Th- Tay::c n-, of th.? .j--.-i (Ho.opt..--.: A *
of the E"st.e:n UTnited States
?' i ',S'. PH JI LS SC-
C :- T
: .1 :- ' j' .'o r
ACKOWLEDG2 .. ''TS
I wish to thank Iiss Louise M. Eussell, USDA
Systematic Entomology Laboratory, Washington, D.C.,
for her suggestions concerning the status of some of
the species included in this paper and for the gen-
erous loan of slide material.
I also wish to thank the following for the gen-
erous loan of material: Dr. J. 0. Pepper, Pennsylvania
State University, State Colle--e, Pennsylvania; Dr. C. F.
Smith, :'orth Carolina State University, Raleigh, north
Carolina; Ir. H. A. Denmark, Division of Plant Industries,
Gainesville, Florida; Dr. H. B. Boudreaux, Louisiana
State University, Baton Rouge, Louisiana; Dr. H. G.
HWalker, Los Angeles State and County Arboretum, Arcadia,
California; and Dr. D. Hille Ris Lambers, Bladuisonder-
zock T.N.O., Bannekom, Holland. I wish to thank Dr. C. F.
!Smith for the loan of pertinent translations, too.
I wish to thank Dr. A. N. Tissot, Professor En critu-,
University of Florida, Gainesville, Florida, for his
;iuiding supervision, for his suggestions concernir-: the
organization of this paper and for the generous lclan of
slide material and renrints.
I also wish to acknowle.Ic-e the assistance of my
supervisory committee in the preparation of this paper:
Dr. D. H. Habeck, Associate Professor of Entomology,
Chairman; Dr. H. L. Cromroy, Associate Professor of
Entomology; Dr. T. J. 'Jalker, Professor of Entomology;
Dr. D. G. Griffin, III, Assistant Professor of Botany;
and Dr. F. W. Zettler, Assistant Professor of Plant
TAB3LE OF CO F '"i3
AC:' Ck '-._'- TS .................................... ii
LTST OF F ES................... ........ ........ vi
ASi TR ACT ........................................... vii
I ~TUTROi'cTION ....................................... 1
P -r:- ino o y ............. ................... 1
Lit rat.r. e review............................. 5
-volution of the yzini................
Characteristics of the Tribe -:. .ini ........... 7
- S A D AT I ALS .............................. 10
-eo .-.-. phical Scone ............... ............. 10
'ountin Technique............................ 10
ir, il )ationn ..................................... 12
S. A O? Y 14
GEUS ZiUS PASS INI ............................. 16
s 71". . c frn 'si ("abricius) ...................... 23
:Li-us hcmer c lis Takahatshi ... .............. 29
'rzus __i thri (Schriank) ........................ 36
v.T- l us or J.tusl ( .......) ......................... 50
S o .- e ( l er)................................
i us oerttis \TWalker) ......................... 71.
1i,1Ls varlians (Davidson) ...................... 79
C ..... 1Hi. .OiJ. PASS N ................ 84
Ph'.~l od.o'is I ~ll i (Sc!hrank) ..................... 35
,";,i-S H izL "i. iUrCISHARDS.S .......................... 92
H:v. : us eriob try ;' (Tisrsot) ............... 94
0i's'i.CY '15 2i2t 1 ................... 100
a 's s at s (iason)'. ............... 104
--- ti.; i e j. ii1 -i ss n d a i Habeck ...... 107
Gi .U OViATUS 'VAi H lO ............................ 11
Ovatus crata.e arils ('alker) .................. 112
"*atus .]: :...7 '. 'a pson) ...................... 118
C.E. I U A:. .:: -O R OA LLTO ....................... 122
'TLr.-,.'1 -l: r hu:Tboldti (Essig).............. 124
GEHTS AULACORTHUM MOHDWILKO............. ...... ... 131
Aulacortihum circumflexum (:Buckton)............ 132
Auilacorthum solani (" 1 tenbch) ......... ...... 136
S-:. l: OD0 IU T HILLE BIS LAM :.................. 145
-co:' -'-. orosum (Sanderson)..... ......... 146
.;:: FT:53 IAPHTS RIC '.'.'.. .. ................ ..... 153
T* aihbr~ a)his f briata Richards ............... 154
Fimbria'oh s scammelli (-!ason) ................. 159
3 i-:- .C APHY ........ ......... .................... 166
BIOGRAPHICAL S .L C;i' ............................... 277
LIST OF FIGURES
1 Diagram of a generalized myzine aphid
(after Bodenheimer and Swirski 1957)......... 2
2 Myzus cerasi (Fab.).......................... 31
3 Kyzus hemerocallis Tak., A-G. M. lythri
(Schrk.), H-P...... ............ .. ......... 35
4 Myzus oliveri (Essig), A-H. M. ornatus
Laing, I-Q...................................... 47
5 Variation in wing venation in Myzus oliveri
(Essig) ...... ...................... .......... 49
6 Iyzus Rersicae (Sulzer), A-L. M. varians
Davidson, M-T... ............. ........... 78
7 Phorodon huruli (Schrk.)..................... 91
8 Hlomyzus eriobotryae (Tissot), A-H. H.
sensoriatus (~iazon) plesiotypes, I-0.......... 101
9 Ovatus cratae-arius (Walker), A-G. 0. phloxae
Tpson), H-O .................. ............. *117
10 UtamrhoroIhora humboldti (Essig) ..............130
11 Aulacorthumr circumflexum (Buckton)............137
12 Aulaco-t'euImi solani (Kalt.)....... ............144
13 Rhodobium poriosum (Sanderson)................ 152
14. Fimbriaohis fimbriata Richards.................158
15 Fimbriaphis scammelli (Mason)....... ........ 165
Abstract of Dissertation Presented to the
Graduate Council of the University of Florida in Partial
Fulfillment of the Requirements for the Degree of
Doctor of Philosophy
The Taxonomy of the Myzini (Homoptera: Aphididae)
of the Eastern United States
Richard Joseph Nielsson
Chairman: Dr. D. H. Habeck
Major Department: Entomology
The tribe Myzini (Homoptera: Aphididae) includes
the genus Myzus and the following related genera in the
Eastern U.S.: Phorodon, Hyalomyzus, Ovatus, Utamp'orophora,
Aulacorthb.im, Rhodobium, and Fimbriaphis.
The 8 species of Myzus which occur in the Eastern U.S.
are described and illustrated. Nectarosiphon is declared
to be invalid as a subgeneric name within Ijyzus, and
persicae and certus are included in yzus without sub-
generic designation. Lectotype, morpholectotype,
paralectotypes, and plesiotypes of varians are designated.
The 13 species in the 7 genera related to Myzus
are described and illustrated. The genus Utamohorophora is
redescribed and plesiotype sexuales of U. humboldti are
designated. Lectotyre, morpholectotype, allolectotype,
and paralectotypes of Fiubriaphis scaurilelll are also
In North America today, Drs. Quednau and Richards in
Canada are studying the Callaphidini; Dr. C. F. Smith at
North Carolina State University, the Eriosomatinae; Drs.
Boudreaux and Rolston at Louisiana State University, the
Macrosiphini; and Dr. J. 0. Pepper, Pennsylvania State
University, and Dr. A. N. Tissot, University of Florida,
the genus Cimara. Aside from the genus Aphi the Myzini
is the only major group of aphids which has not undergone
a recent study. For this reason, I chose to study this
tribe within the eastern United States.
lor:'ho7loz. A drawing of a generalized myzine aphid
is shown in Figure 1. The eyes are many-faceted with 3 fa-
cets in the ocular tubercles. Ocelli are also present in
The front of the head may bear frontal extensions, the
antennal tubercles, which form the bases for the antennae,
The antennae are usually 6-jointed, the joints frequently
numbered from base to apex by Roman numerals; the last joint
is separated into the base, from the proximal end of the seg-
ment to the distal end of the primary ::i;orium, and the ter-
minal filament (processus terniralis), from the distal end
zz -<< CA
I I I. c c-I
=C z Ll m, -J
= j & oa z
- o? Z W 14 S
ac to wo Cs -r 0
a a d CL m
Z UJ 44 u Z S V 1 2
Ii M a- >- z / a
Z Z r 1 / \ U // \ u* -J Q 0.
of the primary sensorium to the end of the segment. Pri-
mary sensoria are located near the terminal end of segment
V and at the distal end of the base of segment VI; these
usually have a ciliated margin. There are also 4-7 small
sensoria without ciliated margins clumped next to the pri-
mary sensorium on segment VI; these are generally considered
to be part of the primary sensorium. Secondary sensoria may
be present on any or all of segments III-V in alatae; they
are only occasionally present in apterae. The secondary
sensoria are generally without ciliated margins.
The rostrum is 5-jointed but the distal segment is so
reduced that segment IV + V is considered to be a single
The legs are composed of the usual parts: coxa, tro-
chanter, femur, tibia, tarsus. Oviparous females usually
have the hind tibiae swollen and bearing various numbers
of pseudosensoria. The tarsi are 2-jointed.
The typical wing venation is shown in Figure 1. Occa-
sionally veins may be reduced or missing on one or both
pairs of wings.
The abdomen is composed of 9 (or according to some au-
thors 10) segments. Eight distinct tergites are visible.
The cauda, a prolongation of the last abdominal tergite, is
usually well developed. Lateral sclerites are located above
the spiracles on segments (I) II-IV; a presiphuncular sclerite
(lateral sclerite on segment V located in front of the corni-
cle base) and a postsiphuncular sclerite (lateral sclerite
on segment VI behind the cornicle base which may extend to
segment VII) are also usually present. Lateral tubercles
may be present on any or all of the sclerites on segments
(I) II-IV and rarely on the presiphuncular sclerite; dorsal
or spinal tubercles may occur on segment VII and/or VIII.
The cornicles (siphunculi) arise dorsolaterally between seg-
ments V and VI. The anal plate is situated below the cauda
on the venter of segment VIII. In viviparae and oviparae
there is a genital plate on the venter of segment VII.
Life cycle. The nymph that hatches from the egg matures
into the fundatrix, or stem-mother, an apterous viviparous
female usually with shorter antennae, legs and cornicles
than subsequent apterae and with antennae frequently 5-seg-
mented. The fundatrix gives birth to fundatrigenic-ac which
may or may not differ morphologically from subsequent gen-
erations. The first generation fundatrigeniae and all sub-
sequent generations produced on the prirmairy host, which may
also be called fundatrigeniae, are primary viviparae (pri-
mary virginoparae, i.e., "produced on the primary host, off-
spring of virgin mothers").
Alate fundatrigeniae migrantse) migrate from the pri-
mary to the se.onaOary host where they give birth to secondary
virginoparae. The alate virginogeniae which migrate to other
plants of the same species or to other secondary hosts are
termed vagrants; this same term may be applied to alatae of
autoecious species infesting other primary hosts.
The alate virgincgeniae which return to the primary host
are called remisrantes. They may also be called gynoparae
if they give birth to oviparous females or sexuparae if they
give birth to both oviparae and males. Frequently however
the males are produced on the secondary host and fly to the
In southern regions virginogeniae frequently live par-
theno&enLctically throughout the year without the formation
The g?.ns Mvzus was erected by Passerini (1860) with
Aphis cerasi Fab, as type. iyzus with its related genera was
included in the Nectarophorini with ect.ioo-o~ohio. Oestlund
(= .':r-..jLr in Passerini) in America (Williams 1910); Eur-
opeans placed it in the Siphonophorini with Siohoonophora
Koch (= Nacrosinhum Passerini) (van der Goot. 1915, 1917).
Passerini's repl.acmimt of Siphononhora by Macrofiin!hi (1860)
became widely used only after Schoutedcn (1901) published
his synonymy of the genus :icostjji lizuis, acrjft rhum
and Amnhoronhora were then generally placed together in the
Macrosiphini (Wilson 1910, Swain 1919, Blanchard 1922, Nevsky
1928, Hottes and Frison 1931, Gillette and Palmer 1934, Eastop
1966). A few authors (Nevsky 1929, Cottier 1953, Bodeuiheimer
and Swirski 1957) placed these genera in the subfamily Aphidinae
and did not attempt further discrimination.
The use of a separate tribe for vzyLu and closely relat.:d
genera was first proposed by Mordwilko (1934) and has since
been adopted by many authors (E&-ner 1952b, Shaposhnikov
1956, Heinze 1960, 1961, Aizenberg 1966, QuednLau 1966).
Borner (1952b) and Heinze (1960, 1961) divided the subfam-
ily Myzinae into the following tribes and genera: Brachycolini
(Brevicoryne, Coloradoa, Hyadaphis, Lipaphis), Crypornyzini
(Capitoohorus, Cryptomyzus), Liosomaphidini (Cavariella),
Myzaphidini (Chaetosiohon, Longcauduls, y.~aohis), Myzini
(Eyzus, Neomyzus, Pentalonia), Nasonoviini (Hyoeromyzus,
Idiooterus, Nasonovia, Rhoralosifphc.ninus) and Phorodontini
(Myzodes, Ovatus, Phorodon, Rhopalomyzus, Scjamyzus). The
Myzini were separated from the Phorodontini principally on
the basis of the presence of spines between the setae on the
hind tibiae of first instar nymphs in the former and the ab-
sence of these spines in the latter. However, this separa-
tion is not valid since it works with European aphids but
not with the Oriental fauna (van Enden et al, 1969). There-
fore, the Myzini, as used in this paper, correspond in gen-
eral to the Myzini and Phorodontini of Borner (1952b) and
Heinze (1960, 1961).
Evolution of the Myzini
According to :ordwilko (1934), Borner (1952b) and Sha-
poshnikov (1956), the Myzini, through the genus Iyz.s, evolved
from the Anuraphidini, the members of both groups retaining
the primitive relationship with the Rosaceae as primary hosts.
Mordwilko (1934) noted that spinal tubercles, which were us-
ually well developed in Myzus, were not developed in Anuranrhi
and. Dvszhis in the Anuraphidini; but, nevertheless he pro-
posed that all three genera should be grouped together since
they share Rosaceae as primary hosts, Kyzaus on Prunoidea and
Anuraphus and Dysaohis on Pomoidea.
Shaposhnikov (1956) considered the Myzini to be inter-
mediate between the Anuraphidini and the 'acrosiphini. He
noted that the spinal tubercles and cuticular sculpturing
were well developed in Myzus but not in Anuraohis and Dysaphis,
supposedly marking the two groups as being distantly removed;
however, D:.sanhis had well developed antennal tubercles simi-
lar to those in Myzus. Furthermore, the cauda in Iyzus was
not as well developed as in the "acrosiphini but the shape of
the body and imbrications on the cornicles were similar.
Aizenberg (1966) considered Myzus and the Myzini to be
closely related to the Macrosiphini based on the structure
of the cauda, the absence of lateral tubercles on abdominal
segment VII, and usually also on segment I, the well devel-
oped antennal tubercles and the structure of the setae on
Characteristics of the Tribe Myzinij
Body size medium to large, usually 1.0-2.6 mm, ovate
to obovate. Color usually green or yellow. Antennal tuber-
cles very prominent, parallel to strongly converging and
scabrous, Antennal segments I and II usually scabrous; I
usually with a variably developed projection on the medial
surface, rarely rounded. Antennae usually 6-segmented;
terminal filament at least 1.5 times the length of the base
of antenna segment VI and usually 3.5-4.5 times that length.
Alate viviparae with secondary sensoria on antennal segment
III only, on III and IV, or on III, IV and V; apterous vivip-
arae usually without secondary sensoria, present only in
Aulacorthum and Rhodobium. Eyes normal; apterae without
ocelli. Rostrum moderately long, with segment V reduced so
that IV + V appears to be a single segment.
Legs normal; first tarsal segments usually with 3 setae,
occasionally 2 on the first hind tarsal segment. Mesothor-
acic furcula of apterae consists of a short stalk with blunt
lateral projections. Wing venation normal, sometimes reduced;
occasionally with bordered wing veins.
Alate viviparae with pigmented lateral abdominal scler-
ites and usually with a pigmented dorsal patch; apterae rarely
with a pigmented patch on the abdomen. Lateral abdominal tu-
bercles always absent on seegent VII and usually also on seg-
ment I; spinal tubercles may be present on segment VII and/or
VIII. Cornicles well developed, 1/5-1/3 the length of the
body, cylindrical or swollen, usually imbricated and rarely
with closed reticulations apically; if so, the dorsum of the
abdomen has a pigmented patch or transverse bars. Cauda mod-
erately developed, usually 1/3-1/2 the length of the cornicles,
conical, with 2-3 pairs of lateral setae.
Oviparae usually resemble apterous viviparae except for
their swollen hind tibiae which bear varying numbers of pseudo-
sensoria. Males similar to alate viviparae, usually alate,
rarely apterous. Fundatrix similar to apterous viviparae
but anten-ae, legs and cornicles shorter; antennae frequently
5-semrented and antennal tubercles usually not developed.
METHODS AND MATERIALS
The eastern United States as defined for this study
consists of those states east of the Mississippi River
plus Louisiana and the District of Columbia. This includes,
in alphabetical order, Alabama, Connecticut, Delaware,
Florida, Georgia, Illinois, Indiana, Kentucky, Louisiana,
Maine, Maryland, Michigan, Mississippi, New Hampshire,
New Jersey, New York, North Carolina, Ohio, Pennsylvania,
Rhode Island, South Carolina, Tennessee, Vermont, Virginia,
West Virginia and Wisconsin.
Aphids were mounted following the procedure of Tissot
and Pepper (personal communication). After storage in 70%
alcohol, the venter of the abdomen was slit,.using minute
pins, from approximately segment I to a point just in front
of the genital plate.
