and elytron not maculate as above, or if similar, not occurring east of
Colorado 21
2 1 (20). Pronotal black area with lateral extension often extending to margin of pronotum,
isolating or partially isolating the pale posterolateral area (Fig. 604j); or postero-
lateral area entirely black (Fig. 6041) 22
Pronotal black area without distinct lateral extension, pronotum with more or
less uniform pale border (Fig. 6121) 26
22(21). Elytron immaculate except for black subbasal band (Fig. 599f); coastal California
glacialis extensa Mulsant
- Elytron with or without subbasal band, additional black maculae present; coastal
California and elsewhere 23
23(22). Elyton with maculae consistently heavy, confluent (Fig. 604j); southern British
Columbia to northern Colorado moesta bowditchi Johnson
Elytron with maculae reduced, if confluent, only narrowly so (Fig. 608f, g) (except
lecontei in east central California) 24
24(23). Basal lobe of male genitalia lacking dorsal crest caseyi Johnson
- Basal lobe of male genitalia with dorsal crest 25
25(24). Basal lobe of male genitalia with dorsal crest bilobed .... glacialis lecontei Mulsant
Basal lobe of male genitalia with dorsal crest entire
quinquesignata qinquesignata (Kirby)
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709
26(21). Species occurring in coastal region from British Columbia to northern Oregon . 27
Species not occurring in above region 30
27(26). Length usually less than 5.50 mm; maculation on elytron, heavy, confluent (Fig.
612j) 28
- Length usually more than 5.50 mm; maculation of elytron reduced, not or only
feebly confluent 29
28(27). Pronotum with strong, convergent pale spots sinuata spuria LeConte
Pronotum without convergent pale spots oregonensis Crotch
29(27). Elytron with median black spots usually united to form transverse band (Fig.
596a); pronotum without or with convergent pale spots
quinquesignata quinquesignata (Kirby)
Elytron without transverse band (Fig. 606f); pronotum always with convergent
pale spots convergens Guerin
30(26). Species occurring from Saskatchewan and Colorado eastward 31
Species occurring from Saakatchewan, Colorado, and New Mexico westward . . 33
3 1(30). Pronotum always with strong, convergent pale spots; elytral maculation light, not
confluent (Fig. 606h) convergens Guerin
- Pronotum with or without convergent pale spots; elytral maculation heavy, usually
at least partly confluent (Fig. 599a) 32
32(31). Base of elytron without transverse black band; common glacialis glacialis (F.)
- Base of elytron usually with transverse black band; rare
quinquesignata quinquesignata (Kirby)
33(30). Pronotum always with strong, convergent pale spots; elytral maculation light, not
confluent (Fig. 606h); male genitalia as in Figure 606a convergens Guerin
- Pronotum with or without convergent pale spots; elytral maculae usually heavy.
somewhat confluent 34
34(33). Length usually less than 5.50 mm; form elongate, narrow (Fig. 612f) 35
Length usually more than 5.50 mm; form broad, robust (Fig. 599a) 36
35(34). Pronotum with convergent pale spots sinuata crotchi Casey
Pronotum without convergent pale spots oregonensis Crotch
36(34). Species occurring from Colorado and New Mexico to Idaho and eastern California;
basal lobe of male genitalia with bilobed crest glacialis lecontei Mulsant
- Species occurring from Alaska and Yukon Territory to New Mexico and southern
California; basal lobe of male genitalia with crest not bilobed
quinquesignata quinquesignata (Kirby)
Hippodamia tredecimpunctata tibialis (Say)
Fig. 58Ia-h; Map, Fig. 582
Coccinella tibialis Say, 1824, p. 94.
Hippodamia tibialis: Mulsant, 1850, p. 10.— Timberlake, 1919, p. 165.— Korschef-
sky, 1932, p. 331.— Timberlake, 1943, p. 51.
Hippodamia 13 -punctata: Crotch, 1873, p. 368.— Crotch, 1874b, p. 94 (in part).—
Wickham, 1894, p. 300. — Casey, 1899, p. 77. — Leng, 1903a, p. 40. — Blatchley,
1910, p. 511. -Leng, 1920, p. 215.
Hippodamia tredecimpunctata tibialis: Chapin, 1946, p. 5.— Wingo, 1952, p. 45.—
Hatch, 1961, p. 168. — Brown and de Ruette, 1962, p. 649.
Hippodamia tredecimpunctata: Belicek, 1976, p. 342. — lablokoff-Khnzorian, 1982,
p. 318.
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Fig. 581. Hippodamia tredecirnpunctata tibialis.
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711
Fig. 582. Distribution. Hippodamia tredecimpunctata tibialis.
Diagnosis. Length 4.50 to 6.40 mm, width 2.65 to 3.85 mm. Pronotum completely
black medially, or with black area broken anterolaterally; elytron with 7 black spots,
spots varying from completely free to somewhat confluent (Fig. 581f-h). Male gen-
italia as in Figure 581a-d. Female genitalia as in Figure 58 le.
Discussion. This is an easily recognized species because of the numerous, always
present elytral spots and small, slender form. The nominate subspecies is holarctic
from Europe to western Asia, with another subspecies in Japan, China, and eastern
Siberia.
Type locality. “Missouri”.
Type depository. Type lost.
Distribution. Figure 582. Newfoundland to South Carolina, west to Alaska and
northern California.
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Fig. 583. Hippodamia americana.
Hippodamia americana Crotch
Fig. 583a-g; Map, Fig. 584
Hippodamia americana Crotch, 1873, p. 368.— Crotch, 1874b, p. IV. — Wickham,
1894, p, 306.— Johnson, 1910, p. 52. — Korschefsky, 1932, p. 338. — Chapin, 1946,
p. 6. — Wingo, 1952, p. 45. — Brown and de Ruette, 1962, p. 549. — Belicek, 1976,
p. 342.
Diagnosis. Length 4.40 to 5.10, with 2.70 to 3.0 mm. Elytron somewhat vittate in
appearance (Fig. 583f, g), apical spots connected or free. Mesepimeron black. Male
genitalia as in Figure 583a-d. Female genitalia as in Figure 583e.
Discussion. This is not a commonly collected species, but is transcontinental in
distribution. The black mesepimeron and vittate appearing elytron distinguish H.
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NORTH AMERICAN COCCINELLIDAE
713
Fig. 584. Distribution. Hippodamia americana (dot); H. falcigera (star); H. washingtoni
(open star).
americana from other species of Hippodamia. I accept a female in the LeConte
collection labeled “H.B./Type 6685(red paper)/americana Cr. Type/Hippodamia
americana Cr.” as the holotype.
Type locality. “Hudson’s Bay”.
Type depository. MCZ.
Distribution. Figure 584. ALBERTA: McMurray. BRITISH COLUMBIA: Fraser
Lake. MANITOBA: Madge Lake. NORTHWEST TERRITORIES: Fort Smith; Nor-
man Wells. ONTARIO: Moose Factory; Ogoki. SASKATCHEWAN: Waskesiu Lake.
MICHIGAN: Lake Superior; White Fish Point. WISCONSIN: Waupaca.
Hippodamia falcigera Crotch
Fig. 585a-f; Map, Fig. 584
Hippodamia falcigera Crotch, 1873, p. 368.— Wickham, 1894, p. 306. — Leng, 1903,
p. 44.— Casey, 1908, p. 399.— Johnson, 1910, p. 55. — Korschefsky, 1932, p. 341.—
Timberlake, 1943, p. 51. — Chapin, 1946, p. 6. — Hatch, 1961, p. 168. — Brown and
de Ruette, 1962, p. 650.-Belicek, 1976, p. 342.
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Fig. 585. Hippodamia falcigem.
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NORTH AMERICAN COCCINELLIDAE
715
Hippodamia sinuata albertana Casey,, 1924, p. 157. — Chapin, 1946, p. 7.
Ceratomegilla cottlei Nunenmacher, 1934a, p. 20.
Hippodamia cottlei: Chapin, 1946, p. 6.
Diagnosis. Length 5.0 to 6.0 mm, width 2.90 to 3.50 mm. Elytron bivittate (Fig.
585f) with broad median vitta sometimes broken apically. Mesepimeron yellow.
Male genitalia as in Figure 585a-d. Female genitalia as in Figure 585e.
Discussion. This is not a commonly collected species and the geographic range is
restricted. The vittate appearance is similar to that of H. sinuata sinuata or H. s.
crotchi, but falcigera is larger and the 3 forms are allopatric. I here designate and
label a female of falcigera bearing the labels “H.B./Type 6684/falcigera Cr. Type/
Hippodamia falcigera Cr.” as the lectotype. Two other specimens in the series labeled
“SI L.” (Slave Lake) are designated and labeled as paralectotypes. The type of H. s.
albertana in the Casey collection is a unique female (holotype). The type of C. cottlei
is a unique male (holotype) labeled “Yellowstone Pk. VII-8-30/E. R. Leach/male
sign/Ceratomegilla cottlei Nun.”
Type locality. Of falcigera. Slave Lake, “Hudson’s Bay” (lectotype here designated);
of albertana, Edmonton, Alberta; of cottlei, Yellowstone Park, Wyoming.
Type depository. Of falcigera, MCZ; of albertana, USNM (3551 1); of cottlei, CAS.
Distribution. Figure 584. ALBERTA: Banff; Edmonton; Hotchkiss; McMurray;
Tofield. BRITISH COLUMBIA: Chilcotin; Summerland. NORTHWEST TERRI-
TORIES: Aklavik; Fort Simpson; Fort Smith. YUKON TERRITORY: Canyon Creek;
Haines Junction; Stewart River. IDAHO: Moscow. WYOMING: Yellowstone Park.
Hippodamia washingtoni Timberlake
Fig. 586a-h; Map, Fig. 584
Hippodamia washingtoni Timh^rX^kQ, 1939, p. 265. — Chapin, 1946, p. 7.— Hatch,
1961, p. 168. — Brown and de Ruette, 1962, p. 650. — Belicek, 1976, p. 343.
Diagnosis. Length 5.40 to 6.70 mm, width 3.25 to 4.0 mm. Dorsal color pattern
variable (Fig. 576f-h), but usually spotted, apical spots free or feebly connected.
Mesepimeron black. Male genitalia as in Figure 586a-d. Female genitalia as in Figure
586e.
Discussion. The black mesepimeron and shiny head distinguish H. washingtoni.
The male genitalia are similar to those of H. falcigera, but in external appearance
H. washingtoni is most similar to H. americana.
Type locality. Longmire Spring, Mount Rainier, Washington.
Type depository. UCR.
Distribution. Figure 584. BRITISH COLUMBIA: Anyox. IDAHO: Moscow Mt.
OREGON: Bear Springs; Blue Mountains; Clackamas Lake; Crater Lake; Mt. Hood.
WASHINGTON: Hoquiam; Monroe.
Hippodamia parenthesis (Say)
Fig. 587a-i; Map, Fig. 588
Coccinella parenthesis Say, 1824, p. 93.
Coccinella tridens Kirby, 1837, p. 229.
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Fig. 586. Hippodamia washingtoni.
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NORTH AMERICAN COCCINELLIDAE
717
Fig. 587. Hippodamia parenthesis.
Adonia tridens: Mulsant, 1850, p. 40.
Hippodamia tridens: Crotch, 1874b, p. 97.
Adonia parenthesis: Mulsant, 1850, p. 40. — Mulsant, 1866, p. 32.
Hippodamia parenthesis: Crotch, 1873, p. 368.— Crotch, 1874b, p. 97.— Wickham,
1894, p. 300. — Casey, 1899, p. 81. — Leng, 1903a, p. 44. — Casey, 1908, p. 399.—
Blatchley, 1910, p. 109.— Johnson, 1910, p. 52.— Timberlake, 1919, p. 165.—
Korschefsky, 1932, p. 343.— Chapin, 1946, p. 8.— Wingo, 1952, p. 45. — Hatch,
1961, p. 167.— Brown and de Ruette, 1962, p. 650. — Belicek, 1976, p. 343.
Hippodamia parenthesis albomacula Fitch, 1862, p. 853.
Hippodamia parenthesis approximata Fitch, 1862, p. 853.
Hippodamia parenthesis confluenta Fitch, 1862, p. 853.
Hippodamia parenthesis connata Fitch, 1862, p. 853.
Hippodamia parenthesis discopunctata Fitch, 1862, p. 853.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 588. Distribution. Hippodamia parenthesis.
Hippodamia parenthesis insulata Fitch, 1862, p. 853.
Hippodamia parenthesis linearis Fitch, 1862, p. 853.
Hippodamia parenthesis lituricollis Fitch, 1862, p. 853.
Hippodamia parenthesis permacrifrons Fitch, 1862, p. 853.
Hippodamia parenthesis triangularis Fitch, 1862, p. 853.
Hippodamia parenthesis tridentifrons Fitch, 1862, p. 853.
Hippodamia parenthesis nimia Fitch, 1862, p. 853.
Hippodamia parenthesis Sih. albomaculata: Leng, 1920, p. 215. — Korschefsky, 1932,
p. 343.
Hippodamia parenthesis ab. approximata: Leng, 1920, p. 215. — Korschefsky, 1932,
p. 343.
Hippodamia parenthesis ah. confluenta: Leng, 1920, p. 215. — Korschefsky, 1932, p.
343.
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NORTH AMERICAN COCCINELLIDAE
719
Hippodamia parenthesis ab. connata: Leng, 1920, p. 21 5. — Korschefsky, 1932, p.
343.
Hippodamia parenthesis 2ib. discopunctata: Leng, 1920, p. 215. — Korschefsky, 1932,
p. 343.
Hippodamia parenthesis ab. insulata: Leng, 1920, p. 2 1 5. — Korschefsky, 1932, p.
343.
Hippodamia parenthesis dih. linearis: Leng, 1920, p. 215. — Korschefsky, 1932, p. 343.
Hippodamia parenthesis ab. lituricollis: Leng, 1920, p. 21 5. — Korschefsky, 1932, p.
343.
Hippodamia parenthesis ab. permacrifrons: Leng, 1920, p. 21 5. — Korschefsky, 1932,
p. 343.
Hippodamia parenthesis ab. triangularis: Leng, 1920, p. 215. — Korschefsky, 1932,
p. 343.
Hippodamia parenthesis ab. tridentifrons: Leng, 1920, p. 21 5. — Korschefsky, 1932,
p. 343.
Hippodamia parenthesis ab. nimia: Leng, 1920, p. 215. — Korschefsky, 1932, p. 343.
Diagnosis. Length 3.75 to 5.60 mm, width 2.25 to 4.50 mm. Elytron spotted, apical
spots often suffused, sutural margin at apex without black spot (Fig. 587f-h). Tarsal
claw with tooth closely appressed (Fig. 587i). Male genitalia as in Figure 587a-d.
Female genitalia as in Figure 587e.
Discussion. This species ranges from Nova Scotia to South Carolina and west to
Alaska and California. East of the Mississippi River it is easily recognized because
no similar appearing species occur there. West of the Mississippi it can be confused
with lunatomaculata. The key characters will usually separate examples of paren-
thesis, but male genitalia must be examined in some instances. A male syntype of
tridens labeled “Syntype/Type/N. Amer./Coccinella tridens Kirby n. amer 5959. Rev.
W. Kirby” is here designated and labeled as the lectotype.
Type locality. Of parenthesis, not stated, “United States”; of all Fitch names, “New
York”; of tridens, “North America” (lectotype here designated).
Type depository. Of parenthesis, type lost; of Fitch types, not located; of tridens,
BMNH.
Distribution. Figure 588. Nova Scotia to South Carolina, west to Alaska and Cal-
ifornia.
Hippodamia apicalis Casey
Fig. 589a-i; Map, Fig. 590
Hippodamia apicalis Casey, 1899, p. 81. — Weise, 1899b, p. 377.— Casey, 1908, p.
399.— Johnson, 1910, p. 54. — Hatch, 1961, p. 167. — Belicek, 1976, p. 344.
Hippodamia parenthesis var. apicalis: Leng, 1903a, p. 44.
Hippodamia lunatomaculata apicalis: Timberlake, 1919, p. 166.
Hippodamia lunatomarginata ab. apicalis: Leng, 1920, p. 215. — Korschefsky, 1932,
p. 342.
Hippodamia apicalis apicalis: Chapin, 1946, p. 9. — Brown and de Ruette, 1962, p.
650.
Hippodamia lengi Johnson, 1910, p. 55.— Timberlake, 1919, p. 1 76. New Synonymy.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93( 1 )
Fig. 589. Hippodarnia apicalis.
Hippodamia lunatomarginata ab. lengi: Leng, 1920, p. 215. — Korschefsky, 1932, p.
342.
Hippodamia apicalis lengi: Chapin, 1946, p. 9.
Adalia nigromaculata Nunenmacher, 1934a, p. 20.
Hippodamia nigromaculata: Chapin, 1946, p. 9.
Hippodamia apicalis rr/co/or Nunenmacher, 1946, p. 72. — Hatch, 1961, p. 167.
Diagnosis. Length 3.50 to 4.70 mm, width 2.25 to 2.90 mm. Color pattern of
elytron variable (Fig. 589f-i), but always with suture of elytron black, at least at
extreme apex. Tarsal claw with tooth not closely appressed. Male genitalia as in
Figure 589a-d. Female genitalia as in Figure 589e.
Discussion. I have not been able to separate this species from H. expurgata except
by examination of the male genitalia. Chapin (1946) illustrated a specimen of H.
apicalis which lacked a black spot at the sutural apex, but on examining the USNM
collection, I find that this specimen is H. parenthesis misidentified as H. apicalis. I
have not seen any H. apicalis that lack this sutural spot. Neither H. parenthesis nor
H. lunatomaculata possess a sutural spot, therefore we have 2 pairs of species, each
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NORTH AMERICAN COCCINELLIDAE
721
Fig. 590. Distribution. Hippodamia apicalis (shaded); H. lunatomaculata lunatomaculata
(star); H. 1. dobzhanskyi (dot).
pair readily separable from the other. There are 4 types of H. apicalis in the Casey
collection, the 4th of which (male) I here designate and label the lectotype, the
remainder as paralectotypes. Hippodamia lengi was treated as a subspecies by Chapin
(1946), but I have seen specimens of typical H. apicalis that grade into the H. lengi
phenotype in a geographic dine, therefore I consider H. lengi a junior synonym of
H. apicalis. Johnson designated a specimen in the Carnegie Museum (Pittsburgh) as
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
the holotype of H. lengi. Two type specimens of A. nigromaculata Nunenmacher are
in the CAS Collection. I designate and label a specimen labeled “San Diego Co. Cal./
G.H. Field collector/ Adalia nigromaculata Nun.” as the lectotype.
Type locality. Of apicalis, Reno, Washoe Co., Nevada (lectotype here designated);
of lengi, California; of tricolor, Lassen Co., California; of nigromaculata, San Diego,
California (lectotype here designated).
Type depository. Of apicalis, USNM (35513); of lengi, CM; of tricolor, CAS; of
nigromaculata, CAS.
Distribution. Figure 590. Montana to New Mexico, west to southern British Co-
lumbia and southern California.
Hippodamia lunatomaculata lunatomaculata Motschulsky
Fig. 591a-f; Map, Fig. 590
Hippodamia lunatomaculata Motschulsky, 1845b, p. 382.— Timberlake, 1919, p.
166 (in part).— Timberlake, 1943, p. 10. — Hatch, 1961, p. 167. — Belicek, 1976, p.
344.
Hippodamia lunatomaculata lunatomaculata: Chapin, 1946, p. 10.
Hippodamia lunatomarginata: Korschefsky, 1932, p. 342.
Hippodamia parenthesis C?L^Qy, 1899, p. 81 (in part).— Johnson, 1910, p. 53 (in part).
Diagnosis. Length 4. 1 5 to 5.60 mm, width 2.70 to 3.70 mm. Pronotal color pattern
heavily maculate; elytron lightly maculate with spots not connected; sutural margin
at apex without black spot (Fig. 5911)- Tarsal claw with tooth not closely appressed
(Fig. 59 li). Male genitalia as in Figure 591a-d. Female genitalia as in Figure 59 le.
Discussion. Hippodamia 1. lunatomaculata is likely to be confused only with H.
parenthesis which has the tooth of the tarsal claw more closely appressed. This
character is difficult to assess without some experience and male genitalia must be
examined in doubtful cases.
Type locality. Vicinity of San Francisco Bay, California (restricted by Chapin,
1946).
Type depository. Type not examined.
Distribution. Figure 590. CALIFORNIA: Half Moon Bay; Willows.
Hippodamia lunatomaculata dobzhanskyi Chapin
Fig. 59 Ig, h; Map, Fig. 590
Hippodamia lunatomaculata dobzhanskyi Chapin, 1946, p. 11. — Hatch, 1961, p.
167. — Brown and de Ruette, 1962, p. 650.
Description as for H. 1. lunatomaculata except pronotum with reduced maculation
and elytron with heavy maculation (Fig. 59 Ig, h).
I am maintaining the 2 subspecies proposed by Chapin (1946), but without any
degree of confidence in the validity of this arrangement. These are mainly coastal
forms and it is most likely that they form a continuous geographic dine that cannot
be divided at any one point. See discussion under /. lunatomaculata for comparative
remarks.
Type locality. Port Angeles, Washington.
Type depository. USNM (57891).
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NORTH AMERICAN COCCINELLIDAE
723
Fig. 591. Hippodamia lunatomaculata lunatomaculata; H. 1. dobzhanskyi.
Distribution. Figure 590. BRITISH COLUMBIA: Nainamo, Victoria. CALIFOR-
NIA: Tacumoa. OREGON: Amity; Corvallis; Dever; Forest Grove; Newport; Wil-
lamette Valley. WASHINGTON: Olympic Peninsula; Medical Lake; Seattle.
Hippodamia expurgata Casey
Fig. 592a-i; Map, Fig. 593
Hippodamia parenthesis expurgata Casey, 1908, p. 400.
Hippodamia lunatomaculata expurgata: Timberlake, 1919, p. 166.
Hippodamia lunatomarginata ab. expurgata: Leng, 1920, p. 215. — Korschefsky, 1932,
p. 342.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 592. Hippodamia expurgata.
Hippodamia expurgata: Chapin, 1946, p. 12. — Hatch, 1961, p. 167. — Brown and de
Ruette, 1962, p. 650.
Diagnosis. Length 3.50 to 5.0 mm, width 2.20 to 3.25 mm. Color pattern as in H.
apicalis except sutural spot on elytron occasionally lacking (Fig. 592f-i). Male gen-
italia as in Figure 592a-d. Female genitalia as in Figure 592e.
Discussion. Examination of the male genitalia is the only certain way I can find to
separate this species from H. apicalis. The 2 species are sympatric in the Rocky
Mountain states, allopatric or nearly so in the Pacific Coast states. Chapin (1946)
stated that in lightly maculate specimens of H. expurgata the apical sutural spot is
usually absent; on examination of the material Chapin saw, I found that the spot is
usually present but extremely reduced. Those specimens with the spot completely
gone will key to H. parenthesis, and again the male genitalia must be examined.
Belicek (1976) synonymized H. expurgata with H. apicalis, apparently basing this
decision on those male specimens seen by Chapin (1946) that have the apex of the
basal lobe somewhat attenuate. I do not accept this synonymy because even in the
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NORTH AMERICAN COCCINELLIDAE
725
Fig. 593. Distribution. Hippodamia expurgata (shaded); H. arctica (dot).
specimens in question it is apparent what species they are, and in normal individuals
there is no question that the genitalia are substantially different. There are 2 types
of H. expurgata in the Casey collection, the second of these, a male, I here designate
and label as the lectotype, the first (female) as a paralectotype.
Type locality. Boulder, Colorado (lectotype here designated).
Type depository. USNM (35514).
Distribution. Figure 593. Saskatchewan to New Mexico, west to Yukon Territory
and Arizona.
Hippodamia arctica (Schneider)
Fig. 594a-i; Map, Fig. 593
Coccinella arctica Schneider, 1792, p. 148.
Adonia arctica: Korschefsky, 1932, p. 346.
Adonia amoena Scott, 1933, p. 126 (not Adonia amoena Falderman).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 594. Hippodamia arctica.
Hippodamia arctica: Crotch, 1874b, p. 97. — Brown and de Ruette, 1962, p. 650.—
Belicek, 1976, p. 344.
Hippodamia (Parippodamia) arctica: lablokoff-Khnzorian, 1979, p. 51.— lablokoff-
Khnzorian, 1982, p. 322.
Diagnosis. Length 4.0 to 4.50 mm, width 2.50 to 2.80 mm. Dorsal color mostly
black with yellowish red marking (Fig. 594g-i). Mesepimeron usually black, occa-
sionally bicolored, rarely yellow. Postcoxal line always complete (Fig. 594a). Male
genitalia as in Figure 594b-e. Female genitalia as in Figure 594f.
Discussion. The short, broad form, mostly black dorsal surface, always complete
postcoxal lines and usually black mesepimeron distinguish H. arctica. Brown and de
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NORTH AMERICAN COCCINELLIDAE
727
Ruette ( 1 962) placed this species in Hippodamia and discussed the generic and specific
relationships.
Type locality. “Lapland.”
Type depository. Type not examined.
Distribution. Figure 593. BRITISH COLUMBIA: Summit Lake, mi. 379 and 392,
Alaska Hwy. LABRADOR: Hebron. QUEBEC: Fort Chimo; Great Whale R.; Green-
ly Island. YUKON TERRITORY: Kirkman Creek; Swim Lakes. ALASKA: Eagle;
Rampart House; Umiat.
Hippodamia quinquesignata quinquesignata (Kirby)
Fig. 595a-e, 596a-d; Map, Fig. 597
Coccinella 5-signata Kirby, 1837, p. 230.
Hemisphaerica quinquesignata: Hope, 1840, p. 157.
Hippodamia quinquesignata: Mulsant, 1850, p. 15. — Mulsant, 1866, p. 10.— Crotch,
1873, p. 366.— Crotch, 1874b, p. 95. — Casey, 1899, p. 78. — Leng, 1903a, p. 40.—
Casey, 1908, p. 395.— Timberlake, 1919, p. 171. — Korschefsky, 1932, p. 344.—
Wingo, 1952, p. 45. -Hatch, 1961, p. I71.-Belicek, 1976, p. 345.
Hippodamia quinquesignata quinquesignata: Chapin, 1946, p. 15. — Brown and de
Ruette, 1962, p. 651.
Hippodamia mulsanti LeConte, 1852, p. 131. — Crotch, 1873, p. 366.
Hippodamia leporina Mulsant, 1856, p. 135. — Crotch, 1873, p. 366.
Hippodamia quinquesignata ab. leporina: Leng, 1920, p. 216. — Korschefsky, 1932,
p. 344.
Hippodamia subsimilis Casey, 1899, p. 79.— Timberlake, 1919, p. 172.— Chapin,
1946, p. 13.
Hippodamia vernix subsimilis: Casey, 1908, p. 396.
Hippodamia ver/i/x Casey, 1899, p. 79. — Timberlake, 1919, p. 171.— Chapin, 1946,
p. 13.
Hippodamia coccinea Casey, 1908, p. 395. — Timberlake, 1919, p. 171.
Hippodamia uteana Casey, 1908, p. 397.— Timberlake, 1919, p. 171.
Hippodamia quinquesignata var. uteana: Chapin, 1946, p. 15. — Belicek, 1976, p.
345.
Hippodamia uteana quadraria Casey, 1924, p. 156.— Chapin, 1946, p. 13.
Hippodamia convergens pugetana Casey, 1924, p. 156. — Belicek, 1976, p. 345.
Hippodamia quinquesignata var. pugetana: Chapin, 1946, p. 15.
Diagnosis. Length 4.0 to 7.0 mm, width 2.80 to 5.0 mm. Pronotum with or without
convergent pale spots; elytral color pattern extremely variable (Fig. 596a-d). Male
genitalia as in Figure 595a-d. Female genitalia as in Figure 595e.
Discussion. This species and some of the other large, robust species {H. glacialis,
H. convergens, H. caseyi, etc.) of Hippodamia are very difficult to distinguish without
referring to male genitalia. Hippodamia quinquesignata is a widespread species but
not particularly common in the east, and extremely variable in the dorsal color
pattern. Hippodamia glacialis, especially the subspecies H. lecontei, is usually con-
fused with H. quinquesignata. In eastern North America H. glacialis and H. quin-
quesignata can be separated on the key characters. Chapin ( 1 946) retained the names
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
H. pugetana and H. uteana as varieties, but I can see no basis for maintaining them
and consider them junior synonyms of H. quinquesignata. There are unique type
specimens (holotypes) of H. vernix, H. subsimilis, H. quadraria and H. coccinea in
the Casey collection. There are 19 types of pugetana, one of which, a male, I here
designate and label as the lectotype, the remainder as paralectotypes. There are 4
type specimens of uteana and I here designate and label a male as the lectotype, the
remainder as paralectotypes. The holotype of quinquesignata is a female labeled
“Type/ n. amer 5960a/Coccinella quinquesignata Kirby n. america. 5960. Rev. Wm.
Kirby.
Type locality. Of quinquesignata, ”Lat. 65“ (near Great Bear Lake); of mulsanti.
Pic River, Lake Superior; of leporina, California; of subsimilis, California?; of vernix,
Wyoming; of coccinea, Boulder, Colorado; of uteana, Nephi, Utah (lectotype here
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NORTH AMERICAN COCCINELLIDAE
729
Fig. 596. Hippodamia quinquesignata quinquesignata\ H. q. ambigua.
designated); of quadraria, Edmonton, Alberta; of pugetana, Fairfield, Washington
(lectotype here designated.
Type depository. Of quinquesignata, BMNH; of mulsanti, not examined; of lepor-
ina, not examined; of subsimilis (35506), vernix (35507), coccinea (35495), uteana
(35499), quadraria (35500), and pugetana (35501), USNM.
Distribution. Figure 597. Prince Edward Island to Yukon Territory and Alaska,
south to New Mexico and California.
Hippodamia quinquesignata ambigua LeConte
Fig. 596e, f; Map, Fig. 597
Hippodamia ambigua LeConte, 1852, p. 131.— Crotch, 1873, p. 366.— Crotch, 1874b,
p. 96. — Casey, 1899, p. 79. — Leng, 1903, p. 41.— Timberlake, 1919, p. 172.
Hippodamia 5-signata ambigua: Timberlake, 1943, p. 12.
Hippodamia quinquesignata ambigua: Chapin, 1946, p. 16. — Hatch, 1961, p. 173.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 597. Distribution. Hippodamia quinquesignata quinquesignata (shaded), eastern lo-
calities dotted); H. q. ambigua (cross hatch).
Hippodamia punctulata LeConte, 1852, p. 131.— Crotch, 1873, p. 366. — Crotch,
1874b, p. 96. -Casey, 1899, p. 79.-Timberlake, 1919, p. 172.
Hippodamia quinquesignata punctulata: Chapin, 1946, p. 16.
Hippodamia obliqua Casey, 1899, p. 79. — Chapin, 1946, p. 13.
Hippodamia quinquesignata obliqua: Timberlake, 1943, p. 12.
Description as for H. q. quinquesignata except elytron almost always immaculate;
pronotum with or without convergent pale spots (Fig. 596e, f)-
Crotch (1873) and Casey (1899) considered H. punctulata a synonym of H. am-
bigua, but Crotch (1874) revived the name H. punctulata and authors since have
considered it a subspecies or race of H. ambigua. Timberlake (1943) pointed out the
difficulty in separating H. ambigua and H. punctulata due to the variability of the
thoracic color pattern. Hippodamia punctulata is the form lacking convergent pro-
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NORTH AMERICAN COCCINELLIDAE
731
notal spots; H. ambigua has those spots; however, most large series from a single
locality will contain examples of both forms. I cannot logically maintain these forms
as subspecies and therefore regard H. punctulata as a junior synonym of H. ambigua.
Hippodamia obliqua Casey is a form of H. ambigua with the pronotum having the
black area reduced and the convergent pale spots large. Hippodamia q. ambigua is
a Pacific Coast subspecies ranging from southern California to northwestern Wash-
ington, but some of the northern series examined contain some specimens not sep-
arable from H. q. quinquesignata. In spite of this, I feel it best to maintain both
subspecies for the present. Hippodamia obliqua is represented by 7 types in the Casey
collection, and I here designate and label a male as the lectotype and the remainder
as paralectotypes. A female of H. ambigua in the LeConte collection labeled ’’(blue
disc)/4007/Type 6680 (red paper)/H. ambigua Lec.“ is here designated and labeled
as the lectotype. Three other specimens in the same series, each bearing a gold disc,
are designated and labeled as paralectotypes. A male of punctulata in the LeConte
collection labeled ’’(gold disc)/Type 6651 (red paper)///, punctulata LeC.“ is here
designated and labeled as the lectotype. Three other specimens in the same series,
each bearing a gold disc, are designated and labeled as paralectotypes.
Type locality. Of ambigua, Oregon (lectotype here designated); of punctulata, Cal-
ifornia (lectotype here designated); of obliqua, Santa Rosa, Sonoma Co., California
(lectotype here designated).
Type depository. Of ambigua and punctulata, MCZ; of obliqua, USNM (35503).
Distribution. Figure 597. Southern British Columbia to southern California (coastal
localities).
Hippodamia glacialis glacialis (F.)
Fig. 598a-e, 599a, b; Map, Fig. 600
Coccinella glacialis F., 1775, p. 80.
Hippodamia glacialis: Mulsant, 1850, p. 18. — Mulsant, 1866, p. 12. — Crotch, 1873,
p. 367.— Crotch, 1874b, p. 95.— Casey, 1899, p. 79. — Leng, 1903a, p. 41. — Blatch-
ley, 1910, p. 512.— Johnson, 1910, p. 19.— Timberlake, 1919, p. 174. — Korschef-
sky, 1932, p. 341. -Timberlake, 1943, p. 12.-Wingo, 1952, p. 45.-Belicek, 1976,
p. 346 (in part).
Hippodamia glacialis glacialis: Chapin, 1946, p. 18. — Brown and de Ruette, 1962,
p. 651.
Diagnosis. Length 5.50 to 8.0 mm, width 3.60 to 5.60 mm. Pronotum with or
without convergent pale spots; elytron not particularly variable, immaculate except
for apical spot and subapical transverse band, these sometimes suffused (Fig. 599a,
b). Male genitalia as in Figure 598a-d. Female genitalia as in Figure 598e.
Discussion. This is primarily an eastern North American subspecies where it is
easily recognized by the dorsal maculation. However, at the western limit of the
range (Manitoba, Saskatchewan, Colorado) specimens occur which are intergrades
with H. g. lecontei. Belicek (1976) stated that Chapin (1946) had synonymized H.
lecontei and H. extensa with H. glacialis, but this is not correct; Chapin merely
reduced them to subspecific rank, a decision with which I presently concur. A single
type specimen of H. glacialis exists and has been compared with eastern specimens
of H. glacialis by John Kingsolver.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Type locality. ’’America boreali.“
Type depository. Fabrician collection, Kiel.
Distribution. Figure 600. Quebec to South Carolina and Alabama, west to Sas-
katchewan and Colorado.
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NORTH AMERICAN COCCINELLIDAE
733
Fig. 599. a, b. Hippodamia glacialis glacialis. c-e. H. g. lecontei. f. H. g. extensa.
Hippodamia glacialis lecontei Mulsant
Fig. 599c-e; Map, Fig. 601
Hippodamia Mulsant, 1850, p. 1010. — Mulsant, 1866, p. 9.— Crotch, 1873,
p. 366.— Casey, 1899, p. 78. — Leng, 1903a, p. 41. — Casey, 1908, p. 396.— Tim-
berlake, 1919, p. 169.— Korschefsky, 1932, p. 341.
Hippodamia quinquesignata var. lecontii: Crotch, 1874b, p. 95.
Hippodamia convergens var. lecontei: Johnson, 1910, p. 23. — Chapin, 1946, p. 17.
Hippodamia convergens var. pseudoglacialis Johnson, 1910, p. 23. — Chapin, 1946,
p. 17.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Hippodamia lecontei abducens Casey, 1908, p. 396. — Leng, 1920, p. 216.
Hippodamia convergens var. defecta Johnson, 1910, p. 21. — Leng, 1920, p. 216.
Hippodamia Nunenmacher, 1934a, p. 21. — Chapin, 1946, p. 21.
Hippodamia glacialis lecontei: Chapin, 1946, p. 18. — Hatch, 1961, p. 173.— Brown
and de Ruette, 1962, p. 651.
Hippodamia glacialis mackenzie Chapin, 1946, p. 19. New Synonymy.
