Table Of ContentPROC. ENTOMOL. SOC. WASH.
93(4), 1991, pp. 845-864
FIELD GUIDE TO RECENTLY INTRODUCED SPECIES OF
COCCINELLIDAE (COLEOPTERA) IN NORTH AMERICA, WITH A
REVISED KEY TO NORTH AMERICAN GENERA OF COCCINELLINI
Robert D. Gordon and Natalia Vandenberg
Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, % U.S.
National Museum ofNatural History, Washington, D.C. 20560.
Abstract.—Six species of predaceous Coccinellidae are being released for biological
control ofthe Russian wheat aphid, Diuraphis noxia (Mordvilko), in North America. The
following3 speciesarenowestablished: Hippodamia (Adonia) variegata(Goeze), Propylea
quatuordecimpunctata (L.), and Coccinella septempunctata L. Hippodamia (Semiadalia)
undecimnotata (Schneider), Oenopia conglobata (L.), and Scymnusfrontalis (F.) are not
known to be established. Two additional species ofpredaceous Coccinellidae are recent
adventive additions to the North American fauna, Harmonia axyridis (Pallas) in Loui-
siana, Mississippi, and Georgia, and Harmonia quadripunctata (Pontopiddian) in New
Jersey and New York. The key to North American genera ofCoccinellini is revised.
Key Words: predaceous Coccinellidae, introduced species, Russian wheat aphid, bio-
logical control, key to genera ofCoccinellini
Discovery of the introduced Russian Hippodamia {Adonia) variegata (Goeze),
wheat aphid, Diuraphis noxia (Mordvilko), Propylea quatuordecimpunctata (L.), and
in the western United States prompted re- Coccinella septempunctata L. are estab-
search on predators and parasites of that lished. Hippodamia {Semiadalia) undecim-
species by Federal and state biological con- notata(Schneider), Oenopiaconglobata(L.)
trol laboratories. This research has resulted and Scymnusfrontalis (F.) are not known
in the introduction, propagation, and re- to be established.
lease ofseveral Old World species ofCoc- In addition, two adventive species, Har-
cinellidae. The USDA Animal and Plant monia quadripunctata (Pontopiddian) and
Health Inspection Service (APHIS) is the Harmonia axyridis (Pallas), have become
organization mainly responsible for rearing established in the United States (Vanden-
and releasing foreign Coccinellidae for bi- berg 1990, J. Chapin and Brou 1991, C. L.
ological control ofthe Russian wheat aphid Smith 1991, pers. comm.). Although it is
through the APHIS National Biological unlikelyeitherofthese specieswillbe found
Control Laboratory, Niles, Michigan. Per- in association with Russian wheat aphid,
sonnel involved with such releases need to theyareillustratedandbrieflycharacterized
identify the introduced species and distin- here.
guish them from native species; hence, the Adultsofthespeciesdiscussedbelowusu-
preparation ofthis "field guide." ally can be distinguished from each other
Six species ofCoccinellidae have thus far and from nativecoccinellidsbydorsal color
beenpropagatedandreleased specifically for pattern alone. Nothing so general can be
controloftheRussianwheataphid. Ofthese. said about the larval stages; therefore, the
.
846 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
larvae of the introduced species are illus- the only species ofHarmonia then known
tratedasabeginningstepinthe preparation to occur in North America. Further, Gor-
of an identification guide to larvae of all don's original key does not work for Neo-
species ofCoccinellidae occurring in North harmonia Crotch, because the lack oftibial
America, native and introduced. The ac- spurs was overlooked during construction
companying larval diagnoses were made of the key. Chapin and Brou (1991) pro-
from live specimens in orderto be useful to duced a modified key that corrected these
field personnel; the illustrations were pre- shortcomings; we have incorporated her
pared from preserved specimens. Because changes along with further modifications to
colors are lost and pigments fade in pre- accommodate newly imported species and
served specimens, the diagnoses differ to improve keyability. The figure numbers in
somedegree from the illustrations. A larval the key refer to illustrations in Gordon
key cannot be prepared now because the (1985).
larvalstagesofmostnativespecieshavenot
Revised Key to the Native and
been described. Habitus illustrations and
Introduced Genera of Coccinellini
"diagnoses" based on color pattern are pre-
OF North America
sented in lieu ofa key because personal ex-
perience suggests that most larvae of the 1. Tarsal claw not toothed or cleft, simply
subfamilyCoccinellinaecanbeidentifiedby widened basally (Fig. 567a) 2
Tarsalclawtoothedorapicallycleft(Figs.
dorsal color pattern.