The specimens were placed in 5% KOH (or NaOH) in a boil-
ing water bath for approximately 5 minutes. It is important
that the KOH not boil. (I found that aphids placed in a
beaker with 5% KOH in running hot tap water for 15 minutes,
or left in cold 5% KOH for 2 days gave satisfactory results
with less chance of Lamaging the specimens,)
The aphids were removed from the KOH, washed gently in
water and transferred to 70% alcohol. Minuten pins were
used to remove embryos, digestive tract, and other soft
tissues from the inside of the abdomen.
The aphids were then successively placed for 15 minutes
each in absolute alcohol, 3 parts absolute alcohol in 1 part
clove oil, 1 part absolute alcohol in 1 part clove oil, and
finally in clove oil.
The aphids were mounted from clove oil directly into
A variation of this procedure (Pepper, personal commun-
ication),which replnicd the clove oil mixtures with a sat-
urated solution of chloral hydrate in absolute alcohol, was
tried. Although this proved to be successful, it could be
used only if the chloral hydrate solution was freshly mixed
because it was not stable and must be replaced at approxi-
mately monthly intervals.
When remounting old slides, the cover slip was soaked
off in xylene or Euparol Essence (Flatters & Garnett Ltd.,
elichester, England) depending on the mounting medium and
the aphids were transferred to 5% KOH in a boiling water
bath for 5 minutes. After cooling, the specirmens were placed
in 70% alcohol and the abdomens were slit and cleaned. The
aphids were returned to 5~. KCH in a boiling i:ater bath and
the procedure followed the same steps as outlined above for
fresh specimens. The original labels were kept and placed
on the remounted slide.
Mounted specimens were borrowed from several sources.
These sources are abbreviated as follows: U.S. National
Museum of Natural History USNM; the Florida State Collec-
tion of Arthropods (including the collection of Dr. A. N.
Tissot) FSCA; Dr. C. F. Smith, North Carolina State Uni-
versity, Raleigh, North Carolina CFS; Dr. J. 0. Pepper,
Pennsylvania State University, State College, Pennsylvania -
JOP; Dr. H. B. Boudreaux, Louisiana State University, Baton
Rouge, Louisiana HBB; Dr. H. G. Walker, Los Angeles State
and County Arboretum, Arcadia, California HGW; and, Dr.
D. Hille Ris Lambers, Bladuisonderzoek T. N. 0., Bennekom,
The descriptions given are based entirely on specimens I
studied and therefore may differ slightly in ranges of mea-
surements given in other publications. The list of speci-
mens studied is summarized in the section "Collections
studied"; this section is followed by abbreviations for the
depositories in which these slides can be found and, when
applicable, references to other host plant lists. The col-
lections are arranged geographically by states, North to
South and East to West, and alphabetically by localities with-
in states. Since Mjzus persicae, M. ornatus and Aulacorthum
circumflexum are so polyphagous, the collections for these
species have been reduced to lists of host plants, alphabet-
ically by families and alphabetically by species within
families. Gray's :Manual of Botany was used as the source
for plant family names.
Even though the geographical scope of this paper is
limited to the eastern United States, specimens from outside
the area were studied so that a proper assessment of varia-
tion could be made; these records from outside the area are
included in the "CGllections studied."
All studies were made with a phase contrast microscope
and therefore the color of mounted specimens appears to be
yellower than with a direct light microscope. All measure-
ments are in iam. Body length was measured from the vertex
of the head to the apex of the anal plate. Head width is
width across the compound eyes. Cauda length is total length
from base to apex.
The drawings are line tracings of 8 X 10 enlargements
of photomicrographs of the specimens. The line drawings
were then reduced to fit within the allowable margins.
The keys apply only to alate and apterous viviparous
females and, unless stated to the contrary, characters will
apply to both morphs.
KEY TO EASTERN U.S. GE!!ERA OF KYZINI
1. Rostral segment III with 2 pairs of lateral
Rostral segment III with at least 3 pairs of
lateral setae ........... .................... 7
2. First hind tarsal segment with 3 setae............. 3
First hind tarsal segment with 2
setae.... ...... ... .... ............. us (in part)
3. Cornicles swollen (Fig. 4, 6)............. .......... 4
Cornicles cylindrical (Fig. 2, 3) or
constricted medially (Fig. 9)................ 5
4. Alate viviparae occasionally with a dorsal
sclerotic patch; apterous viviparae
usually with 1-3 secondary sensoria
basally on antennal segment III.....Utamphorophora
Alate viviparae without a dorsal sclerotic
patch; apterous viviparae without
secondary sensoria on the antennae......Hyalomyzus
5. Cornicles cylindrical; alate viviparae
with secondary sensoria in a more
or less straight line on antennal
segment III; if sensoria scattered
on segments III and IV, abdomen with
dorsal sclerotic patch,....................... 6
Cornicles constricted medially; alate vivip-
arae with numerous secondary sensoria
on antennal segments III and IV; if in
a more or less straight line on segment
III, abdomen without a dorsal sclerotic
patch......................... .......... .. vatus
6. Antennal tubercles with elongate finger-like
projections (Fig. 7) sensoria numerous
and scattered on antenna segment III;
1-4 sensoria also present on segment IV;
wing veins without fuscous borders........Phorodon
Antennal tubercles without elongate finger-
like processes; sensoria in more or
less straight line on antennal segment
III; if numerous and scattered over
segment III and also present on IV, then
wing veins with conspicuous fuscous bor-
ders... ............. ......... Mzus (in part)
7. Cornicles without closed reticulations apically.... 8
Cornicles with closed reticulations apically
(Fig. 11, 12)....... .................. Au7 corthum
8. Alate viviparae with sensoria with ciliated
margins and in a more or less straight
line on antennal segment III; apterae
without secondary sensoria; cauda more
than 1/2 the cornicle length; alate
viviparae occasionally with a dorsal
sclerotic patch'on the abdomen ....... Fimbriaphis
Alate viviparae with scattered sensoria with-
out ciliated margins on antennal seg-
ments III and IV; apterae with 4-19
sensoria on antenna segment III; cauda
less than 1/2 the cornicle length; alate
viviparae without a dorsal sclerotic
patch on the abdomen..................... ghodobium
GENUS NYZUS PASSERINI
Myzus Passerini 1860:27.
Rhodopalosiphum Wilson nec Koch 1910:320.
Myzoides van der Goot 191:167. 1917:47.
yvzodes Mordwilko 1914:69.
1yzu s (lectarosiphon) Hille Ris Lambers nee Schouteden
Wilson (1910) designated Aphis persicae Sulzer as
type of the genus Rhooalosilhum. However, Gerstacker
(1853, in Borner 1951) had designated Aphis nympheae L.
as type for Koch's genus Rhooalosiphum. Therefore, Rho-
palosipjhum Wilson nee Koch is a full synonym of yzus Pass.
Van der Goot (1915) placed species with elongate
capitate setae and sensoria on antenna segment III of
apterae in Myzus while placing other species of ~yzus in
Myzoides n. gen. Therefore, Nyzus van der Goot nee
Passerini is a synonym of Capitophorus van der Goot and
Myzoides is a synonym of Myzus Pass.
The major problem in synonymy of the genus ,yzus, viz.,
jyzuis (rIectarosiphon) vs. Uyzodes, is still prevalent today
and deserves explanation.
Koch (1854) established the genus Sinhonoohora for
those species of Aphis L. which Kaltenbach had ch-aracterized
in 1843 by: "Letzteres Fuhlerglied gorstenformig und langer
al1- das vorletzte. Die Fuhler stehen auf einem hokerartigen
Stirnknapfe. Stirne tief rinnenformig." ("Last antenna
segment bristly and longer than thernext to the last. An-
tennae sit on bumps on the front of the head. Front deeply
groove-like.") Eo:;ever, the name Siphonophora had been used
by Braadt (1840, in Schouteden 1901) for a genus of Myria-
pods, and since his generic name had priority over that of
Koch, Passerini (1860) suggested that Siphonophora Koch be
replaced by the new name Macrosiphum but he himself did not
use the latter designation for several years. Oestlund
(1887), unaware of Passerini's action, replaced Siphonophora
Koch with Nfectaroohora, which was subsequently adopted by
In 1886, Oestlund erected the genus 1acrosiphum for M.
rubiculum, close to Sirhooohora rubi Kalt., a species with
very swollen cornicles. Del Guercio (1900) also used the
name Macrosiphum for species with slightly swollen cornicles.
Schouteden (1901) discovered Passerini's name and sought
to clear up the multiplicity of uses of "Macrosiphum" with
the follo'iing (p. 111) (translation from French; underlining
my own): "In his remarkable work on the Italian aphids
(Prospetto dell' Afidofauna Italiana), which appeared last
year, Professor G. Del Guercio established a new genus con-
tai.-ing the species with slightly swollen cornicles (rubi
Kalt., convolvuli Kalt., viciae Kalt.) in addition to the
genus Si)horiohora; this second new genus he designated under
the nume !lacrosiphum. This genus seems to be identical with
that of Oestlund. Therefore if following the priority we
replace Siplho ot'ora and tc. rlora. by rusihium e
should also reject the denori.o'ation of Macrosiphum for the
penuus established by Del Guercio. I suggest therefore to
replace this name by that of Nectarosiphon,
"In summary then we would have the following synonymy:
Macrosiphum Pass. (nee. Oestl., 1e.. Del G.)
Siphonophora C. Koch, Pass., etc.
Nectarpohora Oestlund, etc.
Nectarociphon n. nom.
5aIrosihtium Del. G.
And finally (p. 113): "I have therefore replaced Mac-
rosiphum Oestl., Del G. by Nectarosiphon and I think this
change is well justified."
Hille His Lambers (1949) quoted only the underlined
portion of Schouteden's remarks above, and concluded that
Schouteden had replaced only Macrosiobum Del Guercio; Hille
Ris L--bers then proposed the name Oestlundia to replace
MNc'c'._iphum Oestlund. If this were the case, the type of
.rcrosirhur, Del G., convolvuli Kalt., would be the type of
Nectarcsiphon Schout.; otherwise, the genotype of the latter
would be that of the older of the genera, viz., rubicola
Oestl., the type of Macrosiphum Oestl.
Hille Ris Lambers (1946) stated that from Baker's (1920)
work, convolvuli Kalt. is clearly Aphis persicae Sulz. which
is then the genotype of .Nectarosiphon Schout., which he places
as a subgenus of .yzus Pass. MacGillivray (in Hille Ris
Lambers and MacGillivray 1959) used this same line of reason-
ing. Although convolvuli Kalt. is generally held to be a
synonym of persicae, Borner (1951) believed that the type of
Macros iohm Del Guercio was a misidentified specimen because
Del Guercio stated that the red spots on the base of the
cornicles are typical characters of other aphids with long
antennae, and this color clearly eliminates persicae,
Baker (1920), however, had placed convolvuli Kalt, in
Arnohororhora and listed Macrosiphum Oestl., Macrosiphum Del
G., and Nectarosiphon Schout. as synonyms of that genus.
Borner (1951, 1952a) stated that IzIsj Passerini is
not available for persicae because persicae is the type of
Myzodes Mord., and this genus is the type of another tribe
of the Myzinae. However, this statement is relevant only
in the light of Borner's (1952b) division of the subfamily
Myzinae into several tribes, one, the Myzini, including Myzus
Pass., and another, the Phorodontini, including Myzodes Mord.
Although the character, presence or absence of spines on the
hind tibia of first instar nymphs, separates the 2 tribes
for European species, it cannot be applied to the Oriental
fauna and therefore cannot be used indiscriminantly for
generic separation (van Emden et al. 1969). Furthermore,
since Myzodes is not the type genus of the Phorodontini,
the assignment of Dersicae is not dependent upon a recognition
of these 2 tribes of IMy.inae.
I believe after a consideration of Schouteden's (1901)
complete paper that he did replace both Macrosiphum Oestl.
and Macrosiohum Dl G. with his genus Nectaro-iphon. There-
fore, lra.ro2Jjn1 Cestl., Nectarosiphon Schout. and Oestlundia
Hille Ris Lrn'bers are synonyms of ATin.hoirophio ra Buckton.
The problems in Borner's (1952b) system of classifica-
tion are further complicated by Shaposhnikov (1964) who in-
cluded oortulacae Macch. (= ornatus Laing), a species with
spines on the hind tibiae of first instar nymphs, in the
genus Myzodes. He based this on the number and arrangement
of "rostral" setae on that segment, which in Myzus are 6 or
7 with the dorsal and ventral ones separated from the rest;
but, in yl-'odes they are either absent or are 5 in number
and the distance between them equal. Borner (1952b) placed
portulacae in Hyzus Pass. but Aizenberg (1966) placed it as
the type of his new genus :e-oivz'odes, differing from Myzodes
Mord. in arternal tubercles parallel rather than converging,
cornicles not swollen, antennae only about 1/2 the length of
the body (rather than 3/4) and the hind tibia of first instar
nymphs with spines between the setae, and differing from Myzus
Pass, in the structure of the forehead, first tarsal segment
of early nymphs with only 2 setae and 3 in adults and the ab-
sence of lateral and spinal tubercles.
The chaetotaxy of aphid nymphs has not yet been shown to
have phylogenetic significance. Richards (1965) developed an
evolutionary sequence in the Callaphidini utilizing setal
patterns on the dorsum of adults; however, his system is much
more complete than that proposed by Borner (1952b). It is
true that the first instar nymphs of persicae differ from
those of cerasi in the station of the hind tibiae. However,
the erection of a new genus based on this character allows
one to establish a new genus for every species which differs
from the genotype; and, thus ignoring variation which occurs
during the evolution of each genus due to the varying selec-
tive pressures imposed on each species depending on its eco-
The real problem is the phylogeny of the Myzini. Aizen-
berg (1966) states that Myzodes and Neomyzodes are derived
from Hyzus, as are all of the Myzinae. Therefore, to place
persicae and ornatus in Myzus Pass. does not make this genus
polyphyletic, the primary consideration of all systematics.
I believe that persicae is properly classified in the
g~nus Hyzus Pass. despite the differences which have been
discussed and have done so in this paper without subgeneric
designation and thus obviate the use of the names Nectaro-
si jobcn and Mlyzodes.
Characteristics. Antennal tubercles moderately to very
prominent, converging, scabrous. Ilelial side of antennal seg-
ment I, and occasionally also II, scabrous. Alate viviparae
with secondary sensoria in a more or less straight line only
on antennal segment III, rarely with sensoria scattered over
segment III and also present on IV (oliveri). Apterous vivip-
arae without secondary sensoria. Rostral segment III with
2 pairs of lateral setae. Setal formula of first tarsal seg-
ments 3:3:3 or 3:3:2. Alate viviparae with a dorsal sclero-
tic patch on the a':. l ,n, rarely absent (hemerocallis). Lat-
eral abdominal tubercles frequently present on segments II-
IV, rarely on I and V. Dorsal tubercles frequently present
on VII or VIII, or both. Cornicles cylindrical or swollen,
always without an apical reticulated portion. Cauda usually
conical and without constriction, rarely with apex knob-like
Type. Aphis cerasi Fabricius.
Key to Eastern U.S. Species of Myzus
1. First hind tarsal segment with 2 apical setae..... 2
First hind tarsal segment with 3 apical setae..... 6
2. Cornicles swollen.......... ....................... 3
Cornicles cylindrical............................. 5
3. Alate viviparae with sensoria on antenna
segment III in a more or less straight
line, rarely with 1-4 also present on IV;
wing veins not bordered; apterous vivi-
parae with antennal tubercles only moder-
ately converging and cauda elongated and
conical ......................, .......... ... 4
Alate viviparae with numerous sensoria scat-
tered over segment III and 11-16 on
IV; wing veins with conspicuous fuscous
borders; apterous viviparae with an-
tennal tubercles strongly converging
and cauda subtriangular....................oliveri
4. Living color yellow to green; apical rostral
segment with 2-3 lateral setae, rarely with
4; alate viviparae never with sensoria
on antennal segment IV; cornicles of
apterae pale or, if dark, only in the
region of the apical flange; terminal
filament of summer apterous viviparae
at least 3 times the length of the base
of antenna segment VI; males alate
............................... persicae (in part)
Living color pink to brown; apical rostral
segment with 2 pairs of lateral setae,
rarely with only 3 setae; alate viviparae
occasionally with sensoria on antenna
segment IV; cornicles of apterae dark;
terminal filament of suimer apterous vivip-
arac subaqual to twice the length of the
.base of antennal segment VI; males
apterous........ ............................ certus
5. Apices of antennal segments IV and V in alatae
and III-V in apterae dark with the rest
of the segments pale; abdominal tubercles
absent...................... ........ ........ vari ns
Antennal segments IV and V in alatae hollyy dark;
III and IV in apterae wholly pale, V wholly
pale or with basal 1/2 pale and apical 1/2
dark; abdominal tubercles usually present,
laterally on segments II-IV and dorsally on
VIII, occasionally also on VII.persicae (in part)
6. Cauda conical, without constriction (Fig. 2F, I)
abdomen of alate viviparae with dorsal
sclerotic patch............................. 7
Cauda with moli:an constriction, apex knob-like
(Fig. AC) abdomen of alate viviparae with-
out a dorsal sclerotic patch.........hemerocallis
7. One of the lateral accessory sensoria on the
base of antennal segment VI much larger
than the others and subequal in size to
the primary sensorium (Fig. 2C)............. 8
All of the lateral accessory sensoria on the base
of antennal segment VI subequal in size
and much smaller than the primary sensor-
ium .................................. ornatus
8. Living color brown to black; dorsal sclerotic patch
on the abdomen of alate viviparae continuous
laterally, or nea-ly so, with the lateral
sclerites (Fig. 2B'; dorsum of the abdomen of
apterous viviparae sclerotic, pigmented; cauda
elongated and conical.....................cerasi
Living color green; dorsal sclerotic patch on the
abdomen of alate viviparae rectangular, on
the mid-4orsum and not extending laterally
(Fig. 31); dorsum of the abdomen of apterous
viviparce membranous, not pigmented; cauda
Myzus cerasi (Fabricus)
Aphis cerasi Fabricius 1775:734.