Diagnosis. Length 5.0 to 7.0 mm, width 3.60 to 4.70 mm. Pronotum usually
without convergent pale spots, or with spots reduced in size; elytron variable but
usually with apical black band or spots, some western specimens heavily maculate
(Fig. 599c-e). Genitalia as in H. g. glacialis.
Discussion. This subspecies is more variable in color pattern than the typical
subspecies, and more likely to be confused with other species. The subspecies H. g.
mackenziei Chapin I consider a junior synonym of H. g. lecontei because the H.
mackenziei color pattern can be found in specimens from Colorado and New Mexico.
In addition, specimens from the type locality of H. mackenziei vary from heavily
maculate (Fig. 599d) to the nearly immaculate appearance of H. g. extensa. There
are 2 type specimens of H. hoppingi in the CAS collection, I here designate and label
a male labeled ”Mt. Stillman, Tulare Co. Calif./ 1000 ft. Aug. 3, 1904/Hippodamia
hoppingi Nun.“ as the lectotype.
Type locality. Of lecontei, Santa Fe, New Mexico (type locality fixed by Chapin,
1946); of pseudoglacialis. New Mexico and northward”; of hoppingi, Mt. Stillman,
Tulare Co., California (lectotype here designated); of mackenziei. Glacier Lodge, Inyo
County, California.
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NORTH AMERICAN COCCINELLIDAE
735
Type depository. Of lecontei, MCZ; of pseudoglacialis, types not preserved; of
hoppingi, CAS; of mackenziei, USNM (57892).
Distribution. Figure 60 1 . Saskatchewan to New Mexico, west to Alberta and Cal-
ifornia.
Hippodamia glacialis extensa Mulsant
Fig. 599f; Map, Fig. 601
Hippodamia extensa Mulsant, 1850, p. 17. — Mulsant, 1866, p. 11. — Crotch, 1873, p.
366.— Casey, 1899, p. 79.— Timberlake, 1919, p. 174. — Korschefsky, 1932, p. 340.
Hippodamia quinquesignata var. extensa: Crotch, 1874b, p. 95. — Leng, 1903, p. 41.
Hippodamia convergens var. extensa: Johnson, 1910, p. 23.
Hippodamia glacialis extensa: Chapin, 1946, p. 19.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 602. Hippodamia quindecimmaculata.
Diagnosis. Length 5.0 to 7.0 mm, width 3.0 to 4.50 mm. Pronotum without pale
convergent spots; elytron immaculate or with basal transverse band (Fig. 599f).
Genitalia as in H. g. glacialis.
Discussion. I originally assumed that H. extensa was simply a junior synonym of
H. lecontei, but on examination of specimens, H. extensa has the apical ridge of the
mesosternum strongly reduced while it is normal in H. g. glacialis and g. lecontei.
The dorsal color pattern of extensa occurs in populations of lecontei and I don’t
consider that pattern a valid basis for distinguishing subspecies, but on the basis of
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NORTH AMERICAN COCCINELLIDAE
737
the different structure of the mesastemum, I maintain extensa as a subspecies of
glacialis. The geographic range of extensa is the San Francisco bay area of California,
a surprisingly restricted range for any species or subspecies ofHippodamia, suggesting
that something is not correct with the classification proposed.
Type locality. “Califomie septentrionale.”
Type depository. Type not examined.
Distribution. Figure 60 1 . CALIFORNIA: Alameda Co.
Hippodamia quindecimmaculata Mulsant
Fig. 602a-i; Map, Fig. 603
Hippodamia quindecimmaculata Mulsant, 1850, p. 20. — Mulsant, 1866, p. 12.—
Korschefsky, 1932, p. 343.— Chapin, 1946, p. 20.— Wingo, 1952, p. 45. — Brown
and de Ruette, 1962, p. 651.— J. Chapin, 1974, p. 59.
Hippodamia 1 5-maculata: Crotch, 1873, p. 367.— Crotch, 1874b, p. 95.— Casey,
1899, p. 81.-Timberlake, 1919, p. 169.
Hippodamia convergens 1 5-maculata: Leng, 1903a, p. 42.— Johnson, 1910, p. 27.
Diagnosis. Length 5.0 to 7.50 mm, width 3.25 to 4.60 mm. Pronotum with con-
vergent pale spots; elytron heavily maculate, spots sometimes feebly confluent (Fig.
602f-i). Male genitalia as in Figure 602a-d. Female genitalia as in Figure 602e.
Discussion. The heavily maculate appearance, strongly developed convergent spots
on the pronotum, and geographic range will distinguish most specimens of this
species. H. convergens is similar in appearance but not as heavily marked and usually
smaller.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 604. f, g. Hippodamia moesta moesta. h-j. H. m. bowditchi. i. H. m. politissima.
Type locality, “le bords du Missouri, dans FAmerique septentrionale”.
Type depository. Type not examined.
Distribution. Figure 603. ONTARIO: Batchawang Bay; Grand Bend. SASKATCH-
EWAN: Carlton; Elbow; Saskatchewan Landing; Saskatoon. Colorado: Denver IL-
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NORTH AMERICAN COCCINELLIDAE
739
LIONIS: “S. 111.”. IOWA: Chelsea; Sioux City. LOUISIANA: Baton Rouge; Con-
cordia Parish; Tensas Parish; West Feliciana Parish. MISSOURI: St. Louis.
NEBRASKA: Badger; Elliott; Lincoln; Omaha. OHIO: Sandusky. OKLAHOMA:
Durant. WISCONSIN: Madison.
Hippodamia moesta moesta LeConte
Fig. 604a-j; Map, Fig. 605
Hippodamia moesta heConlQ, 1854, p. 16. — Crotch, 1874b, p. 97. — Wickham, 1894,
p. 305.— Casey, 1899, p. 78. — Leng, 1903a, p. 41.— Timberlake, 1919, p. 170.—
Korschefsky, 1932, p. 342.-Belicek, 1976, p. 347.
Hippodamia lecontei var. moesta: Crotch, 1873, p. 367.
Hippodamia convergens moesta: Johnson, 1910, p. 45.
Hippodamia moesta moesta: Chapin, 1946, p. 21. — Hatch, 1961, p. 170.— Brown
and de Ruette, 1962, p. 652.
Diagnosis. Length 6.0 to 7.50 mm, width 4.0 to 5.20 mm. Pronotum with or
without pale dots or convergent spots; elytron usually black with pale spot on elytral
margin at apical (Fig. 604f), pattern variable (Fig. 604g) but lateral margin of
elytron always black in basal %. Male genitalia as in Figure 604a-d. Female genitalia
as in Figure 604e.
Discussion. The typical form of H. m. moesta is the most striking example of
Hippodamia in North America, but this black form is not constant. I have seen one
example in a series of black specimens from Humboldt Co., California, which has
the color pattern of H. m. bowditchi except that the lateral elytral margin is black in
the basal two-thirds. Hippodamia m. moesta is a coastal form occurring at low
altitudes from southern British Columbia to northern California. Belicek (1976)
incorrectly stated that Chapin (1946) had synonymized H. bowditchi and H. politis-
sima with H. moesta. A male of H. moesta in the LeConte collection labeled “(blue
disc)/Type 6602 (red paper)/H. moesta LeC.” is here designated and labeled as the
lectotype. One additional specimen in the same series, bearing a blue disc, is des-
ignated and labeled as a paralectotype.
Type locality. Prairie Paso, Oregon (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 605. BRITISH COLUMBIA: Bevan; Chilliwack; Fraser River,
Keremeos; Lillooet; Vancouver Island, Nanaimo. CALIFORNIA: Fort Seward;
Humboldt Co., Redwood Creek; Yreka. OREGON: Cherry ville; Dilley; Parkdale;
Turner. WASHINGTON: Seattle; Mt. Rainier, Longmire Springs; Monroe.
Hippodamia moesta bowditchi Johnson
Fig. 604h-j; Map, Fig. 605
Hippodamia bowditchi Johnson, 1910, p. 45.
Hippodamia moesta bowditchi: Leng, 1920, p. 216.— Chapin, 1946, p. 21.— Hatch,
1961, p. 170. — Brown and de Ruette, 1962, p. 652.
Description as for H. m. moesta except elytron yellow with black maculae (Fig.
604h, j).
The dorsal color pattern of H. bowditchi is very constant in the specimens ex-
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 605. Distribution. Hippodamia moesta moesta (star); H. m. bowditchi (dot); H. m.
politissima (triangle).
amined. The geographic range appears not to overlap that of m. moesta or H. m.
politissima in that I’ve not seen the color pattern of H. bowditchi from any Pacific
Coast locality but I have seen 2 California specimens, Hippodamia bowditchi occurs
in the same region as H. quinquesignata and the 2 species have similar color patterns,
however, //. bowditchi nearly always has the elytral maculae heavy and confluent,
H. quinquesignata rarely does.
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NORTH AMERICAN COCCINELLIDAE
741
Type locality. St. Maries, Idaho.
Type depository. Type not preserved.
Distribution. Figure 605. BRITISH COLUMBIA: Cawston; Cranbrook; Fernie;
Heywood’s Corner; Kaslo; Trinity Valley; Vavenby. CALIFORNIA: Modoc Co.;
Goose Lake; Plumas Co., Meadow Valley. COLORADO: Denver; Fort Collins.
IDAHO: Hayden’s Lake. MONTANA: Bitterroot Mts., w. of Thompson Falls; Flor-
ence; Gallatin Mts.; Kalispell; Missoula; Ravalli Co., Hamilton; Kalispell; Flathead
Co., LaSalle.
Hippodamia moesta politissima Casey
Fig. 604i; Map, Fig. 605
Hippodamia politissima Casey, 1899, p. 80.
Hippodamia ambigua politissima: Leng, 1903a, p. 41.
Hippodamia convergens ab. politissima: Korschefsky, 1932, p. 339.
Hippodamia moesta politissima: Chapin, 1946, p. 22.
Description as for H. m. moesta except pronotum with distinct, convergent pale
spots; elytron immaculate (Fig. 604i).
This subspecies can be separated from the immaculate form of H. convergens and
H. q. ambigua only by examination of genitalia. Thus far H. politissima is known
only from coastal southern California and I’ve not seen intergrades with H. m. moesta.
The type specimen is a male (holotype) in the Casey collection.
Type locality. Santa Cruz, California.
Type depository. USNM (35504).
Distribution. Figure 605. CALIFORNIA: San Luis Obispo.
Hippodamia convergens Guerin
Fig. 606a-i; Map, Fig. 607
Hippodamia convergens Guerin, 1842, p. 321. — Mulsant, 1850, p. 22. — Mulsant,
1866, p. 14.-LeConte, 1852, p. I30.-Crotch, 1873, p. 367. -Crotch, 1874b, p.
96. — Gorham, 1891, p. 153. — Weise, 1895a, p. 135. — Casey, 1899, p. 80. — Leng,
1903a, p. 40. — Casey, 1908, p. 398.— Johnson, 1910, p. 21.— Timberlake, 1919,
p. 168.— Korschefsky, 1932, p. 338.— Timberlake, 1943, p. 1 1. — Chapin, 1946, p.
22.— Wingo, 1952, p. 45. — Hatch, 1961, p. 171. — Brown and de Ruette, 1962, p.
652.— J. Chapin, 1974, p. 60. — Belicek, 1976, p. 347.
Hippodamia juncta Casey, 1899, p. 80.— Timberlake, 1919, p. 168. — Chapin, 1946,
p. 22.
Hippodamia modesta Melsheimer, 1847, p. 178. — LeConte, 1852, p. 130.
Hippodamia convergens var. obsoleta Crotch, 1873, p. 367. — Casey, 1908, p. 398.
Hippodamia praticola Mulsant, 1850, p. 23.— Weise, 1895a, p. 125.
Diagnosis. Length 4.20 to 7.30 mm, width 2.50 to 4.90 mm. Pronotum with
convergent pale spots; elytron typically with full complement of discrete black spots
(Fig. 606f), pattern varying from that to a nearly immaculate form (Fig. 606g-i).
Male genitalia as in Figure 606a-d. Female genitalia as in Figure 606e.
Discussion. This is by far the most abundant and widespread species of Hippodamia
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 606. Hippodamia convergens.
in North America. The specimens having the elytral maculation reduced or absent
are difficult to recognize without examination of the male genitalia. The normally
maculate specimens can usually be recognized without dissection because the elytral
spots are small and nearly always discrete, or if confluent, only feebly so. The other
species possessing convergent pale spots on the pronotum usually have the elytral
spots heavy and with a tendency to coalesce. The range of H. convergens is from
Ontario and British Columbia to the Antilles and Central and South America. The
unique type (holotype) of juncta is a male in the Casey collection.
Type locality. Of convergens, Mexico and California; of juncta, Healdsburg, So-
noma Co., California; of modesta, Pennsylvania; of obsoleta, not stated; of praticola,
not stated.
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NORTH AMERICAN COCCINELLIDAE
743
Fig. 607. Distribution. Hippodamia convergens.
Type depository. Of convergens, UCCC; of juncta, USNM (35505); of modesta,
type not examined; of obsoleta, type not examined; of praticola\ type not examined.
Distribution. Figure 607. Entire United States, northern most Canadian records
from southern Ontario, northern Manitoba, and southern British Columbia.
Hippodamia caseyi Johnson
Fig. 608a-g; Map, Fig. 609
Hippodamia caseyi Johnson, 1910, pp. 21, 33.— Casey, 191 1, p. 250.— Casey, 1924,
p. 155. — Chapin, 1946, p. 24. — Hatch, 1961, p. 170. — Brown and de Ruette, 1962,
p. 652.-Belicek, 1976, p. 348.
Hippodamia lecontei ab. caseyi: Leng, 1920, p. 216. — Korschefsky, 1932, p. 342.
Diagnosis. Length 4.80 to 6.70 mm, width 2.80 to 4.30 mm. Pronotum with or
without convergent pale spots, if present, then usually reduced to dots or small.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
elongate spots, black discal macula extended to, or nearly to, lateral border medially;
elytron with maculation as in H. g. lecontei (Fig. 608f, g). Male genitalia as in Figure
608a-c. Female as in Figure 608e.
Discussion. This species is very similar to H. g. lecontei in color pattern and the
2 species are easily confused. In H. caseyi the pale area of the posterolateral angle
of the pronotum is usually sharply set off from the other pale pronotal areas by the
lateral extension of the discal black spot (//. m. bowditchi also has this characteristic).
In //. lecontei there is a tendency toward this pattern, but the lateral extension of
black rarely reaches the pronotal border or even close to it. Hippodamia caseyi always
has either a subbasal band or elongate scutellar spot on the elytron. Hippodamia
lecontei may have a subbasal band in some specimens, but never an elongate scutellar
spot.
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NORTH AMERICAN COCCINELLIDAE
745
Type locality. Fairfield, Washington.
Type depository. Type not designated by author.
Distribution. Figure 609. BRITISH COLUMBIA: Abbotsford; Copper Mt.; Crows-
nest Pass; Keremeos; Pavilion; Peters Lake; Yale. CALIFORNIA: Dumont; Eldorado
Co., Mt. Tallac; Nevada Co.; Placer Co. COLORADO: Craig; Denver. IDAHO:
Emmett; Lemhi Co, Blue Nose Peak; Mica Peak; Moscow, Cedar Mt.; Soda Springs;
Wardner. MONTANA: Helena; Missoula, Squaw Peak; Ravalli Co., Deer Mt. OR-
EGON: Blue Mts;. Powder Lakes; Unity. UTAH: Alta; Little Baldy Mt.; Ogden; Park
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 610. Hippodamia oregonensis.
City; Spanish Fork; Wasatch. WASHINGTON: Pullman; Metaline Falls; Mt. Rainier.
WYOMING: Centennial; Yellowstone National Park.
Hippodamia oregonensis Crotch
Fig. 610a-h; Map, Fig. 61 1
Hippodamia oregonensis Crotch, 1873, p. 367. — Casey, 1899, p. 81. — Leng, 1903a,
p. 42. — Casey, 1899, p. 395. — Korschefsky, 1932, p. 343. — Hatch, 1961, p. 170.—
Belicek, 1976, p. 348.
Hippodamia oregonensis oregonensis: Chapin, 1946, p. 25. — Brown and de Ruette,
1962, p. 652.
Hippodamia dispar Casey, 1899, p. 79. — Leng, 1903a, p. 44. — Korschefsky, 1932,
p. 340. New Synonymy.
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NORTH AMERICAN COCCINELLIDAE
747
Fig. 611. Distribution. Hippodamia oregonensis (dot); H. sinuata sinuata (star).
Hippodamia oregonensis dispar. Chapin, 1946, p. 25.
Hippodamia puncticollis Casey, 1899, p. 78. — Casey, 1908, p. 397. — Chapin, 1946,
p. 24.
Hippodamia 5-signata puncticollis: Leng, 1903a, p. 41.
Hippodamia quinquesignata ab. puncticollis: Leng, 1 920, p. 2 1 6. — Korschefsky, 1932,
p. 344.
Hippodamia liliputana Casey, 1908, p. 397.— Chapin, 1946, p. 25.
Hippodamia quinquesignata liliputana: Korschefsky, 1932, p. 344.
Hippodamia lilliputana: Casey, 1910, p. 109 (emendation). — Chapin, 1946, p. 25.
Hippodamia cockerelli Johnson, 1910, p. 49.— Timberlake, 1919, p. 167. New Syn-
onymy.
Hippodamia oregonensis cockerelli: Chapin, 1946, p. 26.
Diagnosis. Length 4.0 to 5.0 mm, width 2.70 to 3.60 mm. Pronotum mostly black,
lateral and apical borders narrowly yellow, an occasional specimen with minute trace
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 9 3( 1 )
of convergent pale spots; elytron with black spots heavy with tendency to coalesce
(Fig. 6 1 Of-h). Male genitalia as in Figure 6 1 Oa-d. Female genitalia as in Figure 6 1 Oe.
Discussion. The small size, nearly all black pronotum without convergent pale
spots, and heavy elytral maculation usually will allow H. oregonensis to be recognized
without dissection. Chapin (1946) preserved H. dispar and H. cockerelli as subspecies
of H. oregonensis, but I regard them as junior synonyms. This is a high altitude
species and as such is likely to exhibit varying degrees of melanism. It is presently
rare in collections, but when enough specimens from critical localities become avail-
able, I judge that the variation in elytral pattern previously used to segregate sub-
species will prove to be useless. The types of H. dispar (female), H. puncticollis
(female), and H. lilliputana (male), are all uniques (holotypes) in the Casey collection.
The type (male holotype) of H. cockerelli is also unique as designated by Johnson.
Type locality. Of oregonensis, Oregon; of dispar, Colorado; of puncticollis, “Ca-
nadian Rocky Mts.”; of lilliputana, Colorado; of cockerelli, Sangre de Cristo Range,
Cottonwood Gulch, Saguache Co., Colorado.
Type depository. Of oregonensis, type not located; of dispar (35496), puncticollis
(35497), lilliputana (35498), and cockerelli (21557), USNM.
Distribution. Figure 611. ALBERTA: Banff National Park. BRITISH COLUMBIA:
Kelowna; Mara; Mt. Revelstoke Park; Mt. Todd; Vancouver; Vancouver Island.
COLORADO: Delta Co.; Gothic; Leavenworth Valley; Loveland Pass; Ouray; Rabbit
Ears Pass; Rocky Mt. Nat. Park; Silverton. OREGON: Mt. Hood. UTAH: Silverlake;
Wasatch Mts. WASHINGTON: Mt. Rainier, Paradise Park; Mt. Yakima; Olympic
Mts.
Hippodamia sinuata sinuata Mulsant
Fig. 612a-f; Map, Fig. 611
Hippodamia sinuataMnXsanX, 1850, p. 101 1.— Crotch, 1873, p. 367.— Crotch, 1874b,
p. 96. — Casey, 1899. p. 81. — Leng, 1903a, p. 42.— Casey, 1908, p. 398.— Johnson,
1910, p. 50. — Korschefsky, 1932, p. 344. — Hatch, 1961, p. 169. — Belicek, 1976,
p. 349.
Hippodamia sinuata sinuata: Timberlake, 1919, p. 165.— Chapin, 1946, p 27.
Hippodamia interrogans Mulsant, 1856, p. 139. — Leng, 1920, p. 215.
Hippodamia trivittata Casey, 1899, p. 80. — Korschefsky, 1932, p. 345.— Chapin,
1946, p. 27.
Diagnosis. Length 4.30 to 5.80 mm, width 2.40 to 3.70 mm. Pronotum always
with convergent pale spots; elytron with suture narrowly black from base to apical
%, a broad discal vitta extending from near base nearly to apex (Fig. 6121). Male
genitalia as in Figure 6 1 2a-d. Female genitalia as in Figure 6 1 2e.
Discussion. The uniformly vittate appearance and restricted distribution render
this subspecies one of the most easily recognized Hippodamia. The broad discal vitta
on the elytron shows no tendency to break up as does that vitta in specimens of H.
s. crotchi from Arizona and New Mexico. The type of H. trivittata is a unique
(holotype) male in the Casey collection. Belicek (1976) incorrectly stated that Chapin
(1946) had synonymized H. spuria LeConte and H. crotchi Casey with H. sinuata
Mulsant.
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NORTH AMERICAN COCCINELLIDAE
749
Fig. 612. f. Hippodamia sinuata sinuata. g-i. H. s. crotchi. j-m. H. s. spuria.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Type locality. Of sinuata, California; of trivittata, Sonoma Co., California.
Type depository. Of sinuata, type not examined; of trivittata, USNM (35512).
Distribution. Figure 611. CALIFORNIA: Alameda Co.; Antelope Valley; Benecia;
Birds Landing; Dos Palos; Gilroy; Guadeloupe; Lancaster; Los Angeles; Martinez;
San Jose; Santa Clara; Contra Costa Co.; Vine Hill; Willows.
Hippodamia sinuata crotchi Casey
Fig. 612g-i; Map, Fig. 613
Hippodamia crotchi Casey, 1899, p. 80. — Casey, 1908, p. 399.— Timberlake, 1919,
p. 168.-Korschefsky, 1932, p. 345.
Hippodamia sinuata crotchi: Chapin, 1946, p. 28.
Hippodamia americana fontinalis Casey, 1924, p. 156. — Chapin, 1946, p. 27.
Hippodamia sinuata disjuncta Timberlake, 1919, p. 168. — Chapin, 1946, p. 28.—
Hatch, 1961, p. 169. — Brown and de Ruette, 1962, p. 652. New Synonymy.
Hippodamia sinuata straminea Chapin, 1946, p. 29. New Synonymy.
Hippodamia sinuata Belicek, 1976, p. 349 (in part) (not sinuata Mulsant).
Description as for H. s. sinuata except elytron varies from the typical form with
broken discal vitta (Fig. 6 1 2g) to an immaculate form (Fig. 6 1 2h, i).
The typically vittate form of H. crotchi is easily recognized, but the remaining
phenotypes are readily confused with H. convergens and small examples of other
species possessing strong convergent pronotal spots. I am placing H. s. disjuncta as
a junior synonym of H. s. crotchi because it is not possible to separate the 2 forms
over a broad area encompassing most of Utah and Colorado. Both forms and their
intergrades occur within one local population. Hippodamia s. straminea is based on
teneral specimens of H. crotchi. One of the localities of the type series is Fortuna,
California; I have seen 2 specimens collected on the same date and at the same place,
obviously part of the same series, which are fully mature and marked. Also, as
mentioned by Chapin, some paratypes of H. s. straminea have indistinct markings,
therefore I consider H. s. straminea a junior synonym of H. s. crotchi. Hippodamia
crotchi and H. fontinalis are represented by unique types (male, female) in the Casey
collection.
Type locality. Of crotchi. Lake Co., California; of fontinalis, Jemez Springs, New
Mexico; of disjuncta, Murray, Utah; of straminea, Klamath River, California.
Type depository. Of crotchi (35509), fontinalis (35508), disjuncta (53932), and
straminea (57893), USNM.
Distribution. Figure 613. Northwest Territories south to New Mexico and Cali-
fornia.
Hippodamia sinuata spuria LeConte
Fig. 612j-m; Map, Fig. 613
Hippodamia spuria LeConte, 1861, p. 358. — Mulsant, 1866, p. 15.— Crotch, 1873,
p. 367.— Crotch, 1874b, p. 96.— Casey, 1899, p. 80. — Casey, 1908, p. 399.— John-
son, 1910, p. 50. — Belicek, 1976, p. 349.
Hippodamia sinuata var. spuria: Leng, 1903a, p. 42.
Hippodamia sinuata ab. spuria: Korschefsky, 1932, p. 345.
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NORTH AMERICAN COCCINELLIDAE
751
Fig. 613. Distribution. Hippodamia sinuata crotchi (shaded); H. s. spuria (cross hatch).
Hippodamia sinuata spuria: Timberlake, 1919, p. 165.— Chapin, 1946, p. 28.—
Hatch, 1961, p. 169. — Brown and de Ruette, 1962, p. 652.
Hippodamia complex CdiSty , 1899, p. 80.— Chapin, 1946, p. 27.
Hippodamia spuria var. complex: Timberlake, 1919, p. 168.
Hippodamia spuria ab. complex: Korschefsky, 1932, p. 345.
Description as for 5. sinuata except elytron with heavy black spots varying from
completely discrete to strongly coalescent (Fig. 612j-m).
This subspecies is restricted mostly to the coastal areas from Alaska to Oregon.
No other species occurring in that region has the heavily maculate pattern of H. s.
spuria. Hippodamia 1. dobzhanskyi has a similar elytral pattern, but lacks the con-
vergent pronotal spots. Hippodamia convergens has the pronotal spots, but the elytral
maculation is not heavy and specimens are usually larger. The type of complex Casey
is a unique (holotype) specimen in the Casey collection. A male of spuria in the
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 614. Anatis sp. a. Antenna, b. Prostemum, mesostemum, and metastemum. c. Tarsus,
d. Postcoxal lines.
LeConte collection labeled “(blue disc)/Type 6683(red paper)/H. spuria LeC.” is here
designated and labeled as the lectotype. Three additional specimens in the same
series, each bearing a blue disc, are designated and labeled as paralectotypes.
Type locality. Of spuria, Oregon (lectotype here designated); of complex, Victoria,
Vancouver Island, British Columbia.
Type depository. Of spuria, MCZ; of complex, USNM (35510).
Distribution. Figure 613. Alaska to Oregon.
Genus Anatis Mulsant
Anatis Mulsant, 1846, p. 1 33. — Mulsant, 1850, p. 132. — Crotch, 1873, pp. 364,
374.-Crotch, 1874b, p. 124.-Casey, 1899, p. 97.-Blatchley, 1910, p. 516.-
Korschefsky, 1932, p. 547. -McKenzie, 1936, p. 264.-Wingo, 1952, p. 23.-
Watson, 1956, p. 3.— J. Chapin, 1974, p. 68. — Belicek, 1976, p. 322. — Watson,
1976, p. 935. — lablokoff-Khnzorian, 1982, p. 303. Type-species; Coccinella ocel-
lata L., by monotypy.
Myzia LeConte, 1852, pp. 130, 132 (in part).
Coccinellini with form oval, strongly to weakly convex (Fig. 616a). Antennal club
with 9th and 10th segments obtriangular in contrast to cylindrical 8th segment (Fig.
614a). Margin of elytron variably explanate, subangulate or rounded in front of
middle; apex of elytral suture with distinct patch of hairs on each side; epipleuron
nearly flat, slightly descending externally. Prosternum strongly convex medially, pro-
tuberant at apex (Fig. 6 1 4b), lateral border thickly margined between coxae. Apical
margin of mesostemum deeply, broadly emarginate for reception of prosternal pro-
cess (Fig. 614b). Apex of middle and hind tibia each with 2 spurs. Tarsal claw with
large, subquadrate basal tooth (Fig. 614c). Postcoxal line on first abdominal sternum
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NORTH AMERICAN COCCINELLIDAE
753
incomplete, almost reaching posterior margin (Fig. 614d). Male genitalia symmet-
rical. Female genitalia with well developed infundibulum.
This genus is not closely similar to any other North American genus of Coccinel-
lidae, and is easily recognized by the key characters. Anatis is primarily holarctic
with the geographic range extending to northern Mexico in North America. In ad-
dition to the 4 nearctic species, 2 species occur in the Palearctic Region. Members
oi Anatis are predators on aphids occurring mostly on coniferous and deciduous trees.
Specific host records are; Acyrthosiphum pisum (Harris), Coristoneura pinus Freeman,
Macrosiphum avenue (F.), Myzus cerasi (F.), Nearctaphis crataegifoliae (Fitch), Per-
iphyllus aceris (L.), Phorodon humuli (Schrank), Pinus strobi (Hartig), and Schizo-
lachnus piniradiatae (Davidson). The genus has been reviewed taxonomically by
Watson (1976), and the key and synonymies presented here have been modified from
the latter.
Key to species of Anatis
1. Elytron with lateral border broadly explanate and distinctly angulate in front of
middle (Fig. 612a) 2
Elytron with lateral border weakly explanate, not angulate (Fig. 624a) 3
2(1). Elytron without dark spots; pronotum with lateral border black (Fig. 612a)
lecontei Casey
Elytron with dark spots; pronotum with pale lateral border (Fig. 6 1 9a)
rathvoni (LeConte)
3(1). Dark spots on elytron ringed with white or yellow (Fig. 624a) mali (Say)
- Dark spots on elytron never ringed (Fig. 6 1 9a) labiculata (Say)
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 615. Anatis labiculata (male genitalia).
Anatis labiculata (Say)
Fig. 615a-d, 616a-c; Map, Fig. 617
Coccinella labiculata Say, 1824, p. 288 (in part, var. B).
Coccinella mali Say, 1824, p. 93 (in part, var. b).
Coccinella quindecimpunctata Olivier, 1808, p. 1027 (not quindecirnpunctata DeGeer,
1775).
Anatis quindecimpunctata: Mulsant, 1850, p. 133.— Crotch, 1874b, p. 124 (in part);
Casey, 1899, p. 98. — Leng, 1903b, p. 207. — Korschefsky, 1932, p. 356. — Mckenzie,
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NORTH AMERICAN COCCINELLIDAE
755
Fig. 616. Anatis labiculata (habitus and female genitalia).
1936, p. 268.-Wingo, 1952, pp. 24, 36.-Mader, 1954, p. 125. -J. Chapin, 1974,
p. 68.
Mysia quindecimpunctata: Melsheimer, 1853, p. 130 (in part).
Myzia quindecimpunctata: LeConte, 1859c, p. 192 (in part).
Halyzia quindecimpunctata: Gemminger and Harold, 1876, p. 3760 (in part).
Anatis canadensis Provancher, 1877, p. 696. — Horn, 1880, p. xii.
Anatis caseyi Westcott, 1912, p. 422. — Korschefsky, 1932, p. 556.
Anatis labiculata: Watson, 1976, p. 941.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 617. Distribution. Anatis labiculata (shaded); A. rathvoni (cross hatch).
Diagnosis. Length 7.20 to 9.50 mm, width 5.50 to 8.0 mm. Form rounded, lateral
border of elytron slightly explanate. Color yellow to brownish red with black markings
as in Figure 616a, b. Male genitalia as in Figure 6 1 5a-d. Female genitalia as in Figure
616c.
Type locality. Simcoe, Ontario (neotype designated by Watson, 1976).
Type depository. CNC.
Distribution. Figure 617. Ontario to South Carolina, west to North Dakota, Col-
orado and Texas.
Anatis rathvoni (LeConte)
Fig. 618a-d, 619a, b; Map, Fig. 617
Myzia rathvoni LeConte, 1852, p. 132.
Anatis rathvoni: Crotch, 1873, p. 374.— Crotch, 1874b, p. 124. — Wickham, 1894, p.
306.— Casey, 1899, p. 98. — Leng, 1903b, p. 208. — Korschefsky, 1932, p. 557.—
McKenzie, 1936, p. 266. — Hatch, 1961, p. 182. — Belicek, 1976, p. 323. — Watson,
1976, p. 942.
Halyzia rathvoni: Gemminger and Harold, 1876, p. 3760.
Diagnosis. Length 7.50 to 10.20 mm, width 6.50 to 9.0 mm. Form elongate, lateral
border of elytron strongly explanate. Color yellow to brownish red with black mark-
ings as in Figure 619a. Male genitalia as in Figure 618a-d. Female genitalia as in
Figure 619b.
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NORTH AMERICAN COCCINELLIDAE
757
Discussion. The type is a unique female (holotype) labeled “(gold disc)/ 1633/Type
6690 (red paper)/Myzia rathvoni Lee.”
Type locality. Sacramento, California.
Type depository. MCZ.
Distribution. Figure 617. Southern Alberta to British Columbia, south to northern
California.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 619. Anatis rathvoni (habitus and female genitalia).
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NORTH AMERICAN COCCINELLIDAE
759
Anatis lecontei Casey
Fig. 620a-d, 621a-c; Map, Fig. 622
Anatis lecontei Casey, 1899, p. 98.— Casey, 1908, p. 406. — Korschefsky, 1932, p.
547.-Hatch, 1961, p. 183.-Belicek, 1976, p. 323. -Watson, 1976, p. 943.
Anatis rathvoni lecontei: Leng, 1903b, p. 208. — McKenzie, 1936, p. 268.— Wingo,
1952, p. 46.
Diagnosis. Length 7.75 to 10.50 mm, width 6.50 to 9.00 mm. Form elongate,
lateral border of elytron strongly explanate. Color yellow to brownish red, pronotum
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 621. Anatis lecontei (habitus, pronotum, and spermathecal capsule).
often with 2 basal spots (Fig. 621b), elytron without black marking (Fig. 621a). Male
genitalia as in Figure 620a-d. Female genitalia as in Figure 621c.
Discussion. There are 3 types of A. lecontei in the Casey collection, the first of
these, a female, is here designated and labeled as the lectotype, the other 2, from
Jemez Springs, New Mexico, are designated as paralectotypes.
Type locality. Fort Wingate, New Mexico (lectotype here designated).
Type depository. USNM (35539).
Distribution. Figure 622. Southern Alberta to New Mexico, west to British Colum-
bia and California.
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NORTH AMERICAN COCCINELLIDAE
761
762
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Anatis mali (Say)
Fig. 623a-'-'
f
^ it "
'1‘^rdi
“ ' /Vt\
mi
^. , ■ ■
'iv'0'm
A
r
■ t Wh
la
Fig. 624. Anatis mali (habitus, female genitalia).
Anatis borealis Belicek, 1976, p. 323. New Synonymy.
Anatis mali: Casey, 1899, p. 98. — Watson, 1976, p. 938.
Diagnosis. Length 7.30 to 10.0 mm, width 5.50 to 7.60 mm. Form rounded, lateral
border of elytron slightly explanate. Color yellow to brownish red with black markings
as in Figure 624a. Male genitalia as in Figure 623a-d. Female genitalia as in Figure
624b.
Discussion. The name borealis was proposed because Belicek did not realize that
both A. mali and A. labiculata were available as pointed out by Watson (1976, p.
935). Therefore A. borealis is an unnecessary name and must be placed in synonymy.
Type locality. Of mali. Little Rapids, Ontario (neotype designated by Watson,
1976); of borealis, Edmonton, Alberta.
Type depository. Of mali, CNC; of borealis, CNC.
Distribution. Figure 625. Ontario to British Columbia, south to Virginia and Or-
egon.
Genus Myzia Mulsant
Myzia Mulsant, 1846, p. 277. — LeConte, 1852, p. 130. Type-species; Coccinella
oblongoguttata L., by monotypy.
Mysia Mulsant, 1846, p. 129.— Mulsant, 1850, p. 137.— Crotch, 1873, p. 364.—
Crotch, 1874b, p. 125. -Gorham, 1892, p. 162.-Wickham, 1894, p. 299 (not
Mysia Lamarck, 1818).
Neomysia Casey, 1899, p. 98.— Casey, 1905, p. 161.— Casey, 1908, p. 407— Leng,
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 625. Distribution. Anatis mali.
1903b, p. 194. — Korschefsky, 1932, p. 542.— Timberlake, 1943, p. 21. — Wingo,
1952, p. 23.— J. Chapin, 1974, p. 68. — Belicek, 1976, p. 324. Type-species; Coc-
cinella oblongoguttata L. by subsequent designation of Korschefsky, 1932.
Paramysia Reitter, 1911, pp. 136, 144.— Timberlake, 1943, p. 21 (unnecessary re-
placement name for Mysia Mulsant).
Myzia LeConte: Belicek, 1976, p. 324 (error, name cannot be attributed to LeConte).
Sospita {Myzia): lablokoff-Khnzorian, 1982, p. 158.
