Table Of ContentPROC. ENTOMOL. SOC. WASH.
93(3), 1991. pp. 630-635
HARMONIA AXYRIDIS (PALLAS), THE THIRD SPECIES OF THE
GENUS TO BE FOUND IN THE UNITED STATES
(COLEOPTERA: COCCINELLIDAE)
J. B. Chapin and V. A. Brou
(JBC) Department of Entomology, Louisiana Agricultural Experiment Station, Loui-
siana State University Agricultural Center, Baton Rouge, Louisiana 70803-1710; (VAB)
74320 Jack Loyd Road, Abita Springs, Louisiana 70420.
Abstract.—Harmonia axyridis (Pallas), an Old World Coccinellini, is reported as es-
tablished in southeastern Louisiana. A few specimens were also collected in eastern Mis-
sissippi. The species is described, illustrated, and compared to H. dimidiata (F.) and H.
quadripunctata (Pontopiddian).
Key Words: Coleoptera, Coccinellidae, Harmonia
A lady beetle, not previously known to was made in Shreveport, Bossier [sic] Par-
beestablished in the United States, wascol- ish, on August 24, 1979, and eight releases
lected near Abita Springs, St. Tammany of3781 specimensweremadein Lelandand
Parish, and in othersoutheastern Louisiana Stoneville in Washington County, Missis-
parishes. A few specimens were also col- sippi, from July 12 to October 23, 1980 (P.
lected in eastern Mississippi. Harmoniaax- W. Schaefer, pers. comm.). There were no
yridis (Pallas) occurs in southern Siberia, subsequent reports that this species had be-
Manchuria, China, Formosa, Korea, Japan, come established.
Ryukyu Islands, and the Bonin Islands
(Dobzhansky 1933, Timberlake 1943,
Harmonia axyridis (Pallas)
Chapin 1965, lablokoff-Khnzorian 1982).
Specimens of this species were released in Figs. 1-6
Californiain 1916, 1964,and 1965 (Gordon Diagnosis.—Form oval, convex. Head
1985) and in Nova Scotia, Connecticut, straw-yellow, immaculate, with two longi-
District of Columbia, Delaware, Georgia, tudinal rows of faint spots, or with black
Louisiana, Maine, Maryland, Mississippi, spot on clypeus sometimes extending onto
Ohio, Pennsylvania, and Washington from labrum and head. Antennae and mouth-
1978 to 1982 (Gordon 1985, P. W. Schae- parts yellow-brown, sometimes tinged with
fer, pers. comm.). Specimens were also re- black. Pronotum straw-yellow with four
leasedinConnecticutin 1985 duringastudy black spots and usually a faint mark above
of the potential of H. axyridis to control scutellum, or lateral spots joined to form
Matsucoccus resinosae Bean and Godwin two curved lines or an M-shaped mark ex-
(McClure 1987). tending to anterior margin, occasionally
Japan was the country of origin of the center ofpronotum black. Scutellum straw-
specimens released in Louisiana and Mis- yellow. Elytra yellow-orange, each elytron
sissippi. A single release of 32 specimens in fullymaculateindividualswith faintmark
VOLUME 93, NUMBER 3 631
behind scutellum, two spots at one-fifth,
three spots in semicircle at two-fifths, three
spots at three-fifths, and one spot at four-
fifths, these black spots variously reduced
or absent; each elytron usually with trans-
verse plica anteriorto apex (Fig. 1). Ventral
surfaceyellow-orangetoblack, hypomeron,
epimera, epipleura, and lateral abdominal
margins paler. Intercoxal process of pro-
sternum with carinae. Legs yellow-brown
varying to black with posterior side ofmid-
dle and hind legs and tarsi paler. Length,
4.81-7.47 mm; width, 3.90-5.89 mm.
Localitydata.—The majorityofthe more
than 1000 specimens were collected by V.
A. Brouathishomelocated4.2milesnorth-
east ofAbita Springs (Sec. 24, T6, SR12E)
using six light traps of varying wattages,
ranging from 60-2500 watts per trap. Usu- Fig. 1. Harnionia axvridis, habitus; scale bar = 2
mm.
ally a combination of ultraviolet light
sources were used on the traps, including
fluorescent blacklights and high intensity collected in Mississippi counties: 1, Cal-
discharge lamps, e.g. mercury vapor. The houn (5.5 miles north ofVardaman, black-
traps operated from dusk to dawn and were light trap in sweetpotato field), August 9,
controlledbyphotoelectriccells. Specimens 1990, James Jarrett; 3, Harrison (T5S,
first appeared on July 20, August 8 and 20, R13W, Sec. 1, old field habitat on senescent
October 22, and November 17 in 1988. No Erigeron), November 15, 1990, T. C. Lock-
specimenswereseenpreviouslyeventhough ley; 1, Panola (2 miles south ofCrenshaw),
thelighttrapshadbeenoperating365 nights October 6, 1990, S. Jones (student collec-
per year since 1982 at this same location. tion) (MSU).