Aphis anarines Kaltenbach 1843:46.
Aphis asperulae Walker 1848:2248.
Aphis euthrasiae Walker 1849: appendix 51.
Nyzus cerasi (Fabricius). Passerini 1860:27.
Myzoides cerasi (Fabricius). van der Goot 1913:84.
IEzus auas:iri.r ras Theobald 1929:337.
FUNDATRIX (after Mason 1940). Coloration. General color
of living specimens dark brown to black. General color of
mounted specimens dark brown. Base of antennal segment III
light; rest of antennae dark, Abdomen with dark lateral
patches. Rest of specimen dark brown.
Inoholo1iv. Antennae shorter than body, 5-segmented;
secondary sensoria absent. Leng1?s of antennal segments:
III, 0.29-0.38; IV, 0.14-0.21; V, 0.10-0.14 + 0.14-0.18.
Head width through eyes 0.39-0.46. Rostrum reaches hind
coxae. Cornicles 0.35-0.46 long, cylindrical, slightly
curved, heavily imbricated throughout. Cauda 0.14-0.21
long, conical, not constricted, with 2 pairs of lateral se-
ALATE VIVIPAROUS FEM-'ALE. Coloration. General color
of living specimens dark brown. General body color of
mounted specimens dark amber. Base of antennal segment III
pale brown; rest of antennae and head dark brown. Dorsal
patch on abdomen dark brown on a pale bacVkgrournd; lateral
sclerites dark brown. Cornicles, cauda, and anal and geni-
tal plates brown to dark brown.
Morphology. Body length 1.19-2.00. Antennal tuber-
cles very prominent, conver.gi,, scabrous. Setae on head
and antennae pointed, inconspicuous. Antennae 6-segmented,
subequal to body length, with 12-20 sensoria on segment III
in a more or less straight line; segment VI with 1 lateral
accessory sensorium subequal in size to primary sensorium.
Lengths of antennal segments: III, 0.33-0.49; IV, 0.21-
0.31; V, 0.17-0.26; VI, 0.10-0.15 + 0.34-0.54. Head width
through eyes 0.30-0.44. Rostrum reaches hind coxae; apical
segment 0.106-0.112 long, with 1 pair of lateral setae in
addition td the usual 3 apical pairs. Setal formula of
first tarsal segments 3:3:3. Second hind tarsal segment
0.079-0.101 long. Dorsal patch on abd-,mn on segments II-V;
dorsal patch on VI-VII continuous with postsiphuncular scler-
ites. Lateral tubercles usually present on segments II-IV,
frequently on I, and sometimes on V. Dorsal tubercles us-
ually on VII and occasionally also on VIII. Segments VII
and VIII each with 4 dorsal setae. Cornicles 0.26-0.42 long,
cylindrical, moderately imbricated. Cauda 0.12-0.17 long,
elongated and conical, not constricted, with 4-9 lateral
setae. Anal plate with 14-18 setae evenly distributed. Gen-
ital plate with 2 setae anteriorly and 11-13 along posterior
APTEROUS VIVIPAROUS FEMALE. Coloration. General color
of living specimens dark brown to black. General color of
mounted specimens dark brown to black. Head and antennal
segments I and II dark brown; III, IV, and base of V pale to
light brown; apex of V and VI black. Abdomen black. Corni-
cles black. Cauda and anal and genital plates dark brown.
Morphology. Body length 1.54-2.40. Antennal tubercles
very prominent, strongly converging, scabrous. Setae on
head and antennae blunt to pointed, inconspicuous. Antennae
6-segmented, shorter than body; secondary sensoria absent.
Lengths of antennal segments: III, 0.26-0.55; IV, 0.18-
0.38; V, 0.16-0.26; VI, 0.10-0.16 + 0.32-0.49. Head width
through eyes 0.4L-0.'1. Rostrum reaches hind coxae; apical
segment 0.126-0.135 long, with 1 pair of lateral setae in
addition t6 the usual 3 apical pairs, rarely with an addi-
tional lateral seta. Second hind tarsal segment 0.090-0.117
long. Abdomen with lateral tubercles frequently on segments
I-IV, rarely on V; dorsal tubercles usually on VII, rarely
on VIII. Cornicles 0.36-0.60 long, cylindrical, curved,
strongly imbricated. Cauda 0.10-0.20 long, elongated and
conical, not constricted, with 6-10 lateral setae. Setae
on anal and genital plates as in alate viviparae.
MALE. Generally similar to alate viviparous female..
Antennal segment III with 54-60 scattered sensoria, IV with
20-21, V with 11 in addition to the primary sensorium at
the distal end. Cauda 0.10 long, triangular, with 4 apical
OVIPAROUS FEMALE. Essentially the same as the apter-
ous viviparous female. Lengths of antennal segments: III,
0.19-0.20; IV, 0.13-0.14; V, 0.14; VI, 0.09 + 0.21. Hind
tibiae slightly swollen with 44-49 pseudosensoria scattered
over the middle 2/3 of the segment.
Types. Location of Fabricius' types unknown. Types
of asperulae Walker, euphrasiae Walker, and quasipyrinus
Theobald in the British Museum.
Distribution. Holarctic and Australian. Eastern U.S.:
Maine to North Carolina and westward to the Mississippi River.
Collections studied. MAINE: Caribou, 24 June 1954
(W.A.S., G.W.S.), Prunus nigra Ait.; Veazie, 15 Aug. 1959
(C.F. Smith), Prunus sp. NEW HAMPSHIRE: Durham, 5 June
1923 (P.R. Lowry), cherry. MASSACHUSETTS: Boston, 6 July
1921 (H. Morrison), Prunus sp. and Japanese flowering cherry;
22 July 1921 (H. Morrison), Rubjs ursinus (= vitifolius Cham.
and Schlecht.); Dedham, 3 Aug. 1917 (Mosher), cherry;
::elrose, 11 Sept. 1917 kosherer), chokecherry. NEW YORK:
Babylon, 5 June 1939 (F. S. Elanton), cherry; 13 June 1939
(E. Kurz), cherry; Columbia Co., 11 June 1940 (Turner),
cherry; Ithaca, 27 June 1939 (T. R. Hansberry), Prunus
avium L.; 19 June 1952 (Kerr), near potato fields;
Mattituck, 9 June 1961 (Latham), Lonicera sempervirens L.-
MICHIGAN: Fernville, 26 Tay 1911, sweet cherry; Grand
Haven, 15 June 1936 (L. G. Strom), Prunus sp.; Keeler,
15 July 1946 (J. K. Karlovic). WISCONSIN: Milwaukee,
3 June 1932 (L. G. Strom), Prunus domestic L.: 16 Oct.
1934 (L. G. Strom), Prunus avium L.; Starts, 20 June 1947
(T. Andre), potatoes. :AE! JERSEY: Trenton, 4 June 1888,
cherry; Wenonah, 26 June 1907. PENiLSYLVANIA: Carlisle,
6 June 1929 (E. D. Eaton), oxheart cherry; Harrisburg,
16 Pay 1905; Philadelphia, 5 July 1927 (W. Moffitt),
cherry; Quincy, 20 May 1932 (J. 0. Pepper), peach; Reading,
2 June 1941 (J. O. Pepper), cherry; Scotland, 21 May 1932
(J. 0. Pepper), peach; State College, 4 Sept. 1940
(J. O. Pepper), wild cherry; 16 July 1961 (J. 0. Pepper),
sweeping Camaedaphne calyculata Moench.; York, 26 June
1894, cherry; 14 Iay 1942 (J. O. Pepper), sour cherry,
OHIO: Colqmbus, 25 May 1897, cherry; 21 May 1925 (A. N.
Tissot), cherry; 9 June 1958 (F. W. Mead), Prunus sp.;
20 June 1958, same data; Mowrystown, 25 ::ay 1941 (A. N.
Tissot), sour cherry. INDIANA: Tndianapolis, 14 June 1911
(H. Morrison), cherry; 12 June 1912, same data. ILLINOIS:
Jerseyville, 25 June 1929 (Hottes and Frison), cherry; Mt.
Carmel, 26 May 1928 (Hottes and Frison), peach; Oregora,
28 June 1928 (Hottes and Frison), cherry. WASHINGTON, D.C.:
16 June 1916, cherry; 4 June 1917, cherry. HAEYLAND: Arundel,
23 May 1906;Baltimore, 22 June 1927, cherry. VIRGINIA:
Chatham, 22 May 1942, cherry; Pulaski, 13 June 1916 (J. F.
Strauss), cherry; Vienna, May 1913 (A. C. Baker), cherry,
29 Oct. 1915, same data. NORTH CAROLINA: Boone, 15 Oct.
1963 (C. E. Smith), Prunus sp.; Durham, 4 Apr. 1964,same
data; Laurel Springs, 9 June 1960 (Smith and Tuatay),
sweet cherry; 2 June 1961 (C. F. Smith), cherry; Smoky
Mountain National Park, 30 June 1942 (C. F. Smith),
Prunus pennsylvanica L. f.; Watauga Co., 8 Aug. 1961
(J. Graham), Prunus sp. TENJjESSEE: Hamilton Co., 22 May
1939 (F. Turner), cherry; Roane Co., 26 ".ay 1937, same
data; Sullivan Co., 7 June 1939, same data. UTAH: Brigham,
5 Miay 1959 (G. F. Knowlton), sweet cherry; Providence
5 June 1963 (W. J. Hanson), sour cherry; Salt Lake City,
13 May 1961 (C. Nelson), cherry; 2 June 1961, same data.
BRITISH COLUMBIA: Creston, 4 Aug. 1959 (W. H. A. Widdle
and D. McComb), cherry. ORECOrI: Cornelius, 24 May 1936,
cherry; Corvallis, 25 July 1938 (T. Y. Hsiao), cherry.
CALIFORNIA: Berkeley, 12 May 1916 (E. 0. Essig). USIJM, FSCA,
CFS, HBB, JOP.
Biology. This species overwinters as eggs on Prunus
spp. The fundatrices hatch in late May and alate funda-
trigeniae appear in late June early July. Remigration
and mating occurs in September-October. This species fre-
quently spenis the entire year on its Prunus host without
migration to any secondary hosts (Gillette and Taylor 1908).
Distinguishing characteristics. One of the lateral
accessory sensoria on antennal segment VI subequal in size
to the primary sensorium. Body color dark brown to black.
This species is very similar to Jyzus lythri (Schrank),
which also has 1 large accessory sensorium on antennal
segment VI. However, the latter is green in color and has
the dorsal abdominal patch confined to the middle of the
abdomen, rather than extending to the lateral sclerites.
Myzus hemerocallis Fakahashi
Figure 3, A-G
Mvzus hemerocallis Takahashi 1921:24.
ryz7us hemerocallidis Takahashi 1931:69.
ALATE VIVIPAROUS FEMALE. Coloration. General color
of living specimens green. General color of mounted speci-
mens light amber. Head brown, antennae light amber. Femur
yellow; tibia yellow, its apex and tarsus brown. Abdomen
pale, without a dorsal patch. Cornicles, cauda, and anal
and genital plates yellow to yellowish brown.
ur^bol-.og. Body length 1.82-2.06. Antennal tubercles
moderately prominent, converging, scabrous. Setae on head
and antennae pointed, inconspicuous. Antennae shorter than
body, with 3-12 sensoria in a straight line on segment III.
Lengths of antennal segments: III, 0.34-0.48; TV,
0.20-0.32; V, 0.14-0.20; VI, 0.09-0.12 + 0.29-0.32. Head
Figure 2..--yzus cerasi (Fab.). A-F, alate viviparous
female: A, head and anten-al segments I-III;
B, sclerotic pattern on abdomen (1/2 scale);
C, base of anteniial segrmenlt VI (2 x scale);'
.D, rostrum; E, cornicle; F, cauda; G-I, apter-
ous viviparous female: G, hLead; H, cornicle;
I, cauda; J-K, iale: J, antennal segments
III-V; K, cauda; L-0, oviparous female: L,
head; M, hind tibia; N, cornicle; 0, cauda.
A C L
width through eyes 0.42-0.46. Rostrum reaches middle coxae;
apical segment 0.101-0.119 long, with 2 pairs of lateral setae
in addition to the usual 3 apical pairs. Setal formula of
first trasal segments 3:3:3. Second hind tarsal segment 0.103-
0.121 long. Abdominal tubercles absent. Segments VII and
VIII each with 4 dorsal setae. Cornicles 0.32-0.38 long,
cylindrical, lightly imbricated throughout. Cauda 0.15-0.17
long, strongly constricted medially giving apex a knobbed
appearance, with 3-5 lateral setae. Anal plate with 8-14
setae evenly distributed. Genital plate with 2 setae anteri-
orly and 12-14 along posterior edge, rarely with an additional
seta either near the anterior edge or in the center of the
APT-:RO0US VIVIPAROUS FEMALE. Coloration. General color
of living specimens yellowish green. General color of
mounted specimens yellow. Antennal segments V and VI brown,
head and rest of antennae yellow. i;emur yellow. Tibia
yellow, its apex and tarsus brown. Abdomen pale. Cornicles,
cauda, and anal and genital plates pale to yellow.
orphology. Body length 1.80-2.30. Antennal tubercles
moderately prominent, parallel to slightly converging, scabrous.
Setae on head and antennae blunt to pointed, inconspicuous.
Antennae shorter than body, without secondary sensoria.
Lengths of antennal segments: III, 0.21-0.36, IV, 0.12-0.22;
V. 0.10-0.16; VI, 0.08-0.10 + 0.21-0.27. Head width through
eyes 0.44-00.50 Rostrum surpasses middle coxae; apical
segment 0.114-0.124 long, with 2 pairs of lateral setae in
addition to the usual 3 apical pairs. Second hind tarsal
segment 0.094-0.130 long. Abdomen without sclerotic areas
and tubercles. Segments I-V bilaterally, segmentally
scaly in concentric patterns (Fig. 3E); segments VI-VIII
transversely scaly. Cornicles 0.38-0.65 long, cylindrical,
imbricated throughout. Cauda 0.15-0.19 long, with con-
striction as in alate viviparae, with 3-4 lateral setae
and occasionally an additional subterminal seta. Setae
on anal and genital plates as in alate viviparae.
Sexual forms are not known.
Types. In the Taihoku Agricultural Experiment
Distribution. Oriental and Subtropical Nearctic.
Eastern U. S.: Alabama, Florida, Louisiana..
Collections studied. ALABAMA: Pleasant Grove,
29 Apr. 1969 (F. F. Smith), Hemerocallis sp. FLORIDA:
Gainesville, 6 Jan. 1958 (L. A. Hetrick), daylily; 17 Jan.
1960 (J. 0. Pepper), daylily; 30 Jan. 1969 (A. N. Tissot),
Hemerocallis fulva L.; 6 Mar. 1970 (R. J. Nielsson and
D. Mays), daylily; Holly Hill, 10 Jan. 1964 (J. W. Pott),
Liriope sp.; Jacksonville, 23 Jan. 1958 (R. L. King),
Hemerocallis sp.; Orangedale, 6 Jan. 1958, same data;
Tallahassee, 6 Feb. 1958 (R. H. Miller and K. P. Bragdon),
Hemerocallis sp. LOUISIANA: Baton Rouge, 16 Feb. 1958
(L. D. MT.ewsome), daylily; 22 :ar. 1958, same data; 20 Aug.
1960 (C. AP. Smith), daylily; 4 Apr. 1969 (H. B. Boudreaux),
daylily; 10 Apr. 1969, same data; Lafayette, 21 Mar. 1958
(L. D. Newsome), daylily. US.Jll, FSCA, CFS, HBB, JOP.
Figure 3.--Myzus hemerocallis Tak., A-G. A-C, alate vivi-
parous female: A,head and antennal segments I-
III; B, cornicle; C, cauda; D-G, apterous vivi-
parous female: D, head, E, pattern of scales on
abdomen (1/2 scale); F, cornicle; G, cauda. M.
lythri (Schrk.), H-P. H-K, M, 0, alate vivi-
parous female: H, head and anternal segments I-
.III; I, sclerotic pattern on abdomen (1/2 scale);
J, base of antennal segment VI (2 x scale); K,
rostrum; M, cornicle; 0, cauda; L,N,P, apterous
viviparous female: L, head; N, cornicle; P,
H J K E
I vO P
Bioloz. The biology of this species has yet to be
investigated. It has only been collected on Hemerocallis
spp. and might possibly be anholocyclic on that host.
Aphids have been found on the roots of these plants from
Kay to November (unpublished). Alternate hosts are not
Distinguishing characteristics. Abdomen of alatae
without dorsal sclerotic patch. Abdomen of apterae with
unusual scaly pattern as described above. Abdominal
tubercles absent. Cauda with apex knob-like.
Myzus lythri (Schrank)
Figure 3, H-P
Aphis lythri Schrank 1801:115.
Aohis mahaleb Koch 1854:113.