Coccinellini with length 6.0 to 10.0 mm; form oval, strongly convex. Dorsal color
pattern with pale background, elytron immaculate or with dark vittae. Anterolateral
angle of clypeus produced forward. Lateral margin of elytron variably explanate,
rounded or subangulate in front of middle; epipleuron obliquely descending exter-
nally. Intercoxal process of prosternum with strong lateral ridge, median area de-
pressed. Apical margin of mesosternum broadly, feebly emarginate for reception of
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NORTH AMERICAN COCCINELLIDAE
765
prostemal process. Apex of middle and hind tibia each with 2 spurs. Tarsal claw
cleft (Fig. 626a). Postcoxal line incomplete, of Diomus type (Fig. 626b). Male genitalia
symmetrical. Female genitalia lacking infundibulum; coxal plate short, ovoid, with
strong apical stylus (Fig. 628d).
Myzia is easily recognized by the key characters; in addition, the pale dorsal color,
usually somewhat vittate elytra, and smooth, polished dorsal surface are characteristic
of members of the genus. Myzia is primarily holarctic with the geographic range
extending to northern Mexico in North America. In addition to the species found in
America north of Mexico, 2 occur in the Old World and one in Mexico. Members
of Myzia are predators on aphids associated with a wide variety of woody plants.
Specihc host records are Lachnus pinicola and Chermes sp. The genus was reviewed
by Belicek (1976), who reinstated the generic name Myzia, placing Neomysia as a
junior synonym of Myzia, a decision with which I agree. Belicek designated Coccinella
pullata Say as the type-species of Neomysia’, this action is invalid because Korschefsky
(1932) previously designated Coccinella oblongoguttata L. as the type-species. Tim-
berlake (1943) regarded the North American forms of Myzia as only subspecies of
the European Myzia {Neomysia) oblongoguttata L. The only North American species
with genitalia similar to those of M. oblongoguttata is M. pullata, and even in this
case there are distinct genitalic differences. Therefore I consider the North American
species congeneric with, but specifically distinct from, M. oblongoguttata.
Key to species of Myzia
1. Lateral white border of pronotum with dark spot (Fig. 628a); central and eastern
U.S. and Canada pullata (Say)
- Lateral border of pronotum white or not, never with dark spot; western North
America 2
2(1). Elytron with lateral border broadly explanate, tapered to acute apex; head with dark
area (at least on vertex); pronotum usually with median third black or brown (Fig.
632c) subvittata (Mulsant)
Elytron with lateral border weakly explanate, rounded at apex; head immaculate;
pronotum immaculate, or with pale brown spots (Fig. 6 3 Of) interrupt a (Casey)
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 627. Myzia pullata (male genitalia).
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NORTH AMERICAN COCCINELLIDAE
767
Fig. 628. Myzia pullata (habitus and variation, female genitalia).
Myzia pullata (Say)
Fig. 627a-d, 628a-d; Map, Fig. 629
Coccinella pullata Say, 1826, p. 301.
Mysia pullata: Mulsant, 1850, p. 1023. — Crotch, 1873, p. 375.— Crotch, 1874b, p.
125.-Wickham, 1894, p. 303.
Neomysia pullata: Casey, 1899, p. 99.— Leng, 1903b, p. 209. — Blatchley, 1910, p.
516.— Korschefsky, 1932, p. 546.— Wingo, 1952, p. 47.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Coccinella notans Randall, 1838b, p. 49.
Mysia notans: Mulsant, 1850, p. 1023.
Neomysia randalli Casey, 1899, p. 99. — Leng, 1920, p. 21 7. — Korschefsky, 1932, p.
546.-Wingo, 1952, p. 47.-Belicek, 1976, p. 325.
Neomysia montana Casey, 1899, p. 100. — Leng, 1920, p. 217. — Korschefsky, 1932,
p. 543.-Belicek, 1976, p. 325.
Neomysia pullata var. randalli: Leng, 1903b, p. 209.
Neomysia oblongoguttata pullata: Timberlake, 1943, p. 23.— J. Chapin, 1974, p. 69.
Myzia pullata: Belicek, 1976, p. 325.
Diagnosis. Length 6.50 to 8.0 mm, width 5.20 to 6.0 mm. Form oval, lateral border
of elytron slightly explanate. Dorsal color typically pale brownish yellow; pronotum
dark brown medially, broad lateral border white with a median dark brown spot
often connected to median area (Fig. 628a); northern specimens often with dark
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NORTH AMERICAN COCCINELLIDAE
769
elytral maculae (Fig. 628b, c). Male genitalia as in Figure 627a-d. Female genitalia
as in Figure 628d.
Discussion. Belicek (1976) correctly placed randalli and montana as junior syn-
onyms of pullata. Both species are represented by a single type specimen each in the
Casey collection which must be considered holotypes.
Type locality. Of pullata, “eastern coast of Virginia, and Florida”; of notans, Maine;
of randalli. Lake Superior; of montana, Colorado.
Type depository. Of pullata, type no longer in existence; of notans, type not located;
of randalli, USNM (35541); of montana, USNM (35542).
Distribution. Figure 629. Labrador to South Carolina, west to Alberta and Colorado.
Myzia interrupt a (Casey)
Fig. 630a-g; Map, Fig. 629
Neomysia interrupta Casey, 1899, p. 99. — Korschefsky, 1932, p. 543.
Neomysia hornii var. interrupta: Leng, 1903b, p. 209.
Neomysia horni: Casey, 1899, p. 99. — Leng, 1903, p. 209. — Korschefsky, 1932, p.
543.— Wingo, 1952, p. 47. — Hatch, 1961, p. 184 (not hornii Crotch).
Neomysia oblongoguttata caseyi Timberlake, 1943, p. 21. New Synonymy.
Neomysia oblongoguttata interrupta: Timberlake, 1943, p. 22.
Myzia horni: Belicek, 1976, p. 326 (not hornii Crotch).
Diagnosis. Length 6.50 to 8.0 mm, width 5.0 to 6.0 mm. Form oval, lateral border
of elytron slightly explanate. Dorsal color pale yellowish brown; elytron usually with
light brown vittae and pronotum with 3 light brown, basal maculae (Fig. 630f),
pronotum often immaculate and elytral vittae reduced (Fig. 630g). Male genitalia as
in Figure 630a-d. Female genitalia as in Figure 630e.
Discussion. Myzia interrupta is a valid species previously considered to be a syn-
onym of M. horni Crotch (see remarks under M. subvittata). Myzia interrupta is
represented in the Casey collection by a single type specimen which must be con-
sidered the holotype.
Type locality. Of interrupta. Fort Wingate, New Mexico; of caseyi, Eldorado Co.,
California.
Type depository. Of interrupta, USNM (35540); of caseyi, Koebele collection, Ha-
waii.
Distribution. Figure 629. Alberta to west Texas, west to British Columbia and
California.
Myzia subvittata (Mulsant)
Fig. 631a-d, 632a-e; Map, Fig. 633
Mysia subvittata MulsdoW, 1850, p. 138.— Crotch, 1874b, p. 125. — Wickham, 1894,
p. 303.
Anatis subvittata: Crotch, 1873, p. 375.
Mysia hornii Crotch, 1873, p. 375. New Synonymy.
Neomysia subvittata: Leng, 1903, p. 209.— Korschefsky, 1932, p. 547. — Hatch, 1961,
p. 184.
Neomysia oblongoguttata subvittata: Timberlake, 1943, p. 22.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 630. Myzia interrupta.
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NORTH AMERICAN COCCINELLIDAE
771
Fig. 631. Myzia subvittata (male genitalia).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 632. Myzia subvittata (habitus and variation, female genitalia).
Myzia subvittata: Belicek, 1976, p. 235.
Neomysia oregona Casey, 1924, p. 160. — Hatch, 1961, p. 184.
Diagnosis. Length 5.70 to 8.0 mm, width 4.50 to 6.50 mm. Form somewhat
triangular, tapered to acute apex in apical third, lateral border of elytron broadly
explanate in front of middle. Dorsal color yellowish brown; elytron with dark brown
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NORTH AMERICAN COCCINELLIDAE
773
vittae, pronotum dark brown with broad, yellowish white lateral border (Fig. 632c),
pronotum often nearly immaculate and elytral vittae strongly reduced (Fig. 632a, b).
Male genitalia as in Figure 631a-d. Female genitalia as in Figure 632e.
Discussion. The broadly explanate border of the elytron and posteriorly tapered
form characterize M. subvittata. It is apparent from the original description that M.
horni Crotch is conspecific with M. subvittata Mulsant. Casey (1899) did not detect
this and associated the name M. horni with the form of Myzia having 3 pale pronotal
maculae. He was followed in this by subsequent authors including Belicek (1976).
Casey (1899) described M. interrupta which is the same species as his '"hornr and
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
later (1924) described M. oregona which is a synonym of M. subvittata. The type of
M. subvittata in the Crotch collection is a female holotype labeled “Type (blue paper)/
Type subvittata Reiche.” The type of M. horni is supposed to be in the MCZ col-
lection, but no such specimen is currently there. The 2 specimens under that name
have the wrong locality data to qualify as types. Myzia oregona is represented in the
Casey collection by a single type specimen which must be considered the holotype.
Type locality. Of subvittata, “les parties occidentals de I’Amerique du Nord”
(lectotype here designated); of horni, Oregon; of oregona. Bull Run, Clackamas Co.
(Big Sandy Camp), Oregon.
Type depository. Of subvittata, UCCC; of horni, not located; of oregona, USNM
(35543).
Distribution. Figure 633. British Columbia and Idaho south to southern California.
Genus Calvia Mulsant
Calvia Mulsant, 1850, p. 143. -Crotch, 1874b, p. 143.-Chapuis, 1876, p. 183.-
Mader, 1926, p. 20.-Hatch, 1961, p. I81.-Belicek, 1976, p. 326.-Iablokoff-
Khnzorian, 1982, p. 176. Type-species; Coccinella decemguttata L., by subsequent
designation of Crotch, 1874b.
Anisocalvia Crotch, 1871, p. 329.— Casey, 1899, p. 96. — Leng, 1903b, p. 206.—
Wingo, 1952, p. 24. Type-species; Coccinella quatuordecimguttata (L.), by sub-
sequent designation of Crotch, 1874b.
Calvia {Anisocalvia): Korschefsky, 1932, p. 521. — lablokoff-Khnzorian, 1982, p. 184.
£’oc<2n<3Timberlake, 1943, p. 37. — lablokoff-Khnzorian, 1982, p. 1 76. Type- species;
Eocaria muiri Timberlake, by original designation.
Coccinellini with length 4.0 to 6.0 mm, form oval, not strongly convex. Dorsal
surface polished between punctures, alutaceous sculpture lacking. Dorsal color pattern
extremely variable. Anterolateral angle of clypeus produced forward. Lateral margin
of elytron feebly explanate; epipleuron weakly descending externally. Intercoxal pro-
cess of prostemum with broad lateral ridge extending nearly to apex of prostemum,
median area convex (Fig. 634a). Apical margin of mesostemum triangularly emar-
ginate (Fig. 634a). Apex of middle and hind tibia each with 2 spurs. Tarsal claw with
subquadrate basal tooth. Postcoxal line incomplete, of Diomus type (Fig. 634b). Male
genitalia symmetrical. Female genitalia lacking infundibulum; coxal plate short, sub-
triangular, with strong apical stylus (Fig. 634g).
Calvia is the only genus of North American Coccinellini without at least some
trace of alutaceous sculpture on the pronotum, and this is the only good recognition
character for the genus. Calvia is primarily an Old World genus with approximately
20 species names, mainly in the Palearctic region. Calvia quatuordecimguttata is a
holarctic species and the only North American representative of the genus. Members
of Calvia have been reported as predaceous on aphids, scale insects and other soft
bodied insects, but specihc host data has been lacking until recently. Semjanov (1980)
reported C quatuordecimguttata as a predator of Psyllidae and of aphids, but pref-
erentially a psyllid predator. Specific host data is as follows: Aphididae: Aphis pomi
Degeer; Eucallipterus tiliae (L.); Euceraphis punctipennis (Zetterstedt); Hyalopterus
pruni Geoffroy; Pterocallis alni (Degeer); Rhopalosiphum insertum (Walker); Rho-
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NORTH AMERICAN COCCINELLIDAE
775
palosiphum padi (L.); Psyllidae: Psylla alni L.; P. mali Schmdb. McMullen and Jong
(1967) reported this species as a predator of Psylla pyricola Forster in British Co-
lumbia.
Calvia quatuordecimguttata (L.)
Fig. 634a-g, 635; Map, Fig. 636
Coccinella 14-guttata L., 1758, p. 367.
Coccinella 12-maculata Gebler, 1832, p. 76.
Coccinella incarnata Kirby, 1837, p. 231.
Coccinella cardisce Randall, 1838a, p. 32.
Coccinella obliqua Randall, 1838a, p. 33.
Coccinella similis Randall, 1838b, p. 50.
Harmonia duodecimmaculata: Mulsant, 1850, p. 86.— Wickham, 1894, p. 302.
Harmonia incarnata: Mulsant, 1850, p. 86.
Calvia quatuordecimguttata: Mulsant, 1850, p. 144. — Korschefsky, 1932, p. 524.—
Belicek, 1976, p. 327.
Anisocalvia 14-guttata: Crotch, 1873, p. 373.— Timberlake, 1943, p. 20.
Anisocalvia 14-guttata var. similis: Crotch, 1873, p. 373.
Anisocalvia 14-guttata var. cardisce: Crotch, 1873, p. 373.
Anisocalvia 14-guttata var. hesperica Crotch, 1873, p. 374. — Leng, 1920, p. 217.
Anisocalvia 12-maculata: Crotch, 1873, p. 374.
Coccinella duodecimmaculata: Crotch, 1874b, p. 110.— Weise, 1885a, p. 38.
Anisocalvia quatuordecimguttata: Crotch, 1874b, p. 144. — Wingo, 1952, p. 46.
Anisocalvia cardisce: Casey, 1899, p. 96.
Anisocalvia victoriana Casey, 1899, p. 96. — Belicek, 1976, p. 327.
Anisocalvia 12-maculata: Casey, 1899, p. 97. — Leng, 1903, p. 207.— Wingo,
1952, p. 40.
Anisocalvia duodecim-maculata: Leng, 1920, p. 217.— Wingo, 1952, p. 40.
Anizocalvia duodecim-maculata a. elliptica Leng, 1920, p. 217.
Anisocalvia elliptica Casey, 1899, p. 97. — Belicek, 1976, p. 327.
Anisocalvia quatrodecimguttata: Leng, 1903b, p. 206.
Anisocalvia quatrodecimguttata var. cardisce: Leng, 1903b, p. 206.
Anisocalvia quatrodecimguttata var. similis: Leng, 1903b, p. 206.
Anisocalvia quatrodecimguttata var. victoriana: Leng, 1903b, p. 206.
Anisocalvia incarnata: Leng, 1903b, p. 207.
Agrabia sicardi'lA\xnQnm?ic\\Qr, 1912, p. 448.— Gordon, 1974, p. 170.
Agrabia sicardi var. complexa Nunenmacher, 1912, p. 448. — Gordon, 1974, . 170.
Anisocalvia 12-maculata var. elliptica: Leng, 1903b, p. 207.
Anisocalvia lacustris Casey, 1924, p. 158. — Belicek, 1976, p. 327.
Anisocalvia bicordifera Casey, 1924, p. 1 59. — Belicek, 1976, p. 327.
Anisocalvia vancouveri Casey, 1924, p. 159. — Belicek, 1976, p. 327.
Anisocalvia quadrisignata Casey, 1924, p. 159. — Belicek, 1976, p. 327.
Anisocalvia postplagiata Casey, 1924, p. 159. — Belicek, 1976, p. 327.
Anisocalvia uniformis Casey, 1924, p. 160. — Belicek, 1976, p. 327.
Calvia 12-maculata: Mader, 1931, p. 193.— Korschefsky, 1932, p. 523. — Hatch,
1961, p. 182.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Calvia (Anisocalvia) duodecimmaculata ab. elliptical Korschefsky, 1932, p. 523.
Calvia quatuordecimguttata ab. bicordiferai Korschefsky, 1932, p. 526.
Calvia quatuordecimguttata ab. cardisce: Korschefsky, 1932, p. 526.
Calvia quatuordecimguttata ab. lacustris: Korschefsky, 1932, p. 526.
Calvia quatuordecimguttata ab. obliqua: Korschefsky, 1932, p. 526.
Calvia quatuordecimguttata ab. postplagiatai Korschefsky, 1932, p. 527.
Calvia quatuordecimguttata ab. quadrisignata: Korschefsky, 1932, p. 527.
Calvia quatuordecimguttata ab. similis: Korschefsky, 1932, p. 527.
Calvia quatuordecimguttata ab. uniformisi Korschefsky, 1932, p. 527.
Calvia quatuordecimguttata ab. vancouveri: Korschefsky, 1932, p. 527.
Calvia quatuordecimguttata victoriana: Korschefsky, 1932, p. 527.
Calvia 14-guttata: Hatch, 1961, p. 181.
Diagnosis. Length 4.0 to 5.5 mm, width 3.2 to 4.5 mm. Dorsal color pattern
extremely variable, background color either pale or dark (Fig. 635a-k). Male genitalia
as in Figure 634c-f Female genitalia as in Figure 634g.
Discussion. The many color morphs and wide geographic range of this species have
caused a number of names to be proposed for what is a single polymorphic species.
Most of these names were recently synonymized by Belicek (1976). The synonymy
listed here includes only names involving North American color variants; for com-
plete synonymy, including Old World variants, see Korschefsky (1932). The Casey
names victoriana, elliptica, bicordifera, vancouveri and uniformis were each based on
a single specimen which must be considered the holotype. There are 6 types of lacustris
in the Casey collection, the first of which is here designated and labeled as the lectotype
with the other 5 as paralectotypes. There are 5 types of quadrisignata, the first of
which is here designated and labeled as the lectotype with the other 4 as paralecto-
types. There are 3 types of postplagiata, the first of which is here designated and
labeled as the lectotype, the other 2 as paralectotypes.
Type locality. Of quatuordecimguttata, “Europe”; of duodecimmaculata, Siberia;
of incarnata, “60 N.”; of cardisce, Maine; of obliqua, Maine; of similis, Massachusetts;
of victoriana, British Columbia; of elliptica, Hudson Bay; of sicardi and complexa,
Hornbrook, Siskiyou Co., California; of lacustris, Marquette, Michigan (lectotype
here designated); of bicordifera. Lake George, New York; of vancouveri, British Co-
lumbia; of quadrisignata, Marquette, Lake Superior (lectotype here designated); of
postplagiata, Marquette, Lake Superior (lectotype here designated); of uniformis,
Adirondack Mts., New York.
Type depository. Of quatuordecimguttata, Linnean Society, London; of duodecim-
maculata, not located; of incarnata, not located; of cardisce, obliqua, and similis,
types apparently lost; of elliptica, victoriana, lacustris, bicordifera, vancouveri, quad-
risignata, postplagiata, and uniformis, USNM; oi sicardi, and complexa, CAS.
Distribution. Figure 636. Labrador to New Jersey, west to Alaska and northern
California.
Genus Adalia Mulsant
Adalia Mulsant, 1846, errata addenda p. 2.— Crotch, 1873, p. 372.— Crotch, 1874b,
p. 99. -Gorham, 1891, p. 1 54.- Wickham, 1894, p. 299. -Casey, 1899, p. 82.-
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NORTH AMERICAN COCCINELLIDAE
111
Fig. 634. Calvia quatuordecimguttata. a. Prosternum, mesostemum, and metastemum. b.
Postcoxal lines, c-g. Genitalia.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 635. Calvia quatuordecimguttata (habitus and variation).
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NORTH AMERICAN COCCINELLIDAE
779
Fig. 636. Distribution. Calvia quatuordecimguttata.
Leng, 1903b, p. 194.-Korschefsky, 1932, p. 384.-Wingo, 1952, p. 24.-J. Chap-
in, 1974, p. 65. — Belicek, 1976, p. 328.— lablokoff-Khnzorian, 1982, p. 429. Type-
species; Coccinella bipunctata L., by subsequent designation of Crotch, 1874b.
Idalia Mulsant, 1846, p. 44.— Mulsant, 1850, p. 49 (not Idalia Hubner, 1819).
ArrowellaBxQXYiQS, 1925, p. 147.— Korschefsky, 1932, p. 385. Type-species;
ported Brethes, by monotypy.
Coccinellini with length 3.50 to 5.50 mm; form oval, weakly convex. Polymorphic
in dorsal color pattern. Anterolateral angle of clypeus produced forward. Pronotum
and elytron with lateral margin feebly explanate, usually semitransparent; epipleuron
obliquely inclined. Intercoxal process of prostemum smooth, slightly convex, with
weak lateral ridge extending anteriorly as far as anterior margin of coxa. Apical margin
of mesostemum truncate, ridged. Apex of middle and hind tibia each with 2 spurs.
Tarsal claw with basal, subquadrate tooth. Postcoxal line complete or nearly so, of
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Pullus type (Fig. 637a). Male genitalia symmetrical. Female genitalia with large
infundibulum (Fig. 637f).
Adalia is characterized by the truncate apex of the mesosternum and complete
postcoxal line; in addition, the pronotum always has a pale M-shaped mark at the
middle of the base except in the melanic forms. Members of Adalia are distributed
worldwide with approximately 35 species in the genus, only one of which, A. bi-
punctata, occurs in North America. Adalia bipunctata is extremely widespread, being
very common in Europe and also occurring in South America. Members of Adalia
are aphid and adelgid predators with specific host records as follows: Adelgidae:
Adelges lands (Vallot), Pineus pini (Macquart), Pineus strobi (Hartig). Aphidae: Acyr-
thosiphon carnosum (Buckton), Acyrthosiphon dirhodum (Walker), Acyrthosiphon
pisum (Harris), Acyrthosiphon urticae (Schrank), Acyrthosiphon solani (Kaltenbach),
Amphorophora rubi (Kaltenbach), Anoraphis farfarae (Koch), Aphis fabae Scopoli,
Aphis farinosa (Gmelin), Aphis gossypii Glover, Aphis hederae (Kaltenbach), Aphis
pomi Degeer, Aphis rumicis (L.), Aphis sambuci (L.), Aphis sp.. Aphis urticata (Gme-
lin), Aphis viburni (Scopoli), Betulaphis quadrituberculata (Kaltenbach), Brachycau-
dus helichrysi (Kaltenbach), Brevicoryne brassicae (L.), Capitophorus elongatus
Knowlton, Cavariella aegopodi (Scopoli), Cavariella sp., Chaitophorus capreae {Mos-
ley), Chaitophorus veriscolor (Koch), Chromaphis juglandica (Kaltenbach), Chro-
maphis juglandicola (Kaltenbach), Cryptomyzus ribis (L.), Dactynotus cirsii (L.), Dac-
tynotus eoessigi (Knowlton), Dactynotus sonchi (L.), Drepanosiphum platanoidis
(Schrank), Dysaphis devecta (Walker), Dysaphis sorbi (Kaltenbach), Elatobium abie-
tinum (Walker), Eriosoma lanigerum (Hausmann), Eriosoma pyricola (Baker and
Davidson), Euceraphis punctipennis (Zetterstedt), Euceraphis tiliae (L.), Hyalopterus
pruni (Geoffroy), Laingia psammae (Theobald), Macrosiphum euphorbiae (Thomas),
Macrosiphum rosae (L.), Monellia caryella (Fitch), Monelliopsis California (Essig),
Monelliopsis caryae (Monell), Myzocallis boerneri (Stroyan), Myzocallis carpini (Koch),
Myzocallis castanicola (Baker), Myzocallis coryli (Goeze), Myzus cerasi (F.), Myzus
persicae (Sulzer), Nearctaphis bakeri (Cowen), Nearctaphis crataegifolia (Fitch), Pem-
phigus bursarius (L.), Periphyllus lyropictus (Kessler), Periphyllus negundinis (Thom-
as), Periphyllus testundinaceus (Femie), Phorodon humuli (Schrank), Phorodon men-
thae (Buckton), Phyllaphis fagi (L.), Pterocallis alni (Degeer), Rhopalosiphum insertum
(Walker), Thelaxes dryophila (Schrank), Tuberculoides annulatus (Hartig), Tubero-
lachnus salignus (Gmelin).
Adalia was taxonomically treated by lablokoff-Khnzorian (1982).
Adalia bipunctata (L.)
Fig. 637a-l; Map, Fig. 638
Coccinella 2-punctata L., 1758, p. 364.
Coccinella frigida Schneider, 1792, p. 172.
Coccinella bioculata Say, 1824, p. 94.
Coccinella humeralis Say, 1824, p. 95. — Mulsant, 1850, p. 1049.
Coccinella disjuncta Randall, 1838a, p. 33. — Mulsant, 1850, p. 1049.
Idalia bipunctata: Mulsant, 1846, p. 51.
Adalia opthalmica Mulsant, 1850, p. 56. — Crotch, 1874b, p. 101.— Casey, 1899, p.
86.
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NORTH AMERICAN COCCINELLIDAE
781
Fig. 637. Adalia bipunctata.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Adalia bipunctata: Mulsant, 1850, p. 58.— Crotch, 1873, p. 372.— Crotch, 1874b, p.
102. -Wickham, 1894, p. 302.-Casey, 1899, p. 85.-Leng, 1903b, p. 195.-
Blatchiey, 1910, p. 191. — Palmer, 1911, p. 289. — Palmer, 1917, p. 289.— Kor-
schefsky, 1932, p. 385.— Wingo, 1952, p. 46.— J. Chapin, 1974, p. 65. — Belicek,
1976, p. 328. — lablokoff-Khnzorian, 1982, p. 438.
Coccinella melanopleura LeConte, 1859b, p. 286.
Adalia ludovicae Mulsant, 1866, p. 36. — Leng, 1920, p. 217.
Coccinella annectans Crotch, 1873, p. 371.
Adaliafrigida: Crotch, 1873, p. 372.— Crotch, 1874b, p. 100. — Leng, 1903b, p. 195.—
Korschefsky, 1932, p. 430. — Timberlake, 1943, p. 15. — Wingo, 1952, p. 40.
Adalia bipunctata var. humeralis: Crotch, 1873, p. 373.
Adalia bioculata: Crotch, 1874b, p. 102.
Adalia melanopleura: Casey, 1899, p. 85. — Palmer, 1914, p. 283.
Adalia humeralis: Casey, 1899, p. 85. — Leng, 1903b, p. 196. — Palmer, 1914, p. 285.
Adalia ovipennis Casey, 1899, p. 86. — Lusis, 1947, p. 826.
Adalia transversalis Casey, 1899, p. 86. — Lusis, 1947, p. 826.
Adalia ornatella Casey, 1899, p. 86.
Adalia annectans: Casey, 1899, p. 86. — Leng, 1903b, p. 195. — Palmer, 1911, p.
299.-Palmer, 1914, p. 283.
Adalia frigida var. melanopleura: Leng, 1903b, p. 195. — Korschefsky, 1932, p. 432.
Adalia frigida var. ophthalmica: Leng, 1903b, p. 195. — Korschefsky, 1932, p. 432.
Adalia frigida var. disjuncta: Leng, 1903b, p. 195. — Korschefsky, 1932, p. 432.
Adalia frigida var. ornatella: Leng, 1903b, p. 195. — Korschefsky, 1932, p. 432.
Adalia frigida ab. postica: Korschefsky, 1932, p. 432.
Adalia annectans var. ovipennis: Leng, 1903b, p. 196.— Johnson, 1910, p. 71.—
Korschefsky, 1932, p. 437.
Adalia annectans var. transversalis: Leng, 1903b, p. 196.— Johnson, 1910, p. 71.—
Korschefsky, 1932, p. 437.
Adalia coloradensis Casey, 1908, p. 401. — Palmer, 1914, p. 285.
Adalia annectans var. postica Johnson, 1910, p. 68.
Adalia bipunctata var. sexpustulata Johnson, 1910, p. 71.
Adalia bipunctata var. ocellata Johnson, 1910, p. 71.
Adalia bipunctata var. humeralis Johnson, 1910, p. 71 (not humeralis Say, 1824).
Adalia annectans ab. duplicata Leng, 1920, p. 217. — Korschefsky, 1932, p. 437 (new
name for humeralis Johnson).
Adalia bipunctata ab. bioculata: Korschefsky, 1932, p. 389.
Diagnosis. Length 3.50 to 5.20 mm, width 2.80 to 4.0 mm. Dorsal color pattern
highly variable (Fig. 637g-l). Male genitalia as in Figure 637b-e. Female genitalia as
in Figure 637f
Discussion. The variable color pattern and wide distribution of^. bipunctata have
caused several names to be proposed by a variety of authors, both American and
European. The synonymy listed here is not complete, see Leng (1920) and Korschef-
sky, (1932), for a complete synonymy. The type of A. ophthalmica was supposed to
have been deposited in the BMNH, but R. D. Pope has searched for it without success
so it is probably lost. The type of C. annectans was supposedly in the MCZ collection,
but no such specimen(s) is currently there. The single specimen under that name has
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NORTH AMERICAN COCCINELLIDAE
783
the wrong locality data to qualify as a type. The holotype of C. melanopleum LeConte
is labeled “(gold disc)/Type 6689 (red paper)/C. melanopleura Lee'" Three species
described by Casey, A. ornatella, A. ovipennis, and A. transversalis, are represented
by single type specimens (holotypes). Adalia coloradensis is represented by 3 type
specimens in the Casey collection, the first of which is here designated and labeled
as the lectotype, the other 2 as paralectotypes.
Type locality. Of bipunctata, “Europe”; of frigida, not stated; of bioculata, “United
States”; of humeralis, “Arkansas”; of disjuncta, Maine; of melanopleura, California;
of ophthalmica “TAmerique du Nord”; of annectans, Colorado; of ovipennis, Yount-
ville, Napa Co., California; of transversalis. Las Vegas, New Mexico; of ornatella,
Colorado; of coloradensis, Boulder Co., Colorado (lectotype here designated); of
postica, Springfield, Massachusetts; of sexpustulata and ocellata. Hood River, Oregon.
Type depository. Of bipunctata, Linnean Society, London; oi frigida, not located;
of bioculata, humeralis, and disjuncta, types apparently lost; of melanopleura, MCZ;
of ophthalmica, type apparently lost; of annectans, not known; of ovipennis (34911),
transversalis (349 1 2), ornatella (349 1 3), and coloradensis (349 1 7), USNM; of postica,
sexpustulata, ocellata, and humeralis Johnson, supposedly in USNM but not found.
Distribution. Figure 638. Labrador to Alabama, west to Alaska and California.
Genus Cocci nel la L.
CoccinellaL., 1758, p. 364.— Mulsant, 1850, p. 93.— LeConte, 1852, p. 130.— Crotch,
1873, p. 369.-Crotch, 1874b, p. 105. -Gorham, 1891, p. 154.-Casey, 1899, p.
83. — Dobzhansky, 1925a, p. 241.— Mader, 1926, p. 19. — Dobzhansky, 1931, p.
2.-Timberlake, 1943, p. 13.-Wingo, 1952, p. 23. -Hatch, 1961, p. 171. -Brown,
1962, p. 785. -J. Chapin, 1974, p. 61.-Belicek, 1976, p. 330.-Iablokoff-Khnzo-
rian, 1982, 341. Type-species; Coccinella septempunctata L., by subsequent des-
ignation of Crotch, 1874b.
Coccinella Leach, 1815b, p. 1 16. — Korschefsky, 1932, p. 439.
Coccinella {Coccinella)-. Leng, 1903b, p. 197.
Spilota Billberg, 1820, p. 61.— Timberlake, 1919, p. 163. — Belicek, 1976, p. 331.
Type-species; Coccinella undecimpunctata L., by subsequent designation of Tim-
berlake, 1919.
Coccinella {Neococcinella) Savoyskaya, 1969, p. 104.— lablokoff-Khnzorian, 1982,
p. 341. Type-species; Coccinella undecimpunctata L., by original designation.
Dobzhanskia\s^Aoko^-¥Ainzon?Ln, 1970, p. 89. — lablokoff-Khnzorian, 1982, p. 341.
Type-species; Coccinella undecimpunctata (L.), by original designation.
Coccinellini with length 4.0 to 7.0 mm; form broadly oval, moderately to strongly
convex. Head entirely black, or with 2 pale spots or pale band; pronotum black with
anterolateral angle with white spot varying in size, spots sometimes joined by pale
band along anterior margin, ventral margin of anterolateral angle with pale spot of
varying size; elytron yellow or red; maculate or not. Apex of clypeus nearly truncate,
anterolateral angle produced forward. Lateral margin of elytron narrow, abruptly
reflexed; epipleuron nearly flat. Intercoxal process of prostemum narrow, flat, with
2 convergent or parallel lateral carinae. Apical margin of mesosternum truncate.
Apex of middle and hind tibia each with 2 spurs. Tarsal claw with large tooth, either
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
basal or median (Fig. 1). Postcoxal line incomplete, of Diomus type, with oblique
dividing line (Fig. 1). Male genitalia symmetrical. Female genitalia with infundibu-
lum; coxal plate of the Cycloneda type.
Coccinella is usually separable from other coccinelline genera on the basis of the
pronotal color pattern. This pattern and incomplete postcoxal line with oblique
dividing line will distinguish this genus. Coccinella is mainly a holarctic genus; in
the Western Hemisphere 5 species occur in Mexico, and 4 of these are northern
species with southern range extensions. The generic synonymy presented here in-
cludes only those references pertinent to the genus in North America. Korschefsky
(1932) is to be consulted for synonymy involving other geographic regions. Members
of Coccinella are primarily aphid predators with specific host records as follows:
Acyrthosiphon carnosurn (Buckton), Acyrthosiphon dirhodum (Walker), Acyrthosi-
phon pisum (Harris), Acyrthosiphon solani (Kaltenbach), Acyrthosiphon urticae
(Schrank), Amphorophora rubi (Kaltenbach), Anuraphis farfarae (Koch), Aphis citri-
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NORTH AMERICAN COCCINELLIDAE
785
cola (Van der Goot), Aphis craccivora (Koch), Aphis epilobii (Kaltenbach), Aphis
fabae Scopoli, Aphis forbesi (Weed), Aphis gossypii Glover, Aphis helianthi (Monell),
Aphis jacobaeae (Schrank), Aphis nerii (Boyer de Fonscolombe), Aphis pomi DeGeer,
Aphis rumicis (L.), Aphis sambuci (L.), Aphis urticata (Gmelin), Aphis viburni (Sco-
poli), Brachycaudus cardi (L.), Brachycaudus helichrysi (Kaltenbach), Brevicoryne
brassicae (L.), Cachryphora serotinae (Oestlund), Capitophorus sp., Cavariella ae-
gopodii (Scopoli), Cavariella essigi (Gillette and Bragg), Chromaphis juglandicola
(Kaltenbach), Cryptomyzus ribis (L.), Cuernavaca noxius (Mordvilko), Dactynotus
ambrosiae (Thomas), Dactynotus cirsii (L.), Dactynotus erigeronensis (Thomas), Dac-
tynotus gobonis (Matsumura), Dactynotus sonchi (L.), Drepanosiphum platanoidis
(Schrank), Dysaphis sorbi (Kaltenbach), Eriosoma lanigerum (Hausmann), Eucallip-
terus tiliae (L.), Hyalopterus atriplicis (L.), Hyalopterus pruni (Geoffroy), Hyadaphis
erysimi (Kaltenbach), Hyadaphis foeniculi (Passerini), Hyperomyzus lactucae (L.),
Laingia psammae (Theobald), Macrosiphoniella artemisiae (Boyer de Fonscolombe),
Macrosiphoniella frigidicola (Gillette and Palmer), Macrosiphum avenae (F.), Ma-
crosiphum euphorbiae (Thomas), Macrosiphum rosae (L.), Melanaphis sacchari
(Zehntner), Melanocallis caryaefoliae (Davis), Monellia caryella (Fitch), Monelliopsis
californica (Essig), Monelliopsis caryae (Monell), Monelliopsis nigropunctata (Gra-
novsky), Myzaphis rosarum (Kaltenbach), Myzocallis asclepiadis (Monell), Myzus
cerasi (F.), Myzus persicae (Sulzer), Nearcticaphis bakeri (Cowen), Nearctaphis cra-
taegifoliae (Fitch), Pemphigus brevicornis (Hart), Pemphigus bursarius (L.), Pherioa-
phis trifolii (Monell), Phorodon humuli (Schrank), Pleotrichophorus sp., Rhopalosi-
phum padi (L.), Schizaphis graminum (Rondani), Therioaphis riehmeri (Borner),
Toxoptera aurantii (Boyer de Fonscolombe), Tuberolachnus salignus (Gmelin). Hip-
pa et al. (1978) reported Coccinella hieroglyphica as an effective predator on eggs
and larvae of the chrysomelid beetle Galerucella ""nymphae" (L.) in Finland.