In 1989, specimens were taken on January Comments.—The variability ofthe color
26 and from March 19 to December 27. In pattern has led to the description of more
1990,theywerecollectedonJanuary 13 and than 100 forms (see Korschefsky 1932 for
22 and from February 2 through September synonymy). Twenty-four of these patterns
16 and were present through December. were portrayed by Ayala (1978). Only the
Larvae and adults were collected on crab- pale forms are found in Louisiana and Mis-
apple trees infested with Aphis spiraecola sissippi, including /r/^/<i(3 Mulsant, novem-
Patch in May. This species is now the most decimsignataFalderman,andsuccineaHope
common lady beetle at this site. Additional which are more common in the Maritime
specimens were collected in July 1990, by Province ofRussia, Korea, Manchuria, and
the senior author and V. L. Moseley on China. In southwestern Sibera, black forms
crapemyrtles infested with Tinocallis kaho- with one to several pale spots predominate
waluokalani (Kirkaldy) in the following (Dobzhansky 1933). In Japan, the percent-
southeastern parishes: Jefferson (Metaire), age of the pale form, succinea, decreases
Orleans(New Orleans), St. Helena (Greens- graduallyfromnortheasttosouthwestwhile
burg), St. Tammany (Covington, Folsom), one ofthe dark forms, conspicua, increases
Tangipahoa (Hammond), and Washington conversely; two other dark forms do not
(Franklinton).Thefollowingspecimenswere show such a gradient (Komai 1956). lablo-
632 PROCEEDINGS OFTHE ENTOMOLOGICAL SOCIETY OF WASHINGTON
kofT-Khnzorian (1982) summarized the in- Apex ofeach middleand hind tibia with
formation known about this species, some 2 spurs (Fig. 626a) 12
ofwhich is included in Hodek(1973). Some 10(9). wPiotshtcoobxlailqluienedoinvifdiirsntgalbidneominalsegment 11
ofthis data, including the host information - Postcoxallineonfirstabdominalsegment
given below, may refer to //. yedoemis (Ta- without obliquedividing line
kizawa), aclosely related speciesthatoccurs Mulsantina Weise
in China, Korea, Japan, Ryukyu, and Tai- 11(10). Prosternalcarinaeendingatanteriorcox-
wan. al margin NeoharmoniaCrotch
Prosternal carinae extending anterior to
Two other species ofHarmonia occur in the anteriorcoxal margin orabsent ....
the U.S. Harmonia dimidiata (F.) was in- Harmonia Mulsant
troduced from China into California and 12(9). Prostemum strongly convex medially,
then to Florida in 1926 where it became protuberantatapex(Fig.614b);mesoster-
established (Gordon 1985). Harmonia num deeply emarginatc for reception of
sternal process inatis Mulsant
quadripunctata (Pontopiddian), which may Prostemum normally rounded, not pro-
have been accidentally introduced, was re- tuberant at apex; mesostemum truncate
ported from NewJerseyand New York and or weakly emarginate for reception of
has been in the U.S. since 1924 (Vanden- prosternal process 13
berg 1990). Specimens of //. axyridis are 13(12). Postcoxallineonfirstabdominalsternum
complete, ofthe Fullus type (Fig. 637a)
broadly oval; the pronotal pattern consists idalia Mulsant
offour to five black spots, two curved lines, Postcoxallineonfirstabdominalsternum
an M-shaped mark, orthecenterofthe pro- incomplete,oftheDionntsorNcphustype
notum is black; in addition to the spot be- (Figs. 634b, 682a) 14
14(13). Elytron yellow with black sutural margin
hind the scutcllum, each elytron has nine
and 4 black spots(Fig. 672g), spotsoften
black spots arranged in a 2,3,3,1 pattern, or somewhat coalesced; pronotal hypome-
the spots are variously reduced or absent; ron with shallow fovea; Oriental genus,
and a transverse plica, which is lacking in onespeciespossiblyestablishedinFlorida
theothertwospecies, is usually presentnear Coelophora Mulsant
the elytral apex. Harmonia quadripunctata Elytronwithcolorpatternnotresembling
that o(Coelophora; pronotal hypomeron
is more elongate in shape with 1 1 puncti- without shallow fovea; North American
form black spots on the pronotum, one or or European genera 15
two pairs sometimes faint or absent; the el- 15(14). Tarsalclawwithmediantooth(Fig.626a)
ytron is immaculateorhasapairofelongate Myzia LcConte
Tarsal claw with subquadratebasal tooth
black marks at the lateral margin on either
(Fig. 664b) 16
sideofthemid-line(Vandenberg 1990);and 16(15). Pronotal surfacepolished, shiny, notalu-
the prosternal carinae, present in the other taccous between punctures Calvia Mulsant
. .
two species, are absent (lablokolT-Khnzo- Pronotalsurfacealutaceous,oftendull,not
rian 1982). Specimens of//, dimidiata are polished 17
roundand veryconvex with a bilobcd black 17(16). Pronotum black with large, subtrape-
zoidal or triangular white spot on each
spot at the base ofthe pronotum and seven
anterolateral angle (Fig. 643g) (apical
black spots arranged in a 1,3,2,1 pattern on marginofpronotumsometimesnarrowly
each elytron. pale) Coccinclla L.