:yzus lythri (Schrank). Passerini 1863-26. Mason 1940:
13-4. Heinze 1961:30.
;yzus mahaleb (Koch), Passerini 1863:26. Gillette and
Phorodon humuli var. mahaleb (Koch). Buckton 1876:168.
Myzoides lythri (Schrank). van der Goot 1913:184.
iyzaphis lythri (Schrank). van der Goot 1915:184. Nevsky
ALATE VIVIPAROUS FEMALE. Coloration. General color
of living specimens green. General color of mounted
specimens brownish amber. Base of antenna segment III
pale; head and rest of antennae brownish amber. Basal 1/4
of femur pale, apex brown. Tibia pale to yellowish amber;
apex and tarsus brown. Abdomen with brown central patch and
lateral sclerites on a pale background. Cornicles, cauda,
and anal and genital plates brown.
Morphology. Body length 1.22-2.01. Antennal tubercles
not prominent, scabrous. Setae on head and antennae pointed,
inconspicuous. Antennae considerably shorter than body, with
10-19 sensoria in an irregular row on segment III; segment VI
with one lateral sensorium subequal in size to the primary
sensorium. Lengths of antenna segments: III, 0.31-0.45;
IV, 0.16-0.28; V, 0.11-0.22; VI, 0.07-0.12 + 0.20-0.39.
Head width through eyes 0.35-0.41. Rostrum reaches middle
coxae; apical segment 0.108-0.126 long, with 1 pair of
lateral setae in addition to the usual 3 apical pairs, rarely
with an additional lateral or surface seta. Setal formula
of first tarsal segments 3:3:3. Second hind tarsal segment
0.079-0.101 long. Abdomen with a rectangular central
sclerite on segments III-V, lateral and pleural sclerites
on II-IV, presiphuncular sclerite on V, and postsiphuncular
sclerite on VI-VII which is partially connected to medial
transverse patch on those segments. Lateral tubercles
frequently on segments II-IV. Dorsal tubercles absent.
Segments VII and VIII each with 4 dorsal setae. Cornicles
0.25-0.40 long, cylindrical, weakly imbricated. Cauda
0.12-0.18 long, conical to subtriangular, slightly con-
stricted medially, with 5-7, usually 6, lateral setae.
Anal plate with 10-15 setae evenly distributed. Genital
plate with 2 setae anteriorly, 1 or 2 smaller setae on the
anterior edge or in the rl.ddle of the plate, and 14-19
along the posterior ed.:e.
APTEROUS VIVIPAROUS FEMALE. Coloration. General
color of living specimens light green. Geuerial color of
mounted specimens pale. Head and antennae pale; segment
VI light amber. Femur yellow. Tibia yellow; its apex
and tarsus amber. Abdomen without pigmented sclerites.
Cornicles, cauda, and anal and genital plates yellow to
Morphology. Body length 1.33-1.99. Antennal tubercles
moderately prominent, slightly converging, scabrous. Setae
on head and antennae blunt to pointed, inconspicuous.
Antennae subequal to 1/2 the body length, secondary sensoria
absent; segment VI with 1 lateral sensorium subequal in
size to the primary sensorium. Lengths of antenna seg-
ments: III, 0.19-0.31; IV, 0.10-0.21; V, 0.09-0.16; VI,
0.06-0.10 0.15-0.28. Head width through eyes 0.35-0.46.
Rostrum surpasses middle coxae; apical segment 0.090-0.137
long, with 1 pair of lateral setae in addition to the
usual 3 apical pairs. Second hind tarsal segment 0.072-
0.104 long. Dorsal surface of abdomen sclerotic, rugose;
pigmented areas absent. Cornicles 0.38-0.55 long, cylindri-
cal, tapered and slightly curved, strongly imbricated.
Cauda 0.13-0.20 long, conical, not constricted, with 4-7
lateral setae. Setae on anal and genital plates as in alate
MALE. Generally similar to alate viviparous female.
Antennal segment III with 29-43 scattered sensoria, IV with
14-18, end V with 7-8 in an irregular row in addition to
the primary sensorium.
OVIPAROUS FEIA.LE. No specimens were available for
stuoiy. Gillette and Palmer (1934) figure the hind tibia but
do not discuss any of the morphological characters. Accord-
ing to F;ason (1940), oviparae are similar to apterous vivi-
parae, slightly darker. Cornicles are more slender and
less curved than in apterae.
Types. Location unknown.
Distribution. Holarctic. Eastern U.S.: Widespread.
Collections studied. 4AINE: Veazie, 15 Aug. 1959
(C. F. Smith), Prunus sp. NEW YORK: Ithaca, 21 July 1939
(T. R. Hansberry), Lythrun salicaria L.; Lockport, 4 June
1959 (L. L. Pechuman), Prunus Eahaleb L.; Mt. Kisco, 22 Aug.
1960 (J. Graham), Lythrum salicaria L. MICHIGAN: LaSalle,
15 Sept. 1938 (L. J. Bottimer), Lythrum alatum Prush.
PENNSYLVANIA: Huntingdon, 31 Aug. 1962 (J. O. Pepper),
Ludwi.ia alternifolia L.; Springbrook, 17 July 1945 (Sailer),
Arelanchier sp.; State College, 14 Jan. 1942 (J. 0. Pepper),
Prunus sp.; 31 May 1964 (J. O. Pepper), Pruinus pensylvaica
L. f.; 7 June 1964 (J. 0. Pepper), Prunus virginiana L.
OHIO: Wooster, 16 I.ay 1960 (R. Rings), Prunus mahaleb L.
WASHINGTON, D.C.: 31 May 1916 (D. W.), pear; 4 Apr. 1917
(A. C. Baker), plum. NORTH CAROLINA: Blowing Rock, 29 Sept.
1961 (C. F. Smith), light trap; Blue Ridge Parkway, 29 Sept.
1961 (C. F. Smith), Eilobium sp.; Boone, 11 Sept. 1963
(Smith, Pepper and Tissot), Prunus sp.; 15 Oct. 1963 (C. F.
Smith), PruWuIs sp.; Fletcher, 25 May 1960, light trap;
Wilkesboro, 9 Sept. 1963 (Smith, Pepper and Tissot),
Epilobium sp. FLORIDA: Gainesville, 4 May 1937 (L. J.
Bottimer and A. N. Tissot), L~jthrum lireare L.; Otter Creek,
30 Apr. 1970 (A. N. Tissot and R. J. Nielsson), Lythrum sp.;
Palmetto, 21 Mar. 1941 (A. N. Tissot), Lythrum lineare L.;
Vero Beach, 19 Apr. 1944 (Light), Lythrum sp. USNM, FSCA,
Biology. This species alternates between the primary
hosts Prunus spp. and the secondary hosts Epilobium spp.
and Lythrum spp. There is no published seasonal history
for lythri, but from the collection records it seems that
primary viviparae migrate to secondary hosts in May June
and alate virginoparae remigrate in September October.
The species may also live throughout the year on its primary
Distinguishing characteristics. One lateral sensorium
on segment VI subequal in size to the primary sensorium.
Alate viviparae with a quadrangular medial dorsal patch
on segments III-V. Dorsum of abdomen of apterae sclerotic,
rugose, without pigmentation. Cauda conical to subtriangular.
This species is close to M. cerasi (Fab.). The means
by which they can be separated are discussed under the
Ilyzus oliveri (Essig)
Figure 4, A-H; 5
Ueotoxoptera violae Theobald 1915a:131. (junior homonym)
HicromVzus oliveri Essig 1935:160-2
azns (lioto:-o-ptera) oliveri (Essig). Eastop 1961:36 -
This species, with violae Pergande and formosanus
Takahashi, had been placed in the genera Micromyzus (Essig
1935, 1936), Neotoxoptera (Theoblad 1915a, Pepper 1965),
Fullawayella (Baker 1919, Takahashi 1921, Hardy 1931), or
Idiopterus (Hottes and Frison 1931). Neotoxoptera, type
violae Theobald, was generally considered to be only a
variant of Micromyzus violate (Pergande). The genus
Fullawayella is a synonym of Idiopterus, monotypic genus.
This left only the genus Micromyzus for aphids with numerous
sensoria on the antennal segments of alate viviparae, wing
veins conspicuously bordered and abdomen with a dorsal
Neotoxoptera is closely related to Phordon while
Micromjpu is tropical, with no knotm Palaearctic relatives
and is generally restricted to ferns. Since this species
lacks the finger-like projections on the antennal tubercles
which are characteristic of Phorodon, it was placed in the
genus gyzus, subgenus Neotoxoptera (Eastop 1961).
This species is considered here without subgenerio
designation and is believed to be related to M. persicae.
ALATE VIVIPAROUS FEMALE. Coloration. General color
of living specimens dark red to greenish black. General
color of mounted specirins brown. Head and antenna
segments I and II brorm, rest of antennae pale. Wing veins
brown i with wide fuscous bcrders. Basal 1/3 of femur pale,
apex brown. Tibia amber; its apex and tarsus dark brown.
Lateral sclerites and dorsal patch on the abdomen brown on
a pale background. Cornicles, cauda, and anal and genital
MIorpholozy. Body length 1.18-1.97. Antennal tubercles
very prominent, converging, scabrous. Setae on head and
antennae pointed, inconspicuous. Antennae longer than body,
with 33-54 scattered sensoria on segment III, 11-27 on IV,
and 0-7 in a straight line on V in addition to the primary
sensorium. Lengths of antennal segments: III, 0.46-0.71;
IV, 0.35-0.55; V, 0.29-0.41; VI; 0.13-0.16 + 0.44-0.59.
Head width through eyes 0.35-0.44. Rostrum reaches middle
coxae; apical segment 0.114-0.119 long, with 4-6 lateral
setae in addition to the usual 3 apical pairs. Setal formu-
la of first tarsal segments 3:3:2. Second hind tarsal segment
0.067-0.110 long. Abdomen with lateral sclerites on seg-
ments II-IV, presiphuncular sclerite on V, postsiphuncular
sclerite on VI, dorsal patch with lateral extensions on
III-VI and a median pigmented patch on VII. Segments VII
and VIII transversely wrinkled laterally. Lateral tubercles
sometimes on sclerites on segments III and IV, occasionally
on II. Segment VII with 2 dorsal setae; VIII with 4.
Cornicles 0.24-0.31 long, swollen, nearly smooth. Cauda
0.12-0.15 long, conical, blunt, not constricted, with 2-3
pairs of lateral setae and sometimes with an additional
subterminal seta. Anal plate with 8-12 setae evenly dis-
tributed.' Genital plate with 2 setae anteriorly and 6-11
along posterior edge, rarely with 1-2 additional setae in
center of plate.
APTEROUS VIVIPAROUS FEMALE. Coloration. General color
of living specimens brown. General color of mounted speci-
mens yellowish amber. Head yellowish amber. Antennal seg-
ments I and II brown, III-V yellowish amber with apices
darker, VI yellowish amber with a brown area around the
primary sensorium. Basal 1/6 of femur pale, remainder amber.
Tibia amber; its apex and tarsus brown. Abdomen pale.
Cornicles, cauda, and anal and genital plates light yellowish
Morphology. Body length 1.18-1.95. Antennal tubercles
very prominent,strongly converging, scabrous. Setae on head
and antennae blunt to pointed, inconspicuous. Antenna longer
than body, usually without secondary sensoria (1 specimen
with 1 sensorium basally on segment III on 1 antenna and 3
on the other). Lengths of antennal segments: III, 0.43-0.65;
IV, 0.32-0.51; V, 0.28-0.41; VI, 0.12-0.15 + 0.44-0.59. Head
width through eyes 0.33-0.38. Rostrum surpasses middle
coxae; apical segment 0.110-0.122 long. Abdomen without
pigmented areas; segments VII and VIII strongly wrinkled
transversely. Cornicles 0.26-0.30 long. Cauda 0.09-0.12 long.
Other features as in alate viviparae.
Sexual forms are unknown.
Tjyoes. Location of types unknown.
Distribution. Nearctic, Ethiopian, Australian.
Eastern U. S.: Widespe'oad.
Collections studied. MIASSACHUSETTS: Newton Centre,
29 Nov. 1898 (Pergande), violets. RHODE ISLAND: Providence,
9 Feb. 1900 (F. Roberts), violets. NEW YORK: Ithaca, Dec.
1923 (C. R. Crosby), onion; Long Island, 15 Oct. 1901
(Pergande), violets; Poughkeepsie, 14 Dec. 1898, same data.
PENNSYLVANIA: Lemont, Apr. 1951 (J. 0. Pepper), onion.
MARYLAND: College Park, 15 Jan. 1928 (J. L. Hoerner),
Viola sp. WASHIGITON, D. C.: 11 Nov. 1898 (Pergande),
violets; 18 Nov. 1898, same data. VIRGINIA: Norfolk,
18 Apr. 1939 (L. D. Anderson), onion; Vienna, 30 Nov. 1901
(Pergande), Viola odorata L.; 4 May 1912 (A. C. Baker),
violets; Apr. 1914, same data. NIORTH CAROLINA: Anson
Co., 15 Apr. 1939 (Turner), peach. TENrESSEE: Pleasant
View, 23 May 1946 (L. B. Scott), tobacco. FLORIDA:
Gainesville, 4 Feb. 1927 (!A. N. Tissot), Valeriana
scandens L.; 8 Feb. 1927 (A. N. Tissot), Viola sp.; 25 Feb.
1928 (A. N. Tissot), Valeriana scandens L. and Viola
septemloba LeConte; 5 :*ar. 1928 (A. N. Tissot), Valeriana
scandens L. and Viola walteri House; 9 Feb. 1932 (E. West
and A. N. Tissot), Valeriana scandens L. and Viola
cucullata Ait.; 13 Feb. 1936 (A. N. Tissot), Valeriana
scandens L.; 20 Feb.. 1936, same data; 28 Apr. 1939 (A. N.
Tissot), Viola sp.; 30 May 1939, same data; 15 Mar. 1940
(West, Arnold, and Tissot), Viola floridana Brainerd;
6 Apr. 1941 (A. N. Tissot), Stellaria media (L.) Vill.;
Moultrie, 17 Feb. 1949 (W. G. Genung), Viola sp. ONTARIO:
Toronto, 20 Oct. 1899 (Pergande), violets. KANSAS:
Lawrence, 6 Nov. 1903 (C. E. Sanborn), Viola sp.
CALIFORi!IA: Arcadia, 22 Mar. 1969 (H. G. Walker),
Teucriumn ch'-nj.edrys L.; 8 Apr. 1969 (H. G. Walker), Dianthus
deltoides L.; 13 May 1969 (H G. Walker), Cerastium tomen-
tosum L.; 23 Oct. 1969, same data; 21 Nov. 1969 (H. G.
Walker), Viola odorata L.; 13 Feb. 1970 (H. G. Walker),
Cerastium torentosum L.; 5 Nov. 1970, same-data. USNM, FSCA,
Biology. The biology of this species has not been
thoroughly studied. It apparently lives parthenogenetically
throughout the year on its host plants (Eastop 1961).
The collection from Dianthus deltoides L., 8 Apr.
1969, Arcadia, Calif., contained both alates and apterae;
the latter were normal but the alate viviparae had 7-10
sensoria in a straight line on antennal segment III, no
secondary sensoria on segments IV and V and a very abnormal
pattern of wing venation (Fig. 5 G-J, N-O). Eastop (1961)
had observed similar forms and concluded that they were
Distinguishing characteristics. Alate viviparae
with numerous sensoria on antennal seguents III and IV
and a few usually on V. Wing veins conspicuously bordered.
Abdomen with a dorsal sclerotic patch.
.';yzus ornatus Lain
Figure 4, I-Q
ENzus ornatus Laing 1932:52. Essig 1938:92. -Essig 1947:611.
S ~zus co7l7l-'4 iacchlnati 1883:235-6, in Heinze 1961:32.
13o0w;cdes ornatus (Laing). Aizenbjrg 1966:148-9.
Figure 4.-- M~zus oliveri (Essig), A-H. A-D, alate vivi-
parous female: A, head and antennal segments
I-IV; B, sclerotic pattern on abdomen (1/2
scale); C, cornicle; D, cauda; E-H, apterous
viviparous female: E,head; F, rostrum; G,
cornicle; H, cauda. M. ornatus Laing, I-Q.
I-M, alate viviparous female: I, head and
antennal segments I-III; J, sclerotic pattern
.on abdomen (1/2 scale); K, rostrum; L, cornicle;
M, cauda; N-Q, apterous viviparous female: N,
head; 0, sclerotic pattern on abdomen (1/2
scale); P, cornicle; Q, cauda.
Figure 5.-*Variation in wing venation in r;'zus oliveri
(Essig). A-J, forewings, K-O, hindwings;
A, B, and K typical. A-F and K-M normal;
G-J and N-O from apteriform alatae.
E z ,- o
Heinze (1961) considered ornatus Laing and portulacae
Macch. to be the same species and the latter, due to
priority, would then be the correct name. I examined a
translation of Macchiati's description, which is in Italian
and unfortunately is too brief and without figures. No
specimens were available.for study so that an accurate
judgment of this synonomy cannot be made. However, since
Heinze's work is the first published record of this name
since the original description, as far as I am able to
determine, I declare Kyzus portulacae Eacchiati to be a
nomen oblitum by virtue of Art. 23, Sect. b of the Inter-
national Code of Zoological Nomenclature (50-year rule),
and therefore designate Myzus ornatus Laing as the valid
name for this species.