The North American species of Coccinella have been reviewed by Dobzhansky
(1931) and Brown (1962). Brown’s revision is an excellent treatment, and I herein
use his key to species and species synonymies nearly verbatim, with only slight
modification. For more detailed discussion of distribution and relationships see Brown
(1962).
Key to species of Coccinella
1 . Elytron very largely or entirely black 2
- Elytron with extensive pale areas, at least basally 3
2(1). Elytron entirely black; mesepimeron white; length 5.50 to 6.30 mm; San Jacinto
Mts., Riverside Co., California prolongata bridwelli Nunenmacher
- Elytron with epipleuron largely or entirely pale, sometimes with lateral and basal
margin pale; mesepimeron black; length 4.0 to 4.80 mm; Vancouver Island to
northern California hieroglyphica humboldtiensis Nunenmacher
3(1). Elytral suture very narrowly margined with dark brown or black, at least apically
4
- Elytral suture red, not or only slightly darker than the pale discal areas 9
4(3). Each elytron with a very large black spot which usually encloses a pale spot and
is usually joined to the scutellar spot (Fig. 650g); east central California
prolongata sequoiae Dobzhansky
5
Elytral markings different
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
5(4). Head with 2 pale spots on front 6
Head with broad, pale band between eyes; anterior pronotal margin entirely pale
novemnotata Herbst
6(5). Elytron with small scutellar spot, lacking other spots; anterior pronotal margin
black at middle; southwestern British Columbia to southern California
californica Mannerheim
- Elytron with scutellar spot and at least traces of other spots 7
7(6). Pale area on each side of pronotum extending almost to posterior pronotal angle,
deeply penetrating dark discal area inwardly; anterior margin of pronotum some-
times entirely pale, sometimes dark at middle prolongata prolongata Crotch
Pale area of pronotum smaller, not extending beyond basal % , not penetrating
dark discal area to an unusual depth; anterior margin of pronotum black at middle
8
8(7). Each elytron with 2 moderately large spots in addition to scutellar spot, lacking
humeral spot (Fig. 652f); high elevations, Alberta to Utah, and Fresno and Mono
Cos., California alt a Brown
- Elytral spots small, usually more numerous, the humeral spot usually present (Fig.
645). Occurring near the Pacific coast, southern Alaska to southern California .
johnsoni Casey
9(3). Head pale except for black band across base (male); or dark and with broad pale
band between eyes, band rarely interrupted medially (female) 10
Head dark with 2 well-separated spots on front 11
10(9). Elytron with 3 well-developed transverse fasciae, median and subapical fasciae
interrupted at suture (Fig. 6391); Newfoundland to New Jersey, New Mexico,
California, and Alaska trifasciata perplexa Mulsant
Elytral markings variably reduced (Fig. 639g-j); southwestern British Columbia
to southern California trifasciata subversa LeConte
11(9). Ventral pale spot on each anterior pronotal angle small, usually subtriangular,
extending posteriorly not more than Vi as far as dorsal spot except very rarely . 1 2
Ventral pale spot on pronotal angle large, usually trapezoidal, extending posteriorly
as far or almost as far as dorsal spot except in some specimens of h. kirbyi and
u. undecimpunctata in which the dorsal spot is narrowly prolonged to posterior
pronotal angle 16
12(11). Elytron usually with a subbasal fascia, this sometimes reduced to a scutellar spot
and small spot on each humerus or to a scutellar spot only; the sublateral spot at
basal % lacking when fascia is strongly reduced 13
Elytron never with a subbasal fascia or with spot on humerus; often with a small
sublateral spot at basal % 14
13(12). Length of males 5.0 to 7.50 mm; of females, 5.9 to 7.8 mm; elytron with median
and subapical spots transversely more elongate; Newfoundland to Virginia west
to California and Alaska transversoguttata richardsoni Brown
Length of males, 4.0 to 5.40 mm; of females, 4.70 to 6.0 mm; elytron always with
subbasal fascia; restricted to Greenland transversoguttata ephippiata Zetterstedt
14(12). Occurring at and north of the northern limit of trees; Ungava Bay to Alaska . .
fulgida Watson
- Not occurring north of the St. Lawrence River, Quebec 15
15(14). Length 5.0 to 6.0 mm; tarsal claw with tooth reduced difficilis Crotch
Length 6.50 mm or more; tarsal claw with strong tooth septempunctata (L.)
1 6( 1 1 ). Elytron with 3 transverse fasciae, the median and subapical fasciae interrupted at
suture trifasciata perplexa Mulsant
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NORTH AMERICAN COCCINELLIDAE
787
- Elytra never trifasciate 17
17(16). Mesepimeron black, or pale and more or less infuscate; elytron with tricuspate
subbasal band (Fig. 660g) in specimens with whitish mesepimeron 18
- Mesepimeron largely or entirely white; elytron never with subbasal band 20
18(17). Elytron with more or less strongly tricuspate subbasal band and with transverse
subapical spot 19
- Elytron lacking subbasal band, with scutellar spot, frequently with 2 transverse
spots, frequently with transverse spots broken into round spots or lacking entirely
or in part; Vancouver Island to northern California
hieroglyphica humboldtiensis Nunenmacher
1 9( 1 8). Subbasal band on elytron strongly tricuspate (Fig. 660g); anterior margin of prono-
tum frequently entirely pale; Nova Scotia and New Hampshire to Montana, central
British Columbia, and Yukon Territory hieroglyphica kirbyi Crotch
Subbasal band on elytron less strongly or feebly tricuspate (Fig. 6601); anterior
margin of pronotum broadly dark at middle; subarctic, Alaska to northernmost
Manitoba hieroglyphica mannerheimii Mulsant
20(17). Elytron usually with 4 small, round spots in addition to a humeral and scutellar
spot, the 2 submedian spots sometimes joined, the outer apical or the humeral
spot sometimes lacking (Fig. 657f) undecimpunctata undecimpunctata L.
- Elytral spots less numerous and, except for scutellar spot, entirely lacking in some
western specimens, elytron in eastern and northern specimens with large, oblique
spot near middle and large subapical spot, humeral spot always lacking (Fig.
658f, g) monticola Mulsant
Cocci nel la trifasciata perplexa Mulsant
Fig. 639a-f; Map, Fig. 640
Coccinella perplexa Mulsant, 1850, p. 1022.— Casey, 1899, p. 89.— Johnson, 1910,
p. 57.
Coccinella trifasciata: Mulsant, 1850, p. 1 19 (in part); Mulsant, 1866, p. 98 (in part);
Crotch, 1873, p. 370 (in part); Wickham, 1894, p. 301. — Bowditch, 1902, p. 205.—
Leng, 1903b, p. 200. — Blatchley, 1910, p. 5 14. — Dobzhansky, 1931, p. 22. — Dob-
zhansky, 1933, p. lll.-Wingo, 1952, p. 46.-Belicek, 1976, p. 331.
Coccinella eugenii Mulsant, 1866, p. 95 (see entries under Coccinella trifasciata
subversa LeC.).
Coccinella trifasciata perplexa: Hatch, 1961, p. 179. — Brown, 1962, p. 787.
Diagnosis. Length 4.0 to 5.0 mm. Head pale except for black band across base
(male) or black with 2 pale spots (female); pronotum with anterior margin usually
pale, ventral pale spot large, extending posteriorly as far as dorsal spot; elytron with
3 transverse black fasciae as in Figure 639f. Male genitalia as in Figure 639a-d.
Female genitalia as in Figure 639e.
Discussion. The nominate subspecies is Eurasian and it was separated from perplexa
in a key by Brown (1962, p. 788). The elytral color pattern is usually sufficient to
distinguish this subspecies.
Type locality. “Amerique boreale.”
Type depository. Not located, perhaps in DLM.
Distribution. Figure 640. Labrador to New Jersey, west to Alaska and California.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 639. Coccinella trifasciata trifasciata\ C. t. subversa.
Cocci nel la trifasciata subversa LeConte
Fig. 639g-j; Map, Fig. 640
Coccinella subversa LeConte, 1854, p. 19.— Crotch, 1874b, p. 116.
Coccinella trifasciata subversa-. Crotch, 1873, p. 370. — Leng, 1903b, p. 200. — Dob-
zhansky, 1931, p. 23.-Hatch, 1961, p. 179.-Brown, 1962, p. 789.
Coccinella trifasciata ab. subversa: Korschefsky, 1932, p. 499. — Mader, 1936, p. 375.
Coccinella perplexa subversa: Johnson, 1910, p. 57.
Coccinella Juliana Mulsant, 1856, p. 141.— Casey, 1899, p. 89.
Coccinella trifasciata Juliana: Crotch, 1873, p. 370.— Crotch, 1874b, p. 115. — Leng,
1903b, p. 200. — Dobzhansky, 1931, p. 25.
Coccinella trifasciata ah. Juliana: Mader, 1930, p. 163. — Korschefsky, 1932, p. 499.
Coccinella perplexa Juliana: Johnson, 1910, p. 57.
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NORTH AMERICAN COCCINELLIDAE
789
Coccinella barda LeConte, 1860, p. 286.
Coccinella eugenii Mulsant, 1866, p. 95.— Crotch, 1874b, p. 115.— Casey, 1899, p.
90.
Coccinella trifasciata eugenii: Crotch, 1873, p. 370. — Leng, 1903b, p. 200. — Dob-
zhansky, 1931, p. 24.
Coccinella trifasciata ab. eugenii: Korschefsky, 1932, p. 499. — Mader, 1936, p. 375.
Coccinella perplexa fennica: Johnson, 1910, p. 57.
Coccinella trifasciata ab. praedicta Mader, 1930, p. 163.
Coccinella trifasciata: Dobzhansky, 1933, p. 111.
Description as for C. t. perplexa except head of female black with pale band between
eyes; elytral maculation reduced as in Figure 639g-j. The type of C. Juliana was
supposed to have been deposited in the BMNH, but R. D. Pope has searched for it
without success so it is probably lost. There are 3 specimens of C. t. subversa in the
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
LeConte collection, one of which, female bears a type label. LeConte did not state
how many specimens of C. t. subversa he had, therefore I here designate and label
the specimen labeled “(blue disc)/TYPE 6687(red paper)/C. subversa LeC. Cooper”
as the lectotype. No types of C. barda exist in the LeConte collection and I presume
they are no longer extant.
Type locality. Of subversa, Oregon (lectotype here designated); of Juliana, Califor-
nia; of barda, Punto de los Reyes (Marin Co.), California; of praedicta, Alameda,
California; of eugenii, California.
Type depository. Of subversa MCZ; of barda, type apparently lost; of Juliana, type
apparently lost; of praedicta, not known; of eugenii, not known.
Distribution. Figure 640. British Columbia to California.
Coccinella transversoguttata richardsoni Brown
Fig. 641a-g; Map, Fig. 642
Coccinella quinque-notata Kirby, 1837, p. 230 (not Coccinella quinquenotata Ha-
worth, 1812).
Coccinella 5-notata: Fitch, 1862, p. 849. — Crotch, 1873, p. 370.— Casey, 1899, p.
89.-Palmer, 1914, p. 219.
Coccinella 5-notata interrupta Fitch, 1862, p. 851 (not Coccinella interrupta Four-
croy, 1785).
Coccinella transversoguttata: Mulsant, 1850, p. 1 17 (in part). — Mulsant, 1866, p. 97
(in part). — Crotch, 1874b, p. 1 16 (in part). — Wickham, 1894, p. 301. — Leng, 1903b,
p. 199 (in part).— Johnson, 1910, p. 61 (in part). — Dobzhansky, 1931, p. 14.—
Dobzhansky, 1935, p. 334.-Wingo, 1952, p. 46. -Hatch, 1961, p. 178.-Belicek,
1976, p. 333.
Coccinella transversoguttata transversalis : Wickham, 1894, p. 306.
Coccinella transversoguttata ab. zetterstedti Mader, 1930, p. 151.
Coccinella transversoguttata \3X. nugatoria: Leng, 1903b, p. 199. — Dobzhansky, 1935,
p. 334.
Coccinella transversoguttata richardsoni Brown, 1962, p. 790.
Diagnosis. Length 5.0 to 7.80 mm. Head black with 2 well separated pale spots;
pronotum with anterior margin black at middle, ventral pale spot small, triangular,
extending posteriorly Vs to Vi as far as dorsal spot; elytron variable but usually with
at least a subbasal fascia (Fig. 64 If, g). Male genitalia as in Figure 641a-d. Female
genitalia as in Figure 64 le.
Discussion. This species is extremely widely distributed and commonly collected.
The color pattern is nearly always sufficient for recognizing C t. richardsoni. The
unique type (holotype) of C. quinquenotata is a female labeled “Type (white disc
with red border)/N. Amer 5961 a./Coccinella 5-notata Kirby n. america 5961. Rev.
W. Kirby.”
Type locality. Of quinquenotata. North America; of interrupta. New York; of zet-
terstedti, Lapland and Canada.
Type depository. Of quinquenotata, BMNH; of interrupta, not known; of zetter-
stedti, not known.
Distribution. Figure 642. Labrador to Virginia, west to Alaska and California.
1985
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NORTH AMERICAN COCCINELLIDAE
Fig. 641. Coccinella transversoguttata richardsoni.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 642. Distribution. Coccinella transversoguttata richardsoni (shaded); C. t. ephippiata
(cross hatch, Greenland).
Coccinella transversoguttata ephippiata Zetterstedt
Map, Fig. 642
Coccinella ephippiata Zetterstedt, 1838, p. 186.
Coccinella trifasciata: Fabricius, 1780, p. 186.
Coccinella transversoguttata: Mulsant, 1850, p. 1 17 (in part).— Crotch, 1874b, p. 1 16
(in part). — Henriksen and Lundbeck, 1918, p. 515. — Henriksen, 1939, p. 45.
Smaller than C. t. transversoguttata and C. t. richardsoni, length of males, 4.10 to
5.40 mm, the average about 5.0 mm; length of females, 4.70 to 6.0 mm, the average
about 5.4 mm. Elytron maculate as in C. t. transversoguttata-, the subbasal band
never broken or reduced; the median and subapical spots less elongate transversely
than in C. t. richardsoni, the sublateral spot present in 90 per cent of the specimens,
well-developed or quite distinct in 1 2 per cent, very small or rather indistinct in 8
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NORTH AMERICAN COCCINELLIDAE
793
per cent. Male genitalia much as in the allied forms; the hastate, apical portion of
the median lobe less variable than in the others, more slender than in C. t. trans-
versoguttata or in some specimens of C. t. richardsoni.
Type locality. Greenland.
Type depository. Type not located.
Distribution. Figure 642. Greenland.
Coccinella californica Mannerheim
Fig. 643a-g; Map, Fig. 649
Coccinella californica Mannerheim, 1843, p. 312. — Mulsant, 1850, p. 110.— Mul-
sant, 1866, p. 91.— Casey, 1899, p. 88.— Johnson, 1910, p. 62. — Dobzhansky,
1931, p. 1 1. — Wingo, 1952, p. 46. — Hatch, 1961, p. 178. — Brown, 1962, p. 793.—
Belicek, 1976, p. 335.
Coccinella 5-notata californica: Crotch, 1873, p. 370.
Coccinella transversoguttata californica: Crotch, 1874b, p. 116. — Weise, 1892, p.
25. — Leng, 1903b, p. 200.
Coccinella transversoguttata ab. californica: Mader, 1930, p. 151.
Coccinella californica melanocollis Johnson, 1910, p. 62.
Coccinella transversoguttata ab. melanocollis: Korschefsky, 1932, p. 496.— Mader,
1936, p. 374.
Diagnosis. Length 5.10 to 6.80 mm. Head black with 2 well separated pale spots
(Fig. 643g); pronotum with anterior margin black at middle, ventral pale spot elon-
gate, triangular, extending posteriorly % to % as far as dorsal spot; elytron with small
scutellar spot, sutural margin very narrowly dark brown (Fig. 643 f)- Male genitalia
as in Figure 643a-d. Female genitalia as in Figure 643e.
Discussion. As noted by Brown (1962), C californica is most closely related to C.
johnsoni, but is most likely to be confused with the immaculate form of C. novem-
notata. The latter species has the interocular region and the anterior pronotal margin
entirely pale. This species has been recorded from Oklahoma, Iowa and Missouri
(Wingo, 1952); Brown, 1962) but was probably carried there by commerce. Brown
stated that C californica probably does not breed east of the low region adjacent to
the Pacific Coast. There are 3 types of C. californica in Helsinki, one of which, a
female, labeled “Eschsch./Califomia/coll. Mannerh/Califomica Eschsch” I here des-
ignate and label as the lectotype, the other 2 specimens as paralectotypes.
Type locality. Of californica, California (lectotype here designated); of melanocollis,
Berkley, California.
Type depository. Of californica, UMZH; of melanocollis, no type designated.
Distribution. Figure 649. British Columbia to California.
Coccinella johnsoni Casey
Fig. 645; Map, Fig. 644
Coccinella johnsoni Casey, 1908, p. 403.— Johnson, 1910, p. 61. — Dobzhansky, 1931,
p. 13.-Hatch, 1961, p. 177. -Brown, 1962, p. 794.-Belicek, 1976, p. 336 (as a
synonym of C. californica).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 643. Cocci nella calif or nica.
Coccinella novemnotata johnsoni: Leng, 1920, p. 216.
Coccinella novemnotata dih. johnsoni: Korschefsky, 1932, p. 512.
Differs from C. californica only in having elytral maculae as in Figure 645. Coc-
cinella johnsoni is likely to be confused only with some western forms of C. nov-
emnotata (see remarks under C. californica). The name johnsoni is based on a single
female in the Casey collection (holotype).
1985
NORTH AMERICAN COCCINELLIDAE
795
Fig. 644. Distribution. Coccinella johnsoni (cross hatch, west coast); C. novemnotata.
Type locality. San Diego, California.
Type depository. USNM (35517).
Distribution. Figure 644. Alaska to southern California.
Coccinella septempunctata (L.)
Fig. 646a-e; Map, Fig. 649
Coccinella 7 -punctata L., 1758, p. 365.
Coccinella septempunctata: Mulsant, 1846, p. 79. — Mulsant, 1850, p. 115.— Crotch,
1874b, p. 1 17.— Dobzhansky, 1926a, p. 17. — Hoebeke and Wheeler, 1980, p. 209.
Coccinella {Coccinella) septempunctata: lablokoff-Khnzorian, 1982, p. 370.
Diagnosis. Length 6.50 to 7.80 mm. Head black with 2 well separated pale spots;
pronotum with anterior margin black at middle with ventral pale spot small, ex-
tending posteriorly Vs as far as dorsal spot; elytron with 3 black spots in addition to
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 645. Coccinella johnsoni.
scutellar spot (Fig. 646e). Male genitalia as in Figure 646a-c. Female genitalia as in
Figure 646d.
Discussion. This widespread palearctic species was intentionally introduced into
North America several times from 1956 to 1971 for biological control of aphids. All
of those attempts apparently failed in getting C. septempunctata established, but in
1 973 an established population was found in Bergen Co., New Jersey. This population
is thought to have been the result of an accidental introduction rather than a pur-
poseful one (Angalet and Jacques, 1975). Since 1973, this species has spread naturally
and been colonized and established in Delaware, Georgia, and Oklahoma. It is also
widely distributed in Maine (R. Dysart, pers. comm), but I have not seen any specific
locality data for that state. References with published data on spread and release are:
Angalet and Jacques (1975); Angalet et al. (1979); Cartwright et al. (1979); Tedders
and Angalet (1981); and Hoebeke and Wheeler (1980). Coccinella septempunctata is
established in Quebec as the result of either an accidental introduction or spread
from Maine releases. Locality data listed here is from Larochelle (1979). This species
goes to C. difficilis in Brown’s key (1962), but C. septempunctata is much larger and
as yet does not occur in the same geographic region as C. difficilis. Korschefsky (1932)
presents a complete synonymy of the many names associated with this species.
Type locality. “Suecica”.
Type depository. Type not examined.
Distribution. Figure 649. QUEBEC: Berthier Parish, Berthierville, Lanoraie, Saint-
Jean-de-Matha; Chambly Parish, Saint-Bruno; Chateaugay Parish, Saint-Chrysos-
tome; He de Montreal, Dorval, Montreal, Montreal-Nord; L’ Assumption Parish, La
Plaine, Repentigry; Portneut Parish, Saint- Augustin; Sainte-Maurice Parish, Trois-
Rivieres; Terrebonne Parish, Terrebonne; Vaudruevil Parish, Pincourt, Rigaud.
CONNECTICUT: Fairfield Co.; New Haven Co.; Hammonasett State Park; Mid-
dlesex Co.; New London Co. DELAWARE: Kent Co.; Newcastle Co.; Sussex Co.
GEORGIA: Peach Co., Byron; Houston Co., Baconton. NEW JERSEY: Bergen Co.;
Fig. 646. Coccinella septempunctata.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Essex Co., Gloucester Co.; Hunterdon Co.; Mercer Co.; Middlesex Co.; Monmouth
Co.; Morris Co.; Passaic Co.; Salem Co.; Somerset Co.; Union Co.; Warren Co. NEW
YORK: Bronx Co.; Brooklyn Co.; Far Rockaway; Flushing; Ithaca; Nassau Co.,
Malverne; Orange Co.; Putnam Co.; Queens Co., Rochdale; Richmond Co.; Rockland
Co.; Suffolk Co.; Tompkins Co., Freeville; Ulster Co.; Westchester Co. OKLA-
HOMA: Payne Co. PENNSYLVANIA: Berks Co., Leesport; Bucks Co., Hilltown;
Columbia Co., Catawissa; Lehigh Co., Allentown; Pike Co., Bushkill.
Cocci nella novemnotata Herbst
Fig. 647a-h, 648; Map, Fig. 644
Coccinella 9 -not at a 1793, p. 269. — Fabricius, 1798, p. 78.— Fabricius, 1801,
p. 366. — Mulsant, 1850, p. 123. — Mulsant, 1866, p. 99. — Fitch, 1862, p. 106.—
Crotch, 1873, p. 370.— Crotch, 1874b, p. 1 17. — Wickham, 1894, p. 301.— Casey,
1899,, p. 88. — Bowditch, 1902, p. 205. — Leng, 1903b, p. 198. — Blatchley, 1910,
p. 514.— Johnson, 1910, p. 59. — Palmer, 1914, p. 226. — Dobzhansky, 1931, p.
4.— Korschefsky, 1932, p. 5 12. — Dobzhansky, 1933, p. 111.— Wingo, 1952, p.
46. — Brown, 1962, p. 794.— J. Chapin, 1974, p. 61. — Belicek, 1976, p. 334.
Coccinella franciscana Mulsant, 1853, p. 147.
Coccinella 9- not ata franciscana: Crotch, 1873, p. 370. — Leng, 1903b, p. 198.— John-
son, 1910, p. 59. — Dobzhansky, 1931, p. 111.
Coccinella novemnotata inaequalis Fitch, 1862, p. 107.
Coccinella novemnotata ab. parvamaculata Fitch, 1862, p. 107.
Coccinella novemnotata conjuncta Fitch, 1862, p. 107.
Coccinella novemnotata ab. confluenta Fitch, 1862, p. 107.
Coccinella novemnotata ab. divisicollis Fitch, 1862, p. 107.
Coccinella degener Casey, 1899, p. 88.
Coccinella novemnotata degener: Leng, 1903b, p. 198. — Casey, 1908, p. 404.— John-
son, 1910, p. 59. — Leng, 1920, p. 216. — Dobzhansky, 1931, p. 6. — Dobzhansky,
1933, p. I12.-Hatch, 1961, p. 177.
Coccinella novemnotata oregona Casey, 1908, p. 403. — Dobzhansky, 1931, p. 7.
Diagnosis. Length 4.70 to 7.0 mm. Head with broad, pale band between eyes,
black anteriorly and posteriorly (Fig. 647e); pronotum with anterior margin entirely
pale, ventral pale spot large, trapezoidal, extending posteriorly as far as dorsal spot;
elytron with black spots that decrease in size and number until only the scutellar
spot remains in some specimens, suture narrowly blackish (Fig. 647f-h). Male gen-
italia as in Figure 647a-d. Female genitalia as in Figure 648.
Discussion. This species is extremely widespread in North America and is com-
monly collected. The pale anterior pronotal margin and blackish sutural margin of
the elytron distinguish C. novemnotata from other species of Coccinella. There are
6 types of C. degener in the Casey collection, one of which, a male, I designate and
label as the lectotype, the remaining 5 as paralectotypes. There are no specimens of
C. oregona now in the Casey collection, but it is apparent from the original description
that he had only one specimen (holotype). There are 2 types of C. franciscana in the
Crotch collection, one of which, a female, labeled “Type (blue paper)/Type frauds-
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NORTH AMERICAN COCCINELLIDAE
799
Fig. 647. Coccinella novemnotata.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 648. Coccinella novemnotata (female genitalia).
cana Chevrol.” I here designate and label as the lectotype, the other specimen as a
paralectotype.
Type locality. Of novemnotata. North America; offranciscana, California (lectotype
here designated); of inaequalis, parvamaculata, conjuncta, conjluenta, and divisicollis.
New York; of degener. Fort Wingate, New Mexico; of oregona, southern Oregon.
Type depository. Of novemnotata, not known; offranciscana, UCCC; of inaequalis,
parvamaculata, conjuncta, conjluenta, and divisicollis, not located; of degener and
oregona, USNM.
Distribution. Figure 644. Maine to Georgia, west to British Columbia and southern
California.
Coccinella prolongata prolongata Crotch
Fig. 650a-f; Map, Fig. 651
Coccinella prolongata Crotch, 1873, p. 371. — Casey, 1899, p. 88.— Dobzhansky,
1931, p. 9. -Johnson, 1910, p. 64.-Hatch, 1961, p. 1 77.-Belicek, 1976, p. 337.
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NORTH AMERICAN COCCINELLIDAE
801
Fig. 649. Distribution. Coccinella californica (shaded); C. septempunctata (dot).
Coccinella transversoguttata var. prolongata: Leng, 1903b, p. 199.
Coccinella monticola prolongata: Leng, 1920, p. 216.
Coccinella prolongata prolongata: Brown, 1962, p. 797.
Diagnosis. Length 5.70 to 7.0 mm. Head black with 2 pale spots; pronotum with
anterolateral pale spot larger than in any other species, ventral spot extending pos-
teriorly % to % as far as dorsal spot; elytron with 2 black spots and a scutellar spot,
suture narrowly darkened (Fig. 6501). Male genitalia as in Figure 650a-d. Female
genitalia as in Figure 650e.
Discussion. There are 8 specimens in the LeConte collection in the MCZ under
this name, the first of which bears a “type” label; therefore I designate and label this
specimen labeled “(green disc)/4625/Type 8243(red paper)” as the lectotype.
Type locality. Kansas (lectotype here designated).
Type depository. MCZ.
Distribution. Figure 651. BRITISH COLUMBIA: Aspen Grove; Nicola; Oliver;
Osoyoos; Vernon. CALIFORNIA: Siskiyou Co. COLORADO: Boulder; Clear Creek;
Denver; Garland; Rabbit Ears Pass. IDAHO: McCall; Montpelier; Paris. MON-
TANA: Helena. NEBRASKA: Sioux Co. NEVADA: Franktown. OREGON: Summer
Lake; Upper Klamath Marsh. UTAH: Salt Lake City. WASHINGTON: Chelan;
Pullman; Puyallup; Wenatchee. WYOMING: Yellowstone National Park.
Coccinella prolongata sequoiae Dobzhansky
Fig. 650g; Map, Fig. 651
Coccinella prolongata sequoiae Dobzhansky, 1931, p. 10. — Brown, 1962, p. 797.
Description as for C. p. prolongata except pronotal pale areas not extended to an
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 650. f. Coccinella prolongata prolongata. g. C. p. sequoiae. h. C. p. bridwelli.
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NORTH AMERICAN COCCINELLIDAE
803
Fig. 651. Distribution. Coccinella prolongata prolongata (dot); C. p. sequoiae (star); C. p.
bridwelli (triangle).
unusual degree; elytron with large irregular black spot enclosing small pale spot (Fig.
650g).
Dobzhansky designated 2 specimens as types, one of which, a male, I designate
and label as the lectotype, the other as a paralectotype.
Type locality. Near Camp Wolverton, 7,000 to 9,000 ft.. Sequoia National Park,
California (lectotype here designated).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Type depository. CAS.
Distribution. Figure 651. CALIFORNIA: Alpine Co.; Inyo Co., Deep Spring Valley;
Mono Co., McGee Greek; Tulare Co. NEVADA: Winemucca Lake.
Cocci nel la prolongata bridwelli Nunenmacher
Fig. 650h; Map, Fig. 651
Coccinella Nunenmacher, 1913, p. 76. — Leng, 1920, p. 216.
Coccinella prolongata bridwelli: Dobzhansky, 1931, p. 11. — Brown, 1962, p. 798.
Description as for C. p. sequoiae except elytron entirely black (Fig. 6 5 Oh). There
are 2 type specimens of C. p. bridwelli in the CAS, one of which, a male, I designate
and label as the lectotype, the other as a paralectotype.
Type locality. Tahquitz Valley, San Jacinto Mountains, California (lectotype here
designated).
Type depository. CAS.
Distribution. Figure 651. CALIFORNIA: Riverside Co., San Jacinto Mts., Idyll-
wild, Santa Rosa Peak.
Coccinella aha Brown
Fig. 652a-f; Map, Fig. 653
Coccinella alta Brown, 1962, p. 798. — Belicek, 1976, p. 332.
Coccinella suturalis Casey,, 1899,, p. 89. — Dobzhansky, 1931, p. 21.— Brown, 1962,
p. 798. — Belicek, 1976, p. 332 (not Coccinella suturalis Olivier, 1791).
Coccinella monticola var. suturalis: Leng, 1903b, p. 198.
Diagnosis. Length 4.80 to 5.30 mm. Head black with well separated pale spots;
pronotum with anterior margin black at middle, ventral pale spot small, triangular,
extending posteriorly 'A to Vi as far as dorsal spot; elytron with sutural margin blackish,
2 black spots present (Fig. 6521). Male genitalia as in Figure 652a-d. Female genitalia
as in Figure 652e.
Discussion. This species most nearly resembles monticola which does not have the
elytral suture darkened. The name suturalis is based on a single female (holotype) in
the Casey collection.
Type locality. Of alta. Salt Lake Co., Utah; of suturalis, Colorado.
Type depository. Of alta, USNM; of suturalis, USNM (35521).
Distribution. Figure 653. ALBERTA: Morley, Mount Rae. CALIFORNIA: Fresno
Co.; Kings River (Bubbs Creek Canyon), Mount Gold, Mount Kaiser; Inyo Co.,
Coyote Ridge; Mono Co., White Mtn.
Coccinella difficilis Crotch
Fig. 654a-f; Map, Fig. 653
Coccinella difficilis Crotch, 1873, p. 370. — Leng, 1903b, p. 200.— Johnson, 1910, p.
64. — Dobzhansky, 1931, p. 20. — Hatch, 1961, p. 178. — Brown, 1962, p. 799.
Coccinella monticola difficilis: Leng, 1920, p. 216.
Coccinella vandykei ^ur\cnmachcr, 1909, p. 161.
Coccinella transversoguttata vandykei: Leng, 1920, p. 216.
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NORTH AMERICAN COCCINELLIDAE
805
Fig. 652. Coccinella aha.
Diagnosis. Length 5.0 to 6.0 mm. Head black with 2 well separated pale spots;
pronotum with anterior margin black at middle, ventral pale spot small, subtrian-
gular, extending posteriorly V3 to as far as dorsal spot; elytron as in Figure 654f.
except some spots rarely lacking. Male genitalia as in Figure 654a-d. Female genitalia
as in Figure 654e.
Discussion. C. difficilis can be confused with examples of C. richardsoni in which
the subbasal elytral band is reduced to a scutellar spot. In such specimens the size
is usually greater; the sublateral spot of each elytron is lacking; and the discal spots
are transversely more elongate. There are 9 specimens under the name C. difficilis
presently in the MCZ collection, 6 of which could possibly be type material. It is
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
impossible to tell from the original description how many type specimens Crotch
had. Therefore I designate and label as the lectotype the hrst specimen (female) in
the series labeled “Utah/Coccineila difficilis/Horn Co. H-2057/C. difficilis Cr.” There
are 2 type specimens of C. vandykei in the CAS, and I designate a male labeled
“Goldheld/Esmeralda Co. Nev. VI. 29.07/coH’d by F. W. Nunenmacher” as the
lectotype.
Type locality. Of difficilis, Utah (lectotype here designated); of vandykei, Goldheld,
Nevada (lectotype here designated).
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NORTH AMERICAN COCCINELLIDAE
807
Fig. 654. Coccinella difficilis.
Type depository. Of difficilis, MCZ; of vandykei, CAS.
Distribution. Figure 653. ARIZONA: Flagstalf, San Francisco Mts. CALIFORNIA:
Adin; Alturas, Chilcoot. COLORADO: Colorado Springs; Glenwood Springs. IDA-
HO: Boise; Jerome; Pocatello. MONTANA: Helena. NEVADA: Elko; Lovelock.
OREGON: Grant Co.; Unity. UTAH: Fort Douglas; Logan; Milford; Salt Lake Valley.
WYOMING: Cheyenne; Green River City.
Coccinella fulgida Watson
Fig. 655a-f; Map, Fig. 656
Coccinella fulgida Watson, 1954, p. 45. — Brown, 1962, p. 799. — Belicek, 1976, p.
335.
Coccinella nugatoria: Leng, 1919, p. 17 (misidentification).
Coccinella undecimpunctata: Dobzhansky, 1931, p. 28 (in part)
(misidentification).— Wheeler and Hoebeke, 1981, p. 213.
Coccinella difficilis: Chapin, 1956, p. 152 (in part) (misidentification).
Diagnosis. Length 4.50 to 5.60 mm. Head black with 2 well separated pale spots;
pronotum with anterior margin black at middle, ventral pale spot triangular, small.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 655. Coccinella fulgida.
extending posteriorly U to V5 as far as dorsal spot; elytron with small subbasal spot
which may be lacking or may be joined to transverse median spot, a transverse spot
at apical in all specimens (Fig. 65 5e, f). Male genitalia as in Figure 655a-c. Female
genitalia as in Figure 655d.
Type locality. Cape Henrietta Maria, north central Ontario at 55 N., 28 15 W.
Type depository. CNC.
Distribution. Figure 656. BRITISH COLUMBIA: Summit Lake. NORTHWEST
TERRITORIES: Bathurst Inlet; Kater Point, Langton Bay; Reindeer Depot. QUE-
BEC: Fort Chimo. ALASKA: Mead River, south of Point Barrow; Rampart House,
Alaska-Yukon border; Rampart House, 60 to 70 miles north; Toms Lake.
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NORTH AMERICAN COCCINELLIDAE
809
Fig. 656. Distribution. Coccinella fulgida (star); C. undecimpunctata (dot).
Coccinella undecimpunctata undecimpunctata (L.)
Fig. 657a-f; Map, Fig. 656
Coccinella 1 1-punctata L., 1758, p. 366.
Coccinella undecimpunctata: Schaeffer, 1912, p. 104. — Dobzhansky, 1931. p. 27 (in
part).-Davis, 1932, p. 101. -Brown, 1940, p. 72.-Chapin, 1956, p. 155. -Be-
licek, 1976, p. 335.
Coccinella undecimpunctata undecimpunctata: Brown, 1962, p. 800.
Coccinella {Neococcinella) undecimpunctata: Savoyskaya, 1969, p. 104.— lablokoff-
Khnzorian, 1982, p. 357.
Diagnosis. Length 4.0 to 5.0 mm. More elongate and less convex than any other
species of Coccinella. Head black with 2 well separated pale spots; pronotum with
anterior margin black at middle, ventral pale spot large, extending posteriorly nearly
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 657. Coccinella undecimpunctata.
as far as dorsal spot; elytron usually with 5 black spots and a small scutellar spot
(Fig. 657f). Male genitalia as in Figure 657a-d. Female genitalia as in Figure 657e.
Discussion. This species is an accidentally introduced Eurasian species first reported
from North America by Schaeffer (1912). The most recent surveys of distribution
are by Watson (1979) and Wheeler and Hoebeke (1981). It is highly distinctive in
our fauna and should not be confused with any other species of Coccinella.
Type locality. “Europa.”
Type depository. Type not examined.