587i, 614c) 5
Larvae bear various rounded or conical 2(1). Pronotal base with fine, entire marginal
fleshy protuberances armed with setae. For bead 3
oursimplecolorpatterndiagnoses, wehave Pronotalbase not margined 4
abandoned complex morphological cate- 3(2). Metastemumwithpostcoxalline;elytron
gories and refer to these structures collec- with large black spots (Fig. 570g)
Naemia Mulsant
tivelyaslobes. Eachabdominalsegmenthas Metastemumwithoutpostcoxalline;ely-
six lobes visible in dorsal view, three on tron vittate (Fig. 567g) Paranaemia Casey
. .
each side (dorsal, dorsolateral, lateral). The 4(2). Apex ofmiddleand hind tibiaeach with
pleuralregionsofthemeso-andmetathorax 2 spurs; elytron vittate (Fig. 5650; epi-
areeachequippedwith a smallanteriorand pleuron slopingdownward internally. .
Macronaemia Casey
a larger posterior lobe. Each thoracic seg-
- Apex ofmiddleand hind tibiaeach with
ment has a pair ofdorsal plates or tergites. singlespur; elytron spottedorvery irreg-
The dorsal plates ofthe pronotum may be ularly vittate(Fig. 560f); epipleuron hor-
laterally constricted or completely subdi- izontal Anisosticta Dejean
vided by membranous areas. Morphologi- 5(1). Eachtarsalclawcleftnearapical 1/3(Fig.
587i);form slender, legsdistinctlyvisible
cal details ofthe various species were treat-
beyond body in dorsal view
ed by Savoiskaya (1983). Hippodamia Dejean
All ofthe Russian wheat aphid predators Each tarsal claw with subquadrate basal
except S.frontalis belong to the tribe Coc- tooth(Fig. 614c); oriftoothmedian then
cinellini. Adultsofthe respectivegeneraare form rounded, legsbarely visible beyond
diagnosed in the following key. Scymnus body in dorsal view (genus Myzia) 6
6(5). Apex of middle and hind tibia without
frontalis belongs to the Scymninae and is spurs 7
treated separately. Apexofmiddleandhindtibiawith2spurs
Gordon's (1985) key to genera ofCocci- (Fig. 626a) 10
nellini includes the genus Harmonia Mul- 7(6). Postcoxallineon 1stabdominal sternum
recurvedtowardbaseofsternum, ofPul
sant; however, H. axyridis and H. quadri-
lustype (Fig. 679a) Aphidecta Mulsant
punctata will not key out because couplet 9 Postcoxallineon 1stabdominalsternum
wasconstructedtodistinguishH. dimidiata, not recurved, ofDiomusorNephustype.
.
VOLUME 93, NUMBER 4 847
butwithobliquedividinglinesometimes Pronotalsurfacealutaceous,oftendull,not
present(Figs. 674, 677, 682) 8 polished; anterior margin of mesoster-
8(7). Postcoxal lineon 1stabdominal sternum numtruncate,withshallowemargination,
without oblique dividing line (Fig. 682a) orwith deepbut narrowemargination . . 17
Mulsantina Weise 17(16). Pronotum black with large, subtrapezoi-
Postcoxallineon 1stabdominal sternum dal or triangular white spot on each an-
with obliquedividingline 9 terolateralangle;hindpronotalanglemuch
9(7). Lateral margin of elytron transparent, morebroadlyroundedthananteriorangle
withoutmarginalbead;prostemalcarinae Coccinella L.