Aizenberg (1966).placed ornatus Laing as the type of
his new genus Neomyzodes, differing from Ayzus, in which
it had been placed by Eorner (1952b), in the structure of
the vertex, first hind tarsal segment with only 2 setae in
early nymphs and 3 in adults and the absence of abdominal
tubercles. From Myzodes it differs in antennal tubercles
less developed and more parallel, antennae only about
1/2 the body length (rather than more than 3/4) and
hind tibiae of first instar nymphs with spines between
the setae. I consider Aizenberg's
genus to be based on evolutionary differences of a minor
magnitude and therefore retain ornatus in the genus Myzus
ALATE VIVIPAROUS FEMALE. Coloration. General color
of living specimens dark green. General body color of
mounted specimens brownish amber. Head brown. Base of
antennal segment III pale, rest of antennae amber. Basal
1/4-1/3 of femur pale, its apex dark brown. Tibia yellowish
amber, its apex and tarsus brown. Central abdominal
sclerite and lateral sclerites brown on a pale background.
Cornicles, cauda, and anal and genital plates amber-brown.
oroholocy.. Body length 1.19-1.50. Antennal tubercles
not prominent, parallel to slightly converging, weakly
scabrous. Setae on head and antennae pointed, inconspicuous.
Antennae shorter than body, with 4-8 sensoria in a straight
row on segment III. Lengths of antennal segments: III,
0.34-0.37; IV, 0.24-0.25; V, 0.18-0.20; VI, 0.10-0.13 +
0.23-0.27. Head width through eyes 0.30-0.31. Rostrum
not reaching middle coxae; apical segment 0.074 long, with
1 pair of lateral setae in addition to the usual 3 apical
pairs. Setal formula of first tarsal segments 3:3:3. Second
hind tarsal segment 0.072-0.079 long. Abdomen with an
irregular central sclerotic area on segments III-V, a row
of pleural spots on each side on I-IV, large lateral
sclerites on II-IV, presiphuncular sclerite on V, and well-
developed postsiphuncular sclerite on VI-VII. Abdominal
tubercles absent. Sagmeqts VII and VIII each with 2 dorsal
setae. Cornicles 0.21-0.23 long,cylindrical, slightly
tapered, moderately imbricated. Cauda 0.13 long, conical,
not constricted, with 5-7 lateral setae. Anal plate with
12-14 setae evenly distributed. Genital plate with 2 setae
anteriorly and 8-10 along posterior edge.
APTEROUS VIVIPAROUS FEMALE. Coloration. General color
of living specimens green. General body color of mounted
specimens pale to yellow. Antennal segment VI light amber,
head and rest of antennae yellow. Legs yellow; apex of
tibia and tarsus amber. Abdomen pale with brown markings.
Cornicles, cauda, and anal and genital plates yellow.
Morphology. Body length 1.12-1.45. Antennal tubercles
not very prominent, slightly converging, scabrous. Setae
on head and antennae blunt to pointed, inconspicuous.
Antennae subcqual to 1/2 the body length; secondary sensoria
absent. Lengths of antennal segments; III, 0.20-0.25;
IV, 0.13-0.16; V, 0.11-0.13; VI, 0.07-0.09 + 0.17-0.20.
Head width through eyes 0.29-0.35. Rostrum surpasses hind
coxae; apical segment 0.090-0.097 long, with 1 pair of lateral
setae in addition to the usual 3 apical pairs. Second hind
tarsal segment 0.068-0.079 long. Dorsal surface of abdomen
sclerotic, wrinkled, with transversely elongated pigmented
patches dorsolaterally on seg-ments I-VI and with transverse
rows of pigmented spots medially on V and VI. Cornicles
0.26-0.30 long, cylindrical, tapered, moderately imbricated,
Cauda 0.11-0.14 long, conical with obtuse apex, not con-
stricted, with 4-6 lateral setae. Setae on anal and genital
plates as in alate viviparae.
Sexual forms are not known.
Types. In the British Museum. Paratypes in the USNM.
Distribution. Holarctic. Eastern U.S.: Widespread.
Collections studied. I studied specimens collected
from Maine to Florida and because the species is so poly-
phagous, I have reduced the collection records to a list of
plants, by families, on which it was collected; presumably
many of these collections were on greenhouse plants.
Ai'iARAT:HACEAE: Achyrantes sp., Celosia sp.; ARACEAE:
Dieffenbacchia sp.; ARMERIACEAE: Limonium sp.; BALSAMINACEAE:
Impatiens sp.; BEGONIIACEAE: Begonia sp.; BORAGINACEAE:
Anchusa sp.; CARYOPHYLLACEAE: Dianthus caryophyllus L.,
Myosotis sp.; CELASTRACEAE: Euonymous sp.; CISTACEAE:
IFelia]nther.um sp.; COMMELINACEAE: Tradescantia sp.; COM-
POSITAE: An.cgclus sp., Artemisia vulgaris L., Aster sp.,
Bellis sp., Bidelns bipinnata L., Chrysanthernum leucanthe-
rilun L., Chrysanthemumi sp., Cineraria sp., Eupatorium
glandulosum HBK, E. riparium Regel, Gaillardia picta
Gray var. lorenziana Hort., Gazania sp., Helianthus anuus
L., Tutisia sp., Parthenium sp., Senecio sp., Sonchus asper
(L.) All., Sonchus sp.; CRASSULACEAE: Sedum sp.; CRUCIFERAE:
cabbage Brassica oleracea L., cauliflower B. oleracea
otris DC, Hutch1 nsia sp., horseradish Roria armoracia
A. Hitchc., Teesdalia nudicaulis (L.) R. Br.; CUCURBITACEAE:
Cucurbita sp., Echinocystis sp.; ERICACEAE: Azal.ea procaurabens
L., Azalea.sp., heather Calluna vul.j is (L.) Hull, heath -
Erica 3p.;* EUHORBIACEAE1: Croton sp.; GENTIAI ACEAE: Gentiana
acaulis L., Gentiana sp.; GESIEIEIACEAE: Gloxinia sp.,
Saintpaulia sp.; GRATIIIAE: barley Hordeum sp.; HEDERACEAE:
Hedera helix L.; JUNIPERACEAE: Junioerus sp.; LABIATAE:
Coleus sp., Lavandula sp., Mentha sp., Salvia azurea Lam.,
S. greEggi A. Gray, S. mellifera Hort., S. microphylla Hort.,
TeucrJum chamaedrys L., T. scorodonia L., thyme Thymus
serphyllum L.; LEGUMINOSAE: Lathyrus sp., shamrock -
Trifolium dubium Sibth., Trifoliun sp.; LILIACEAE: garlic -
Allium muscari L., Convallaria sp., Heloniopsis breviscapa
Maxim.; iMALVACEAE: Malvaviscus sp.; MORIIACEAE: Dipsacus
fullonum L.; ONAGRACEAE: Fuchsia sp.; ORCHIDACEAE: Bletia
sp., Cymbidium sp.; PAPAVERACEAE: Dicentra spectabilis Lem.;
PITTOSPORACEAE: Pittosporum sp.; POLYGOITIACEAE: sorrel -
Rumex acetosella L., R. obtusifolius L.; PRIMULACEAE:
Androsace lanuginosa Wall., gycla~3n sp., Primula sp.,
Ramonrdia sp.; RANUNCULACEAE: Clercais seiboldii D. Don.,
Helleborus sp.; ROSACEAE: Contoneaster pyracantha (L.)
Spach., Fragaria sp., Potentilla sp., Rosa sp., Spiraea sp.;
RUBIACEAE: Gardenia sp.; SAXIFRAGACEAE: Hydrangea sp.,
Saz.ifraga sp.; SCROPHULARIACEAE: Mazus pumilo R. Br.,
Veronica sp.; SOLAI1ACEAE: potato Solanum tuberosum L.;
THEACEAE: Camellia sp.; ULMACEAE: Ulmus campestris L.,
Zelkova sp.; UMBELLIFERAE: celery Aium graveoles L.,
Azorella sp., parsley Petroselinum crisoum (Mill.) Mansf.;
Valeriana sp.; VERBENACEAE: Duranta plumieri Jacq., Duranta
sp., Lantana camera. L.; VIOLACEAE: Viola sp. IUS.;M, FSCA,
CFS, JOP, HG'-.
B iola This species lives parthenogenetically
throughout the year on plants in greenhouses and, in
subtropical climates, in the field. Alates are only
occasionally produced and sexual forms are not known.
Distinguishing characteristics. Abdominal tubercles
absent. Apterae with a bilateral series of transversely
elongate pigmented patches located pleurally on the abdomen.
Myzus persicae (Sulzer
Figure 6, A-L
Aphis Eersicae Sulzer 1776: 105.
Aphis dianthi Schrank 1801: 114.
Aphis vulgaris Kyber 1815: 9.
Achis furcioes Rafinesque 1817: 361, in Hottes 1931: 63.
Cinara r._'?haui ;;osley 1841: 747-8.
Aphis r 5:.e Curtis 1842: 53.
Achis dubia Curtis 1842: 54.
ADhis convolvuli Kaltenbach 1843: 40-1.
Aohis v.2.-ntor S'3e3 1846: 137, in Meier 1954: 348.
Aphis cvno-:!s3i Walker 1848: 2217.
Aohis particeps Walker 1848: 2217.
Aphis sodalis Walker 1848: 2217.
Aplis2 p2rsola Walker 1848: 2246.
Aphis roli.'nin Walker 1849: 32.
Anhis auct. .,alk-,r 1849: 33 in part.
Aphis gEgessa Walker 1849: 38.
Anhis malvae Walker 1849: 47.
Arhiis ~mtiJ:. Walker 1849: 49.
Aohis dlijt;i Walker 1849: 50.
Rhj.plosinhu^r dianthi (Schrank), Koch 1854: 55-6. --
Williams 1910: 69 in part.
Aphiks jrne.; ic j, 1- Passerini 1860: 36.
l11OT-!nlin lm aesrm Lsi.' e (Sulzer), Passerini 1863: 20.
Aps s -~i alcBco.'. Boisduval 1867: 251.
ZUs ..'rsic.e (Sulzer), Buckton 1877: 178-80. -- Gillette
and Pal-.rnir 1934: 204-6.
Siohononhori aThyr ,- ntc-s Monell 1879: 18-9.
Sitooopj.,ra clo,,dulella nonoll 1879: 19.
Si.o.onoon!i.-'a an~irrhi ~r i .I.~ochiati 1883: 228.
;'- u .v' O.:-' l nd 'i-~18'36: 31-32
-, .'.-'nos (Monell), Oestlu;rd 1.887: 74.
.r -os .i i eorivolvuli (Kalt.), Del Guercio 1900: 159.
vI;,is ;.p'n .Jii ndderson 1901: 72-3.
Ehooz.!osichur,, t u lir:o Davidson 1910: 377.
Phonalosinh'um dianthi poae Williams 1910: 70.
Fnoroion cy;,.olossi Williams 1911: 172-3.
Rho-alosiphum solani Theobald 1912: 165-9.
Rhhialoosiohu. betae Theobald 1913: 918, in Eeier 1954: 349.
inacrosio:lum betae (Theobald), Theobald 1913: 153.
iacrosiphium antirrhinii (-iacchiati), Theobald 1913: 157.
l'yzoides oersicae (Sulzer), van der Goot 1913: 84. --
van der Goot 1915: 170. -- 1917: 48.
Myzodes tabaci Mordwilko 1914: 52.
Rhonalosiphum lactucellum Theobald 1915: 115.
Myzus godetiae Shinji 1917: 49-50.
Rhopalosiphum solanella-Theobald 1918: 12, in Meier 1954:349.
Rhiooalosio'hui tuberosellae Theobald 1919: 161.
I,. zus tuberosellae (Theobald), Theobald 1922: 1.
Myzus impacts Theobald 1926: 342-3 in part.
Thoralosipooninus dianthi (Schrnm.nk), Borner 1926: 22.
i:yzodes oersicae (Sulzer Mordwilko 1928: 192. -- Borner
1951: 101-11. -- Borner 1952a: 122-7. -- Heinze
Phorodon (yzodes) persicae (Sulzer), Borner 1930: 139.
Myzus (Nectarosiphon) persicae (Sulzer), Hille Ris Lambers
1946: 197. -- 1952a: 119-21.
This long list of synonyms is due to the very poly-
phagous habit of the species and to its wide distribution.
Schrank (1801) described Aphis dianthi from carnation.
In Europe there is an anholocyclic strain of persicae on
carnation (Hille Ris Lambers 1966) which is sometimes given
subspecific status. However, the dianthi of most authors
is not from carnation and therefore refers to persicae s.s.,
or, if from carnation, are actually records for Myzus certus
(Walker), a close relative of persicae, which does d~.;:'ge
carnation. I have examined specimens from carnation which
were persicae; therefore I know that persicae can be found
on this host, but it causes little or no damage and cannot'
propagate continuously on this host.
The question of Myzodes and Nectarosiphon was taken up
undsr the generic discussion. Borner (1930) considered
flyzcdes to be a subgenus of Phorodon since the second hind
tarsal segment has only 2 setae and the hind tibiae of first
instar nymphs are without spines between the setae. In set-
ting up this association he ignored the structure of the
antennal tubercles. Later zodes was considered to be a
separate genus (Borner 1951, 1952a, b, Heinze 1960).
According to Aizenberg (1966), Myzodes has been placed
in Myzus because authors considered the character of the
antennal tubercles and not the differences in this character
in each group. Aizenberg considered Nyzodes to be near the
Pentalonini, and particularly the genus Fullawayella Del G.
(Idiopterus Davis), due to similarities in setae, the
structure of the antennal tubercles, and an approximate
similarity in the form of the radial sector. However, I
would point out that the apterae of Idiooterus have
elongate capitate setae on the head and typically bear
sensoria on the base of antennal s~reenit III. These
characters would seem to place the genus closer to Capito-
phorus. However, persicae does not have these sebae on
the head, never has secondary sensoria in the apterae, and
has Prunus as its primary host, all typical characters of the
ALATE VIVIPAROUS FEI1ALE. Coloration. General color
of living specimens yellowish green to green. Color of
mounted specimens light amber. Base of antennal segment
III pale, head and rest of antennae light brown. Basal
1/3 of femur pale, rest light amber; tibia pale, its
apex brown; tarsus brown. Abdomen with a light amber central
sclerotic area and lateral sclerites on a pale background.
Cornicles, cauda, and anal and genital plates amber.
Mornholopy. Body length 1.45-2.25. Antennal tubercles
moderately prominent, converging, scabrous. Setae on head
and antennae pointed, inconspicuous. Antennae subequal to
body length or somewhat longer, with 8-19 sensoria in a more
or less straight row on segment III. Lengths of antennal
segments: III, 0.40-0.61; IV, 0.29-0.50; V, 0.10-0.19 +
0.35-0.78. Head width through eyes 0.37-0.45. Rostrum
reaches hind coxae; apical segment 0.094-0.135 long, with
2-3, rarely 4, lateral setae and 0-3 surface setae in
addition to the usual 3 apical pairs. Setal formula of
first tarsal segments 3:3:2. Second hind tarsal segment
0.092-0.140 long. Abdomen with lateral sclerites on
segments (I) II-IV, presiphuncular sclerite on V, post-
siphuncular sclerite on VI-VII, and a doral sclerotic patch
on II-V connected to pleural patches on these segments.
Lateral tubercles frequently on any or all of the sclerites
II-IV, rarely on V and never on I. Dorsal tubercles fre-
quently on VIII, sometimes on VII. Segment VII with 2-3
dorsal setae, VIII with 4-5. Cornicles 0.29-0.48 long,
cylindrical (fundatrigeniae) to slightly swollen (virgin-
geniae), smooth. Cauda 0..17-0.23 long, conical, not
constricted, with 3 pairs of lateral setae, sometimes with
only 2 setae on 1 side and 3 on the other or with an addi-
tional terminal seta. Anal plate with 12-20 setae evenly
distributed. Genital plate with 2 setae anteriorly and
8-12 along the posterior edge, rarely with an additional
seta either on the anterior edge or in the center of the
APTEROUS VIVIPAROUS FEMALE. Coloration. General
color of living specimens light yellow to light green.
Mounted specimens pale; apex of V, VI, tarsi, and sometimes
apex of cornicles brown.
Forvpholocy. Body length 1.56-2.35. Antennal tuber-
cles moderately prominent, parallel, scabrous. Setae on
head and antennae blunt to pointed, inconspicuous. Antennae
subequal to body length, secondary sensorial absent.
Lengths of antennal segments: III, 0.32-0.48; IV, 0.24-
0.40; V, 0.19-0.30; VI, 0.09-0.19 + 0.32-0.78. Head width
through eyes 0.40-0.48. Rostrum roaches hind coxae; apical
segment 0.108-0.155 long, with 2-3, rarely 4, lateral setae
and 0-3 surface setae in addition to the usual 3 apical
pairs. Abdomen without sclerites. Lateral tubercles some-
times on segments III and IV, rarely on V. Dorsal tubercles
often on segment VIII and rarely on VII. Segment VII with
2-3 dorsal setae, VIII with 4-5. Cornicles 0.30-0.63 lo: g,
cylindrical (fundatrigeniae) to slightly swollen (virgino-
geniae), lightly imbricated. Cauda 0.17-0.26 long, coni-
cal, not constricted, with 3 pairs of lateral setae, rarely
with only 2 setae on 1 side and 3 on the other. Anal plate
with 9-20 setae evenly distribute. Genital plate with 2
setae anteriorly and 9-17 along posterior edge, rarely with
an additional seta either on the anterior edge or in the
center of the plate.
FUVDATRIX. Essentially the same as the apterous viviparous
female, except for 5-segme.ted antennae.