Distribution. Figure 656. BRITISH COLUMBIA: Vancouver. MIQUELON IS-
LAND. NEW BRUNSWICK: Fundy National Park. NEWFOUNDLAND: Cow
Head; Fogo; Grand Bank; Piccadilly; Port au Port Peninsula; St. Pauls; Tilting;
Twillinggate. NOVA SCOTIA: Halifax. ONTARIO: Collingwood; Guelph; Harrow;
London; Manitoulin Island; Ottawa; Port Stanley; Toronto; Waterloo. PRINCE ED-
WARD ISLAND: Brackley Beach. QUEBEC: Baie St. Paul; Becancour; Berthierville;
Richelieu; St. Augustin; St. Louis; St. Foy; Sorasboro. CONNECTICUT: Rocky Hills
St. Pk. MASSACHUSETTS: Arlington; Chelsea; Falmouth Heights; Lynn Beach;
Medford; Nahant; Provincetown; Saugus; Stoneham; Wollaston. NEW JERSEY:
Camden; Rutherford. NEW YORK: East Marion; Flanders; Ghent; Great Kills;
Greenport; Ithaca; Jefferson Co.; Long Island; Ludlowville; Mexico; Newburgh; New
York City; Orient; Riverhead; Rossie; Staten Island. OHIO: Bowling Green. ORE-
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NORTH AMERICAN COCCINELLIDAE
811
GON: Benton Co., Corvallis. PENNSYLVANIA: Philadelphia. RHODE ISLAND:
Providence Co., Cumberland. WASHINGTON: Seattle.
Coccinella monticola Mulsant
Fig. 658a-g; Map, Fig. 659
Coccinella monticola Mulsant, 1850, p. 1 15. — Mulsant, 1866, p. 96.— Crotch, 1873,
p. 371.— Crotch, 1874b, p. 1 1 5.— Wickham, 1894, p. 301.— Casey, 1899, p. 89.—
Leng, 1903b, p. 198.— Johnson, 1910, p. 63. — Palmer, 1914, p. 222. — Brown,
1962, p. 802.-Belicek, 1976, p. 333.
Coccinella nivicola monticola: Dobzhansky, 1931, p. 17. — Dobzhansky, 1933, p.
111. — Wingo, 1952, p. 46. — Hatch, 1961, p. 178.
Coccinella lacustris LeConte, 1852, p. 131.
Coccinella alutacea Casey, 1899, p. 89.
Coccinella transversoguttata alutacea: Leng, 1 903b, p. 200.
Coccinella monticola alutacea: Johnson, 1910, p. 63.
Coccinella nivicola alutacea: Dobzhansky, 1931, p. 19.— Dobzhansky, 1933.— p.
111. -Hatch, 1961, p. 178.
Coccinella impressa Casey, 1899, p. 89 (not Coccinella undecimpunctata impressa
Verhoeff, 1891).
Coccinella transversoguttata impressa: Leng, 1903b, p. 199.
Coccinella monticola impressa: Casey, 1908, p. 404.
Coccinella monticola ab. impressa: Johnson, 1910, p. 216.
Coccinella nevadica Casey, 1899, p. 88.— Casey, 1908, p. 402.
Coccinella transversoguttata var. nevadica: Leng, 1903b, p. 201.
Coccinella transversoguttata nevadica: Leng, 1920, p. 216.
Coccinella transversoguttata ab. nevadica: Mader, 1936, p. 374.
Coccinella californica nevadica: (?) Dobzhansky, 1931, p. 12.
Coccinella monticola sellica Johnson, 1910, p. 63.
Coccinella monticola postica Johnson, 1910, p. 63 (not Coccinella postica Mulsant,
1850).
Coccinella monticola confluenta Johnson, 1910, p. 63 (not Coccinella novemnotata
confluenta Fitch, 1862).
Coccinella monticola biguttata Johnson, 1910, p. 63 (not Coccinella biguttata Fa-
bricius, 1787).
Diagnosis. Length 5.20 to 7.0 mm. Head black with 2 pale spots; pronotum with
anterior margin black at middle; ventral pale spot large, trapezoidal, extending pos-
teriorly nearly as far as dorsal spot; elytron with scutellar spot, other spots heavy,
reduced or lacking depending on locality (Fig. 658f, g). Male genitalia as in Figure
658a-d. Female genitalia as in Figure 658e.
Discussion. C. monticola is widespread and variable in color pattern. It is not a
commonly collected species but the key characters will enable it to be recognized.
The types of the Casey names alutacea (female), impressa (female), and nevadica
(male), are all unique (holotypes). The type of C. monticola is supposed to be in the
LeConte collection, but no specimens presently exist there that could be type material.
Three specimens under that name bear a dark blue disc signifying Oregon or Wash-
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 658. Coccinella monticola.
ington as the locality, but have no other data. Two specimens have pale blue, cut
edged labels signifying the north shore of Lake Superior. The third specimen in the
series is labeled as a type of C. lacustris. I here designate and label as the lectotype
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NORTH AMERICAN COCCINELLIDAE
813
this specimen bearing the labels “(pale blue disc)/4624/TYPE 6688 (red paper)/C.
lacustris LeC.”. I also consider the 2 specimens with pale blue, cut edged labels, type
material of lacustris and label them as paralectotypes.
Type locality. Of monticola, “les Montagnes Rocheuses” (Rocky Mountains); of
lacustris. Lake Superior (lectotype here designated); of alutacea, New Mexico; of
impressa, California; of nevadica, Reno,, Nevada; of sellica, California, of postica,
California; of confluenta, California; of biguttata, Colorado.
Type depository. Of monticola, not known; of lacustris, MCZ; of alutacea (35519),
impressa (35520), and nevadica (35522), USNM; of sellica, postica, confluenta, and
biguttata, types not designated.
Distribution. Figure 659. Nova Scotia to Massachusetts and New York, west to
Northwest Territories and California. Peripheral localities: Baddeck, Nova Scotia;
Fredericton, New Brunswick; Sudbury, Ontario; Treesbank, Manitoba; Waskesiu
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 660. Coccinella hieroglyphica Kirbyi\ C. h. mannerheimi.
Lake, Saskatchewan; Fort Smith and Fort McPherson, Northwest Territories; Spring-
field, Massachusetts; Duluth, Minnesota; Rincon, New Mexico; Prescott, Arizona;
Mono Co., California.
Coccinella hieroglyphica kirbyi Crotch
Fig. 660a-e, g; Map, Fig. 661
Coccinella kirbyi Crotch, 1874, p. 37 (new name for tricuspis Kirby, not Thunberg).
Coccinella hieroglyphica kirbyi: Timberlake, 1943, p. 14. — Brown, 1962, p. 804.
Coccinella hieroglyphica L., 1758, p. 365 (in part). — Hatch, 1961, p. 180. — Belicek,
1976, p. 334.
Coccinella tricuspis Kirby, 1837, p. 231 (not Coccinella tricuspis T\mr\hQYg, 1794).—
Mulsant, 1850, p. 107. — Mulsant, 1866, p. 88. — Crotch, 1873, p. 371.— Weise,
1892,, p. 25 (in part). — Wickham, 1894, p. 301.— Casey, 1899, p. 90. — Leng,
1903b, p. 201 (in part); Johnson, 1910, p. 59.
Coccinella hieroglyphica tricuspis: Dobzhansky, 1931, p. 26. — Wingo, 1952, p. 46.
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NORTH AMERICAN COCCINELLIDAE
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Fig. 661. Distribution. Coccinella hieroglyphica Kirbyi (shaded); C h. mannerheimi (tri-
angle); C. h. humboldtiensis (dot).
Coccinella mannerheimi tricuspis: Mader, 1930, p. 160.
Coccinella mannerheimi: Crotch, 1874b (in part).
Diagnosis. Length 3.70 to 4.70 mm. Head black with 2 well separated pale spots;
pronotum with anterior margin narrowly pale at middle in nearly all males and some
females, black in most females and some males, ventral pale spot large, trapezoidal,
extending posteriorly nearly as far as dorsal spot; elytron with heavy, tricuspate
subbasal band and large transverse spot at apical V4 (Fig. 660g). Male genitalia as in
Figure 660a-g. Female genitalia as in Figure 660e.
Discussion. The nominate subspecies is Eurasian, and Brown (1962) provides a
key to separate it from kirbyi. One female type specimen of tricuspis labeled “Type
(white disc with red border)/n. amer. 5962 a./Coccinella tricuspis Kirby n. amer.
5962 Rev. W. Kirby” has been examined and is here designated and labeled as the
lectotype.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Type locality. North America (lectotype here designated).
Type depository. BMNH.
Distribution. Figure 661. Nova Scotia and New Hampshire to Yukon Territory,
south to British Columbia and Montana.
Coccinella hieroglyphica mannerheimi Mulsant
Fig. 660f; Map, Fig. 661
Coccinella mannerheimii Mulsant, 1850, p. 106. — Mulsant, 1866, p. 88.— Crotch,
1874, p. 1 15 (in part); Mader, 1930, p. 160.
Coccinella tricuspis mannerheimi: Weise, 1892, p. 26.
Coccinella hieroglyphica mannerheimi: Dobzhansky, 1926, p. 24. — Brown, 1962, p.
805.
Description as for C. h. kirbyi except anterior margin of pronotum always black,
subbasal band of elytron rarely tricuspate (Fig. 660f).
This is a subarctic subspecies reported from North America for the hrst time by
Brown (1962).
Type locality. Siberia.
Type depository. Not known.
Distribution. Figure 661. MANITOBA: Churchill. NORTHWEST TERRITO-
RIES: Aklavik; Reindeer Depot. ALASKA: Matanuska.
Coccinella hieroglyphica humboldtiensis Nunenmacher
Map, Fig. 661
Coccinella humboldtiensis Nunenmacher, 1912, p. 448.
Coccinella hieroglyphica humboldtiensis: Dobzhansky, 1931, p. 37. — Hatch, 1961,
p. 180. — Brown, 1962, p. 805.
Diagnosis. Length 4.0 to 4.80 mm. Head black with 2 well separated spots or
entirely black; pronotum with anterior margin black at middle; elytron varies from
maculation extremely reduced to surface almost entirely black except for obscure
reddish streak at middle of lateral margin. Other characters as in C. h. kirbyi.
Discussion. This variable species occurs mainly in the Pacific coastal regions from
northern and east central California to Vancouver Island. The known specimens of
the dark phase are from Vancouver Island. There are 2 type specimens in the CAS,
I designate and label a male labeled “C. City, Del Norte Co. Cal. V.17.10/coird by
F. W. Nunenmacher/male sign/Coccinella humboldtiensis Nun” as the lectotype.
Type locality. Crescent City, Del Norte Co., California (lectotype here designated).
Type depository. CAS.
Distribution. Figure 661. VANCOUVER ISLAND: Courtenay; Duncan. CALI-
FORNIA: Crescent City; Mammoth; Plumas Co., Siskiyou Co. WASHINGTON:
Olympia.
Genus Propylaea Mulsant
Propy/acfl Mulsant, 1846, p. 1 52. — Mulsant, 1850, p. 212. — Mulsant, 1866, p. 150.—
Crotch, 1874b, p. 1 57.-Korschefsky, 1932, p. 530.-Iablokoff-Khnzorian, 1982,
p. 164. Type-species; Coccinella quatuordecimpunctata L., by monotypy.
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NORTH AMERICAN COCCINELLIDAE
817
Coccinellini with length 3.0 to 5.50 mm; form oval, slightly depressed. Antero-
lateral angle of clypeus produced forward. Pronotum and elytron with lateral margin
weakly explanate; epipleuron descending externally. Intercoxal process of prostemum
with distinct, narrow, lateral ridge extending nearly to apex of prostemum. Apical
margin of mesostemum arcuately notched for reception of prostemal process. Apex
of middle and hind tibiae with 2 spurs. Tarsal claw with basal, subquadrate tooth
(Fig. 662b). Postcoxal line of Nephus type (Fig. 662a). Male genitalia symmetrical.
Female genitalia with infundibulum (Fig. 66 3e).
Propyleae is an Old World genus with one species established in eastern Canada.
Propyleae quatuordecimpunctata has been intentionally released in the United States
several times (New Jersey, Oklahoma) but is not established. The origin of the
established population in Canada is unknown and was reported for the first time by
Chantal (1972). Members of this genus are primarily aphid predators with specific
host records as follows; Acyrthosiphon dirhodum (Walker), Acyrthosiphon pelargonii
(Kaltenbach), Acyrthosiphon pisum (Harris), Acyrthosiphon solani (Kaltenbach), An-
oecia corni (F.), Aphis craccivora Koch, Aphis donacis Passerini, Aphis fabae Scopoli,
Aphis gossypii Glover, Aphis nerii Boyer de Fonscolombe, Aphis pomi de Geer, Aphis
spiraephila Patch, Brachycaudus helichrysii (Kaltenbach), Brachycaudus lychnidis (L.)
Brachycaudus persicae (Passerini), Brevicoryne brassicae (L.), Dactynotus cirsii (L.),
Dactynotus jacea (L.), Dactynotus sonchi (L.), Drepanosiphum platanoidis (Schrank),
Dysaphis plantaginea (Passerini), Dysaphis reamuri (Mordvilko), Eriosoma laniger-
um (Hausmann), Eucallipterus tiliae (L.), Euceraphis punctipennis (Zetterstedt), Gly-
phina betulae (L.), Hyadaphis erysimi (Kaltenbach), Hyalopterus pruni (Geoffroy),
Macrosiphum avenae (F.), Macrosiphum euphorbiae (Thomas), Megoura vicia Buck-
ton, Microlophium evansi (Theobald), Myzus malisucta Matsumura, Myzus persicae
(Sulzer), Nasonovia lactucae (L.), Prociphilus xylostei (Degeer), Pterocallis alni (De-
geer). P. quatuordecimpunctata has also been recorded as feeding on larvae of Lema
melanopus (L.) (Chrysomelidae). Propyleae has been systematically treated by la-
blokoff-Khnzorian (1982).
Propyleae quatuordecimpunctata (L.)
Fig. 662a-f, 663a-e; Map, Fig. 665
Coccinella quatuordecimpunctata L., 1758, p. 366.
Propylea quatuordecimpunctata: Mulsant, 1846, p. 152.— Mulsant, 1866, p. 150.-
Crotch, 1874b, p. 157.— Korschefsky, 1932, p. 532. — Sasaji, 1971, p. 264.— Chan-
tal, 1972, p. 243. -Sasaji, 1975, p. 13.-Iablokoff-Khnzorian, 1982, p. 167.
Diagnosis. Length 3.50 to 5.20 mm, width 2.80 to 4.0 mm. Male head yellow
except vertex black, female head with black spot on clypeus; pronotum with large,
irregular, black area medially; elytron yellow with variable black maculation (Fig.
663a-d). Male genitalia as in Figure 662c-f. Female genitalia as in Figure 663e.
Discussion. The dorsal color pattern of this species is unlike that of any native
North American species of Coccinellini, and the combination of color pattern and
the key generic characters make it easy to recognize.
Type locality. “Suecica”.
Type depository. Type not examined.
Distribution. Figure 665. QUEBEC: Ste-Foy.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Voi. 93(1)
Fig. 662. Propyleae quatuordecimpunctata. a. Postcoxal lines, b. Tarsus, c-f. Male genitalia.
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NORTH AMERICAN COCCINELLIDAE
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Fig. 663. Propyleae quatuordecimpunctata (habitus and variation, female genitalia).
Genus Cycloneda Crotch
Cycloneda Crotch, 1871, p. 6.— Crotch, 1873, p. 371.— Crotch, 1874b, p. 162.—
Gorham, 1891, p. 58. — Casey, 1899, p. 84. — Leng, 1920, p. 216.— Korschefsky,
1932, p. 282.— Timberlake, 1943, p. 23.— Wingo, 1952, p. 24. — Mader, 1958, p.
238.-Hatch, 1961, p. 181. -J. Chapin, 1974, p. 62.-Belicek, 1976, p. 330. Type-
species; Coccinella sanguinea L., by subsequent designation of Crotch, 1874b.
Daulis Mulsani, 1850, p. 296 (not Erichson, 1842).— Crotch, 1874b, p. 162.—
Berg, 1874, p. 290.— Chapuis, 1876, p. 201. Type-species; not designated.
Coccinella {Cycloneda), Leng, 1903b, p. 202.
Coccinellini with length 3.0 to 9.0 mm; form rounded, convex (Fig. 664h). Elytron
pale, immaculate; pronotum black with pale markings (North American species).
Apex of clypeus truncate, anterolateral angle produced forward. Lateral margin of
elytron feebly explanate; epipleuron obliquely descending externally. Intercoxal pro-
cess of prostemum narrow, ridged medially, lateral ridges obsolete. Apical margin
of mesostemum truncate or barely perceptibly emarginate. Apex of middle and hind
tibia each with 2 spurs. Tarsal claw with large, subquadrate basal tooth (Fig. 664b).
Postcoxal line incomplete, of Diomus type, without oblique dividing line (Fig. 664a).
Male genitalia symmetrical. Female genitalia with infundibulum; coxal plate irreg-
ularly elongate with distinct apical stylus (Fig. 668e).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Cycloneda is very similar to Olla (see comments under Olid) but at least in the
limited North American fauna the two genera can be readily separated by the key
characters. In addition, C sanguinea and allies have a strong infundibulum in the
female genitalia; O. v- nigrum lacks an infundibulum. Cycloneda is a New World
genus with more than 50 names presently included. Most of these are neotropical
with 3 species occurring north of Mexico. Casey (1899) described C. ater from an
unlabeled specimen.. This species is not a member of the North American fauna and
is most likely a neotropical member of the genus. Timberlake (1943) expressed the
opinion that the genus Cycloneda should be restricted to C sanguinea and allied
species having immaculate elytra. Mader (1958) published a key to the species of
Cycloneda but did not examine genitalia. He did not alter the generic classification
appreciably and a complete study of this group is still needed. Members of Cycloneda
are primarily aphid predators with specific host records as follows: Acyrthosiphon
dirhodum (Walker), Acyrthosiphon pisum (Harris), Aphis gossypii Glover, Aphis nerii
Boyer de Fonscolombe, Aphis pomi Degeer, Aphis viburni Scopoli, Brevicoryne bras-
sicae (L.), Carolinaia cyperi Ainslie, Chapitophorus eleagni (del Guercio), Eriosoma
lanigerum (Hausmann), Hyadaphis erysimi (Kaltenbach), Macrosiphum avenae (F.),
Macrosiphum euphorbiae (Thomas), Myzus cerasi (F.), Myzus persicae (Sulzer),
Nearctaphis crataegifoliae (Fitch), Periphyllus negundinis (Thomas), Phorodon hu-
muli (Schrank), Rhopalosiphum maidis (Fitch), Sipha flava (Forbes), Sipha maydis
Passerini, Toxoptera aurantii (Boyer de Fonscolombe). The genus has not been tax-
onomically studied as a whole since Crotch (1874b), except for Mader’s (1958) key
to species.
Key to species of Cycloneda
1. Pronotum with pale lateral spot enclosed by black coloration (Fig. 664h) 3
- Pronotum with pale lateral spot not entirely enclosed, or with isolated lateral black
spot (Fig. 6671) 2
2(1). Elytron usually red; California and the Pacific Northwest polita Casey
- Elytron orange; eastern United States munda (Say)
3(1). Lateral border of elytron black sanguinea limbifer Casey
- Lateral border of elytron pale sanguinea sanguinea (L.)
Cycloneda sanguinea sanguinea (L.)
Fig. 664a-h; Map, Fig. 665
Coccinella sanguinea L., 1763, p. 10.
Daulis sanguinea: Mulsant, 1850, p. 326.
Cycloneda sanguinea: Crotch, 1871, p. 6. — Crotch, 1873, p. 372.— Crotch, 1874b,
p. 164. — Blatchley, 1910, p. 515. — Palmer, 1914, p. 232. — Korschefsky, 1932, p.
286. -Timberlake, 1943, p. 23.-Wingo, 1952, p. 46. -Mader, 1958, p. 241. -J.
Chapin, 1974, p. 62.
Coccinella immaculata F., 1792, p. 267.
Daulis immaculata: Mulsant, 1850, p. 327.
Cycloneda immaculata: Casey, 1899, p. 92.
Cycloneda rubripennis Casey, 1899, p. 92. — Korschefsky, 1932, p. 285 (as synonym
of mwA2(7<3). — Mader, 1958, p. 241.
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NORTH AMERICAN COCCINELLIDAE
821
Fig. 664. Cycloneda sanguinea sanguinea. a. Postcoxal lines, b. Tarsus, c-f. Male genitalia,
g. Female genitalia, h. Habitus.
Coccinella {Cycloneda) sanguinea: Leng, 1903b, p. 202.
Coccinella {Cycloneda) sanguinea var. immaculata: Leng, 1903b, p. 203.
Coccinella {Cycloneda) sanguinea var. rubripennis: Leng, 1903b, p. 203.
Diagnosis. Length 3.20 to 6.50 mm, width 2.90 to 5.10 mm. Pronotum mostly
black with lateral pale spot enclosed by black area; elytron orange to red (Fig. 664h).
Male genitalia as in Figure 664c-f. Female genitalia as in Figure 664g.
Discussion. The pronotal color pattern will distinguish C. sanguinea from the other
North American species in almost all instances. The male and female genitalia are
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 665. Distribution. Cycloneda sangidnea sanguinea (shaded, southern); C. 5. limbifer
(star); C. munda (shaded, northern); C. polita (shaded, western).
distinctive for all species and should be ehecked where external characters are doubt-
ful. This species is found from the southern United States to Argentina and varies
greatly in size and coloration. The synonymy above deals only with the geographic
area north of Mexico and Korschefsky (1932) is to be consulted for the synonymy
of C. sanguinea in the neotropieal region. Cycloneda rubripennis Casey was consid-
ered a synonym of C. munda (Say) by Korschefsky, and I agree that it is a junior
synonym, but of C. sanguinea. There are 7 types of C. rubripennis in the Casey
collection and I designate and label the first of these as the lectotype, the remainder
as paralectotypes.
Type locality. Of sanguinea, Surinam; of immaculata, “Americae Insulis” (Amer-
ican Islands); of rubripennis, San Diego, California (lectotype here designated).
Type depository. Of sanguinea and immaculata, not known; of rubripennis, USNM
(35525).
Distribution. Figure 665. North Carolina to Florida, west to southern California.
Cycloneda sanguinea limbifer Casey
Fig. 660a-f; Map, Fig. 665
Cycloneda limbifer Casey, 1899, p. 92.
Coccinella {Cycloneda) limbifer. Leng, 1903b, p. 204.
Cycloneda sanguinea ab. limbifera: Korschefsky, 1932, p. 286. — Mader, 1958, p.
241.
Cycloneda sanguinea limbifer: Chapin, 1949, p. 23. — Chapin, 1957, p. 89.
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NORTH AMERICAN COCCINELLIDAE
823
Description as for C. 5. sanguinea except lateral border of elytron black (Fig. 666f).
Male genitalia as in Figure 666a-d. Female genitalia as in Figure 666e.
This is a West Indian subspecies recorded from Key West, Florida, by Chapin
(1949). Only 2 specimens were collected and it may not be established on the main-
land. There are 4 types of C. limbifer in the Casey collection, the first of which is
designated and labeled as the lectotype and the remainder as paralectotypes.
Type locality. Egg Island, Bahama Islands (lectotype here designated).
Type depository. USNM (35526).
Distribution. Figure 665. FLORIDA; Key West.
Cycloneda munda (Say)
Fig. 667a-f; Map, Fig. 665
Coccinella munda Say, 1835, p. 202. — Crotch, 1874b, p. 107.
Daulis munda: Mulsant, 1850, p. 324.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 667. Cycloneda munda.
Cycloneda munda: Crotch, 1871, p. 6. — Casey, 1899, p. 93. — Leng, 1920, p. 216.—
Korschefsky, 1932, p. 284. — Timberlake, 1943, p. 23. — Wingo, 1952, p. 46.— J.
Chapin, 1974, p. 63.
Coccinella {Cycloneda) sanguinea var. munda: Leng, 1903b, p. 203.
Diagnosis. Length 3.70 to 5.70 mm, width 3.10 to 4.20 mm. Pronotum mostly
black with lateral pale spot not completely enclosed by black area, or with separate
black spot laterally (Fig. 6671); elytron reddish yellow. Male genitalia as in Figure
667a-d. Female genitalia as in Figure 667e.
Discussion. This species is widespread in the eastern United States and overlaps
the distribution of C. sanguinea (see comments under sanguinea).
Type locality. Type material was from various localities in North America,
Type depository. Type lost.
Distribution. Figure 665.
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NORTH AMERICAN COCCINELLIDAE
825
Cycloneda polita Casey
Fig. 668a-f; Map, Fig. 665
Cycloneda polita Casey, 1899, p. 93.— Timberlake, 1943, p. 24. — Hatch, 1961, p.
I81.-Belicek, 1976, p. 330.
Coccinella {Cycloneda) sanguinea var. polita: Leng, 1903b, p. 203.
Cycloneda munda ab. polita: Leng, 1920, p. 216. — Korschefsky, 1932, p. 285.
Cycloneda polita flava Timberlake, 1943, p. 24. New Synonymy.
Diagnosis. Length 3.50 to 6.20 mm, width 3.0 to 4.0 mm. Color as for munda
except elytron usually red (Fig. 6681). Male genitalia as in Figure 668a-d. Female
genitalia as in Figure 668e.
Discussion. This species will be confused with C. munda but the two species are
allopatric and both the male and female genitalia are distinctive for each species.
There are 8 type specimens of C. polita in the Casey collection, the first of which is
designated and labeled as the lectotype and the remainder as paralectotypes. The
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
only difference between C. polita and C. p. flava is the yellow elytral color of the
latter. The genitalia are the same. Therefore I do not recognize it as a valid subspecies
and place it as a junior synonym of polita.
Type locality. Of polita, Marin Co., peninsula north of San Francisco, California
(lectotype here designated); oi flava, Alameda, California.
Type depository. Of polita, USNM (35527); of flava, HSPA.
Distribution. Figure 655.
Genus Olla Casey
Olla Casey, 1899, p. 93. — Leng, 1920, p. 216. — Korschefsky, 1932, p. 288.— Tim-
berlake, 1943, p. 55.— Wingo, 1952, p. 24.— J. Chapin, 1974, p. 64. — Belicek,
1976, p. 329. Type-species; Coccinella abdominalis Say, by subsequent designation
of Korschefsky, 1932.
Coccinella {Olla)\ Leng, 1903b, p. 197.
Coccinellini with length 3.0 to 9.0 mm; form rounded, convex (Fig. 669). Color
pattern polymorphic in the North American species. Dorsal surface mostly polished
between punctures, fine alutaceous sculpture present. Apex of clypeus truncate, an-
terolateral angle produced forward. Lateral margin of elytron feebly explanate; epi-
pleuron strongly, abruptly descending externally. Intercoxal process of prostemum
narrow, medially flattened, with broad lateral ridges parallel or slightly convergent
anteriorly. Apical margin of mesostemum broadly, feebly emarginate for reception
of prostemal process. Apex of middle and hind tibia each with 2 spurs. Tarsal claw
with large, subquadrate basal tooth (Fig. 669b). Postcoxal line incomplete, with
oblique dividing line (Fig. 669a). Male genitalia symmetrical. Female genitalia lacking
infundibulum; coxal plate irregularly elongate with distinct apical stylus (Fig. 670).
Olla may not be distinct from Cycloneda, but a comprehensive study of all the
included species of both genera is needed to decide this. In any event, the two genera
as represented in North America are readily separated on the basis of color pattern.
Olla is a New World genus with approximately 6 species names, mainly in the
Neotropical region. Members of Olla are aphid predators with specific host records
as follows: Aphis pomi Degeer, Chromaphis juglandica (Kaltenbach), Monellia car-
yella (Fitch), Monelliopsis californica, (Essig), Monelliopsis caryae (Monell), Phoro-
don humuli (Schrank). Olla has not been taxonomically treated as a whole, but was
discussed in part by Timberlake (1943).
Olla V- nigrum (Mulsant)
Fig. 669a-i; Map, Fig. 671
Harmonia v-nigrum Mulsant, 1866, p. 64.
Coccinella abdominalis Say, 1824, p. 95 (not Coccinella abdominalis Thunberg,
1794).
Cycloneda sayi Crotch, 1871, p. 6 (new name for abdominalis Say).
Cycloneda oculata var. abdominalis: Crotch, 1873, p. 372.
Cycloneda abdominalis: Crotch, 1874b, p. 163.— Gorham, 1892, p. 172.
Olla abdominalis: Casey, 1899, p. 93. — Blatchley, 1910, p. 514. — Palmer, 1914, p.
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NORTH AMERICAN COCCINELLIDAE
827
Fig. 669. Olla v-nigrum. a. Postcoxal lines, b. Tarsus, c-f. Male genitalia, g-i. Habitus and
variations.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 670. Olla v-nigrum (female genitalia).
232.-Leng, 1920, p. 216.-Korschefsky, 1932, p. 288.-Wingo, 1952, p. 46.-J.
Chapin, 1974, p. 64.
Olla plagiata Casey, 1899, p. 94. — Belicek, 1976, p. 329.
Olla sobrina Casey, 1899, p. 94. — Blatchley, 1918, p. 419. — Belicek, 1976, p. 329.
Olla fenestralis Casey, 1899, p. 95. — Belicek, 1976, p. 329.
Coccinella {Olla) oculata var. plagiata: Leng, 1903b, p. 204.
Coccinella {Olid) oculata var. sobrina: Leng, 1903b, p. 204.
Coccinella {Olla) oculata Yds. fenestralis: Leng, 1903b, p. 204.
Coccinella {Olla) abdominalis: Leng, 1903b, p. 205.
Olla minuta Casey, 1908, p. 406. — Casey, 1924, p. 1 57. — Belicek, 1976, p. 329.
Olla semilunaris Johnson, 1910, p. 66.
Olla abdominalis arizonae Casey, 1924, p. 158. — Korschefsky, 1932, p. 289.— Be-
licek, 1976, p. 329.
Olla abdominalis ab. minuta: Korschefsky, 1932, p. 289.
Olla abdominalis ab. plagiata: Korschefsky, 1932, p. 289.
Olla abdominalis ab. sobrina: Korschefsky, 1932, p. 289.
Olla abdominalis ab. v-nigrum: Korschefsky, 1932, p. 289.
Olla v-nigrum: Timberlake, 1943, p. 24. — Belicek, 1976, p. 329.
Olla v-nigrum var. plagiata: Timberlake, 1943, p. 24.
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NORTH AMERICAN COCCINELLIDAE
829
Diagnosis. Length 3.70 to 6.10 mm, width 3.0 to 5.0 mm. Dorsal color pattern
with background black, maculae pale, or pale yellow with black maculae (Fig. 669g-
i). Male genitalia as in Figure 669c-f. Female genitalia as in Figure 670.
Discussion. This species has two basic color variants that are strikingly different
and I have seen little evidence of intergradation between the variants. The dark form
occurs mostly in the United States. The pale form is very widespread occurring from
southern Canada to Central America and on the islands of Guam, Hawaii, and
Midway, where it has been introduced for biocontrol purposes. Several names have
been proposed for this species, most of which were recently synonymized by Belicek
(1976). The name oculata F. has been used for the dark form of v-nigrum by many
authors and is included in the Korschefsky catalog (1932). However, the description
of oculata is that of a species which does not occur in North America; therefore the
name must be removed from North American literature. The Casey names O. minuta
and O.fenestralis are based on single specimens which must be considered holotypes.
The other Casey names are based on more than one specimen as follows: O. a.
arizonae 2, O. plagiata 10; and sobrina, 3. In each instance the first specimen is
designated and labeled as the lectotytpe and the remainder as paralectotype(s). One
type specimen (female) of v-nigrum labeled “Oaxaca/Type/22 13(green paper)/Mex-
ico, Salle Coll./B.C.A. Col., VII, Cycloneda abdominalis Say/Harmonia v-nigrum
Muls Salle. Type” has been located and here designated and labeled as the lectotype.
Type locality. Of v-nigrum, Oaxaca, Mexico (lectotype here designated); of abdom-
inalis, “Arkansa”; of plagiata, Brownsville, Texas (lectotype here designated); of
sobrina, Florida (lectotype here designated); of fenestralis. Las Vegas, New Mexico;
of minuta, Brownsville, Texas; of semilunaris, Arizona and Texas; of arizonae, Tuc-
son, Arizona (lectotype here designated).
Type depository. Of v-nigrum, BMNH; of abdominalis, type lost; of plagiata (35529),
sobrina (35530), of fenestralis (35531), minuta (35532), and arizonae (35533), USNM;
of semilunaris, no types designated.
Distribution. Figure 671. Southeastern Canada to Florida, west to southern British
Columbia and southern California.
Genus Coelophora
Coelophora Mulsant, 1850, p. 390. — Mulsant, 1866, p. 257.— Crotch, 1874b, p.
148. — Korschefsky, 1932, p. 290. — Timberlake, 1943, p. 31.— Chapin, 1965b, p.
214. — lablokoff-Khnzorian, 1982, p. 265. Type species; Coccinella inaequalis F.,
by subsequent designation of Crotch, 1874b.
Coccinellini with length 3.70 to 9.0 mm; form rounded, convex. Elytron yellow
with black markings; pronotum yellow with base black (North American species).
Apex of clypeus truncate, anterolateral angle produced forward. Lateral margin of
elytron feebly explanate; epipleuron abruptly, strongly descending externally. Inter-
coxal process of prostemum broad, with strong lateral ridges. Apical margin of
mesostemum truncate. Apex of middle and hind tibia each with 2 spurs. Tarsal claw
with large, subquadrate basal tooth (Fig. 672b). Postcoxal line incomplete, of Nephus
type (Fig. 672a). Male genitalia symmetrical. Female genitalia without infundibulum;
coxal plate with lateral projection and apical stylus (Fig. 6721).
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Coelophora is similar in appearance to Cycloneda and Olla, but the dorsal color
pattern of the only North American representative of this genus is very different from
that of either Olla or Cycloneda. Coelophora is an Australian and Oriental genus,
one member of which, C. inaequalis, is possibly established in Florida. This was
apparently not the result of an intentional introduction, and the source of the stock
is not known. Members of Coelophora are mostly aphid predators, but specific host
records are scarce. It is known to prey on the following hosts; Aphis craccivora Koch,
Aphis gossyppii Glover, Aphis nerii Boyer de Fonscolombe, Hyadaphis erysimi (Kal-
tenbach), Melanaphis sacchari (Zehntner), Myzus persicae (Sulzer), Neophyllaphis
araucariae Takahashi, Rhopalosiphum rnaidis (Fitch), Sipha flava (Forbes), Thora-
caphis fici (Takahashi), Toxoptera aurantii (Boyer de Fonscolombe), and Toxoptera
citricidus (Kirkaldy). Chazeau (1981) reported Coelophora quadrivitta Fauvel as a
predator on the coccid Coccus viridis Green in New Caledonia. The genus was treated
briefly by lablokoff-Klinzorian (1981).
Coelophora inaequalis (F.)
Fig. 672a-g; Map, Fig. 673
Coccinella inaequalis F., 1775, p. 80.
Coelophora inaequalis Mulsant, 1850, p. 404. — Mulsant, 1866, p. 266.— Crotch,
1874b, p. 153. — Korschefsky, 1932, p. 292. — Timberlake, 1943, p. 31. — Chapin,
1965b, p. 215. — Leeper, 1976, p. 288. — lablokoff-Khnzorian, 1982, p. 266.
Diagnosis. Length 3.70 to 5.20 mm, width 3.50 to 4.30 mm. Pronotum yellow,
base with irregular black area, elytron yellow with 4 or 5 black maculae (Fig. 672g),
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NORTH AMERICAN COCCINELLIDAE
831
pattern variable. Male genitalia as in Figure 672c-e. Female genitalia as in Figure
672f.
Discussion. The Florida specimens of C. inaequalis correspond to the Philippine
specimens described by Timberlake (1943) as C. inaequalis comperei. I prefer not to
use subspecific definitions in this group at present because much more extensive
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 673. Distribution. Coelophora inaequalis (triangle); Hannonia dimidiata (dot); Neo-
harmonia venusta venusta (shaded); N. v. ampla (star).
taxonomic research is necessary to accurately define specific and subspecific limits.
The synonymy given here is limited and lablokoff-Khnzorian ( 1 982) is to be consulted
for the complete history. The first North American records of this species are spec-
imens collected at Clewiston, Florida, 1939. In 1978, 1979, and 1982 specimens
were again collected in Florida, but whether it is firmly established or not is open to
question.