With the addition oftwo more species of Pronotum not as above 18
Harmonia to our fauna, the key to the gen- 18(17). Postcoxallineonfirstabdominalsternum
with obliquedividing line Olla Casey
era of Coccinellini (Gordon 1985, p. 681) Postcoxallineonfirstabdominalsternum
needs to be modified as follows: without obliquedividing line 19
19(18). Apex ofmesostcrnum notched forrecep-
9(6). Apexofeach middleand hind tibiawith- tion ofprosternal process; European ge-
out spurs 10 nus, one species presently established in
VOLUME 93, NUMBER 3 633
Figs. 2-6. Harmonia axyridis. male and female genitalia. Figs. 2, 3, ventral and lateral viewsofphallobase
without and with trabes. Figs. 4, 5, entire sipho and enlarged view ofdistal end. Fig. 6, spcrmathecal capsule,
sperm duct and infundibulum.
634 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON
southeasternCanada,Vermont,and New Toxoptera piricola Matsumura [= Schiza-
York' Propylaca Mulsant phispiricola (Matsumura)]. Yasumatsu and
- Apexofmcsosternum truncate;occurring Watanabe (1964) listed 13 hosts including
overmostofNorth America from south-
ern Canada to Mexico ...CyclonedaCrotch Cinara kochi Inouye, Hyalopterus pinni
Matsumura, and Chrysomela vicintipunc-
tata Scopoli. Fuyuan and Dongxi (1989) re-
Koebele's field notes (Timberlake 1943)
ported the control of Nesticoccus sinensis
indicated that H. axyridis was common in
Tang (Pseudococcidae) and Rhizococcus
Japan on various aphid-infested trees, such
transversus (Eriococcidae) after coccinel-
as maple, walnut, and willow, and also on
lids, predominately //. axyridis and H. ob-
rose. It fed on Lachmis sp. on Podocarpus
scurosignata (Liu-Chung-Lo), were attract-
and Kermaphis pint (Koch) [= Pineiis pint
edtoaphid-infestedrapeinterplanted in the
(Koch)] on pine in Japan, on Lachmis sp. bamboo
forest.
on Pinus sinensis at Hongkong, China, and
was collected on Lachnus infested pine in Acknowledgments
Formosa. lablokoff-Khnzorian (1982) also
The authorswish to thank R. D. Gordon,
stated that itwasarboreal, that it was found USDA Systematic Entomology Laboratory,
on rosaceous arborescents and also on oak
Washington, D.C., for confirming the iden-
and other trees, and that it was chiefly an tification of H. axyridis; M. B. Stoetzel,
aphid feeder although it fed on Icerya pur- USDA SystematicEntomology Laboratory,
chasiMaskell (Margarodidae), Phenaccocus
Beltsville, Md., for identifying A. spirae-
pergandei Cockerell (Pseudococcidae) and
cola; Terence Schiefer, Mississippi State
Anomoneura mori Schwarz (Psyllidae), at
University (MSU), Starkville, Miss., for
least in the laboratory. Preferred aphid hosts
permission to use the label data on the Mis-
given by lablokoff-Khnzorian (1982), some
sissippi specimens; K. A. Westphal for the
ofwhich were listed by Hodek (1973), are
illustrations; V. L. Moseley, D. A. Rider,
as follows: Acyrthosiphon pisum (Harris),
and L. H. Rolston, Louisiana State Uni-
Amphorophora oleracea v.d. Goot [= Hy- versity, and P. W. Schaeffer, USDA Bene-
peromyzus carduellinus (Theobald)], Aphis
pomi De Geer, Cinara laricicola (Bomer) ficial Insects Research Laboratory, for re-
viewing earlier drafts of this manuscript.
[= C. cuneomaculata (del Guercio)] (?), C. Approved for publication by the Director
todocola (Inouye) (?), Cryptosiphum gallar- of the Louisiana Agricultural Experiment
iim(Kaltenbach)[= C. artemisiaeBuckton],
Station as manuscript number 90-17-4432.
Eriosoma lanigerum (Hausmann), Hyalop-
terus pnini (Geoffrey), Macrosiphum rosae
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