]AL`E. Alate, generally similar to alate viviparous
female. Antennal segment III with 28-48 scattered sensoria,
IV with 11-30; V with 9-23 in addition to the primary sensorium.
Cauda 0.09-0.13 long, with 6 setae.
OVIPAROUS FEIHALE. Red, but otherwise essentially the
same as apterous viviparous female. Hind tibiae swollen, with
16-71pseudosensoria scattered over the middle 2/3 3/4 of
Types. Location of types uni:rkown. Types of Myzus
pergandii Sanderson in the USiiM.
Distribution. Worldwide. Eastern U.S.: Widespread.
Collections studied. I studied specimens collected
from every state within the area. Since this species is so
widespread and so polyphagous, I have reduced the collection
records for this species to a list of plants, by families,
on which this species was collected. ACANTHACEAE:
Beloperone sp., Jacobina coccinea Hiern.; ALLIOI-ACEAE:
Boerhaavia riscosa Lag. & Rodr. Small Fl., four-o'clock -
Mirabilis jilapa L.; AMARANTHACEAE: alligator weed -
Alternanthera philoxeroides (1-iairt.) Griseb., Arraranthus
blitoides S. Wats., A. hybridus L., A. retroflexus L.,
Arriranthus sp., Celosia nitida Vahl.; AAiARYLLIDACEAE:
Afave sp. Alstroneria sp., Tarlcissus sp., Polianthes
tuberosa L., Sprekelia sp.; AiliIJlACEAE: dill Anethum
graveolens L., parsley Apium petroselenium L., Aralia
seiboldii Hort.; APOCYNACEAE: oleander Nerium oleander L.,
Vinca major L., V. minor L., Vinca sp.; AUTIUFOLIACEAE: Ilex
rar~',ariLnsis St. Hil., Ilex sp.; ARACEAE: Caladium sp.,
Philodendron selloum C. Koch; Philodendron sp., Syngonium
sp.; ARALIACEAE: Polyscias q ilfoylei Bailey, Polyscias
sp., Schefflera sp.; ARISTOLOCHIACEAE: Dutchman's pipe -
Aristolochia macrophylla Lam., Aristolochia sp.; ASCLEPIA-
DACEAE: Asclepias incarnata L., Asclepias sp., Matelea
floridana Woodson; BALSATITIACEAE: Impatiens sp.; BIGiONIACEAE:
Catalpa sp.; BORAGINACEAE: Amsinckia barbata Greene, A.
douglasiana A. DC, A. intermedia F. & M., Amsinckia sp.,
Begonia sp., forget-me-not Myosotis sp., Pavonia hastata
Cav., Tournefortia sp.; BROHiELIACEAE: Spanish moss -
Tillandsia usneoides L.; LW:iiiLh'i!ti"ACEBAE: Riedlea sp.;
CACTACEAE: Eiphyllum sp.; CAHPANULACEAE: Camoanula sp.;
CANNABINACEAE: hop- H.nL:.lus lupulus L.; CAPPARIDACEAE:
Cleome serrulata Pursh., C. spinosa Jac.; CAPRIFOLIACEAE:
honeysuckle Lonicera japonica Thuni., Lonicera sp.,
elderberry Sambucus sp., Viburnum sp.; CARYOPHYLLACEAE:
Cerastium tomentosum L., mouse-ear C. vulgatum L.,
carnation Dianthus barbatus L.; D. caryophyllus L., D.
chinensis L., Dianthus sp., Oxalis stricta L., Oxalis sp.,
Polycaron !.c:trlanyl.lum L. f., Sileno sp., chickweed -
Stellaria media (L.) Vill.; CELASTRACEAE: Eaonymous japonica
L., E ~ymous sp., Shaeffleria sp.; CHEiiOPODIACEAE: Atriplex
sp., Russian pigweed Axris amaranthoides L., Baccharis sp.,
beet Beta vulgaris L., pigweed Chenopodium album L.,
C. ambrosiodes L., Jerusalem oak C. botrys L., Chenopo-
dium sp., spinach Soinacia oleracea L.; COMPOSITAE: Achil-
lea millefolium L., Actinea odorata 0. Ktze., ragweed -
Ambrosia trifida L., Ambrosia sp., Antennaria sp., Anthe-
mis sp., Aster sp., daisy Bellis integrifolia Michx.,
Bellis sp., Bidens sp., Calendula sp., safflower Cart-
hamus sp., cornflower Centaurea cyanus L., Chrysanthemum
morifolium Ram., Chrysothamnus sp., Cineraria sp., Cirsium
lanceolatum (L.) Hill, Dahlia sp., Echinops sohaerocephalus
L., Emilia coccinea (Sims) Sweet., Erechtites hieracifolia
(L.) Raf., Erechtites sp., Erigeron philadelphicus L.,
E. strigosus Muhl., Eupatorium serotinum Michx., Gail-
lardia sp., Gazania sp., Gerbera sp., Granhalium obtusi-
folium L., G. spathulatun Lam., Gynura aurantiaca (Blume)
DC, Gynura sp., artichoke Helianthus tuberosus L., Hel-
ianthus sp., straw-flower Helichrysum sp., HidalFoa
wercklei Hook. f., hauknieed Hieracium sp., lettuce -
Lactuca sativa L., L. scariola L., Lactuca sp., Iaaruta
cotula (L.) DC,Myriocephalus sp., Pyrrhoappus carolinia-
nus (Walt.) DC, Scruentum sp., Senecio aureus L., S. con-
fusus J. Britton, S. decorus Greens., S. glabellus Poir.,
Senecioides cinerea (L.) Kuntze, Sonchus arvensis L., S.
asper (L.) All., S. oleraceus L., Sonchus sp., marigold -
Tagetes sp., Tansy sp., TaraYacum sp., Verbesina sp.,
cockleburr Xanthium sp., Zinnia sp.; CONVALLARIACEIAE:
Asparajus 1srengeri Regel.; CONVOLVULACEAE: dodder -
Convolvulus sp., sweetpotato Ipomoea batatas (L.) Lam.,
morning glory Ipomoea sp., bindweed Strophocaulos
sp.; CRASSULACEAE: Bryophyllum pinnatum (Lam.) S. Kurz.,
Bryophyllum sp., Kalanchoe flamrea Stapf., Sedum sp.;
CRUCIFERAE: Barbarea vulgaris R. Br., mustard Bras-
sica alba Boiss., rutabaga B. campestris L., Chinese
cabbage B. chinensis L., B. juncea (L.) Coss., rape -
B. 8Eaus L., B. nigra (L.) Koch, cabbage B. oleracea
L., collards (Kale) B. oleracea acephala DC, broccoli
(cauliflower) B. oleracea botrytis DC, Kohlrabi B.
sinapistrum Boiss., Brassica sp., Cakile edentula (Big-
el.) Hook., Canoides halei Small, shepherd's-purse -
Capsella bursapastoris (L.) :Iedic., Cardaria pubescens
(C.A. Meyer) Jarmolenko, Cardaria sp., Dentaria integri-
folia Nutt., flixweed Diplot~xis tenuifolia (L.) DC,
Lepidium draba L., L. perfoliatum L., Lepidium sp., Mat-
thiola sp., watercress Nasturtium officinale R. BR.,
Norta altissjma (L.) Britton, Radicula sp., radish -
Raph;.nus sativus L., R. re.nani-strui L., horseradish -
Roripa armoracia (L.) A. Hitche., Sisymbrium altissimum
L., S. officinale (L.) Britton, Soohia pi nata (Walt.)
Britton, Thelypodium laciniatum L.; CUCURBITACEAE:
cantaloupe Cvcrlmic melo L., gourd Cucurbita lagen-
aria L., squash C. mrxira Duchesne, crook-n3ck squash
- C. moschata Duchesne, cucujmber C. sativus L., Erod-
ium sp., Luffa sp.; CYP'RACEAE: CGperus esculentus L.;
ERICACEAE: Azalea sp.; Rhodendron sp., blueberry Vac-
cinium myrsinites Lam.; EUPHORBIACEAE: Acalypha sp.;
FOUQUiRIACEAE: Fouqueria splendens Engelm.; GENTIAI!ACEAE:
Buddleia sp.; GERAlIiAC--AE: Erodium sp., Geranium caro-
linianum L., G. dissectum L., Geranium sp.; GRAMINAE:
oats Aveina sativa L., foxtail Chaetochloa sp., John-
son grass Holcus halpensis L., barley Hordeum sp.,
wheat Triticum vulgare L., corn Zea mays L.; HEDEBA-
CEAE: Hedera helix L.; IRIDACEAE: Freesia sp., Gladiolus
sp., Iris tingitana Boiss. & Reut., Iris sp.; LABIATAE:
Coleus sp., Galeopsis tetrahit L., gill-over-the-ground
- Glechoma hederacea L., Lamium amplexicaule L., Mentha
sp., catnip Nepeta cataria L.; LEGUIII-OSAE: Acacia
sp., Caesalpinia sp., Cassia sp.,, Cercis canadensis L.,
Lupinus sp., alfalfa Iedicago sativa L., sweet clover
- Melilotus sp., Phaseolus coccineus L., P. vulgaris L.,
Phaseolus sp., pea Pisum sativum L., black locust -
Robinia pseudoacacia L., red clover Trifolium prataense
L., white clover T. reopens L., Trifolium sp., cowpea
Vigna sinensis L., Wisteria sp.; JUGLtANDACEAE: pecan -
Carya illinoisensis (Wang) K. Kock; LABIATAE: Salvia
azurea Lam., S. greggii A. Gray, S. mellifera Hort., S.
nmicrophylla Hort., S. nutans L., Teucriun chamaedrys L.,
T. flavum Hort., T. scorodonia L.; LILIACEAE: Agaanthus
sp., Aae virginica L., leek Allium porrum L., onion
Allium sp., Aloe variegata L., Aloe sp., Asparagus of-
ficinalis L., Chlor'chytum sp., Lilium auratum Lindl., L.
candidum L., Easter lily L. lonTiflorum Thunb., L. long-
iflorum formosanum Hort., L. soeciosum Thunb., L. specio-
sum rubrum Hort., tulip L. sylvestris L., tiger lily -
L. tigrinum Thunb., Smilax rotundifolia L., Ornithogalum
sp., Yucca sp.; LYTHRACEAE: crape-myrtle Lagerstroemia
indica L.; MALVACEAE: okra Abelmoschus esculentus (L.)
Moench., Abutilon sp., hollyhock Althaea rose Cav.,
Crotalaria spectabilis Both., Gossypium sp., Hibiscus can-
nabinus L., H. syriacus L., Hibiscus sp., Malva borealis
Wallm., M. ne-lecta Wallr., M. parviflora L., M. rotundi-
folia L., 7alva sp., Malvastrum coromandelianum (L.)
Garcke., Malvaviscus drummondii T. & G., Sphaeralcea em-
oryi Torr., Sphaeralcea sp.; VORINGACEAE: Moringa oleifera
Lam.; MUSACEAE: banana Musa sp.; MYRICACEAE: Myrica
sp.; MYRTACEAE: guava Psidium uajava Raddi; NYCTAGINA-
CEAE: Bougainvillea glabra Choisy., Bougainvillea sp.;
NYMPHAEACEAE: Nelumbo sp.; OLEACEAE: lilac Syringa
vulgaris L., OSIAIUTHUS sp.; ONAGRACEAE: Fuschsia sp.,
evening primrose Oenothera laciniata Hill, 0. sneciosa
Nutt., Oenothera sp.; ORCHIDACEAE: Cattaleya sp., Cymbidium
sp.-, Cripedium sp., Epidendrum secundum Jacq.,Ibidium odora-
turn (Nutt.) House, Saccolabium sp.; PALILAE: Sabal sp.;
PAPAVERACEAE: bleeding heart Dicentra sp., Papaver
somniferum L., Pacaver sp.; PAPAYACEAE: papaya Carica
ua1.va L.; PASSIFLORACEAE: Passiflora edulis L.; PEDIA.-
LACEAE: Sesamum orientale L., Sesamum sp.; PHYTOLACCACEAE:
pokeweed Phytolacca decandra L., Phytolacca sp.; PIPER-
ACEAE: Piper sp.; PITTOSPORACEAE: Pittosoorum undulatum
Vent., Pittosporum sp.; PLANTAGINACEAE: Plantago virgin-
ica L.; PLUMBAGINACEAE: Limonium sp.; POLYGONACEAE: buck-
wheat Fagopyrum fagopyrum (L.) Karst., Nephrolepis sp.,
smartweed Persicaria sp., Polygonum convolvulus L., P.
lacathifolium L., P. pennsylvanicum L., rhubarb Rheum
sp., curled-dock (yellow-dock) Rumex cripus L., R.
obtusifolius L., reddock E. sanzuineus L., Rumex sp.;
PORTULACEAE: Portulaca sp.; PROTEACEAE: Protea sp.;
PRINULACEAE: Cyclamen sp., Primula sp.; RANUNCULACEAE:
Anemone sp., Clematis recta L. var. mandshurica -'a.im,,
Delphinium sp., Paeonia sp., Ranunculus sp.; ROSACEAE:
Chaenomeles sp., Crataegus compitalis Beadle, C. limno-
philoides Murrill, C. newelliana Hurrill, C. subpaludosa
Murrill, C. subviridis Beadle, Crataegus sp., Cydonia sp.,
strawberry Fragaria sp., apple Malus sylvestris Rydb.,
Malus sp., plum Prunus americana Marsh., apricot P.
armeniaca L., sour cherry P. cerasus L., P. erarginata
L., P. fasciculata L., P. hortulana Bailey, P. nira Ait.,
pin cherry P. ennsylvanica L. f., P. serotina Ehrh.,
chockcherry P. virginiana L., Prunus sp., pear Pyrus
cGmmunis L., Rosa sp., Rubus alleghaniesis Porter, red
raspberry R. carolinianus Rybd., blackberry R. lac-
inIatus Willd., R. occidentalis L., R. phoenicolasius
Penstemon sp., mullen Verbascum sp., Veronica sp.;
SOLANACEAE: red popper Capsicum anuum L., C. baccatum
L., C. betacea L., C. frutescens L., Capsicum sp., Cestrum
nocturnum L., Datura meteloides Dunal., Jimoson weed D.
stramonium L., Datura sp., Hyoscyamus nier L., Lycium
halimifolium Mill., tomato Lycopersicon lycopersicon_(L.)
Karst., Nicotiana glauca Graham, tobacco N. tabacum L.,
Petunia sp., Physalis turbinata Medic., Physalis sp.,
horse-nettle Solanum carolinense L., white horse-nettle -
S. elaeagnifolium Cav., eggplant S. melonena L., S.
nigrum L., Jerusalem-cherry S. pseudocapsicum L., S.
seaforthianum Andr., S. tuberosum L., Solanum sp.; SPONDIA-
CEAE: mango Maanif era indica L.; TAIABICACEAE: Teagrrix
sp.; THEACEAE: Camellia sp.; TROPAEOLACEAE: nasturtium -
Tropaeolom majus L.; UMBELLIFERAE: celery Apium graveo-
lens L., Chaerophyllum teinturieri Hook., coriander -
Coriandrum sativum L., carrot Daucus carota L., Eryngium
sp., parsley Torilis nodosa (L.) Gaertn.; 'URTICACEAE:
Urtica sp.; VERBENACEAE: Caryopteris sp., Cierodendron
bungeii Hort., hracoiort., ra Hort., C. nutans Hort., _.
trichotomum Hort., Cochranea anachusaefolia (Poir.) Guerke.,
Lippia sp., Tectona grandis L., Verbena sp.; VIOLACEAE:
Viola arvensis Murr., V. odorata L., pansy V. tricolor
L., Viola sp.; VITACEAE: Virginia creeper Parthenocissus
Quinguefolia (L.) Planch. USNM, FSCA, CFS, HBB, HGW, JOP.
See also Horsfall (1924), Gillette and Palmer (1934),
Cottier (1953), Bodenheimer and Swirski (1957) and Leonard,
et al. (1970).
Eiolog.y. -iordwilko (1907) first combined the individual
work of earlier authors into a pattern of host alternation for
M. persicae between peach and nany herbaceous plants.
Gillette and Taylor (1908) independently made a similar report.
Eggs overwinter on Prunus persica Sieb. & Zucc.,
P. persica nucipersica Schneid., and older trees of P.
serotina Ehrh. (Borner 1951, Meier 1954). The eggs hatch in
early spring producing nymphs which mature into fundatrices.
There are up to 8 fundatrigenious generations on the primary
hosts. Alates produced in April-June migrate to many
different herbaceous secondary hosts; in general they alight,
leave a few progeny, and then fly to another plant and repeat
the process (van Emden, et al. 1969). During September and
October gynoparae fly to various Prunus spp. :~ales are
produced in October-early November and fly to the primary
hosts and mating occurs. Eggs are then laid from late October
to early December (Horsfall 1924). Berry and Simpson (1967)
found that heavy flights in mid-June and again in early or
mid-September corresponded with peak periods of potato
Hille Ris Lambers (1946) pointed out that the gynoparae
are not host selective but males prefer peach although they
do frequently fly to other hosts; and the nymphs hatching from
eggs mature only on peach and nectarine. This lack of
selectivity in host selection has resulted in several
erroneous reports concerning the primary host plants for
this aphid. Gillette and Taylor (1908) reported overwintering
on peach (Prunus persica), plum (P. domestic L. and americana
Marsh.), apricot (P. armeniaca L.), nectarine (P. persica
nucipersica), cherry (P. cerasus L.), prune (P. insitita L.),
and oviposition on chokecherry (P. virginiana L.), and
sandcherry (P. melanocarpa Shafer). Gillette and Palmer (1934)
listed Prunus persica, domestic, americana, melanocarpa,
cerasus, armeniaca, and besseyi Bailey (sandcherry) as winter
hosts. These reports were interpreted by Borner (1951) as
a confusion between primary and secondary colonization.