Type locality, “nova Hollandia”.
Type depository. BMNH (not examined).
Distribution. Figure 673. FLORIDA; Broward Co., Fort Lauderdale; Plantation.
Genus Harmonia Mulsant
Harmonia Mulsant, 1850, p. 75. — Mulsant, 1866, p. 55. — Crotch, 1873, p. 373.—
Crotch, 1874b, p. 105.-Mader, 1926,p. 19.-Timberlake, 1943,p. 17. -lablokoff-
Khnzorian, 1982, p. 456. Type-species, Coccinella marginepunctata Schaller, by
subsequent designation of Timberlake, 1943.
Coccinella {Harmonia): Korschefsky, 1932, p. 439.
Lc/5 Mulsant, 1850, p. 241. — Mulsant, 1866, p. 174. — Crotch, 1874b, p. 119.— Chap-
uis, 1876, p. 200. — Sicard, 1909, p. 68. — Korschefsky, 1932, p. 273. lablokoff-
Khnzorian, 1982, p. 456. Type-species; Coccinella dimidiata F., by subsequent
designation of Crotch, 1874.
Coccinellini with length 4.60 to 11.0 mm; form rounded, convex. Color variable
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NORTH AMERICAN COCCINELLIDAE
833
a
b
Fig. 674. Harmonia dimidiata. a. Postcoxal lines, b. Tarsus.
but dorsum pale with dark maculae. Apex of clypeus truncate, anterolateral angle
produced forward. Lateral margin of elytron weakly to strongly explanate; epipleuron
abruptly descending externally. Intercoxal process of prostemum broad, flat, with
fine Carina on each side. Apical margin of mesostemum weakly emarginate medially.
Apex of middle and hind tibia each without spurs. Tarsal claw with large, subquadrate
basal tooth (Fig. 674b). Postcoxal line incomplete, of Nephus type, with irregular,
oblique dividing line (Fig. 674a). Male genitalia symmetrical. Female genitalia with
infundibulum; coxal plate irregularly elongate with distinct apical stylus (Fig. 676b).
Harmonia is an Oriental and Australian genus of approximately 1 7 species, one
of which, Harmonia dimidiata (F.) {quindecimspilota Hope), was introduced in 1924
into California (not established) and then from California into Florida where it is
established, for biological control of aphids. The large size and dorsal maculation of
this representative of the genus easily distinguishes it from all other North American
Coccinellini. Harmonia axyridis (Pallas) has been released in Delaware, Georgia, and
Washington, but is apparently not established.
Members of this genus have been recorded as mainly aphid predators, but they
are also predators of Psyllidae and scale insects. Specific host records are listed as
follows. Aphids; Acyrthosiphum pisum (Harris), Aphis gossypii Glover, Aphi nerii
Boyer de Fonscolombe, Aphis pomi de Geer, Aphis punicae Passerini, Amphorophora
oleraceae (Van der Goot), Cinara laricicola (Matsumura), Cinara todocola (Ihouye),
Cryptosiphum gallarum (Kaltenbach), Eriosoma lanigerum (Hausmann), Hyadaphis
erysimi (Kaltenbach), Hyalopterus pruni (Geoffroy), Macrosiphum ibarae Matsu-
mura, Megoura crassicauda Mordvilko, Melanaphis sacchari (Zehntner), Myzus mal-
isucta Matsumura, Myzus persicae (Sulzer), Neophyllaphis podocarpi Takahashi, Nip-
polachnus piri Matsumura, Periphyllus californicus David, Rhopalosiphum padi (L.),
Toxoptera piricola Matsumura. Scale insects; Chrysomphalus aonidum (L.), Dacty-
lopius sp., Icerya purchasi Maskell, ""Lecanium"' sp., Phenacoccus pergandei Cock-
erell, Pulvinaria sp., Saccharicoccus sacchari (Cockerell). Psyllidae; Anomoneura mori
Schwarz. The genus has been treated in part by lablokoff-Khnzorian (1982) who has
synonymized Leis with Harmonia.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Harmonia dimidiata (F.)
Fig. 675a-c, 676 a, b; Map, Fig. 673
Coccinella dimidiata F., 1781, p. 94.
Leis dimidiata: Mulsant, 1850, p. 242. — Mulsant, 1866, p. 174.— Crotch, 1874b, p.
I19.-Mader, 1934, p. 307.-Korschefsky, 1932, p. 273.
Leis dimidiata ab. quindecimmaculata: Korschefsky, 1932, p. 274.
Coccinella 15-maculata Hope, 1831, p. 30.
Leis quindecimmaculata: Mulsant, 1850, p. 246.
Coccinella 15-spilota Hope, 1831, p. 30.
Leis quindecimspilota: Mulsant, 1850, p. 248. — Sicard, 1912, p. 500.— Korschefsky,
1932, p. 274.
Harmonia dimidiata: lablokoff-Khnzorian, 1982, p. 484.
Diagnosis. Length 7.40 to 10.0 mm, width 6.20 to 9.0 mm. Dorsum reddish yellow
with black maculae as in Figure 676a. Male genitalia as in Figure 675a-c. Female
genitalia as in Figure 676b.
Discussion. The status of the names included as subspecies or synonyms of H.
dimidiata is doubtful at best, and I’ve not attempted to examine the types of these
names. Korschefsky (1932) and lablokoff-Khnzorian (1982) should be consulted for
synonymy.
Type locality. Nepal.
Type depository. Type not examined.
Distribution. Figure 673. FLORIDA: Aubumdale; Ocoee; Orlando; Winter Garden;
Winter Park.
Genus Neoharmonia Crotch
Neoharmonia Crotch, 1871, p. 2. — Rye, 1873, p. 329.— Timberlake, 1943, p. 20.—
Gordon, 1974b, p. 166.— J. Chapin, 1974, p. 60. Type-species: Harmonia viridi-
pennis Mulsant, by subsequent designation of Rye, 1873.
Agrabia Casey, 1899, p. 87.— Leng, 1903b, p. 196. — Korschefsky, 1932, p. 438.—
Gordon, 1974b, p. 166. Type-species; Harmonia cyanoptera Mulsant, by mono-
typy.
Neoharmonia Casey, 1899, p. 90 (not Neoharmonia Crotch). — Leng, 1920, p. 216.—
Timberlake, 1943, p. 20.— Wingo, 1952, p. 23. Type-species; Coccinella venusta
Melsheimer, by subsequent designation of Timberlake, 1943.
Harmoniaspis Casey, 1908, p. 404. — Korschefsky, 1932, p. 575. — Blackwelder, 1945,
p. 455.— Gordon, 1974b, p. 166. Type-species: Harmonia ampla Mulsant, by
subsequent designation of Gordon, 1974b.
Coccinella {Neoharmonia): Leng, 1903b, p. 201. — Korschefsky, 1932, p. 440.
Coccinellini with length 4.50 to 7.00 mm; form elongate, oval, depressed. An-
terolateral angle of clypeus produced forward. Pronotum and elytron with lateral
margin weakly explanate, usually semitransparent or at least pale in color; epipleuron
descending externally. Intercoxal process of prostemum with narrow lateral ridge
extending anteriorly as far as anterior margin of coxa, median area nearly flat. Apical
margin of mesosternum ridged, arcuately emarginate medially for reception of pros-
ternal process. Apex of middle and hind tibia each without spurs. Tarsal claw with
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NORTH AMERICAN COCCINELLIDAE
835
basal, subquadrate tooth. Postcoxal line incomplete, of Scymnobius type, but with
median intersecting line (Fig. 677a). Male genitalia symmetrical. Female genitalia
lacking infundibulum, base of spermathecal capsule terminating in well developed
ramus (Fig. 677e).
Neoharmonia is easily recognized by the key characters; in addition, the dorsal
color pattern, although variable, is usually distinct from that of other North American
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 676. Harmonia dimidiata (habitus and female genitalia).
Coccinellini. Neoharmonia is a genus restricted to the Western Hemisphere with N.
venusta venusta and N. venusta ampla occurring north of Mexico. The total number
of species in the genus is a matter of conjecture at present, but there are probably
less than 5. Members of Neoharmonia have been presumed to be predators of aphids
and possibly scale insects, but specific host data has been lacking. Whitehead and
Duffield (1982) reported N. venusta venusta as a major predator of the willow leaf
beetle, Plagiodera versicolora (Laicharteg). The genus north of Mexico was reviewed
by Gordon (1974b).
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NORTH AMERICAN COCCINELLIDAE
837
Key to subspecies of Neoharmonia venusta (Melsheimer)
1 . Color of pronotum either mostly black (Fig. 677d), or pale with black spots as in Figure
677b; eastern U.S. to eastern Texas venusta venusta (Melsheimer)
- Color of pronotum either entirely pale or pale with red or brown spots as in Figure
678e-g; southwestern U.S. and northern Mexico venusta ampla (Mulsant)
Neoharmonia venusta venusta (Melsheimer)
Fig. 677a-e, 678; Map, Fig. 673
Coccinella venusta Melsheimer, 1847, p. 178.— Crotch, 1874b, p. 108.— Gorham,
1891, p. 156. — Blackwelder, 1945, p. 454.
Harmonia notulata Mulsant, 1850, p. 83.
Harmonia venusta: Mulsant, 1856, p. 141. — Mulsant, 1866, p. 61.
Neoharmonia venusta: Crotch, 1871,. p. 2. — Casey, 1899, p. 71. — Leng, 1920, p.
216.-Wingo, 1952, p. 45.
Neoharmonia notulata: Crotch, 1871, p. 3.— Casey, 1899, p. 91.
Coccinella notulata: Crotch, 1874b, p. 108.
Neoharmonia venusta var. dissimila Blatchley, 1914, p. 65. — Leng, 1920, p. 216.—
Gordon, 1974b, p. 169.
Neoharmonia venusta var. fattigi Blatchley, 1920, p. 43. — Gordon, 1974b, p. 169.
Neoharmonia venusta var. centralis Casey, 1924, p. 127.— Gordon, 1974b, p. 169.
Coccinella {Neoharmonia) notulata: Leng, 1903b, p. 202. — Korschefsky, 1932, p.
514.
Coccinella {Neoharmonia) venusta: Leng, 1903b, p. 202. — Korschefsky, 1932, p. 514.
Coccinella {Neoharmonia) venusta ab. dissimila: Korschefsky, 1932, p. 541.
Coccinella {Neoharmonia) venusta ab.fattigi: Korschefsky, 1932, p. 541.
Coccinella {Neoharmonia) venusta ab. centralis: Korschefsky, 1932, p. 541.
Neoharmonia venusta venusta: Gordon, 1974b, p. 169.— J. Chapin, 1974, p. 60.
Color pattern variable as in Figure 677b-d. Male genitalia as in Figure 678a-d.
Female genitalia as in Figure 677e. For a more detailed discussion see Gordon
(1974b).
There are 3 types of centralis in the Casey collection, the first of these is here
designated and labeled as the lectotype, the other 2 as paralectotypes.
Type locality. Of venusta, Pennsylvania; of notulata, Louisiana; of dissimila. Lake
Okeechobee, Florida; of fattigi, Pahokee, Florida; of centralis, Illinois, 20 miles south
of St. Louis (lectotype here designated).
Type depository. Of venusta, not located; of notulata, not located; of dissimila and
fattigi, PU; of centralis, USNM (35523).
Distribution. Figure 673. Maine to Florida, west to Michigan, Nebraska and eastern
Texas.
Neoharmonia venusta ampla (Mulsant)
Fig. 678e-g; Map, Fig. 673
Harmonia ampla Mulsant, 1850, p. 81. — Mulsant, 1866, p. 61.
Harmonia cyanoptera MuXsanX, 1850, p. 82. — Mulsant, 1866, p. 61.— Gordon, 1974b,
p. 169.
Harmonia viridipennis Mulsant, 1866, p. 60.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 611 . Neoharmonia venusta venusta. a. Postcoxal lines, b-d. Habitus and variations, e.
Female genitalia.
Harmonia soularyi Mulsant, 1866, p. 63.
Coccinella cyanoptera: Crotch, 1873, p. 373. — Crotch, 1874b, p. 108.— Gorham,
1891, p. 155.
Coccinella viridipennis: Crotch, 1874b, p. 108.
Coccinella soularyi: Crotch, 1874b, p. 109. — Gorham, 1891, p. 156.
Coccinella ampla: Crotch, 1874b, p. 108.— Gorham, 1891, p. 156. — Blackwelder,
1945, p. 454.
Agrabia cyanoptera: Casey, 1899, p. 87. — Leng, 1903, p. 196. — Leng, 1920, p. 217.—
Korschefsky, 1932, p. 438. — Blackwelder, 1945, p. 454.
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NORTH AMERICAN COCCINELLIDAE
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Neharmonia cyanoptera: Timberlake, 1943, p. 20.
Neoharmonia ampla: Leng, 1920, p. 216.
Agrabia cyanoptera ab. viridipennis: Korschefsky, 1932, p. 438.
Coccinella {Neoharmonia) ampla: Leng, 1903b, p. 202.— Korschefsky, 1932, p. 509.
Coccinella {Neoharmonia) soularyi: Korschefsky, 1932, p. 509.
Coccinella ampla var. rufa Nunenmacher, 1944, p. 146.— Gordon, 1974b, p. 169.
Neoharmonia venusta ampla: Gordon, 1974b, p. 169.
Color pattern variable as in Figure 678e-g. Male and female genitalia identical to
those of A^. V. venusta (Pig. 678a-d). For more detailed discussion see Gordon (1974b).
One type specimen of viridipennis (female, lacking head and thorax) labeled “Type/
Harmonia viridipennis/2209(green paper)/Mexico, Salle Coll./B.C. A., Col., VII. Coc-
cinella cyanoptera m/Harmonia viridipennis Muls Salle Type” has been located and
here designated and labeled as the lectotype.
Type locality. Of ampla, Mexico; of cyanoptera, Mexico; of soularyi, Playa Vicente,
Mexico; of viridipennis, Mexico (lectotype here designated); of rufa, Oaxaca, Mexico.
Type depository. Oi ampla, cyanoptera, and soularyi, UCCC; of viridipennis, BMNH;
of rufa, CAS.
Distribution. Figure 673. ARIZONA: Clemenceau; Douglas; Globe; Huachucha
Mts.; Cochise Co.; Palmerlee; San Bernardino Ranch; Sabino Canyon. NEW MEX-
ICO: Albuquerque; Grant Co.; Santa Rosa. TEXAS: Davis Mts.; Brownsville; Pres-
idio.
Genus Aphidecta Weise
AphidectaV^tiSQ, 1899b, p. 375 (emendation). — Korschefsky, 1932, p. 373.— lablo-
koff-Khnzorian, 1982, p. 307. Type-species; Coccinella obliterata L., by monotypy.
Aphideita Weise, 1893, p. 107 (incorrect spelling).
Coccinellini with length 3.60 to 5.60 mm; form elongate, somewhat flattened.
Dorsal color variable but usually yellow with suffused darkened areas. Apex of clypeus
broadly, feebly emarginate, anterolateral angle produced forward. Lateral margin of
elytron feebly explanate; epipleuron flat. Intercoxal process of prostemum feebly
convex, slightly ridged laterally, without carina. Apical margin of mesosternum trun-
cate. Apex of femur extending beyond margin of elytron. Apex of middle and hind
tibia each without spurs. Tarsal claw with large, subquadrate basal tooth (Fig. 679b).
Postcoxal line on first abdominal sternum complete (Fig. 679a). Male genitalia sym-
metrical. Female genitalia with small infundibulum; coxal plate rectangular in apical
half, stylus distinct (Fig. 680c).
Aphidecta is a palearctic genus containing one species, A. obliterata (L.), according
to the current classification. This species has been introduced into Canada and North
Carolina for control of Adelges piceae (Ratzeburg) (balsam woolly adelgid).
It is apparently established in North America only on Mt. Mitchell in North
Carolina (Amman, 1966) in spite of attempts to establish it in various other places
in Canada and the United States. The only genus in North America with which
Aphidecta might be confused is Mulsantina, but Mulsantina has the postcoxal line
incomplete, in Aphidecta the postcoxal line is complete, and of the Pullus type. Aphid
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 678. Neoharmonia venusta ampla.
and adelgid species listed as hosts of A. obliterata are Adelges piceae (Ratzeburg),
Cinem pinicornus Hartig, Elatobium abietinum (Walker), Pineus pini (Gmelin).
Aphidecta obliterata (L.)
Fig. 679a-f, 680a-c; Map, Fig. 681
Coccinella obliterata L., 1758, p. 367.
Adalia obliterata: Mulsant, 1850, p. 49. — Crotch, 1874b, p. 99.
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Fig. 679. Aphidecta obliterata. a. Postcoxal lines, b. Claw. c-f. Male genitalia.
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Fig. 680. Aphidecta obliterata (habitus and variation, female genitalia).
Aphideita obliterata: Weise, 1893, p. 106.
Aphidecta obliterata: Weise, 1899b, p. 375. — Korschefsky, 1932, p. 373.
Color pattern as in Figure 680a, b. Male genitalia as in Figure 679c-f. Female
genitalia as in Figure 680c.
There are several names listed by Korschefsky (1932) as synonyms or aberrations
of A. obliterata, refer to Korschefsky for the complete synonymy.
Type locality. “Europa”.
Type depository. Type not examined.
Distribution. Figure 681. NORTH CAROLINA: Mt. Mitchell.
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NORTH AMERICAN COCCINELLIDAE
843
Fig. 681. Distribution. Aphidecta obliterata (triangle); Mulsantina luteodorsa (dot); Mul-
santina cyathigera (star).
Genus Mulsantina Weise
Mulsantina 1906, p. 34.— Korschefsky, 1932, p. 564.— Timberlake, 1943, p.
54.-Hatch, 1961, p. 183. -J. Chapin, 1974, p. 65.-Belicek, 1976, p. 351.
C/mMulsant, 1850, p. 208. — Mulsant, 1866, p. 148.— Crotch, 1871, p. 5.— Crotch,
1874b, p. 142.-Gorham, 1892, p. 168. -Casey, 1899, p. 84.-Leng, 1920, p.
217.— Wingo, 1952, p. 23 (not C/m Mulsant, 1850, nor C/m Guerin, 1832). Type-
species; Cleis mirifica Mulsant, by subsequent designation of Crotch, 1874b.
Pseudocleis C?i?,Qy , 1908, p. 406.— Leng, 1920, p. 217.— Korschefsky, 1932, p. 564.—
Timberlake, 1943, p. 19. Type-species: Coccinella picta Randall, by original des-
ignation.
Coccinellini with length 3.15 to 5.31 mm. Form elongate to oval, somewhat flat-
tened dorsoventrally. Dorsal color yellow with black or brown maculation. Apex of
clypeus truncate, anterolateral angle feebly produced forward. Lateral margins of
pronotum and elytron feebly explanate; epipleuron slightly descending externally.
Intercoxal process of prostemum narrow, convex, without lateral carina. Apical
margin of mesostemum weakly emarginate. Apex of femur barely extending beyond
margin of elytron. Apex of middle and hind tibia each without spurs. Tarsal claw
with small, basal, subquadrate tooth (Fig. 682b). Postcoxal line on first abdominal
sternum incomplete (Fig. 682a). Male genitalia symmetrical with extremely long
flagellum. Female genitalia without infundibulum, sperm duct extremely long in most
species (Fig. 683e).
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Mulsantina is a New World genus of 9 species, 4 of which occur north of Mexico.
The combination of incomplete postcoxal line and absence of apical spurs on the
middle and hind tibiae will distinguish Mulsantina from other genera of Coccinellini.
Mulsantina is a replacement name for Cleis. Belicek (1976) stated that Pseudocleis
Casey was synonymized by Timberlake ( 1 943), but the synonymy was first established
by Leng (1920). The preferred prey of members of this genus is uncertain. Both scale
insects and aphids have been implicated but I have not seen the results of any
definitive studies. Hosts recorded in the literature are listed below. Adelgidae; Chermes
sp., Adelges cooleyi (Gillette), Adelges piceae (Ratzeburg), Pineus strobi (Hartig).
Aphididae; Dilachnus spp., Eulachnus rileyi (Williams), Mindarus abietinus Kock,
Schizolachnus piniradiatae (Davidson). Scale insects; Chionaspis pinifoliae (Fitch),
Lecanium sp., Matsucoccus resinosae Bean and Godwin, Saissetia oleae (Olivier),
Coccus hesperidum L. This genus has been recently revised by J. Chapin (in press).
Key to species of Mulsantina
1. Elytron with 4 to 6 discrete brown spots (Fig. 6880; southern Arizona
cyathigera (Gorham)
- Elytron variably marked or immaculate, but never with 4 to 6 discrete spots; widely
distributed 2
2(1). Pronotum with slightly curved black vitta on each side of middle (Fig. 6870; elytron
immaculate luteodorsa J. Chapin
Pronotum not as described above; elytron rarely immaculate 3
3(2). Pronotum with median, black, M-shaped mark and lateral spot attached to M-shaped
mark or broken into component spots (Fig. 682d); elytron variable but rarely with
median vitta, transverse vitta usually joined across suture picta (Randall)
Pronotum usually with median, black, M-shaped mark broken into several irregular
spots with dash above each lateral leg near anterior pronotal margin (Fig. 6851);
elytron with median vitta, and spot near vitta near middle plus one behind middle
near anterior margin hudsonica (Casey)
Mulsantina picta (Randall)
Fig. 682a-e, 683a-e; Map, Fig. 684
Coccinella picta Randall, 1838, p. 51. — LeConte, 1850, p. 238. — LeConte, 1852, p.
131. -Crotch, 1874b, p. 105. -Gorham, 1891, p. 154.
Harmonia picta: Mulsant, 1850, p. 1017.— Mulsant, 1856, p. 141. — Mulsant, 1866,
p. 65. -Crotch, 1873, p. 373. -Wickham, 1894, p. 303. -Leng, 1903b, p. 205.
Neoharmonia picta: Crotch, 1871, p. 2.
Cleis picta: Casey, 1899, p. 95. — Bowditch, 1902, p. 206. — Schaeffer, 1905, p. 143.—
Johnson, 1910, p. 72.-Leng, 1920, p. 217.-Stehr, 1930, p. 40.-Wingo, 1952,
p. 46.
Pseudocleis picta: Casey, 1908, p. 406.
Mulsantina picta: Korschefsky, 1932, p. 565.— Timberlake, 1943, p. 19.— Black-
welder, 1945, p. 453. — Hatch, 1961, p. 183.— J. Chapin, 1974, p. 66. — Belicek,
1976, p. 350.-Larochelle, 1979, p. 55, 81.
Coccinella concinnata Melsheimer, 1847, p. 177.
Harmonia contexta Mulsant, 1850, p. 87, 1017.
Coccinella picta var. impictipennis Weise, 1895, p. 125.
Mulsantina picta ab. impictipennis: Korschefsky, 1932, p. 565.
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NORTH AMERICAN COCCINELLIDAE
845
a
b
Fig. 682. Mulsantina picta. a. Postcoxal lines, b. Tarsus, c-e. Habitus and variation.
Cleis picta var. blanchardi Johnson, 1910, p. 72.
Cleis mmor Casey, 1899, p. 95.
Cleis picta var. minor. Bowditch, 1902, p. 206. — Schaeffer, 1905, p. 143.— Johnson,
1910, p. 72.-Leng, 1920, p. 217.
Harmonia picta var. minor. Leng, 1903b, p. 205.
Cleis picta minor. Dobzhansky, 1935, p. 335. — Malkin, 1943, p. 197.
Mulsantina picta ab. minor. Korschefsky, 1932, p. 565. — Blackwelder, 1945, p. 453.
Mulsantina picta minor. Timberlake, 1943, p. 19. — Hatch, 1961, p. 183.
Cleis picta nubilata Casey, 1924, p. 158.
Mulsantina picta nubilata: Korschefsky, 1932, p. 565.
Mulsantina picta ab. nubilata: Blackwelder, 1945, p. 453.
Cleis concolor. Gaines, 1933, p. 263 (misidentihcation).
Diagnosis. Length 3.32 to 5.31 mm, width 2.24 to 3.98 mm. Color yellow except
head with black spot on each side of clypeus, spot narrowly connected to black vertex;
pronotum with black, median, M-shaped mark, lateral spot connected to M-shaped
mark; elytron extremely variable from heavily maculate to immaculate (Fig. 682c-
e). Male genitalia as in Figure 683a-c. Female genitalia as in Figure 683e.
Discussion. The elytral color pattern is extremely variable, but the pronotal mark-
ings are quite constant and afford an easy means of recognition.
Type locality. Of picta, Chelsea Beach, Massachusetts; of concinnata, Pennsylvania;
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 683. Mulsantina picta.
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NORTH AMERICAN COCCINELLIDAE
847
Fig. 684. Distribution. Mulsantina picta.
of contexta, “Mexique”; of impictipennis and blanchardi, not given; of minor, Sis-
kiyou Co., California; of nubilata, Mexico.
Type depository. Of picta and concinnata, types not located; of contexta, UCCC;
of impictipennis and blanchardi, types not designated; of minor (35534) and nubilata
(35535) USNM.
Distribution. Figure 684. New Brunswick to Florida, west to Yukon Territory and
California.
Mulsantina hudsonica (Casey)
Fig. 685a-f; Map, Fig. 686
Cleis hudsonica Casey, 1899, p. 95. — Schaeffer, 1905, p. Leng, 1920, p. 217.
Cleis picta var. hudsonica: Leng, 1903b, p. 206.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 685. Mulsantina hudsonica.
Pseudocleis hudsonica: Casey, 1908, p. 406.
Mulsantina hudsonica: Korschefsky, 1932, p. 564. — Hatch, 1961, p. 183.— Belicek,
1976, p. 351.
Diagnosis. Length 3.49 to 4.98 mm, width 3.0 to 3.60 mm. Color yellow except
head with 2 interrupted black lines on face connected to black vertex; pronotum
typically with irregular, black spots medially forming M-shaped design, irregular
black blotch laterally, markings variable; elytron with median brown vitta and 2
black spots, one medially, often connected to vitta, one near lateral margin in apical
*/2 (Fig. 6850- Male genitalia as in Figure 685a-d. Female genitalia as in Figure 685e.
Discussion. The dorsal color pattern is variable, but the vitta on the elytron is
constant as is the broken M-shaped mark on the pronotum. This species was described
from a unique specimen (holotype).
Type locality. “H. B.” (Hudson Bay).
Type depository. USNM (35536).
Distribution. Figure 686. Labrador to North Carolina, west to British Columbia.
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Mulsantina luteodorsa J. Chapin
Fig. 687a-f; Map, Fig. 681
Mulsantina luteodorsa J. Chapin, 1973, p. 1073.— J. Chapin, 1974, p. 67.
Diagnosis. Length 3.80 to 4.60 mm, width 2.99 to 3.40 mm. Form rounded,
somewhat elongate. Male head yellow except vertex black, female head yellow with
vertex and clypeus black or brown; pronotum yellow with slightly curved black vitta
on each side of middle, vittae sometimes joined basally and a black spot in each
lateral pale area (Fig. 68 7f); elytron yellowish orange, immaculate; ventral surface
black or brown except mouthparts, pro- and mesostemum, epipleuron, and leg yellow
(in female). Male genitalia as in Figure 687a-d. Female genitalia as in Figure 687e.
Discussion. The immaculate elytron of M. luteodorsa is unique within the genus
except for some specimens of M. picta, and causes this species to resemble members
of Cycloneda.
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Fig. 687. Mulsantina luteodorsa.
Type locality. Baton Rouge, Louisiana.
Type depository. USNM.
Distribution. Figure 681. ALABAMA: Hagen; Madison Co., Monte Sano St. Pk.
LOUISIANA: Baton Rouge; Many; Shreveport. MISSISSIPPI: Starkville. NORTH
CAROLINA: Macon Co. VIRGINIA: Vienna.
Mulsantina cyathigera (Gorham)
Fig. 688a-f; Map, Fig. 681
Coccinella cyathigera Gorham, 1891, p. 158. — Gordon, 1974b, p. 165.
Harmoniaspis cyathigera: Casey, 1908, p. 404.
Coccinella (Neoharmonia) cyathigera: Korschefsky, 1932, p. 510.
Mulsantina cyathigera: Blackwelder, 1945, p. 453.— J. Chapin (in press).
Diagnosis. Length 3.32 to 4.65 mm, width 2.57 to 3.40 mm. Color yellow except
pronotum with median, brown, M-shaped marking; elytron with 6 brown spots (Fig.
6881), spots sometimes reduced and with median and lateral or one or both missing;
ventral surface and leg reddish yellow. Male genitalia as in Figure 688a-d. Female
genitalia as in Figure 688e.
Discussion. This species occurs from Guatemala to southern Arizona. The finely
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851
defined M-shaped mark on the pronotum and discrete elytral spots are characteristic
of M. cyathigera which resembles large species of Psyllobora or Olla more than it
does other species of Mulsantina.
Type locality. Guatemala: Quezaltenango and Quiche Mts. (lectotype not desig-
nated).
Type depository. BMNH.
Distribution. Figure 681. ARIZONA: Chiricahua Mts.; Graham Mts.; Huachucha
Mts.; Santa Catalina Mts.; Santa Rita Mts.
Tribe Psylloborini
Psylloborini Casey, 1899, pp. 73, 100.— Leng, 1920, p. 215. — Korschefsky, 1932, p.
558.-Wingo, 1952, p. 22.-Sasaji, 1968, p. 21, 31. -J. Chapin, 1974, p. 69.
Halyziini Capra, 1927, p. 158. — Korschefsky, 1932, p. 558.
Coccinellinae with body length usually 3.0 mm or less. Head with gena not ex-
tending onto eye (Fig. 558a); eye coarsely faceted; mandible with 5 teeth, 2 apical,
3 internal; clypeus wider than frons, apex without produced anterolateral angle (Fig.
558a). Anterior border of pronotum feebly emarginate, partially concealing head.
One genus represents this tribe north of Mexico, but others occur worldwide, mainly
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in tropical regions. In North America the Psylloborini are likely to be confused only
with the Coccinellini (see discussion under the latter tribe).
Genus Psyllobora Dejean
Psyllobora Dejean, 1836, p. 458. — LeConte, 1852, p. 130.— Crotch, 1873, p. 375.—
1874b, p. 135.— Gorham, 1892, p. 165.— Wickham, 1894, p. 299.— Casey, 1899,
p. 100. — Leng, 1903b, p. 210. — Blatchley, 1910, p. 517. — Leng, 1920, p. 215.—
Korschefsky, 1932, p. 565. — Timberlake, 1943, pp. 41, 59.— Wingo, 1952, p. 22.—
J. Chapin, 1974, p. 69. — Belicek, 1976, p. 352. — lablokolf-Khnzorian, 1982, p.
299. Type-species; Coccinella lineola F., by subsequent designation of Timberlake
(1943).
Psyllobora {Psyllobora) Mulsant, 1850, p. 169. — Mulsant, 1866, p. 128.
Mulsant, 1846, p. 159. — Mulsant, 1850, p. 162. — Casey, 1899, p. 100.— Gor-
ham, 1892, p. 165. — Timberlake, 1943, pp. 41, 59. — Fursch, 1966, p. 90. Type-
species; Coccinella vigintiduopunctata L., by subsequent designation of Crotch,
1874b.
Coccinellini with length 1.80 to 6.50 mm; broadly oval to elongate oval, dorso-
ventrally flattened. Color pale yellow with brown maculation on pronotum and
elytron. Head pubescent; basal segment of antenna enlarged, somewhat flattened (Fig.
689a). Lateral margin of pronotum broadly explanate, transparent. Lateral margin
of elytron usually narrowly explanate, transparent; epipleuron nearly flat. Intercoxal
process of prostemum flat, feebly ridged laterally, truncate apically. Apical margin
of mesostemum truncate. Apex of middle and hind tibiae each without spurs, claw
with basal tooth (Fig. 689b). Postcoxal line incomplete, of Diornus type (Fig. 689c).
Male genitalia with basal lobe either symmetrical or asymmetrical. Female genitalia
with or without infundibulum; coxal plate irregularly elongate with distinct apical
stylus (Fig. 69 le).
Most of the Western Hemisphere species of this genus are neotropical, but 6 species
occur north of Mexico. Because there are many undescribed species in South America,
it is not possible to give an accurate estimate of the total number of species in this
hemisphere. There are probably over 50. The genus Psyllobora as presently consti-
tuted is not a monophyletic unit, and systematic research in the group will result in
the establishment of one or more additional genera. The North American species
are very similar in external appearance but are divergent in genitalic structure. Two
groups are evident based on genitalia, the vigintimaculata group, composed of the
nominate species and possibly P. nana, and the borealis group, eomposed of the
remaining 4 species. Psyllobora vigintimaculata has an extemely long, convoluted
spermathecal capsule with a correspondingly long, threadlike siphonal apex in the
male. The borealis group has a more normal type of spermathecal capsule and the
siphonal apex is not long and threadlike. Within the borealis group, the female
genitalia of P. borealis and P. plagiata eaeh have an infundibulum, the genitalia of
P. renifer and P. parvinotata lack that structure. Members of Psyllobora are known
to feed on fungus, particularly the mildew type (Davidson, 1921). Other hosts such
as mites, aphids, and scale insects have been recorded in the literature, but much of
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NORTH AMERICAN COCCINELLIDAE
853
this data is a result of inaccurate observation and assumption. Specific fungal host
data are listed as follows: Erysiphe communis (Wallr.) Fries, Erysiphe polygoni D.C.,
Erysiphe polygoni D.C. ex St. Amans, Erysiphe tortilis Wallr. ex Fr., Microsphaera
alphitoides Griff, and Maubl, Phyllactinia suffulta (Rabent.) Sacc., Podosphaera leu-
cotricha (Ellis & Everh.) Salm., Podosphaera oxyacanthae (D.C.) DeBy, Sphaerotheca
pannosa (Wallr. ex Fr.) Lev., Uncinula nector (Schw.) Burill.
The genus has not been taxonomically treated as a whole, but Timberlake (1943)
constructed a key to the North American species based on male genitalia.
Key to species of Psyllobora
1 . Elytron with large, common, sutural spot at apical third (Fig. 689g); Greater Antillean
species recorded from Florida, rare nana Mulsant
Elytron without large sutural spots at apical third; Florida and elsewhere 2
2(1). Elytron with spots strongly coalescent, suffused, at least in apical half, marginal spots
usually absent (Fig. 696f) renifer Casey
- Elytron with spots not strongly coalescent, marginal spots usually present 3
3(2). Form broadly oval; elytron with maculae reduced, only subapical spot large, strongly
evident (Fig. 694f); known only from southern Arizona plagiata Schaeffer
Form elongate; elytron with numerous small spots 4
4(3). Elytron with strongly developed lateral spot behind middle, spot either free or nar-
rowly connected to large inner spot, widely separated from apical spots (Fig. 69 3f)
borealis Casey
Elytron with lateral spot behind middle usually not strongly developed, narrowly
separated from apical spot or spots, or joined to it (Figs. 69 If, 695 f) 5
5(4). Pronotum often without spots (Fig. 695f, g); genitalia as in Figure 695a-d; Rorida
to Louisiana parvinotata Casey
Pronotum always with spots (Fig. 69 If); genitalia as in Figure 691a-d; occurring
over most of North America, not known from Rorida vigintimaculata (Say)
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Fig. 689. Psyllobora sp. a. Antenna, b. Tarsus, c. Postcoxal lines, d-f. Psyllobora nana.
Psyllobora nana Mulsant
Fig. 689d-g; Map, Fig. 690
Psyllobora nana Mulsant, 1850, p. 181. — Mulsant, 1866, p. 137.— Crotch, 1873, p.
376.-Crotch, 1874b, p. 141. -Casey, 1899, p. 102.-Leng, 1903b, p. 211.-Kor-
schefsky, 1932, p. 568.
Diagnosis. Length 2.30 to 2.70 mm, width 1.50 to 1.90 mm. Color pattern as
described for P. vigintimaculata except elytral spots less coalescent, one spot on
suture at apical V3 united with mate on opposite elytron, subapical spot near suture
free, large (Fig. 689g). Male genitalia as in Figure 689d-f.
Discussion. This is a distinctive Greater Antillean species that has been recorded
from Florida in the literature; one specimen in the Casey collection is labeled “Dry
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NORTH AMERICAN COCCINELLIDAE
855
Fig. 690. Distribution. Psyllobora nana (open star); P. borealis (shaded); P. plagiata (star);
P. parvinotata (cross hatch).
Tortugas, Fla.” A specimen in the Crotch collection labeled “P. nana Cuba/TYPE
(blue paper)” is here designated and labeled as the lectotype.