endingat anteriorcoxal margin - Pronotal color pattem not as described
NeoharmoniaCrotch above, or ifso, then hind pronotal angle
Lateralmarginofelytronnottransparent, notmuchmorebroadlyroundedthanan-
with more orlessdistinctmarginal bead; teriorangle 18
prostemal carinae extending anterior to 18(17). Pronotum with pale spot on each side of
frontcoxal margin orabsent; Old World middle, spot entirely enclosed by black
genuswith 3 speciesestablishedinNorth area or spot connected to pale anterior
America Harmonia Mulsant border Cycloneda Crotch
10(6). Pronotal base with marginal bead 11 Pronotum not asdescribed above 19
Pronotal base without marginal bead . . 12 19(18). Second tarsal segment elongate, about
11(10). Metastemum, 1stabdominalsternumwith twice as long as wide, or hind pronotal
distinct postcoxal line (Fig. 1) angle not much more broadly rounded
Ceratomegilla Crotch thananteriorangle;OldWorldgenus,one
Metasternum, 1st abdominal sternum species released but not known to be es-
without postcoxal line tablished in North America
Coleomegilla Timberlake Oenopia Mulsant
12(10). Prostemum strongly convex medially, Second tarsal segment short, triangular,
protuberant at apex (Fig. 614b); length not more than 1.5 timesas longas wide;
7.20 mm orgreater AnatisMulsant hind pronotal angle much more broadly
Prostemum normally rounded, not pro- rounded than anteriorangle 20
tuberant at apex; length variable 13 20(19). Distal antennal segment elongate, oval;
13(12). Postcoxallineon 1stabdominalstemum scutellum with base slightly longer than
complete, ofPullustype (Fig. 637a) . . side;maculationonelytronusuallyform-
Adalia Mulsant ingyellowandblack"checkerboard"pat-
Postcoxallineon 1stabdominal stemum tem; European genus, one species estab-
incomplete, of Diomus or Nephus type lished in North America Propylea Mulsant
. .
(Figs. 634b, 682a) 14 Distalantennalsegmentshort,robust,ob-
14(13). Hind margin ofmesepimeron with me- triangular; scutellum with side slightly
dian,triangularprojection;elytronorangy longer than base; elytron black with red
yellowwithblacksuturalmargin,4irreg- spotorpalewithminutedarkspotsnever
ularblackspots(C inaequalisonly)(Fig. forming "checkerboard" pattem; native
672g); Orientalgenus, onespecies(C in- North American species Olla Casey
aequalis F.) possibly established in Flor-
ida, Hawaii, and Puerto Rico
Coelophora Mulsant Coccinella septempunctata L.
Hind margin ofmesepimeron straight or Figs, (adult) 1, 13a-d; (larva) 20
curved, without projection; elytron with — mm
colorpattem notas above; North Amer- Adult diagnosis. Length 6.5 or
ican orOld Worldgenera 15 more; headblackwith 2 well-separatedpale
15(14). Tarsalclawwithmediantooth(Fig.626a); spots; pronotumwithanteriormargin black
elytron vittate or immaculate
at middle with ventral pale spot small, ex-
MyziaMulsant
Tarsalclawwith subquadratebasaltooth tending posteriorly '/3 as far as dorsal spot;
(Fig.664b);elytronvariable,nevervittate elytron with 3 black spots in addition to
16 scutellar spot; tarsal claw with large basal
16(15). Pronotalsurfacepolished,shiny, notalu-
tooth.
gtianceoofumsebseotwseteenmpuumncwtiutrhesd;eeapn,tebrriooard,matrri-- Diagnosis of 4th larval instar.—Body
angularemargination Calvia Mulsant mostly black or dark bluish gray; head yel-
PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON
Figs. 1^. Habitusviews. 1.Coccinellaseptempunctata.2,3.Propyleaquatuordecimpunctata.4.Hippoda
unaecimnotata.