A report of hibernation on P. nana Stokes by Wahlgren
(1939) was verified by Hille Ris Lambers (1952a), but Borner
(1952b) and Keier (1954) were unable to find eggs on this
host. Gorham (1942, in Hille Ris Lambers 1946) listed P.
nigra Ait. as an overwintering host in Cananda; a similar
report was lade by Shands and Simpson (1948) in :aine.
There reports differentiate between primary colonization
and autumn wandering flights of virginogeniae. Because
Heinze (1948a, b) never found persicae on P. nigra in the
Dahlem Botanical Garden, Borner (1951) concluded that this
is a subspecies of nigra endemic to North America and not
found in Europe.
Reports of overwintering on cabbage (Huckett 1925,
Theobald 1926) and roses (Fenjves 1945, Volkhart 1939, in
Hille Ris Lambers 1946) are believed to be the result of
misidentifications (Borner 1951).
The green peach aphid also overwinters parthenogenetically
on herbaceous plants; in greenhouses and in the field if the
climate is mild, particularly in tropical and subtropical
areas (van der Goot 1917, Mason 1922, Heinze and Profft 1938,
Jacob 1941, Moericke 1941, 1949, Heie 1954). Gillette and
Bragg (1915) and Mordwilko (1935a, b) assumed that these
represented an independent anholocyclic race, having lost
its generations on peach. Heinze and Profft (1938) and
Klinkowski and Leius (1943) could not however demonstrate
Distinguishing characteristics. Color yellow to green.
Second hind tarsal segment with 2 setae. Cornicles swollen
M. persicae is very close morphologically to Myzus
certus (Walker), a pink or brown aphid found on carnation
and other Caryophyllaceae. The color of the living aphids
is sufficient to distinguish between the two. However,
since most specimens studied are either mounted on slides
or in alcohol and the living color is not evident, the
taxonomist must resort to morphological differences which
are not always exactly differentiating. Hille Ris Lambers
(1959, 1966) pointed out that certus usually has 2 pairs
of lateral setae on the apical rostral segment while persicae
only rarely has 2 complete pairs. The ventral sclerites on
the abdomen of certus are usually very distinct and in a
straight row, uwhreas, in persicae they appear to be absent;
also, certus alatae may have sensoria on antennal segment IV.
The apterae of certus always have swollen coriicles which are
wholly dark, and the dorsum of the abdomen is slightly
sclerotic and pinentUcd; the fundatlrces frequently have
6-segmented antennae; and, the males are apterous (Hille
Bis Lambers 1946, 1966). Additionally the terminal
filament of summer apterae is subequal to twice the length
of the base in certus and always greater than 3 times the
length of the base in persicae (Hille Ris Lambers 1966).
I have also found the following characters to be
useful for separating the two species. In certus the
sensoria on antennal segment III are frequently doubled
at 1 or rore locations, whereas in oersicae this is only
rarely the case. Abdominal segment VII in alate certus
frequently has 3 setae on the postsiphuncular sclerite;
the usual number in persicae is 2. And, the dorsal tubercles
on abdominal segment VIII in certus are subequal to the
diameter of the setal membranes of the dorsal setae on that
segment, while in persicae they are freqnetly as much as 3
times that diameter.
As Hille Ris Lambers (1966) stated, no one character
will always separate the 2 species, but an analysis of a
combination of characters is necessary in order to arrive
at an identification; and, all specimens will not be able
to be identified positively.
The other species of rjjzjis which are very similar mor-
phologically to persicae (ajugae Schouteden, 1903, ascalonicus
Doncaster, 1946, myosotidis Borner, 1950, polaris Hille
Ris Lambers, 1952b, linariae Holman, 1965, and dianthicola
Hille Ris Lambers, 1966) are not found in t.e Eastern United
Eyzus certus (Walker)
Aohis certa Walker 1849: appendix 32.
RhoalosiDhum dianthij (Schrank) in part. Williams 1910:69.
Hizus rersicae var. cerastii Theobald 1926: 323.
Iyzjs caryiooylla.cea-rurm Hille Ris Lambers 1946: 198.
v-yzus certus (W alkefr). Hille Ris Lambers 1946: 198-9. --
yvzodes auctus Borner nec Walker 1951: 103.
Nyzodes certus (Walker.- Borner 1951: 103. Heinze
Myzodes auctus pseudopersicae Borner 1952b: 469.
Borner (1951, 1952b) considered auctus (Walker 1849)
and certus to be separate species and placed persicae
cerastil Theobald and caryophyllacearum Hille Ris Lambers
as synonyms of auctus. However, Walker described the
apterae of auctus as green, and Doncaster (1961) concluded
from transfer experiments that caryophyllacearum and certus
are the same species. This is the view taken in this paper.
Because this aphid is so similar to Myzus persicae
(Sulzer) I have not figured it separately and will not
take up the space with a complete description. I will
however list the chief distinguishing characteristics.
ALATE VIVIPAROUS FEMALE. Coloration. General color
of living specimens dark brown to black.
Morphology. Antennal segment III with 8-17 sensoria
in a more or less straight line, usually more than 10 in
number and with 2-3 clustered at 1 or more locations;
segment IV with 0-2. Apical rostral segment with 2
pairs of lateral setae and 0-3 surface setae in addition
to the usual 3 apical pairs, rarely with only 3 lateral
setae in addition to the surface and apical setae. Corni-
cles always swollen, lightly imbricated. Postsiphuncular
sclerite on abdominal segment VI usually with 3 setae,
occasionally with 2. Lateral tubercles usually on any or
all sclerites on segments II-IV, never on I and V. Dorsal
tubercles usually on abdominal segment VIII, sometimes on
VII; in size, subequal to the diameter of the membrane
surrounding the base of the dorsal setae on that segment,
rarely twice as much as that diameter or more. Ventral
sclerites distinct, arranged in 2 parallel rows ventro-
APTEROUS VIVIPAROUS FEMALE. Coloration. General color
of living specimens pink or brown. Cornicles wholly dark,
or dark other than at extreme apex only.
-oro~olory. Terminal filament of summer apterae
subequal to twice the length of the base of antennal segment
VI; in other morphs as ii oersicae. Setae on apical rostral
segment as in alate viviparous female. Dorsum of abdomen
sclerotic, pigmented. Cornicles always swollen, lightly
imbricated, usually with the basal portion significantly
wider than the mid-portion of the hind tibia. Dorsal
tubercles, when present, as in alate vivparous female.
MALE. Apterous; 34-46 sensoria on antennal segment III,
16-20 on IV, and 8-13 on V in addition to the primary
sensorium at the distal end.
OVIPAROUS FE1:ALE. Hind tibia slightly swollen, with 20-32
pseudosensoria located in the middle 1/2-2/3 of the segment.
Ty1:.; In the British Museum.
Distribution. Holarctic and Oriental oin Caryophyllaceae
and Violaceae. Eastern U. S.: Widespread.
Collections studied. 1;A.[rE: Presque Isle, 11 Oct.
1961 (Shands), Stellaria media (L.) Vill. .!ASSACHUSETTS:
England at Boston, 5 June 1932 (C. A. Eavis), carnation;
Gerr.any at Boston, 13 Apr. 1956 (J. D. Crump, Jr.),
carnation; Ireland at Boston, 17 June 1964 (VanValkenburgh),
Dianthus caryophyllus L.; England at Boston, 17 June 1965
(J. M. VanValkenburgh), Dianthus sp.; Waltham, 12 Mar. 1954
(F. Smith), carnation. NEW YORK: Bellmore, 21 Iar. 1958
(G. V. Johnson), chickweed; Farmingdale, 6 Apr. 1946 (F. F.
Smith), carnation; 3 Jan. 1961 (G. V. Johnson), carnation;
England at New York, 7 Iar. 1949 (Laddey), carnation; Germany
at New York, 4 July 1958 (C. Post), Dianthus sp.; Mexico at
New York, 3 Dec. 1958 (G. Vanech), carnation; France at New
York, 14 Jan. 1959 (E. Erickson), Dianthus sp.; Mexico at
;:e York, 30 Jan. 1959 (T. Reid), Dianthus sp.; 5 Feb. 1959
(Snowden and Reid), Dianthus sp.; 7 Feb. 1959 (T. Reid),
Dianthus sp.; Holland at New York, 15 Oct. 1959 (A. L. Brown),
on Gypsophilla sp.; Mexico at New York, 12 May 1960 (L.
Walden), carnation; England at New York, 3 June 1960
(L. Walden), rose; Israel from New York, 11 June 1960 (L.
Walden), carnation; Germany at New York,ll June 1960
(L. Walden), carnation; Switzerland at fiew York, 12 July
1962 (Shiroishi), Dianthus sp. MICHIGAU: Scotland at
Detroit, 10 July 1968 (Husnik), Dianthus caryophyllus L.
PEIHISYLVAniA: Philadelphia, 20 Aug. 1969 (Arehart),
Dianthus caryophyllus L.; Lancaster, 1 Apr. 1946 (F. F.
Smith), Viola sp. ILLINOIS: Germany at Chicago, 6 Apr.
1957 (G. E. Brenneke), carnation; Mexico at Chicago, 11
Mar. 1958 (R. E. Boyd), carnation; 29 Mar. 1959 (G. E.
Brenneke), Diant}h;s sp.; 4 Far. 1960 (J. Rood), carnation;
23 :a-y 1960 (R. Aolman), Dianthus sp.; 14 ;'ar. 1961 (J. Rood),
Dianthus caryoohyllus L.; 11 May 1961 (Rood and Brenneke),
carnation; 16 May 1961 (Crane), Dianthus sp.; Italy at Chicago,
19 Apr. 1962 (J. Rood), Dianthus caryophyllus L.; Mexico at
Chicago, 5 Jan. 1963 (J. Rood), Dianthus caryophyllus L.;
11 Feb. 1963 (Kennedy), Dianthus sp.; 17 ;iar. 1963 (Imai),
carnation; 26 Mar. 1963 (J. Rood), Dianthus caryophyllus L.
U:EW JERSEY: England at Hoboken, 30 Apr. 1947 (Olsen),
Dianthus sp.; 11 May 1958 (Bennett), Dianthus sp.; Denmark
at Hoboken, 17 Oct. 1956 (R. D. ;unkittrick), Dianthus sp.;
Weston, 5 Apr. 1946 (F. F. Smith), carnation. WASHINGTON,
D.C.: 7 May 1898 (T. Pergande), pink. NORTH CAROLINA:
Atlantic Beach, 7 Apr. 1968 (C. F. Smith), pansy; Raleigh,
14 Apr. 1961 (J. Graham), pansy; 29 Dec. 1961, same data.
GCORGIA: Moultrie, 17 Feb. 1949 (W. G. Genung), violet.
FLORIDA: Gainesville, 21 Feb. 1950 (A. N. Tissot), pansy;
10 Feb. 1956 (R. W. Blacklock), pansy; Mexico at Tampa,
29 Nov. 1961 (F. W. Patton), carnation; Mexico at Miami,
2 Dec. 1959 (Faircloth and Higgins), carnation; 5 Feb.
1960 (Faircloth and Higgins), carnation; 23 Oct. 1961'
(Rowan and Uaka.n.rd), carnation; 17 :ar. 1962 (R. P. Higgins),
carnation; 19 Oct. 1964 (L. A. McClain), Dianthus .sp.; 7 Dec.
1964 (H. S. Harrison), carnation. ALASKA: Japan at
Anchorage, 27 Jan. .-967 (D. V. Akins), Dianthus sp.
WASHINGTON: England ? at Seattle, 24 Apr. 1961 (Bryan
and Nelson), carnation. MISSOURI: Columbia, 29 1ar. 1905
(Hayhurst), pansy. iIEW IEXICO, Bayard, 20 Oct. 1961 (Nielsen
and Heninger), carnation. CALIFORNIA: Palo Alto, 15 Nov.
1956 (Ecucette), carnation; Arcadia, 13 May 1969 (H. G.
Walker), Cerastium tomentosum L.; 23 Oct.1969, same data; 13
.Feb. 1970 (H. G. Ualker), Cerastium tomentosum L. and Viola
tricolor L. hortensis Hort.; 13 Y'ay 1970 (H. G. Walker)
Hymenanthera sp.; 5 Nov. 1970 (H. G. Walker), Cerastium
tomentosum L. HAWAII: Japan at Hawaii, 15 Mar. 1962
(H. A. Woolford), Dianthus caryophyllus L.; 11 Mar. 1963,
same data; 18 :;ay 1965 (W. Chun), Dianthus caryciylluss L.
USNM, FSCA, CFS, HGW.
Although most of these records are for introduced
aphids, I do not believe that this accurately represents
the occurrence of this species in the Eastern U.S.
Biology. This species may hibernate as eggs on Cerastium
spp., Dianthus deltoides L. or Viola arvensis Iurr. Funda-
trices hatch in early April. Alates are produced by the
third generation in late May early June which may migrate to
Stellaria media (L.) Vill. or to other species of Dianthus
and Viola. Oviparae and males develop in mid-October. M.
certus may also live parthenogenetically throughout the year
on its host plants (Hille Ris Lambers 1946, 1959, Meier
The means by which certus can be separated from
persicae have been discussed under the latter species.
Figure 6.--Myzus persicae (Sulzer), A-L. A-F, alate vivi-
parous female: A, head and antenna segments
I-III; B, sclerotic pattern on abdomen (1/2
scale); C, rostrum; D, cauda; E, cornicle of
fundatrigenious alate; F, same of virginogenious
alate; G-J, apterous viviparous female: G, head
H, cauda; I, cornicle of fundatrigenious aptera;
J, same of virginogenious aptera; K-L, male: K,
cauda; L, antennal segments III-V. M. varians
Davidson, M-T. M-O, S, alate viviparous female:
M, head and antenna segments I-III; N, sclerotic
pattern on abdomen (1/2 scale); 0, cornicle; S,
'cauda; P-R, T, apterous viviparous female: P,
cornicle; Q, head; R, rostrum; T, cauda.
E F O
L S T
'yzus varians Davidson
Figure 6, M-T
L~.:us varies Davidson 1912: 409. -- Meier 1954: 232.
L',sj tronicalis Ta:ahashi 1923: 24.
Lyzus cleatifoliae Shinji 1924: 369.
jyzodes varians (Davidson). Borner 1952b: 123. Heinze
Phorodon varians (Davidson). Borner 1952b: 123. Heinze
Borner (1952b) placed this species in Phorodon because
the antennae of first instar nymphs are indistinctly 5-
segucnted, rather than distinctly 5-seg ented as in the
genus yzodes6 (Nlzus). Heinze (1960) followed this scheme,
but at the same time placed the species in Mlzodes because
all other aspects, aside from the segmentation of the
antennae of first instar nymphs, agree with the characteri-
zation of the latter group. Since Ilyzodes is considered
to be a synonym of Myzus Pass., varians is placed in the
latter genus in this paper.
ALATE VIVIPAROUS FEMALE. Coloration. Color of living
specimens green with a reddish tinge. General color of
mounted specimens yellowish amber. Head light amber.
Ant-nrmal az:-.ents I, II, most of III, extreme base and
a-,:< of IV and V, and all of VI amber; base of III and
rest of IV and V pale. (Paralectotype with segments IV and
V wholly amber.) Tibia pale; its apex and tarsus yellowish
amber. Dorsal patch and lateral sclerites on abdomen amber
on a pale ba.kgroound. Cornicles amber. Ccuda and anal
-and ..'n-'cJ.tal plates light yellow.
.-oriThholozy. Body length 1.28-1.73. Antennal tubercles
,l;.c~ately prominent, converging, scabrous. Setae on head
and antennae pointed, inconspicuous. Antennae longer than
body, with 7-8 (12) sensoria in a straight row on segment
III. Lengths of antenna segi3nnts: III, 0.43-0.53; IV,
0.34-0.42; V, 0.29-0.37; VI, 0.11-0.14 + 0.63-0.73. Head
width through eyes 0.39-0.43. Rostrum does not reach middle
coxae; apical segment 0.108-0.117 long, with 1, rarely 2,
pair of lateral setae in addition to the usual 3 apical
pairs. Setal formula of first tarsal segments 3:3:2. Second
hind tarsal segment 0.079-0.097 long. Abdomen with a dorsal
sclerotic patch on segments III-V, lateral sclerites on
II-IV, small presiphuncular sclerite on V and postsiphun-
cular sclerite on VI. Segment VII sclerotic laterally but
not continuous with the postsiphuncular sclerite. Abdominal
tubercles absent. Segment VII with 2 dorsal setae, VIII
with 4. Cornicles 0.35-0.42 long, cylindrical, imbricated.
Cauda 0.16-0.17 long, conical, apex acute, with 7-9 lateral
setae. Anal plate with 12-16 setae evenly distributed.
Genital plate with 2 setae anteriorly and 10-16 along
posterior edge, rarely with an additional seta in the center
of the plate.
APTEROUS VIVIPAROUS FEMALE. Coloration. General color
of living specimens light green. General color of mounted
specimens pale yellow. Apex of antennal segments III-V,
apical part of the base of VI and apex of cornicles amber.
Tarsi light brown. Pest of specimen pale yellow.