Type locality. Cuba (lectotype here designated).
Type depository. UCCC
Distribution. Figure 690. FLORIDA: Dry Tortugas.
Psyllobora vigintimaculata (Say)
Fig. 691a-i; Map, Fig. 692
Coccinella 20-maculata Say, 1824, p. 96.
Psyllobora viginti-maculata: Mulsant, 1850, p. 183. — Mulsant, 1866, p. 137.— Crotch,
1874b, p. 141. — Gorham, 1892, p. 167 (in part). — Timberlake, 1943, p. 42.
Psyllobora 20-maculata: Crotch, 1873, p. 375.— Wickham, 1894, p. 303.— Casey,
1899, p. lOl.-Leng, 1903b, p. 210.-Blatchley, 1910, p. 5 17. -Timberlake, 1943,
p. 59.
Psyllobora viginti-maculata: Leng, 1920, p. 215.
Psyllobora vigintimaculata: Korschefsky, 1932, p. 569.— Wingo, 1952, p. 45.— J.
Chapin, 1974, p. 70.— Belicek, 1976, p. 353.
Psyllobora taedata LeConte, 1860, p. 70.— Wickham, 1894, p. 306. — Casey, 1899,
p. 102.-Leng, 1903b, p. 21 1.- Timberlake, 1943, pp. 42, 59.
Psyllobora 20-maculata var. taedata: Crotch, 1873, p. 376. — Korschefsky, 1932, p.
570.-Hatch, 1961, p. 184.
Psyllobora obsoleta Casey, 1899, p. 101.— Casey, 1908, p. 407.
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Fig. 691. Psyllobora vigintimaculata.
Psyllobora 20-maculata var. obsoleta: Leng, 1903b, p. 210.
Psyllobora separata Casey, 1899, p. 102. — Leng, 1903b, p. 211.
Psyllobora taedata separata: Casey, 1908, p. 407.
Psyllobora vigintimaculata ab. separata: Korschefsky, 1932, p. 569.
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Diagnosis. Length 1.75 to 3.0 mm, width 1.40 to 2.35 mm. Pronotum with 4 dark
spots; elytron usually with 9 spots, spots partially confluent (Fig. 69 If), pattern
variable (Fig. 691g-i). Male genitalia as in Figure 691a-d. Female genitalia as in
Figure 69 le.
Discussion. The dorsal color pattern will usually separate vigintimaculata from all
other North American species of Psyllobora except strongly maculate specimens of
P. parvinotata, where genitalia must be used. Psyllobora vigintimaculata is found
throughout the United States (except Florida) and southern Canada with the range
extending into Mexico. Psyllobora taedata has been considered a synonym or variety
ofP. vigintimaculata by some previous authors, and most recently, Timberlake ( 1 943)
regarded P. taedata as a valid species. I consider P. taedata a junior synonym of P.
vigintimaculata because the morphological differences between the eastern and west-
ern population are gradually clinal and it is impossible to delimit well characterized
geographic races.
The types (females) of P. obsoleta and separata are unique (holotypes). LeConte
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had several type specimens of P. taedata, therefore I designate and label a specimen
labeled “(gold disc)/46 3 9/Type 6691 (red paper)/?, taedata LeC.” as the lectotype.
Three additional specimens bearing gold discs are designated and labeled as para-
lectotypes.
Type locality. Of vigintimaculata, “Missouri”; of taedata, San Francisco, California
(lectotype here designated); of obsoleta, Keokuk, Iowa; of separata, Siskiyou Co.,
California.
Type depository. Of vigintimaculata, type lost; of taedata, MCZ; of obsoleta (35545)
and separata (35547), USNM.
Distribution. Figure 692. Northern range limit. New Foundland to Alaska; southern
range limit, Virginia to Tennessee and Louisiana, west to southern California.
Psyllobora borealis Casey
Fig. 693a-f; Map, Fig. 690
Psyllobora borealis Casey, 1899, p. 102.— Casey, 1908, p. 407.— Timberlake, 1943,
pp. 42, 60.-Hatch, 1961, p. 185.-Belicek, 1976, p. 353.
Psyllobora taedata: Leng, 1903b, p. 211.
Psyllobora vigintimaculata ab. borealis: Korschefsky, 1932, p. 569.
Psyllobora deficiens CsLsey, 1899, p. 102. — Casey, 1908, p. 407.— Timberlake, 1943,
p. 60.
Psyllobora vigintimaculata ab. deficiens: Korschefsky, 1932, p. 569.
Diagnosis. Length 2.40 to 3.10 mm, width 1.90 to 2.40 mm. Description as for P.
vigintimaculata except form more robust; lateral, submedian spot large, widely sep-
arated from subapical spots (Fig. 693f). Male genitalia as in Figure 693a-d. Female
genitalia as in Figure 693e.
Discussion. The robust form and elytral color pattern will separate P. borealis from
P. vigintimaculata which it most nearly resembles. The type is a unique (holotype)
male. Timberlake (1943) regarded P. deficiens as a valid species, but I cannot separate
it from P. borealis and consider P. deficiens a junior synonym of P. borealis. The
female type of P. deficiens is unique (holotype).
Type locality. Of borealis, Coeur d’Alene, Idaho; of deficiens, California.
Type depository. Of borealis (35549) and deficiens (35550), USNM.
Distribution. Figure 690. Montana to New Mexico, west to southern British Co-
lumbia and southern California.
Psyllobora plagiata Schaeffer
Fig. 694a-f; Map, Fig. 690
Psyllobora plagiata Schaeffer, 1908, p. 125. — Korschefsky, 1932, p. 568.
Psyllobora Nunenmacher, 191 1, p. 7 1 . — Korschefsky, 1932, p. 567.— Tim-
berlake, 1943, p. 60. New Synonymy.
Diagnosis. Length 2.50 to 3.10 mm, width 2.20 to 2.50 mm. Description as for P.
vigintimaculata except form more robust; maculation on elytron strongly reduced
(Fig. 694f). Male genitalia as in Figure 694a-d. Female genitalia as in Figure 694e.
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NORTH AMERICAN COCCINELLIDAE
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Fig. 693. Psyllobora borealis.
Discussion. This species is not at all unlike P. borealis in spite of the seemingly
obvious dissimilarity in color pattern. There are, however, significant differences in
the genitalia of the 2 species. Psyllobora koebelei is a junior synonym of plagiata.
Almost all of the Schaeffer types are now in the USNM, but there are no specimens
of P. plagiata labeled as type material. However, there are 4 specimens in the USNM
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labeled “Brooklyn Museum Coll. 1929” which I believe are part or all of the original
series. One of these also bears the label “Huach Mts., Arz, VII, 9000 ft.”. I here
designate and label one of these specimens, a male, as the lectotype. The type of
koebelei is a unique male (holotype). It is labeled “Nogales/St. Cruz Co. VI. 02 Ariz/
2426/A. Koebele collector/Psyllobora Koebelei Nun.”.
Type locality. Of plagiata, Huachucha Mts., Arizona (lectotype here designated);
of koebelei, Nogales, Santa Cruz Co., Arizona.
Type depository. Of plagiata, USNM; of koebelei, CAS.
Distribution. Figure 690. ARIZONA: Huachucha Mts.; Santa Catalina Mts.
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Fig. 695. Psyllobora parvinotata.
Psyllobora parvinotata Casey
Fig. 695a-g; Map, Fig. 690
Psyllobora parvinotata Casey, 1899, p. 101. — Blatchley, 1918, p. 420.— Timberlake,
1943, pp. 42, 59. -J. Chapin, 1974, p. 70.
Psyllobora 20-maculata var. parvinotata: Leng, 1903b, p. 210.— Casey, 1908, p.
407.-Korschefsky, 1932, p. 569.
Psyllobora pallidicola: Blatchley, 1918, p. 420.
Psyllobora 20-maculata var. pallidicola Blatchley, 1914, p. 65. — Blatchley, 1930, p.
38.-Korschefsky, 1932, p. 569.
Diagnosis. Length 2.75 to 3.40 mm, width 1.40 to 1.80 mm. Color pattern as in
P. vigintimaculata except pronotum often without spots, or spots pale; elytron with
spots less confluent and overall appearance paler than P. vigintimaculata (Fig. 69 5f-
g). Male genitalia as in Figure 695a-d. Female genitalia as in Figure 695e.
Discussion. Examples of this species that lack pronotal spots or have them strongly
reduced are readily distinguished from P. vigintimaculata because the latter species
almost always has well defined pronotal maculation. Specimens of P. parvinotata
with maculate pronota are difficult to distinguish and genitalia often must be ex-
amined. These 2 species are at least partially allopatric in that I have not seen any
specimens of P. vigintimaculata from peninsular Florida, and very few from other
Gulf Coast localities where P. parvinotata occurs. The type of P. parvinotata is a
unique (holotype) male.
Type locality. Of parvinotata. Palm Beach, Florida; of pallidicola, Dunedin, Florida.
Type depository. Of parvinotata (35546), USNM; of pallidicola, PU.
Distribution. Figure 690. Florida to Louisiana.
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Psyllobora renifer Casey
Fig. 696a-g; Map, Fig. 697
Psyllobora renifer Casey, 1899, p. 102. — Leng, 1903b, p. 210.— Casey, 1908, p. 407.—
Timberlake, 1943, pp. 42, 59. -J. Chapin, 1974, p. 71.
Psyllobora vigintimaculata ab. renifer. Korschefsky, 1932, p. 569.
Psyllobora renifera: Timberlake, 1943, pp. 42, 59.— J. Chapin, 1974, p. 71.
Diagnosis. Length 1.75 to 2.40 mm, width 1.40 to 1.80 mm. Color pattern as
described for P. vigintimaculata except most elytral spots coalescent, suffused (Fig.
696f, g). Male genitalia as in Figure 696a-d. Female genitalia as in Figure 696e.
Discussion. Most examples of this species are recognized because of the strongly
coalescent elytral spots. The range of P. renifer is from Louisiana west to southern
California and into northern Mexico. The holotype of renifer is a unique male.
Type locality. Brownsville, Texas.
Type depository. USNM (35548).
Distribution. Figure 697. Lousiana to Nevada and California.
Subfamily Epilachninae
Epilachninae Ganglbauer, 1899, p. 947. — Leng, 1920, p. 217. — Korschefsky, 1931,
p. 16. — Black welder, 1945, p. 440.— Wingo, 1952, p. 25.— J. Chapin, 1974, p.
71. — Gordon, 1976a, p. 16.
Epilachniens Mulsant, 1846, p. 190. — Mulsant, 1850, p. 696.
Trischosomides Mulsant, 1850, p. 696 (in part).
Coccinellidae with dorsal surface pubescent. Head retracted under pronotum so
hind margin of eye usually covered by pronotum. Eye oblique, finely faceted, inner
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NORTH AMERICAN COCCINELLIDAE
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margin broadly, feebly curved. Labium with palpal insertion median or subterminal.
Apical segment of maxillary palpus weakly securiform (Fig. 698b). Mandible without
basal tooth but with several teeth of various types in apical half (Fig. 698d). Antenna
inserted in depression on inner side of eye, 1 1 -segmented, last 3 segments forming
loose club (Fig. 698c), outer margin of club segments appearing slightly serrate.
Prostemal process narrow, apex truncate or bluntly rounded. Tarsus cryptotetra-
merous. Male genitalia symmetrical. Female genital plate short, broad, stylus visible
or not (Fig. 703e).
Members of this subfamily occur worldwide, mostly in the tropical regions. There
are 2 tribes, Epilachnini and Madaini, represented in the Western Hemisphere, with
only the former represented north of Mexico. The feeding habits of the Epilachninae
are unusual because most Coccinellidae are predators, but epilachnines are plant
feeders. A few species are serious pests of cultivated crops, such as E. varivestis
Mulsant on beans of various types. The subfamily was revised by Gordon (1976a)
for the Western Hemisphere.
Tribe Epilachnini
Epilachnini Costa, 1849, p. 69.— Casey, 1899, p. 102. — Korschefsky, 1931, p. 16.—
Gordon, 1976a, p. 17.
Epilachninae with form oval, cordate, or elongate oval, widest anterior to middle
of elytra, lateral margin of elytron usually explanate. Leg with tibia slender, nearly
as long as femur and trochanter combined, tarsus received in tibial grooves (Fig.
6981); front tibia with one or 2 large, acuminate spurs at apex on inner margin, middle
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 698. Epilachna sp. a. Head. b. Maxillary palpus, c. Antenna, d. Mandible, e. Postcoxal
lines, f. Tibia.
and hind tibia each with 2 acuminate spurs. Epipleuron flat or descending externally
with no depressions for reception of leg.
Two genera represent this tribe north of Mexico. One genus, Epilachna, is native
to the region and the other, Subcoccinella, is an introduction from Europe. The
complex mandibular structure, which is an adaptation for plant feeding, characterizes
members of this tribe. See Gordon (1976a).
Key to genera of Epilachnini
1. Postcoxal line strongly recurved apically (fig. 698e); length more than 6.0 mm
Epilachna Dejean
- Postcoxal line not recurved apically (Fig. 705c); length less than 5.0 mm
Subcoccinella Huber
Genus Epilachna Dejean
Epilachna Dejean, 1837, p. 460. — Hope, 1840, p. 157. — Mulsant, 1850, p. 700.—
LeConte, 1852, p. I30.-Crotch, 1874b, p. 53. -Casey, 1899, p. 103.-Leng, 1920,
p. 217.-Korschefsky, 1931, p. 17. -Black welder, 1945, p. 440.-Wingo, 1952, p.
25.— J. Chapin, 1974, p. 72. — Gordon, 1976a, p. 37. Type-species; Coccinella
borealis (F.), by subsequent designation of Hope, 1840.
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Solanophila ^ 1898a, p. 99.— Korschefsky, 1931, p. 18. — Blackwelder, 1945,
p. 440.— Dieke, 1947, p. 7.— Li and Cook, 1961, p. 33. Type-species; Solanophila
gibbosa Weise, by subsequent designation of Li and Cook, 1961.
Afissa Dieke, 1947, p. 106.— Li and Cook, 1961, p. 33. Type-species; Coccinella
Jlavicollis Thunberg, by original designation.
Epilachnini with labrum usually truncate or feebly emarginate, sometimes strongly
emarginate, usually concealing at least basal V3 of mandible. Mandible with apex
usually trifid, sometimes bifid, apical teeth acuminate, usually with 2 large teeth and
several minor teeth or dentules below apex. Apex of anterior tibia with single acu-
minate spur; apices of middle and hind tibiae each with 2 acuminate spurs. All tibiae
grooved for reception of tarsus.
Most species of Epilachna are neotropical with 3 species occurring in the United
States. Two plant families serve as hosts for the North American species. Epilachna
borealis and E. tredecimnotata feed on members of the Cucurbitaceae, E. varivestis
feeds on members of the Leguminaceae. See Gordon (1976a).
Key to species of Epilachna
1. Each elytron with 8 dark spots (Fig. 699e); pronotum immaculate . . varivestis Mulsant
- Each elytron with 7 dark spots (Fig. 7011); pronotum usually with 4 dark spots . . 2
2(1). Species occurring from eastern United States to central Texas (Fig. 702) . . borealis (F.)
- Species occurring from West Texas to Arizona and south (Fig. 704)
tredecimnotata (Latreille)
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 699. Epilachna varivestis.
Epilachna varivestis Mulsant
Fig. 699a-e; Map, Fig. 700
Epilachna varivestis Mulsant, 1850, p. 815. — Crotch, 1874b, p. 62. — Casey, 1899,
p. 103.-Leng, 1920, p. 217.-Korschefsky, 1931, p. 58.-Wingo, 1952, p. 47.-J.
Chapin, 1974, p. 73.
Epilachna corrupt a Mulsant, 1850, p. 815. — Casey, 1899, p. 103.— Korschefsky,
1931, p. 58.
Epilachna varivestis var. cervina Mulsant, 1850, p. 817.
Epilachna varivestis var. genuina Mulsant, 1850, p. 817.
Epilachna corrupta ab. cervina’. Korschefsky, 1931, p. 58.
Epilachna corrupta ab. genuina: Korschefsky, 1931, p. 58.
Epilachna cervina: Blackwelder, 1945, p. 442.
Epilachna maculiventris Bland, 1864, p. 256.— Crotch, 1874, p. 64 (as a synonym
of borealis F.). — Korschefsky, 1931, p. 38 (as a synonym of corrupta Mulsant).—
Blackwelder, 1945, p. 442.
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NORTH AMERICAN COCCINELLIDAE
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Epilachna cuprea Cockerell, 1918, p. 153.— Blackwelder, 1945, p. 442.
Epilachna toweri Johnson, 1910, p. 78.— Leng, 1920, p. 217.
Epilachna juncta Johnson, 1910, p. 79.— Blackwelder, 1945, p. 442.
Epilachna corrupta ab. cuprea: Korschefsky, 1931, p. 58.
Epilachna corrupta dih. juncta: Korschefsky, 1931, p. 58.
Diagnosis. Length 6.43 to 8. 10 mm, width 4.85 to 6.38 mm. Color brownish yellow;
metastemum and median area of abdominal sterna slightly darker; elytron with 8
dark brown spots arranged in transverse rows (Fig. 699e), but pattern variable. Male
genitalia as in Figure 699a-c. Female genitalia as in Figure 699d.
Discussion. See Gordon (1976a) for detailed discussion.
Type locality. Mexico.
Type depository. UCCC.
Distribution. Figure 700. Eastern distribution; Maine to Florida, west to Nebraska
and Louisiana. Western Distribution; South Dakota and Idaho south to west Texas
and Arizona.
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Fig. 701. Epilachna borealis.
Epilachna borealis (F.)
Fig. 701a-f; Map, Fig. 702
Coccinella borealis Fabricius, 1775, p. 82.
Coccinella punctatissima Weber, 1801, p. 48.
Epilachna borealis: Hope, 1840, p. 1 57. — Mulsant, 1850, p. 826 (in part).— Crotch,
1874b, p. 64.— Weise, 1898a, p. 98.— Casey, 1899, p. 103. — Leng, 1920, p. 217.—
Korschefsky, 1931, p. 55.— Wingo, 1952, p. 47.— J. Chapin, 1974, p. 72.— Gordon,
1976a, p. 133.
Epilachna borealis ab. punctatissima: Korschefsky, 1931, p. 56.
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Diagnosis. Length 7.35 to 9.80 mm, width 5.50 to 7.63 mm. Color yellow; prono-
tum with 4 black spots; elytron with 7 large, dark brown spots (Fig. 7011). Male
genitalia as in Figure 701a-d. Female genitalia as in Figure 70 le.
Discussion. As discussed by Gordon (1976a), this species and E. tredecimnotata
are very similar in appearance. The elytral spots on the disc are always confluent in
E. borealis, rarely so in E. tredecimnotata, and the male genitalia are different in
these 2 species.
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Fig. 703. Epilachna tredecimnotata.
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Type locality. North America, probably Jamaica, Long Island.
Type depository. Fabrician collection, Kiel (not examined).
Distribution. Figure 702. Massachusetts to Florida, west to Kansas and east Texas.
Epilachna tredecimnotata (Latreille)
Fig. 703a-g; Map, Fig. 704
Coccinella tredecimnotata Latreille, 1833, p. 67.
Epilachna borealis 3b. 13-notata: Mulsant, 1850, p. 827.
Epilachna borealis db. tredecimnotata: Korschefsky, 1931, p. 56. — Blackwelder, 1945,
p. 441 (for detailed synonymy see Gordon, 1976a, p. 135).
Diagnosis. Length 6.75 to 10.0 mm, width 5.10 to 8.05 mm. Description as for
E. borealis except elytron with spots usually smaller and sutural spot on disc of
elytron separated from suture (Fig. 703f, g); pronotum often with less than 4 spots
or with none. Male genitalia as in Figure 703a-d. Female genitalia as in Figure 703e.
Discussion. For further discussion see under E. borealis and Gordon (1976a).
Type locality. “America equinox”.
Type depository. DLM.
Distribution. Figure 704. West Texas to Arizona.
Genus Subcoccinella Huber
Subcoccinella Huber, 1842, p. 376. — Crotch, 1874b, p. 90. — Korschefsky, 1931, p.
68. Type-species; Coccinella vigintiquatuorpunctata L., by monotypy (for detailed
synonymy, see Korschefsky, 1931, p. 69).
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Fig. 705. Subcoccinella vigintiquatuorpunctata.
Epilachnini with form oval, lateral margin of elytron feebly explanate. Labrum
broadly, feebly emarginate. Mandible with 2 strong apical teeth and 2 strong subapical
teeth, each having multiple denticles (Fig. 705a). Anterior tibia with 2 acuminate
spurs at apex, laterally flattened with outer margin angulate at apical (Fig. 705b);
middle and hind tibiae with 2 apical spurs, slender, not angulate (Fig. 705b). Postcoxal
line incomplete, not recurved apically (Fig. 705c).
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This is a monobasic European genus which has accidentally been introduced into
North America. It was first reported in 1974 from specimens collected near Hack-
ensack, Bergen Co., New Jersey, in 1973, but has obviously been long established
because subsequent collecting has shown it to be widely distributed. In Europe S.
vigintiquatuorpunctata is a major pest on alfalfa, but so far it has shown no inclination
to feed on alfalfa in North America. The principal host in North America is “bouncing
bet”, Saponaria officinalis L., also a European import widely distributed along rail-
road tracks and highways. In Europe it has been recorded on 70 species of plants in
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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
12 families and can live on several species of legumes, particularly clover. Nearly all
previous information on this species in the United States is to be found in various
volumes and numbers of the “Cooperative Plant Pest Report (USDA)” from 1974
to April, 1976. I incorrectly placed this genus in the tribe Madaini (Gordon 1976a,
p. 206). I here place it in the tribe Epilachnini.
Subcoccinella vigintiquatuorpunctata (L.)
Fig. 705a-i; Map, Fig. 706
Coccinella 24-punctata L., 1758, 366.
Subcoccinella vigintiquatuorpunctata: Huber, 1842, p. 476.— Crotch, 1874b, p.90.—
USDA, 1974, p. 731. -USDA, 1975, p. 184.-Gordon, 1976a, p. 206. -Wheeler
and Henry, 1981, p. 197 (for detailed synonymy see Korschefsky, 1931, p. 69).
Diagnosis. Length 3.50 to 4.40 mm, width 2.85 to 3.15 mm. Color yellowish red;
elytron with varying number of black spots (Fig. 705h, i). Male genitalia as in Figure
705d, e. Female genital plate as in Figure 705g.
Discussion. This species is easily recognized on appearance alone in the depauperate
epilachnine fauna of North America. The dorsal color pattern is variable but the
small size and oval form distinguish it from Epilachna varivestis and E. borealis
which occur in the same geographic region.
Type locality. Europe.
Type depository.— Type not examined.
Distribution. Figure 706. ILLINOIS: “East of St. Louis, Mo.”. MARYLAND:
Allhegeny Co.; Washington Co. NEW JERSEY: Hunterdon Co.; Morris Co.; Passaic
Co. NEW YORK: Tioga Co. OHIO: Columbiana Co.; Coshocton Co.; Cuyahoga
Co.; Trumbull Co., Custer, Kelly. PENNSYLVANIA: Beaver Co.; Berks Co.; Brad-
ford Co.; Bucks Co.; Chester Co., Malvern; Cumberland Co.; Dauphin Co., Harris-
burg; Franklin Co.; Lackawanna Co.; Lancaster Co.; Lawrence Co.;; Lebanon Co.;
Lehigh Co., Allentown; Luzerne Co.; Mercer Co.; Montgomery Co.; Northampton
Co., Easton; Schuylkill Co. WEST VIRGINIA: Berkeley Co.; Jefferson Co.; Morgan
Co.; Paoli; Taylor Co., Grafton.
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Index to family-group and genus-group names
Names in italics are synonyms; boldface page numbers indicate page on which each
taxon is described.
Adalia 681,776
Anatis 681,752
Anisocalvia 774
Anisostica 682, 681
Anovia 665, 667
Aphidecta 681,839
Arawana 619, 603
Axion 603, 611
Azya 674
Azyini 655,671
Blaisdelliana 353
Brachiacantha 3 5 3 , 556
Brumoides 603
Calvia 681,774
Cephaloscymnini 36, 66
Cephaloscymnus 68
Ceratomegilla 681, 702
Chilocorinae 34, 602
Chilocorini 602
Chilocorus 602, 641
Cleis 843
Coccidophilus 37, 44
Coccidula 655,656
Coccidulinae 34, 654
Coccidulini 655
Coccinella 682, 783
Coccinellidae 33
Coccinellinae 678
Coccinellini 679
Coelophora 829, 681
Coleomegilla 696, 681
Cryptognatha 600
Cryptognathini 74, 599
Cryptolaemus 100, 105
Cryptoweisea 44
Cycloneda 682, 819
Delphastus 61
Depressoscymnus 287,314
Didion 107, 102
Diomus 315, 102
Dobzhanskia 783
Eocaria 774
Epilachna 864
Epilachninae 34, 862
Epilachnini 863
Epismilia 37
894
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
Exochomus . . .
Exoplectra ....
Exoplectrini . . .
Gnathoweisea .
Halmus
Harmonia ....
Helesius
Hippodamia . .
Hyperaspidius .
Hyperaspini ...
Hyperaspis ....
Leis
Lindorus
Macronaemia .
Microweisea . .
Microweisini . .
Mulsantina ...
Mysia
Myzia
Naemia
Neoharmonia .
Neomysia
Nephaspis
Nephopullus . . . .
Nephus
Nipus
Noviini
Olla
Oxynychus
Paramysia
Paranaemia ...
Pentilia
Propylaea
Pseudoazya ...
Pseudocleis . . . .
Pseudoscymnus
Pseudoweisea . . .
Psyllobora
Psylloborini . . . ,
Pullus
Rhyzobius
Rodolia
Scymnillus
Scymninae
Scymnini
Scymnobius . . . .
Scymnus
Selvadiini
Selvadius
Serangiini
Sidis
. ... 603,621
668
. ... 655,668
36,49
. . . . 640, 603
. ... 681,832
. ... 353,396
. ... 681,706
. ... 353,357
74,352
. . 353,401
832
659
. ... 686, 681
37
36
. ... 681,843
763
. ... 763,681
. ... 693,681
. ... 682,834
763
. . . . 100, 102
88
102, 286, 287
36,56
. ... 655,662
. ... 682,826
402
764
. ... 689, 681
23
. ... 682,817
. ... 674,677
843
24
37
852
. ... 679,851
. ... 115,139
. ... 655,659
665
75
34,74
74,99
. . . . 296, 287
113, 115, 102
74,347
347
36,58
. . . . 287, 291
1985
NORTH AMERICAN COCCINELLIDAE
895
Srnilia 37
Spiladelpha 704
Spilota 783
Stethorini 74, 88
Stethorus 88
Sticholotidinae 34, 34
Subcoccinella 864, 871
Thalassa 353, 400
Thea 852
Turboscymnus 287, 293
Zagloba 75,81
Zilini 74
Zilus 75
Index to species-group names
Names in italics are synonyms; boldface page numbers indicate page on which each
taxon is described.