VOLUME 93, NUMBER 4 849
lowexceptbasal, lateral margins black; pro- ten with spots rectangular, forming "check-
notum with apical margin narrowly yellow, erboard" pattern.
lateral margin broadly yellow; metapleuron Diagnosis of 4th larval instar.—Body
with posterior '/a of posterior lobe bright mostly black or at least very dark brown;
orange; 1st and 4th abdominal segments head yellow with posterolateral margin
with dorsolateral, lateral patches, including brown; pronotum pale yellow except dorsal
lobes, bright orange. tergite dark brown; mesonotum, metano-
Current American distribution.—All of tum pale yellow between tergites; meso-
the United States and southern Canada, pleuron, metapleuron with anterior, poste-
marginally established in California and riorlateral lobespale yellow; 1st abdominal
Nevada. segmentwith dorsum between tergites, dor-
—
Comments. Coccinellaseptempunctata, solateral, lateral lobes pale yellow; abdom-
or C-7 as it is commonly known, is a wide- inalsegments2, 3, 5-8withnarrow, median
spread palearctic species that was inten- dorsal area, lateral lobe pale yellow; 4th ab-
tionally introduced and released in North dominal segment with median area includ-
America several times from 1956 to 1971. ingdorsallobe,dorsolaterallobe, laterallobe
Those attempts were apparently unsuccess- pale yellow.
ful but an established population was dis- Current American distribution.—From
coveredin BergenCo., NewJersey, in 1973. the vicinity of Montreal, Quebec, south
This population is thought to have resulted along the St. Lawrence River to northern
from an accidental introduction (Angalet NewYork, Maine, and Vermont. One spec-
andJacques 1975). Since 1973, C-7 hasbeen imen examined from Massachusetts.
colonized and released in every state and in Comments.—Propv/^a quatuordecim-
southern Canada, and it is now the most punctata, or P-Q, is another palearctic in-
commonly collected species of Coccinella tentionally released in North America sev-
east of the Rocky Mountains. The advent eraltimeswithoutsuccessfulestablishment.
of the Russian wheat aphid caused an in- It is almost certainly an adventive species
crease in rearing and distribution efforts in in North America. Chantal (1972) was the
the western states that have resulted in at first to report on an established population
least marginal establishment of C-7 inQuebec,andDysart(1988)presentednew
throughout the west. locality records extending into the northern
References.—Angalet andJacques (1975); United States. This species is being reared
Angalet et al. (1979); Tedders and Angalet and released for Russian wheat aphid con-
(1981); Hoebeke and Wheeler (1980); Gor- trol in the western United States and Can-
don (1985); Schaefer et al. (1987); Schaefer ada, but thusfarthere is noevidenceofnew
and Dysart (1988); Hodek (1973) (larva il- establishment. InadditiontoCanadian ma-
lustrated in color). terial, P-Q has been brought in for culture
from France, Turkey, and several locations
in the USSR.
Propylea quatuordecimpunctata (L.) References.—Chantal (1972); Gordon
Figs, (adult) 2, 3, 14a-<i, 15d; (larva) 21 (1985); Dysart (1988); Hodek (1973) (larva
—
Adult diagnosis. Length 3.50 to 5.20 illustrated in color); Schaefer and Dysart
mm; maleheadusuallyyellowexceptvertex (1988); Wheeler (1990).
black, prostemalplategrayishwhite; female
head usually with black spot on clypeus, Hippodamia variegata (Goeze)
prostemalplateblack;pronotumyellowwith Figs, (adult) 5, 6, 15a-c; (larva) 22
large, irregular, blackarea medially; elytron Adultdiagnosis.—Length4.40to 5.0 mm;
yellow with variable black maculation, of- base ofpronotum with fine marginal bead;
850 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
Figs. 5-8. Habitus views. 5, 6. Hippodamia variegata. 7. Oenopia conglobata. 8. Scymnusfrontalis.
VOLUME NUMBER
93, 4 851
headyellowwith vertexblack (male) oryel- 1981 without successful establishment.
low with vertex and large frontal spot black Gordon (1987) reportedthe 1984 establish-
(female); pronotum blackwith anterior, lat- mentin Quebec. Since 1984, and especially
eralborders, smallspotoneachsideofmid- sincetheadventoftheRussianwheataphid,
dle yellow (female) or with anterior border many releases, most involving Canadian
of black area deeply emarginate medially stock, havetakenplaceintheUnitedStates,
with yellow, spot on each side of middle thus far without evidence ofnew establish-
broadlyconnectedtoyellowanteriorborder ment. In addition to Canadian material,
(male); elytron orange with 5, 6 or 7 black specimens have been brought into culture
spots: scutellar, posthumeral, 2 postdiscal, from France, Morocco, and several loca-
apical spot constant; humeral, postscutellar tions in the USSR.
spots or both often absent; ventral surface This species is currently classified in the
blackexceptpropleuron, meso-andmetepi- Old World as Hippodamia (Adonia) varie-
meron yellow, anterior coxa white. gata by lablokoff-Khnzorian (1982).