Morphology. Body length 1.28-1.73. Anteimal tubercles
moderately prominent, converging, scabrous. Setae on head
and antennae blunt to pointed, inconspicuous. Antennae
longer than body; secondary sensoria absent. Lengths of
antennal segments: III, 0.31-0.50; IV, 0.25-0.36; V, 0.25-
0.35; VI, 0.11-0.13 + 0.58-0.66. Head width through eyes
0.34-0.41. Rostrum reaches hind coxae; apical segment
0.112-0.115 long, with 1 pair of lateral setae in addition
to the usual 3 apical pairs. Second hind tarsal segment
0.076-0.097 long. Abdominal sclerites and tubercles absent.
Segment VII with 2 dorsal setae, VIII with 4. Cornicles
0.35-0.50 long, cylindrical, curved, moderately imbricated.
Cauda 0.17-0.20 long, elongated and conical, not constricted,
with 2-3 pairs of lateral setae and frequently an additional
preapical seta. Setae on anal plate as in alate viviparae.
Genital plate with 2 setae anteriorly and 9-11 along posterior
No sexual forms were available for study. The following
descriptions are from Meier (1954).
1ALE. Generally similar to alate viviparous female.
Antennal segment III with 40-55 scattered sensoria; IV with
16-25; and V with 3-21 in addition to the primary sensorium.
OVIPAROUS FE;ALE. Essentially the samo as apterous
viviparous female. Hind tibiae somewhat swollen and bearing
Types. Two slides in the USNM contained 1 apterous and
4 alate viviparous females. One of the latter which corres-
ponds to Davidson's "light form" is Iacrosiphum euphorbiae
(Thos.). The apterous viviparous female was designated
lectotype and the other alate viviparous females, morpho-
lectotype and 2 par-lectotypes. These latter 3 specimens
(Davidson's "dark form") agree with Shinji's (1924) des-
cription of clematifoliae, a synonym of varians, but differ
from the majority of the specimens studied in the colora-
tion of antenna segments IV and V. I therefore designated
the following plesiotypes which have antennal segments IV
and V dark only at the extreme base and apex of each seg-
ment; the rest of these segments is pale: 4 alate vivi-
parous females; Myakka State Park, Fla., 8 Apr. 1969 (D. H.
Habeck), Clematis sp., in the USNM and FSCA.
Distribution. Nearctic, Oriental. Eastern U.S.:
North Carolina, Florida.
Collections studied. In addition to the lectotypic
specimens above, NORTH CAROLINA: Raleigh, 28 May 1964
(C. F. Smith and J. Graham), Clematis sp.; Wake Co., 1 June
1964 (J. Graham), Clematis sp. FLORIDA: Gainesville, 19
Aug. 1970 (R. J. ::ielsson), Clematis catesbyana Pursh;
Myakka State Park, 8 Apr. 1969 (D. H. Habeck), Clematis
sp. CALIFORNIA: Davis, 20 Mar. 1970 (T. Kono), Clematis
sp. FSCA, CFS.
Biology. This species overwinters on peach, Prunus
persica. The fundatrices hatch from mid-March to early
April and by the end of April are fully grown producing
offspring. There are 3 fundatrigenious generations on
peach and by mid-June the aphids can be found on Clematis
spp. Meier (1954) reported that fuidatrigeniae reared on
peach in the laboratory had no alates in the first genera-
tion, 7.5% alate in the second, and 84% in the third; and,
the maturation of this third generation coincided with the
appearance of alates on Clematis in the field. Males and
gynoparae are produced in late October-early November and
migrate to peach where the gynoparae give birth to ovi-
parae. The sexuales mate and eggs are laid in late
Meier (1954) reported that the species may also live
parthenogenetically throughout the year inside rolled
Distinguishing characteristics. Alate viviparae
usually with the extreme base and apex of antennal seg-
ments IV and V dark and the rest of these segments pale.
Apterous viviparae with the apex of segment III, base and
apex of IV and V and apex of the base of VI dark; rest
of anteumae pale. Cornicles straight to slightly curved,
with apex dark in apterae. Abdominal tubercles absent
(in the forms on Cle'matis).
GE:;US PHORODON PASSERINI
Phorodon Passerini 1860: 27.
Ovat.us (Phorodon) van der Goot. Quednau 1966: 426.
Hille Ris Lambers (personal communication, in Leonard
1963) stated that the genus Ovatus is primarily associated
with the Pomoideae and migrates to Labiatae as summer hosts,
or has developed an independent cycle with sexuales on
Labiatae; however, the genus Phorodon is associated with
Prunoideae and migrates to Urticaceae, or has developed a
complete cycle on the latter. This, then, defines two
biologically distinct groups which should not be joined
together, as was done by Quednau (1966).
Mordwilko (1914) placed Phorodon, with Ovatus, in the
Rhopalosiphea because the antennal tubercles, aside from
the projecting processes, are low and only slightly pro-
truding. Borner (1930) and BSrner and Schilder (1932)
placed Phorodon and Eyzodes in the genus Phorodon, based on
the lack of spines on the hind tibia of first instar nymphs
and the presence of only 2 setae on the first segment of the
hind tarsus of early numphs and in so doing ignored the dif-
ferences in the nature of the antennal tubercles. Consider-
ing the latter character Phorodon and Myzodes were then
recognized as separate genera and placed in the tribe
Phorodontini (Burner 1952b).
Characteristics: Antennal tubercles with very pfominnent
projections, especially evident in apterae. Alate viviparae
with sensoria on internal segments III and IV, absent in
apterae. Rostral segment III with 2 pairs of lateral setae.
Wing venation normal. Alate viviparae with a dorsal pigmented
patch on the ab?1omen, absent in apterae. Cornicles cylindri-
cal; cauda elongate, acutely conical.
Tpe: Aphis humuli Schrank, 1801
One species occurs in the Eastern United States, Phoro-
don humuli (Schrank).
Phorodon humuli (Schrank)
Ahis pruni Geoffrey 1762: 497. (nomen nudum)
Aohis pruni Scopoli 1763. (junior homonym)
Aphis humuli Schrank 1801: 110.
Phorodo'i humuli (Schrank). Buckton 1876: 166-8 Theobald
1926: 273-8. Gillette and Palmer 1934: 207. -
Sampson 1939: 173.
MIyzus hurmuli (Schrank). Pepper 1965: 210.
Phorodon pruni (Geoffrey). Aizenberg 1966: 137.
Ovatus (Phorodon) hu:ui (Schrank). Quednau 1966: 426.
Aizenberg (1966) stated that Aphis pruni Geoffrey 1762
pruni Scopoli 1763 = humuli Schrank 1801 and therefore the
name of the hop aphis should be Phorodon pruni (Geoffrey).
However, pruni Geoffrey is synonyi.ous with Hyalopterus
arundinis (Fab.). A. pruni Scop. which is a junior homonym
is then replaced by humull Schra-uk (Hottes 1930).
ALATE VIVIPAROUS FEMALE. Coloration. General color of
living specimens light greenish yellow. General color of
mounted specimens light brown. Head brown, antennae light
brown, base of segment III pale. Thorax brown. Wings
hyaline, veins amber. Apex of femur and tibia and tarsi
light brown, base of femur and tibia pale to light yellowish
brown. Abdomen pale. Pigncc-ted spots on abdomen, cornicles,
cauda, and anal and genital plates brown.
Mor-pholog. Body length 1.50-1.97. Antennal tubercles
very prominent, slightly converging, tapering, scabrous.
Setae on head and antennae pointed, inconspicuous. Antennae
slightly shorter than body, with 23-31 scattered sensoria on
segment III, and 1-5 in a straight line on IV. Lengths of
antenna segments; III, 0.44 -0.56; IV, 0.25-0.35; V, 0.26-
0.29; VI, 0.10-0.12 + 0.42-0.47. Head width through eyes
0.38-0.40. Rostrum reaches middle coxae; apical segment
0.094-0.108 long, distinctly longer than the second hind
tarsal segment, with 2-3 lateral setae and sometimes an addi-
tional seta on the dorsal surface in addition to the usual 3
apical pairs. Setal formula of first tarsal segments 3:3:3.
Second hind tprsal segment 0.081-0.090 long. Lateral sclerites
on abdominal segments I-VI; pigmented areas pleurally on
I-IV, connected medially on III-IV forming a loose dorsal patch,
together with pigmented transverse areas on VI-VII. Abdominal
segments VII and VIII each with 4 dorsal setae. Abdominal
tubercles absent. Cornicles 0.30-0.41 long, cylindrical,
weakly imbricated. Cauda 0.15-0.16 long, acutely conical,
not constricted, with 6-7 lateral setae. Anal plate with 14
setae evenly distributed. Genital plate with 2 setae anter-
iorly, 2-3 on the midline of the plate and 9-10 along the
APTEROUS VIVIPAROUS FEMALE. Coloration. General color
of living specimens pale green. General color of mounted
specimens pale-light amber. Head light amber; antennal seg-
ment II and base of III pale, rest of antennae light amber-
brown. Legs brown. Abdomen pale, pigmented areas absent.
Cornicles, cauda, and a.-al and genital plates light amber.
.craYiI:T l Ludatrix. Eody length 2.08-2.22. Antennal
tubercles not prominent, without frontal projections. Setae
on head and antennae blunt to pointed, inconspicuous.
Antennae shorter than body, secondary sensoria absent.
Antennae 5-segmented; lengths of antennal segments; III,
0.35-0.36; IV, 0.17-0.19; v, 0.09-0.10 + 0.08-0.10.
Head width through eyes 0.42-0.48. Rostrum reaches middle
coxae; apical segment 0.109-0.119 long, with 1 pair of lateral
setae in addition to the usual 3 apical pairs. Setal formula
of first tarsal segments 3:3:3. Second hind tarsal segment
0.083-0.099 long. Abdominal sclerites and tubercles absent.
Abdominal segment VII with 2 .dorsal setae, VIII with 4.
Cornicles 0.46-0.47 long, cylindrical, tapering and curved,
strongly imbricated. Cauda 0.22-0.23 long, conical, with 7-10
lateral setae. Anal plate with 14 setae evenly distributed.
Genital plate with 2 setae anteriorly, 3-7 along midline of
plate, and 10-14 along posterior edge.
Spring Viviparae. Similar to fundatrix. Antennal
tubercles well developed, converging, with well-developed
frontal projections. Antennae 6-segmented; lengths of
antenna segments: III, 0.47-0.52; IV, 0.30-0.33;
V, 0.28-0.32; VI, 0.11-0.13 + 0.36-0.40. Cornicles
0.66-0.70 long. Other features as in fundatrix.
Fall Viviparae. :uch smaller than summer apterous
viviparae. Body length 1.00-1.37. Lengths of antennal
seSgints: III, 0.18-0.26; IV, 0.13-0.18; V, 0.13-0.18;
VI, 0.06-0.08 4 0.21-0.28. Head width through eyes
0.26-0.29. Apical rostral segment 0.083-0.088 long, with 1
pair of lateral setae in addition to the usual 3 apical pairs.
Cornicles 0.24-0.40 long. Cauda 0.12-0.15 long, with 6-7
lateral setae. Other features as in fall viyiparae.
MALE. Generally similar to alate viviparous female.
Body length 1.50-1.63. Antennae slightly longer than body,
with 41-55 sensoria on segment III, 15-25 on IV, and 6-10
on V in addition to the primary sensorium. Cornicles
0.28-0.31 long, distinctly tapered and slightly enlarged
apically. Cauda 0.08-0.10 long. Anal plate with 9-12
setae evenly distributed.
OVIPARA. Essentially the same as fall apterous vivi-
parae, except antennal segments III and IV sometimes fused
or incompletely separated. Hind tibiae slightly swollen,
with 42-63 pseudosensoria along almost the entire length.
Types. Location of types unknown.
Distribution. Holarctic. Eastern U.S.: Maine, New
York, Washington, D.C., Virginia.
Collections studied. -AIINE: Ashland, 29 Sept. 1953
(W. A. Shands), Prunus nigra Ait.; Houlton, 29 Sept. 1953
(W. A. Shands), Prunus nigra Ait. NEW YORK: Richfield
Springs, 28 Sept. 1887 (T. Pergande), plum; 6 June 1888,
same data. WASHINGTON, D.C.: 16 Nov. 1903 (T. Pergande),
prune. VIRGINIA: Vienna, 10 May 1915 (A. C. Baker), plum.
MANITOBA: Winnipeg, 22 Aug. 1965 (K. A. Hamilton),
Humulus lupulus L. WASHINGTON: Ellensburg, 13 Oct. 1940
(Christenson), chokecherry; Wenatchee, 25 June 1915 (New-
comer), prune. 0 EGON: Albany, 20 Sept. 1933 (N.P.L.),
prune; 7 Oct. 1933 (J. Reaf), prune; Corvallis, 19 Sept. 1933
(J. Reaf), hops; Riddle 15 May 1938 (S. C. Jones) prune.
CALIFORNIA: San Francisco, 25 May 1916 (E. 0. Essig), plum.
USNM, FSCA, CFS.
Biology. Generally, the hop aphids overwinter as eggs
on plum and prune. The eggs hatch in April-early May pro-
ducing the apterous viviparous stem-mothers, or fundatrices.
Alate viviparae appear in late May-July and migrate to hops,
Humulus lupulus L. or they may remain on plum throughout the
year (Gillette and Taylor 1908). The viviparae go through
3-4 generations on hops during the summer and in August-
September gynoparae and males return to the primary host.
Eggs are laid in October-November. Clarko (1904) reported
that the aphids may overwinter as eggs on or in hop vines,
or in the ground near the vines because oviparae were found
on hops in September and the first generations in the spring
were apterous viviparous females in early May. Parker (1913)
noted that the hop aphid may overwinter as viviparae on the
roots of the hop plants; and, that fall migrant viviparae
sometimes flew to cherry, alder, peach, and apple, but no
eggs were laid indicating that in mild winters they might
survive parthenogenetically on these hosts.
Distinguishing characteristics. Elongate finger-like
projections on the antenna tubercles of apterous viviparae;
antennal tubercles of alate viviparae very prominent, al-
though without projections as obvious as in apterae.
Figure 7.--Phorodon humuli (Schrk.). A-E, alate viviparous
female: A, head and antennal segments I-IV; B,
sclerotic pattern on abdomen (1/2 scale); C,
rostrum; D, cornicle; E, cauda; F-H, apterous vivi-
parous female: F, head; G, cornicle; H, cauda;
.I-K, male: I, head and antennal segments I-V;
J, cornicle; K, cauda; L-0, oviparous female: L,
head; M, cornicle; N, cauda; 0, hind tibia.
GENUS HYALOMYZUS RICHARDS
Hvalovnzus Richards 1958: 169
Ov;atus (nH-yaloryzus) van der Goot. Que,.nau 1966: 426.
Eastop (1966), in his discussion of the genus Ovatus,
stated that he suspected Utamohoroohora Knowlton and Hyalo-
myzus Richards to be North American suj)genera of Ovatus
van der Goot. Quednau (1966) listed Hyalomyzus as a sub-
genus of Ovatus. The genus Hyalomrzus is without doubt
closely related to Ovatus, particularly eriobotryae which
has a pattern of host alternation similar to species of
Ovatus. In Hyalomyzus, the general body color is br'c-wnish
amber, the cornicles are swollen and the cycle is between
Crataegus (and other Rosaceae) and Labiatae (Verbenaceae) or
Onagciaccac (Hypericaceae), while in Ovatus the body color is
green, the cornicles are cylindrical and the cycle is
between Crataegus and Labiatae. Because swollen cornicles
are more advanced than cylindrical ones (Hille Ris Lambers,
1949, discussion of IHyperomyzus and Nasonovia) and because
of the greater host range of its species yvalomnyzus is
considered to be more advanced than Ovatus and probably
derived from it. However, since the Onagraceae are in no
way related to the Labiatae, I regard Hyalomyzus to be a
group biologically and riicrphologically distinct from
Ovatus. H. eriobotryae, whichlis biologically similar in
some respects to species of Ovatus, is retained in
Hvaloinymus because of the obvious morphological similarities.
Characteristics. Frontal tubercles very prominent,
parallel to strongly converging and scabrous. Ventral
tubercles, scabrous evaginations of the frons laterad to
the base of the rostrum, well developed (absent in sensor-
iatus). Alate viviparae with many secondary sensoria on
segments III, IV and also usually on V. Rostral segment
III with 2 pairs of lateral setae. Veins of fore;.ing with
narrow fuscous borders; venation normal, except hindwing of
tissoti Nielsson and Habeck in which Cu2 is usually partially
or wholly absent. Setal formula of first tarsal segments
3:3:3. Alate viviparae without a dorsal sclerotic patch
on the abdomen. Abdomen of apterous viviparae strongly
wrinkled on segments I-VI. Cornicles well developed,
cylindrical to swollen, smooth (in alate viviparae) to
strongly imbricated (in apterous viviparae).
TZE. Myzun eriobotryae Tissot.
Key to Eastern U.S. Species of Hyalomyzus
1. Terminal filament less than 3 times as long as base of
antennal segment VI; base of antennal segment VI
more than 1.5 times as long as rostral IV + V;
ventral tubercles absent from head.......sensoriatus
*Terminal filament more than 3 times as long as base of
antennal segment VI; base of anternal segment VI
less than 1.3 times as long as rostral IV + V;
ventral tubercles present on head (Fig. 8H )........2
2. Antennal segment III of alate viviparae with 35 or
more sensoria; abdominal segment VIII with 4
dorsal setae; alternating between Rosaceae and
Labiatae or Verbenaccae................riQbj.tr