abbreviatus LeConte (Scymnus) 141, 246
abdominalis Say (Olla) 826
adulans Casey (Diomus) 325
advena Casey (Scymnus) 266
aeger Casey (Diomus) 347
aemulator Casey (Hyperaspis) 407, 463
aeneipennis (Sicard) (Zagloba) 28
aethiops (Bland) (Exochomus) 623, 631
affinis Crotch (Cryptolaemus) 17
affinis Randall (Hyperaspis) 423
Casey (Scymnus) 274
albertana Casey (Hippodamia) 715
albifrons (Say) (Brachiacantha) 560, 561, 594
algodonus, n. sp. (Hyperaspidius) 359, 385
aha Brown (Coccinella) 786,804
alutaceum Casey (Axion) 619
aluticollis Casey (Scymnus) 122
amabilis (LeConte) (Diomus) 318, 321
ambigua LeConte (Hippodamia) 708, 729
americana Crotch (Hippodamia) 707, 712
americanus Mulsant (Scymnus) 119, 131
amnicola Wingo (Nephaspis) 102
amoena Scott (Hippodamia) 725
ampla (Mulsant) (Neoharmonia) 837
andrewsi, n. sp. (Hyperaspidius) 359, 392
angolensis Crotch (Chilocorus) 15
angustula Casey (Hyperaspis) 516
angustus Casey (Scymnus) 25
annectans Crotch (Adalia) 782
annexa LeConte (Hyperaspis) 410, 524
antipodum (White) (Harmonia) 19
apacheanus Casey (Scymnus) 220
apetzi Mulsant (Scymnus) 25
896 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
apicalis Casey (Hippodamia) 708, 719
apicanus J. Chapin (Scymnus) 119, 133
apiciventris Casey (Scymnus) 169
apithanus Gordon (Scymnus) 147, 188
appalacheus Casey (Diomus) 343
aquilonarius Gordon (Scymnus) 1 50, 262
arctica (Schneider) (Hippodamia) 707, 725
arcuatus (LeConte) (Hyperaspidius) 358, 360, 369
ardelio Horn (Scymnus) 145, 220, 148, 150
aridoides Gordon (Scymnus) 147, 226
aridus Casey (Scymnus) 224, 144, 145, 148, 150
arizonae Casey (Olla) 828
arizonica (Casey) (Arawana) 620
arizonica Dobhzansky (Hyperaspis) 412, 525
arizonica Schaeffer (Brachiacantha) 559, 585
arizonicus Gordon (Diomus) 317, 344
asphaltina Casey (Hyperaspidius) 378
aterrima Casey (Hyperaspis) 504
aterrimus (Horn) (Zilus) 76
atlantica Dobzhansky (Hyperaspis) 488
atomus Casey (Stethorus) 90
atramentarius (Boheman) (Nephus) 296, 304
atronitens (Casey) (Coccidophilus) 45, 47
aurantii Blackburn (Rhyzobius) 24
australasiae (Boisduval) (Parapriasus) 23
australasiae Blackburn (Scymnus) 25
australis Gordon (Cephaloscymnus) 68, 70
austrinus Gordon (Diomus) 334, 3 1 7
axyridis (Pallas) (Harmonia) 19
balteatus (LeConte) (Diomus) 317, 319
barberi Gordon (Scymnus) 206, 145, 148
barberi, n. sp. (Brachiacantha) 559, 572
bar da LeConte (Coccinella) 789
barovskii S. & D. (Ceratomegilla) 704
barri Hatch (Hyperaspis) 505
basalis Melsheimer (Brachiacantha) 591
bellus (Blackburn) (Rodolia) 25
bensonica Casey (Hyperaspis) 413, 502
bicentralis Casey (Hyperaspis) 432
bicolor Casey (Zagloba) 85, 83
bicolor Kamiya (Sukunahikona) 28
bigeminata (Randall) (Hyperaspis) 407, 440
bigemmeus (Horn (Diomus) 332, 3 1 7
bilunulatus (Boisduval) (Priasus) 24
binaevatus (Mulsant) (Nephus) 31,22, 292
binaria Casey (Hyperaspis) 409, 411, 498
binotata (Say) (Hyperaspis) 409, 423
bioculatus Mulsant (Nephus) 297
biornatus Nunenmacher (Hyperaspis) 469
biplagiata (Swartz) (Lemnia) 21
biplagiatus Casey (Nipus) 56
1985 NORTH AMERICAN COCCINELLIDAE 897
bipunctata (L.) (Adalia) 780
bipunctata Malkin (Hyperaspis) 509
bipunctatus Kugelann (Nephus) 23
bipustulatus (L.) (Chilocorus) 13, 15, 644, 653
hisignatus Horn (Nephus) 294
bitriangularis (Say) (Anisosticta) 683
bivulnerus (Horn) (Nephus) 296, 299
bivulnerus Mulsant (Chilocorus) 651
blaisdelli Casey (Scymnus) 200
blaisdelli Nunenmacher (Brachiacantha) 558, 568
blanchardi Johnson (Mulsantina) 845
blatchleyi, n. name (Hyperaspidius) 358, 376
blumi (Nunenmacher) (Brumoides) 603, 611
bollii Crotch (Brachiacantha) 560, 590
bolteri LeConte (Hyperaspis) 410, 554
borealis (F.) (Epilachna) 866, 865
borealis Casey (Psyllobora) 853, 858
borealis Dobzhansky (Hyperaspis) 410, 412, 412, 412, 540
borealis Gordon (Scymnus) 134
borealis Hatch (Scymnus) 194
borealis Timberlake (Anisosticta) 683, 684
bowditchi Johnson (Hippodamia) 708, 739
breiti Mader (Harmonia) 19
brevis Casey (Stethorus) 96
bridwelli Nunenmacher (Coccinella) 785, 804
brullei (Scymnus) 143, 283, 141, 142, 144
brunnescens Casey (Diomus) 343
brunnescens Dobzhansky (Hyperaspis) 410, 544
bryanti Gordon (Scymnus) 278, 148
bryanti Nunenmacher (Hyperaspidius) 360, 373
cacti (L.) (Chilocorus) 646, 644
caecus Blackburn (Rhyzobius) 24
caffer Gordon (Scymnus) 151,146
Calaveras Casey (Scymnus) 144, 147, 149, 209
californica Dobzhansky (Hyperaspis) 420
caiifomica Mannerheim (Coccinella) 786, 793
Boheman (Scymnus) 193
califomicus Casey (Exochomus) 624, 634
Dobzhansky (Hyperaspis) 541
canterius Casey (Scymnus) 176
cardinalis (Mulsant) (Rodolia) 32, 25, 666
Carolina (Casey) (Brachiacantha) 593
carri Gordon (Scymnus) 234, 149
carri Nunenmacher (Hyperaspidius) 362
caseyi Gordon and Chapin (Stethorus) 90
caseyi Johnson (Hippodamia) 708, 743
caseyi Timberlake (Myzia) 769
caseyi Weise (Diomus) 343
caseyi Westcott (Anatis) 755
caseyi, n. sp. (Hyperaspis) 411, 493
caseyianus Leng and Mutchler (Diomus) 343
898
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
catalinae (Horn) (Delphastus) 62, 63
caudalis LeConte (Scymnus) 142, 145, 272
caurinus Horn (Scymnus) 121, 118
centralis Casey (Neoharmonia) 837
centralis plagiata Dobzhansky (Hyperaspis) 426
cervicalis Mulsant (Scymnus) 141, 143, 146, 174
chalybeus (Boisduval) (Halmus) 31, 19, 641
chapini Dobzhansky (Hyperaspis) 407, 458
chapini Kapur (Catana) 15
children) Mulsant (Exochomus) 624, 626
chromopyga Casey (Scymnus) 175
cincta LeConte (Hyperaspis) 411, 492
LeConte (Scymnus) 191
c/rcw/arA Sharp (Rhyzobius) 661
circumdatus (Schoenherr) (Chilocorus) 15
circumspectus Horn (Scymnus) 1 16, 128
citricola Brethes (Coccidophilus) 16
coccidivora (Ashmead) (Microweisea) 38, 42
coccinea Casey (Hippodamia) 727
coccivora Ramakrishna (Scymnus) 26
cockerelli Casey (Scymnus) 145, 147, 148, 232
cockerelli Johnson (Hippodamia) 747
coeruleus Mulsant (Curinus) 17
collaris Melsheimer (Scymnus) 207
coloradana Casey (Hyperaspis) 511
coloradensis Horn (Nephus) 301
coloradensis Nunenmacher (Hyperaspidius) 362
compar Casey (Scymnus) 240, 140
comparatus Casey (Hyperaspidius) 360
complex Casey (Hippodamia) 751
concavus Watson (Hyperaspis) 409, 445
concinnata Melsheimer (Mulsantina) 844
concurrens Casey (Hyperaspis) 484
conformis (Boisduval) (Harmonia) 20
CO Mulsant (Brachiacantha) 591
confusor Casey (Chilocorus) 646
congeminata Watson (Hyperaspis) 444
conglobata (L.) (Oenopia) 23
conglomerata (L.) (Adalia) 14
congressis Watson (Hyperaspis) 437
congruens Casey (Brachiacantha) 580
coniferarum Crotch (Scymnus) 142, 145, 148, 151
connectens (Thunberg) (Hyperaspis) 409, 473
connexa Mulsant (Eriopis) 18
consimilis LeConte (Hyperaspis) 410, 541
consobrinus LeConte (Scymnus) 142, 228
conspirans Casey (Hyperaspis) 407, 479
conspiratus Casey (Hyperaspidius) 366
contexta Mulsant (Mulsantina) 844
convergens Guerin (Hippodamia) 709, 741
conviva Casey (Hyperaspis) 408, 409, 437
1985 NORTH AMERICAN COCCINELLIDAE 899
coosi Hatch (Scymnus) 120, 117
corrupta Mulsant (Epilachna) 866
cottlei Nunenmacher (Hippodamia) 715
creperus Mulsant (Scymnus) 142, 145, 147, 276
crotchi Casey (Hippodamia) 707, 709, 750
cruenta LeConte (Hyperaspis) 409, 485
cruentoides Dobzhansky (Hyperaspis) 499
cultratus Wingo (Scymnus) 269
cuprea Cockerell (Epilachna) 867
cyanoptera Mulsant (Neoharmonia) 837
cyathigera (Gorham) (Mulsantina) 844, 850
davisi (Leng) (Brumoides) 608, 603
debilis (LeConte) (Diomus) 317, 347
debilis Blackburn (Rhyzobius) 24
decempustulata (Melsheimer) (Brachiacantha) 560, 579
decepta Blatchley (Naemia) 694
decipiens Casey (Scymnus) 220
decora Casey (Brachiacantha) 558, 559, 561
deficiens Casey (Psyllobora) 858
deludens, n. sp. (Hyperaspis) 407, 460
dentipes (F.) (Brachiacantha) 558, 559, 564
Fall (Scymnus) 183
desertorum (Casey) (Brumoides) 604
desertorum Casey (Scymnus) 194
difficilis Casey (Scymnus) 1 16, 120
difhcilis Crotch (Coccinella) 786, 804
dimidiata (F.) (Harmonia) 834, 20, 3 1
discoideus Crotch (Chilocorus) 15
disconotata Mulsant (Hyperaspis) 410,532
discreta LeConte (Hyperaspis) 534
Timberlake (Hippodamia) 750
disjunctus Casey (Hyperaspis) 556
dispar Casey (Hippodamia) 746
disrupta Dobzhansky (Hyperaspis) 413,507
dissimila Blatchley (Neoharmonia) 837
dissoluta Crotch (Hyperaspis) 411, 511
distigma Klug (Chilocorus) 15
diversa Mulsant (Brachiacantha) 591
dobzhansky! Chapin (Hippodamia) 708, 722
dobzhanskyi, n. sp. (Hyperaspis) 409, 476
dorsalis Blackburn (Rhyzobius) 24
z/w/m Casey (Diomus) 321
durangoensis Casey (Hyperaspis) 556
effeta Casey (Hyperaspis) 526
elali Nutting (Hyperaspis) 540
elegans Mulsant (Hyperaspis) 536
eleutherae (Casey) (Zilus) 76, 79
elliptica Casey (Hyperaspis) 516
elusivus Gordon (Scymnus) . 252, 149
emarginata Chapin (Telsimia) 28
emertoni Casey (Diomus) 323
900 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol, 93(1)
enochrus Gordon (Scymnus) 177, 143, 146
ephippiata Zetterstedt (Coccinella) 786, 792
episcopalis (Kirby) (Macronaemia) 687
erythronotum Gordon (Scymnus) 149, 222
esclavium Dobzhansky (Hyperaspis) 406, 407, 468
essigi Malkin (Hyperaspis) 516
eucharis (Mulsant) (Harmonia) 20
eugenii Mulsant (Coccinella) 787, 789
excelsa Fall (Hyperaspis) 453, 406
exiguus Weise (Pharoscymnus) 23
expurgata Casey (Hippodamia) 708, 723
extensa Mulsant (Hippodamia) 708, 735
extricatus Casey (Scymnus) 203
falcigera Crotch (Hippodamia) 707, 713
falli Gordon (Scymnus) 199, 148
fain Nunenmacher (Hyperaspis) 504
fasciatus Casey (Exochomus) 624, 628
fastidiosa Casey (Hyperaspis) 406, 480
fattigi Blatchley (Neoharmonia) 837
feiina (F.) (Brachiacantha) 560, 577
felschei (Weise) (Microweisea) 43
femoralis LeConte (Diomus) 338
fenderi Malkin (Scymnus) 121, 117
fenestralis Casey (Olla) 828
fenyesi Leng (Brachiacantha) 582
ferox, n. sp. (Gnathoweisea) 49, 55
festatus Wingo (Scymnus) 180, 141
fidelis Casey (Hyperaspis) 521
filiola Casey (Hyperaspis) 413, 498
fimbriolata Melsheimer (Hyperaspis) 411, 488
flammula Nunenmacher (Hyperaspis) 450
flava timberlake (Cycloneda) 825
flavescens (Motschulsky) (Serangium) 27
flavescens Casey (Scymnus) 143, 146, 155
flavifrons (Melsheimer) (Nephus) 296
flavifrons Blackburn (Diomus) 336
flavifrons Mulsant (Brachiacantha) 560, 592
flavipes (Thunberg) (Exochomus) 30, 18, 639, 623
flavocephalus Blatchley (Hyperaspidius) 358, 375
flavomaculata (DeGeer) (Lioadalia) 21
Horn (Scymnus) 192
floralis (Motschulsky) (Exochomus) 18
floridana Leng (Coleomegilla) 700
floridanus (Mulsant) (Diomus) 318,320
floridensis Blatchley (Brachiacantha) 560, 589
ybwimafo Casey (Hippodamia) 750
forestieri (Mulsant) (Rhyzobius) 32, 24, 660, 661
franciscana Mulsant (Coccinella) 798
fratemus LeConte (Chilocorus) 644, 649
fratemus LeConte (Scymnus) 142,144,183
frigida Schneider (Adalia) 780
1985 NORTH AMERICAN COCCINELLIDAE 901
frosti Casey (Nephus) 289
fugax Blackburn (Rhyzobius) 24
fulgida Watson (Coccinella) 786, 807
fulvopustulata Melsheimer (Brachiacantha) 577
fuscilabris (Mulsant) (Coleomegilla) 698, 700
galbula (Mulsant) (Ileis) 21
garlandicus Casey (Scymnus) 145, 147, 148, 202
gemellata Mulsant (Cryptognatha) 17
gemina LeConte (Hyperaspis) 407, 438
gemma Casey (Hyperaspis) 478, 407
georgei (Weise) (Nephus) 295
gilae Casey (Scymnus) 169, 147, 149
gilvifrons (Mulsant) (Stethorus) 27
glacialis (F.) (Hippodamia) 709, 731
globosa Casey (Hyperaspis) 20
globula Casey (Hyperaspis) 408, 418
gordoni (Dozier) (Nephus) 314, 296
granum (Gorham) (Stethorus) 27
guexi LeConte (Exochomus) 627, 624
guexi Mulsant (Hyperaspis) 440
guttifera Weise (Hyperaspis) 536
guttiger Mulsant (Nephus) 297
guttulatus (LeConte) (Nephus) 296, 301
haematosticta Fall (Hyperaspis) 408, 425
haemorrhous LeConte (Scymnus) 183
hageni, n. sp. (Gnathoweisea) 49, 54
hardyi, n. sp. (Hyperaspidius) 359, 395
harneyi Hatch; (Scymnus) 120
hemorrhous divisus Casey (Scymnus) 284
hemorrhous laurenticus Casey (Scymnus) 284
hemorrhous subaeneus Casey (Scymnus) 284
hercules Belicek (Hyperaspidius) 359, 391
hesperius Gordon (Scymnus) 150, 265
hexacyclus Smith (Chilocorus) 644, 652
hirtuosum Blackburn (Serangium) 27
histrio (Fall) (Brumoides) 603, 610
hogei (Gorham) (Brumoides) 604, 605
Crotch (Myzia) 769
homi Gorham (Scymnus) 216, 147, 149
horni Nunenmacher (Hyperaspidius) 364
homi Weise (Pharoscymnus) 23
homi, n. sp. (Zilus) 77, 76
hortensis Wingo (Scymnus) 277
hottentota Mulsant (Hyperaspis) 519
houstoni Casey (Diomus) 343
howdeni Gordon (Scymnus) 280, 147
huachuca Gordon (Scymnus) 282, 148
hubbardi Gordon (Scymnus) 280, 147
hudsonica (Casey) (Mulsantina) 844, 847
humboldti Casey (Scymnus) 204, 150
humboldtiensis Nunenmacher (Coccinella) 785,787,816
902
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
humeralis Say (Adalia) 780
humilis Gordon (Diomus) 327, 3 1 7
hystrix Casey (Zagloba) 86, 84
iceryae Janson (Rodolia) 25
idae Nunenmacher (Hyperaspis) 450
ignarus Gordon (Scymnus) 212, 148
illustris Casey (Brachiacantha) 561,595
imitator, n. sp. (Hyperaspis) 413, 551
immaculata F. (Cycloneda) 820
immaculata Hatch (Hyperaspis) 410, 528
immaculatus Hatch (Hyperaspidius) 368
impexus (Mulsant) (Scymnus) 140, 151, 26, 32
impictipennis Weise (Mulsantina) 844
impletus Gordon (Scymnus) 242, 144, 147, 150
impunctus Wingo (Scymnus) 240
inaequalis (F.) (Coelophora) 1 7, 20, 830
inaequalis Casey (Pharoscymnus) 23
Nunenmacher (Axion) 615
indianensis Weise (Scymnus) 117,126
indubitabilis Crotch (Brachiacantha) 560, 596
indutus Casey (Scymnus) 274
inedita Mulsant (Hyperaspis) 409, 426
Casey (Scymnus) 169
inflexa Casey (Hyperaspis) 411, 490
infuscatus Boheman (Scymnus) 119
ingenitus Casey (Hyperaspidius) 359, 387
innocens Casey (Scymnus) 195
innocuus Casey (Scymnus) 268
inops Casey (Scymnus) 310
insignis Casey (Hyperaspidius) 358, 359, 382
insolens Casey (Hyperaspis) ' 437
insulatus Gordon (Cephaloscymnus) 69, 72
intermedia (Crotch) (Coccinella) 16
interrogans Mulsant (Hippodamia) 748
interrupta (Casey) (Myzia) 765, 769
interruptus Goeze (Scymnus) 26
intrusoides Hatch (Nephus) 308
intrusus (Horn) (Nephus) 310, 296
iowensis Casey (Scymnus) 207, 142, 145, 145, 150
irregularis Weise (Anisosticta) 683
jacinto Casey (Scymnus) 203
jacobianus Casey (Scymnus) 149, 203
japonica (Crotch) (Hyperaspis) 21
japonica (Thunberg) (Propyleae) 24
jasperensis Belicek (Hyperaspis) 409, 552
jejunus Casey (Stethorus) 27
johnsoni Casey (Coccinella) 786, 793
jovialis Fall (Hyperaspis) 408, 420
jucunda LeConte (Hyperaspis) 446
jucunda Mulsant (Hyperaspis) 21
Juliana Mulsant (Coccinella) 788
1985 NORTH AMERICAN COCCINELLIDAE 903
juncta Casey (Hippodamia) 741
juncta Johnson (Epilachna) 867
juniperus Nunenmacher (Hyperaspidius) 360
kansanus Casey (Scymnus) 162, 141, 143
kincaidi Hatch (Nephus) 290
kingi Macleay (Egleis) 18
/cmzW/ Casey (Scymnus) 195
kirbyi Crotch (Coccinella) 787, 814
koebelei (Ollifl) (Rodolia) 25
koebelei Nunenmacher (Psyllobora) 858
kunzii Schaeffer (Hyperaspis) 415
kuwanae Silvestri (Chilocorus) 15, 13, 644, 652
labiculata (Say) (Anatis) 754, 753
lacustris LeConte (Scymnus) 144, 145, 150, 268
laevipennis Casey (Hyperaspis) 450
laevis Gordon (Cephaloscymnus) 68, 73
lanei Hatch (Hyperaspis) 530
lanosus Blackburn (Erithionyx) 18
lateralis Mulsant (Hyperaspis) 406, 448
laticollis Casey (Zagloba) 84
latiusculus Casey (Exochomus) 627
leachi Nunenmacher (Hyperaspis) 408, 422
lecontei Casey (Anatis) 759, 753
lecontei Crotch (Hyperaspis) 446
lecontei Crotch (Scymnus) 191
lecontei Mulsant (Hippodamia) 708, 709, 733
lengi Johnson (Hippodamia) 719
lengi Nunenmacher (Brachiacantha) 582
lengi Schaeffer (Hyperaspis) 473
lengi Timberlake (Coleomegilla) 698
lepida LeConte (Coccidula) 656
lepida Mulsant (Brachiacantha) 599
leporina Mulsant (Hippodamia) 727
leucopsis Melsheimer (Hyperaspis) 423
levaillanti Mulsant (Scymnus) 26
levrati (Mulsant) (Hyperaspis) 459, 406
lewisi Crotch (Hyperaspis) 407, 448
lichatschovi Hummel (Buleae) 29
liebecki (Horn) (Diomus) 325, 3 1 8
liliputana Casey (Hippodamia) 747
limbalis Casey (Hyperaspis) 411, 497
limbata (Motschulsky) (Jauravia) 21
limbifer Casey (Cycloneda) 820, 822
lindi Blackburn (Rhyzobius) 24
linearis Gordon (Scymnus) 136
lineata (Thunberg) (Micraspis) 22
lineola (F.) (Micraspis) 22
litigiosa Mulsant (Naemia) 693, 696
lituratus Gorham (Exochomus) 18
lodi Stehr (Scymnus) 284
loewii Mulsant (Scymnus) 140, 143, 146, 148, 189
904
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
longicoxitis Nutting (Hyperaspis) 406, 482
longulum Casey (Didion) Ill, 110
lophanthae (Blaisdell) (Rhyzobius) 32, 25, 660
louisianae J. Chapin (Scymnus) 142, 144, 146, 187
luctuosus Casey (Scymnus) 266, 1 50
ludovicianus Casey (Nephus) 298
lugubris (Randall) (Hyperaspis) 406, 446
lunaris Weise (Diomus) 332
lunata (F.) (Cheilomenes) 15
lunatomaculata Motschulsky (Hippodamia) 708, 722
luteipes Mulsant (Azya) 14
luteodorsa J. Chapin (Mulsantina) 844, 849
luteopicta Mulsant (Coccinella) 16
mackenziei Chapin (Hippodamia) 734
maculatus Hatch (Nephus) 304
maculifera Melsheimer (Hyperaspis) 536
maculigerum Blackburn (Serangium) 27
maculiventris Bland (Epilachna) 866
maderi (Nunenmacher) (Selvadius) 349, 350
major Dobzhansky (Hyperaspis) 409, 444
majus, n. name (Scymnus) 211, 142
majusculus Wingo (Scymnus) 211
mali (Say) (Anatis) 762, 753
maneei Casey (Hyperaspis) 448
mannerheimi Mulsant (Coccinella) 816, 787
marginata (LeConte) (Coccidophilus) 45, 48
marginatus (Gaines) (Hyperaspidius) 358, 380
marginellus Mulsant (Nephus) 298
marginicollis Mannerheim (Scymnus) 140, 193
marginipennis (LeConte) (Exochomus) 624
martini Gordon (Scymnus) 264, 1 50
mckenziei Nutting (Hyperaspis) 408, 472
medialis Casey (Hyperaspis) 407, 462
medionotans Casey (Scymnus) 277
megacephalus (Fall) (Selvadius) 349, 351
melanocephalus Zoubk. (Exochomus) 18
melsheimeri Weise (Scymnus) 207
mendocino Casey (Scymnus) 230, 149
metallicus Korschefsky (Exochomus) 31, 18, 623, 640
metator Casey (Hyperaspis) 459
microsticta (Casey) (Brachiacantha) 597
micula, n. sp. (Gnathoweisea) 49, 52
militaris (LeConte) (Hyperaspidius) 358, 365
mimoides Gordon (Scymnus) 171, 148
mimus Casey (Hyperaspidius) 360, 362
mimus Fall (Scymnus) 232
minor Casey (Mulsantina) 845
minor Korschefsky (Diomus) 347
mmor Leng (Brachiacantha) 561
minuta (Casey) (Microweisea) 38, 39
minutissimus Casey (Diomus) 347
1985 NORTH AMERICAN COCCINELLIDAE 905
minutus Blackburn (Cycloscymnus) 17
misella (LeConte) (Micro weisea) 38, 40
moerens (LeConte) (Hyperaspis) 546, 4 1 2
moesta LeConte (Hippodamia) 707, 739
molliculus Casey (Diomus) 347
montana Casey (Myzia) 768
montanica Casey (Hyperaspis) 449
montezumae Mulsant (Thalassa) 400
monticola Casey (Scymnus) 213, 144
monticola Mulsant (Coccinella) 787,811
montrouzieri Mulsant (Cryptolaemus) 17,105
morelletti Mulsant (Scymnus) 26
mormon Casey (Scymnus) 218, 144, 147
mormonicus Casey (Exochomus) 631
mulsanti LeConte (Hippodamia) Ill
multiguttata Randall (Anisosticta) 683
munda (Say) (Cycloneda) 823, 820
myrmidon (Mulsant) (Diomus) 317, 326
nana Mulsant (Psyllobora) 854, 853
nanellus, n. sp. (Hyperaspidius) 358, 374
nanum (LeConte) (Didion) 111,110
natchezianus Casey (Scymnus) 211
nebulosus LeConte (Scymnus). . 1 16, 119
neglecta Mulsant (Lotis) 21
nemorivagus Wingo (Scymnus) 141, 182
neomexicanus Gordon (Scymnus) 146, 237
nemdensis (Leng) (Brumoides) 605
nevadensis Weise (Scymnus) 149, 268
nevadica Casey (Hyperaspis) 411, 512
nigellum Blackburn (Cyrema) 17
niger Casey (Nipus) 58, 56
nigerrima Casey (Lotis) 21
nigricollis Gordon (Scymnus) 146, 161
nigripennis (LeConte) (Helesius) 397
nigripes Kapur (Stethorus) 27
nigrocauda Dobzhansky (Hyperaspis) 450
Nunenrnacher (Hippodamia) 720
nigropennis Blatchley (Hyperaspis) 426
nigrosuturalis Blatchley (Hyperaspis) 406, 435
nodiceps Marshall (Cryptognatha) 30, 17, 600
normata Say (Hyperaspis) 423
notans Randall (Myzia) 768
Casey (Hyperaspis) 521
notescens Blackburn (Scymnus) 26
notulata Mulsant (Neoharmonia) 837
novascotiae Chapin (Hyperaspis) 534
novemnotata Herbst (Coccinella) 786, 798
nubes Casey (Scymnus) 192
nubilans Casey (Helesius) 398, 397
nubilata Casey (Mulsantina) 845
nubilatus (Casey) (Hyperaspidius) 378, 358
906
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY VoL 93(1)
nubilis Mulsant (Scymnus) 26
nugator Casey (Scymnus) 143, 146, 235
nummeralis (Boisduval) (Parapriasus) 23
nunenmacheri Casey (Hyperaspis) 411, 514
nunenmacheri Gordon (Selvadius) 349, 351
nupta Casey (Hyperaspis) 511
nuttingi Gordon (Scymnus) 239, 1 50
obliqua Casey (Hippodamia) 730
obliterata (L.) (Aphidecta) 14, 13, 840
oblongus Casey (Hyperaspidius) 359, 386
obscum Malkin (Hyperaspis) 540
obsoleta Casey (Psyllobora) 855
occidentalis Dobzhansky (Hyperaspis) 464
occidentalis Horn (Cephaloscymnus) 69, 71
occidentalis Horn (Coccidula) 657
occiduus Casey (Didion) Ill
occiduus Gordon (Nipus) 58, 56
octavia Casey (Hyperaspis) 413, 538
octomaculata (F.) (Harmonia) 20
octonotata Casey (Hyperaspis) 406, 428
oculaticauda Casey (Hyperaspis) 412, 526
oculatus (Blatchley) (Nephaspis) 102
oculifera Casey (Hyperaspis) 408, 457
ohioensis Stehr (Diomus) 325
omma Casey (Hyperaspis) 450
opaculus Horn (Scymnus) 138, 117
orbiculatus (Leng) (Brumoides) 605
orbigera Mulsant (Azya) 13, 29, 676
orbipennis Casey (Zagloba) 84
orbus Casey (Chilocorus) 648, 644
oregona Casey (Coccinella) 798
oregona Casey (Myzia) 772
oregona Dobzhansky (Hyperaspis) 410, 412, 413, 530
oregonensis Crotch (Hippodamia) 709, 746
omata (Horn) (Zagloba) 84, 83
omatella, n. sp. (Hyperaspis) 408, 454
omatus naviculatus (Casey) (Nephus) 287, 290
omatus omatus (LeConte) (Nephus) 287, 289
osculans LeConte (Hyperaspis) 408, 469
ovalis (LeConte) (Micro weisea) 38, 43
ovoideus (Casey) (Brumoides) 604
pacific a Casey (Brachiacantha) 594
pacificus Crotch (Scymnus) 142, 145, 148, 154
pallens LeConte (Scymnus) 166, 143, 146, 148
pallescens Casey (Hyperaspidius) 360, 372
pallescens Dobzhansky (Hyperaspis) 509
/7a///6(icc»/^z Blatchley (Psyllobora) 861
pallidula Dobzhansky (Hyperaspis) 509
pallidus (LeConte) (Delphastus) 62
pallidus Casey (Hyperaspidius) 364
paludicola Schwarz (Hyperaspis) 413,539
1985 NORTH AMERICAN COCCINELLIDAE 907
papago Casey (Scymnus) 144, 147, 255
paracanus J. Chapin (Scymnus) 119, 134
parcesetosa (Sicard) (Catana) 15
parenthesis (Say) (Hippodamia) 708, 715
parthenica Meyrick (Coelophora) 17
partitus Casey (Diomus) 338
parva Watson (Ceratomegilla) 704
parviceps Casey (Didion) Ill
parvicollis (Casey) (Brumoides) 610
parvinotata Casey (Psyllobora) 853, 861
paspalis Watson (Hyperaspis) 423
pauculus Gordon (Scymnus) 164, 143, 146
pellio Blatchley (Diomus) 320
peninsularis Gordon (Scymnus) 142, 275
perdistinctus Kapur (Catana) 15
perpallida Dobzhansky (Hyperaspis) 420
perplexa Mulsant (Coccinella) 786, 787
phelpsii Crotch (Scymnus) 120
phosphorus Lewis (Nephus) 23
picta (Randall) (Mulsantina) 844
pilatii Mulsant (Axion) 617
pinachi Gordon and Chapin (Stethorus) 90, 93
pinguis Casey (Hyperaspis) 407, 452
pinorum Casey (Hyperaspis) 426
pistillata Watson (Hyperaspis) 408, 409, 434
plagiata Casey (Olla) 828
plagiata Schaeffer (Psyllobora) 853, 858
plagiatum (Olivier) (Axion) 615, 617
planatus Gordon (Nipus) 58, 56
planiceps (Casey) (Gnathoweisea) 49, 51
pleuralis Casey (Hyperaspis) 410, 41 1, 504
pleuralis LeConte (Axion) 619
ploribundus (Nunenmacher) (Hyperaspidius) 359, 368
pluto Fall (Hyperaspis) 454, 406
polita Casey (Cycloneda) 825, 820
politissima Casey (Hippodamia) 741
postica LeConte (Hyperaspis) 412, 516
postpictus (Scymnus) 143, 146, 149, 194
praetextatus Melsheimer (Exochomus) 624
pratensis LeConte (Hyperaspis) 407, 461
Mulsant (Hippodamia) 741
proba (Say) (Hyperaspis) 414, 407
prolongata Crotch (Coccinella) 786, 800
protensa Casey (Hyperaspis) 411, 547
pseudapicanus, n. name (Scymnus) 1 18, 134
pseudotaedatus Gordon (Diomus) 317, 329
psyche Casey (Hyperaspis) 503, 4 1 2
pugetana Casey (Hippodamia) 121
pulchella (Mulsant (Coccinellina) 17
pulcher Blackburn (Rhyzobius) 25
pullata (Say) (Myzia) 767, 765
908
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
pulvinatus Wingo (Scymnus) 140, 181
pumilio Weise (Diomus) 336, 17, 18, 30, 317
punctata LeConte (Hyperaspis) 413, 549
punctatum (Melsheimer) (Didion) 109, 110
puncticollis (LeConte) (Delphastus) 64
puncticollis Casey (Hippodamia) 747
puncticollis LeConte (Scymnus) 142, 259
punctillum Weise (Stethorus) 33, 27, 29, 90, 96
punctulata LeConte (Hippodamia) 730
punctum picipes Casey (Stethorus) 90, 96
punctum punctum (LeConte) (Stethorus) 90, 95
purit anus Casey (Scymnus) 167
pusillus (LeConte) (Delphastus) 62, 64
pusio Casey (Diomus) 347
pygmaeus Blackburn (Midus) 22
quadraria Casey (Hippodamia) 727
quadrarius (Casey) (Nephus) 296, 305
quadrillum LeConte (Brachiacantha) 559, 569
quadrimaculatus (Blackburn) (Sticholotis) 28
quadrioculata (Motschulsky) (Hyperaspis) 412, 413, 521
quadriplagiatus (Swartz) (Menochilus) 22
quadripunctata (Melsheimer) (Brachiacantha) 560, 591
quadripustulatus (L.) (Exochomus) 31,19, 624, 636
quadristicta (Telsimia) 28
quadritaeniatus LeConte (Diomus) 320
quadrivittata LeConte (Hyperaspis) 410, 545
quadrivittatus Mulsant (Scymnus) 26
quatuordecimguttata (L.) (Calvia) 14, 775
quatuordecimpunctata (L.) (Propyleae) 24, 29, 31, 817
querceti Schwarz (Brachiacantha) 559, 588
querquesi Nutting (Hyperaspis) 412,529
quindecimmaculata Hope (Harmonia) 834
quindecimmaculata Mulsant (Hippodamia) 708, 737
quindecimpunctata Olivier (Anatis) 754, 762
quindecimspilota Hope (Harmonia) 834
quinquesignata (Kirby) (Hippodamia) 708, 709, 727
randalli Casey (Myzia) 768
rathvoni (LeConte) (Anatis) 753, 756
rectus Casey (Selvadius) 350, 349
regalis Casey (Hyperaspis) 426
reitteri Dodge (Coccidula) 657
renifer Casey (Psyllobora) 853, 862
renoicus Casey (Scymnus) 215, 144, 149
reversa (Fall) (Coccidophilus) 47
revocans Casey (Hyperaspis) 406, 464
rhesus Casey (Scymnus) 253
richardsoni Brown (Coccinella) 786, 790
rivularis Dobzhansky (Hyperaspis) 409, 4 1 7
roseicollis (Mulsant) (Diomus) 317,341
rossi Nunenmacher (Hyperaspidius) 369
rotunda Casey (Hyperaspis) 409, 475
1985 NORTH AMERICAN COCCINELLIDAE 909
rotunda, n. sp. (Brachiacantha) 560, 575
rubidus tristis Falderman (Chilocorus) 15
rubricaudus Casey (Scymnus) 141, 143, 175
rubripennis Casey (Cycloneda) 820
rufa Nunenmacher (Neoharmonia) 839
rufescens Dobzhansky (Hyperaspis) 485
ruficollis Blackburn (Rhyzobius) 25
rufomarginata Mulsant (Hyperaspis) 488
rufopilosa Mulsant (Rodolia) 25
rusticus Casey (Scymnus) 126
saginatus Casey (Scymnus) 209
sanctaeritae Dobzhansky (Hyperaspis) 411, 487
sanctus Weise (Scymnus) 220
sanguinea (L.) (Cycloneda) 820
sanguinifer Casey (Nephus) 289
Casey (Scymnus) 192
satana, n. sp. (Zagloba) 86, 84
satellus Blackburn (Rhyzobius) 35
saucia Mulsant (Lemnia) 21
sauzeti Mulsant (Oenopia) 23
sayi Crotch (Olla) 826
schaefferi, n. sp. (Exoplectra) 671
schaefFeri, n. sp. (Hyperaspis) 411, 496
schuberti Nunenmacher (Nephus) 305
schuhi Hatch (Hyperaspis) 540
schwarzi Gordon (Gnathoweisea) 49, 50
schwarzi Gordon (Nephus) 315
schwarzi, n. sp. (Brachiacantha) 559, 585
scitus Casey (Nephus) 302
Dobzhansky (Hyperaspis) 521
scotti Nunenmacher (Scymnus) 192
securus J. Chapin (Scymnus) 141, 285
semilunaris Johnson (Olla) 828
semiruber Horn (Scymnus) 167, 141
senegalensis hottentota Mulsant (Hyperaspis) 31,21
separata Casey (Hyperaspis) 447
separata Casey (Psyllobora) 856
separata Leng (Brachiacantha) 564
septempunctata L. (Coccinella) 16, 786, 795, 29, 13
septentrionis (Weise) (Brumoides) 604
septentrionis Dobzhansky (Hyperaspis) 480
sequoiae Dobzhansky (Coccinella) 785, 801
serena Casey (Hyperaspis) 490
seriata (Melsheimer) (Naemia) 693, 694
sexmaculatus (F.) (Menochilus) 22
sexualis (Casey) (Blaisdelliana) 355
shauli Nunenmacher (Hyperaspidius) 359, 389
sidneyensis Blackburn (Scymnus) 26
signata bicentralis Casey (Hyperaspis) 408, 432
signata signata (Olivier) (Hyperaspis) 408, 429, 409
signifera (Reiche) (Lioadalia) 21
910
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 93(1)
significans Casey (Hyperaspis) 409, 484
similis Casey (Paranaemia) 692
simulans Casey (Hyperaspis) 410, 41 1, 550
simulans Gordon (Scymnus) 244, 145
simulatrix Dobzhansky (Hyperaspis) 413, 520
simulatus, n. sp. (Hyperaspidius) 360, 370
simuloides Hatch (Hyperaspis) 540
sinuata Mulsant (Hippodamia) 707, 748
sobrina Casey (Olla) 828
socer LeConte (Scymnus) 195, 142
socialis Casey (Brachiacantha) 564
solidus Casey (Scymnus) 200, 145, 148, 150
soltaui, n. sp. (Brachiacantha) 558, 566
sonomae Casey (Scymnus) 266
sonorana Casey (Brachiacantha) 582
sordidus (Horn) (Nephus) 308, 296
soulary i Mulsant (Neoharmonia) 838
speculifer Blackburn (Rhyzobius) 35
spiculinota Fall (Hyperaspis) 413, 543
spuria LeConte (Hippodamia) 709, 750
stellata Casey (Brachiacantha) 579
stephani, n. sp. (Brachiacantha) 559, 571
stigma (Say) (Chilocorus) 651, 644
stigma Casey (Diomus) 332
strabus Horn (Scymnus) 154
straminea Chapin (Hippodamia) 750
strenua (Casey) (Coleomegilla) 698, 699
St renuus Casey (Scymnus) 210
stygicus Casey (Scymnus) 210
suavis Casey (Nephus) 302
subdepressa Casey (Hyperaspis) 526
subfasciata Mulsant (Brachiacantha) 558, 559, 574
subrotundus Casey (Exochomus) 624, 630
subsimilis Casey (Hippodamia) 727
subsimilis Casey (Scymnus) 218
subtropicus (Casey) (Zilus) 76, 80
subtropicus Casey (Scymnus) 277
sub versa LeConte (Coccinella) 786, 788
subviridis (Blackburn) (Telsimia) 28
subvittata (Mulsant) (Myzia) 765, 769
suturalis (F.) (Brumoides) 14
suturalis (Schwarz) (Microweisea) 38
suturalis Casey (Coccinella) 804
suturalis Thunberg (Scymnus) 32, 26, 29, 151
suturalis Weise (Coccidula) 657
taedata LeConte (Hyperaspis) 429
taedata LeConte (Psyllobora) 855
taedatus (Fall) (Diomus) 330, 3 1 7
taeniata LeConte (Hyperaspis) 412,413,509
tahoensis Casey (Scymnus) 270, 144, 150
tau LeConte (Brachiacantha) 559, 562
1 985 NORTH AMERICAN COCCINELLIDAE 9 1 1
tenebricus Gordon (Scymnus) 147, 150, 248
tenebrosus Mulsant (Scymnus) 141
tenebrosus Mulsant (Scymnus) 144, 197
tenuivestis Casey (Scymnus) 210
terminatus (Say) (Diomus) 337, 3 1 7
testudinaria Mulsant (Neda) 22
testudo Casey (Brachiacantha) 560, 585
tetraneura Casey (Hyperaspis) 545
tetraspilota (Hope) (Adalia) 14
tetrasticta Casey (Telsimia) 28
texana, n. sp. (Gnathoweisea) 49, 52
texanus Casey (Scymnus) 175
texanus Gordon (Diomus) 339, 3 1 7
texanus LeConte (Axion) 617
tibialis (Say) (Hippodamia) 707, 709
timberlakei, n. sp. (Nephus) 296, 312
toowoombae Blackburn (Rhyzobius) 660
toweri Johnson (Epilachna) 867
townsendi Casey (Exochomus) 623, 631
transfugatus Casey (Hyperaspidius) 358, 363
transversalis F. (Coccinella) 16
tredecimnotata (Latreille) (Epilachna) 865, 871
triangulum Casey (Hyperaspis) 407, 466
tricolor Nunenmacher (Hippodamia) 720
tricuspis Kirby (Coccinella) 814
tricyclus Smith (Chilocorus) 644, 652
tridens Kirby (Hippodamia) 717
trifurcata Schaeffer (Hyperaspis) 410, 556
trimaculata (L.) (Hyperaspidius) 390
trinifev Casey (Hyperaspis) 414
trinitatis (Marshall) (Pseudoazya) 24, 678
triplicans (Casey (Brachiacantha) 597
tripustulatum (Degeer) (Axion) 615
tristis (LeConte) (Hyperaspidius) 359, 366
trivittata Casey (Hippodamia) 748
troglodytes Mulsant (Brachiacantha) 579
troglodytes Mulsant (Hyperaspis) 410, 534
tuckeri Casey (Hyperaspis) 408, 4 1 9
tumidus Leng (Chilocorus) 647, 644
ulkei Crotch (Ceratomegilla) 704
uncus Wingo (Scymnus) 256, 1 42
undecimpunctata L. (Coccinella) 16, 809, 30, 29, 787
undulata (Say) (Hyperaspis) 413, 536
uniformis Casey (Hyperaspis) 409, 500
uropygialis Mulsant (Exochomus) 19
ursina (F.) (Brachiacantha) 560, 580
utahensis Gordon (Scymnus) 145, 150, 214
uteana Casey (Hippodamia) 727
uteana, n. sp. (Hyperaspis) 407, 471
uteanus Casey (Scymnus) 253, 144, 147
uteella Casey (Brachiacantha) 560, 582
912
JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY VoL 93(1)
utilis (Horn) (Stethorus) 90
v-nignim (Mulsant) (Olla) 826
vagans (Blackburn) (Stethorus) 27
vandykei Nunenmacher (Coccinella) 804
variegata (Goeze) (Hippodamia) 20
varivestis Mulsant (Epilachna) 30, 29, 865, 866
ventralis (Erichson) (Rhyzobius) 35, 662
venusta (Melsheimer) (Neoharmonia) 837
venustulus (Mulsant) (Hyperaspidius) 358, 381
vernix Casey (Hippodamia) 727
verrucatus (Melsheimer) (Axion) 615
vicksburgicus Casey (Scymnus) 240
vigintiduopunctata (L.) (Psyllobora) 24
vigintimaculata (Say) (Psyllobora) 853, 855
vigintiquatuorpunctata (L.) (Subcoccinella) 32, 20, 874
virginalis (Wickham) (Anovia) 668
viridipennis Mulsant (Neoharmonia) 837
vittigera (Mannerheim) (Paranaemia) 691
vittigerus (LeConte) (Hyperaspidius) 359, 390
wahlbergi Mulsant (Chilocorus) 16
Wallace! Crotch (Cryptolaemus) 17
Washington! Timberlake (Hippodamia) 707, 715
weidti Casey (Scymnus) 260, 147, 149
weisei Schaeffer (Hyperaspis) 407, 415
wellmani Nunenmacher (Hyperaspis) 450
whittonensis Blackburn (Scymnus) 26
wickhami Casey (Hyperaspis) 408, 442
wickhami Gordon (Nephus) 307, 296
wickhami Gordon (Scymnus) 250, 144, 150
wingoi Gordon (Scymnus) 245, 142
wolcotti (Nunenmacher) (Hyperaspidius) 359, 385
xanthaspis (Mulsant) (Diomus) 317, 343
zimmermanni Crotch (Cephaloscymnus) 68, 69
/
■k*'.
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Journal of the
New York Entomological Society
VOLUME 93 JANUARY 1985 NO. 1
CONTENTS
The Coccinellidae (Coleopiera) of America North of Mexico
Robert D. Gordon 1-912
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