Diagnosis of 4th larval instar.—Body References.—Gordon (1987); lablokoff-
mostly light grayish blue; head brown ex- Khnzorian (1982); Hodek (1973) (larva il-
cept median area posterior to mouthparts lustrated in color); Schaefer and Dysart
yellow; pronotum pale yellow except dorsal (1988).
tergite black; mesonotum and metanotum
Hippodamia (Semiadalia) undecimnotata
pale yellow between tergites; mesopleuron
(Schneider)
withanterior, posteriorlobesfaintlyyellow;
metapleuron with posterior lobe pale yel- Figs, (adult) 4, 16a-c; (larva) 23
low, with medianareaoflobebrightyellow; Adult diagnosis.—Length 5.0 to 7.0 mm;
1stabdominal segmentpaleyellowbetween pronotum with raised basal margin; head
dorsal and dorsolateral lobes, with dorso- yellow with vertex black (male) or yellow
lateral and lateral lobes bright yellow; all with vertex, clypeus black, black area on
abdominal segments with faint yellow area clypeus often connected to vertex (female);
between dorsal and dorsolateral lobes; 4th pronotum black with anterolateral angle
segmentwith areabetween dorsolateral and broadly yellow (female) or with anterior
lateral lobes, lateral lobe yellow; leg black. bordercompletelyyellow(male);elytronor-
Current American distribution.—Vicini- ange with 4, 5 or 6 black spots, scutellar,
ty ofMontreal, Quebec, and scattered areas humeral, postdiscal spots constant, apical,
in eastern Canada. lateral spots often absent; ventral surface
Comments.—Adults ofHippodamia var- blackexceptpropleuron, epipleuronyellow.
iegata, or H-V as it is commonly known, Diagnosis of 4th instar larva.—Body
can be immediately distinguished from na- mostly orangy with rosy tints; head entirely
tive American species and H. undecimno- dark brown; pronotum with dorsal tergite
tata by the distinctly raised margin at the dark brown; mesonotum, metanotum with
base ofthe pronotum. Thedorsal colorpat- tergites dark brown, anterior margin gray;
tern is also unlike any ofthe other species mesopleurondarkgrayexceptanteriorlobe,
except for a superficial resemblance to H. posterior Vi ofposterior lobe yellow; meta-
convergens Guerin, which has the anterior pleuron gray except posterior lobe mostly
coxa black. A curious parallel exists be- yellow; 1st, 4th abdominal segmententirely
tween this species and P-Q in that both yellowexceptdorsal lobedarkbrown, small
were first found established in Quebec and area anterior to dorsal lobe, narrow area
bothareapparentlyadventive. Hippodamia between dorsolateral, lateral lobes gray; ab-
variegata was also introduced into North dominal segments 2, 3 mostly gray except
America many times between 1957 and dorsum with basal margin, areas between
PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
852
%»%«
V
1 -
I
I
11
Figs. 9-12. Habitus views. 9-11. VariationsofHarmonia axyridis. 12. Harmonia quadripunctata.
VOLUME 93, NUMBER 4 853
Fig. 13. a-d. Genitalia ofCoccinellaseptempunctala. a-c. Malegenitalia, d. Femalegenitalia.
PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
854
I:^vf
J1^$$J<^
Fig. 14. a-d. Malegenitalia ofPropylea quatuordecimpunctata.
dorsal lobes, dorsal, dorsolateral lobes yel- Comments.-This species was received
low;abdominalsegments 5-8 mostlyyellow by the Agriculture Research quarantine fa-
except all lobes dark brown, apical margin cility in Newark, Delaware, as a contami-
narrowly gray. nant in shipments of C-7 from the Soviet
Current American distribution.—Not Union in 1989. It has been reared and re-
known to be established in America. leased in the western United States against
