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The taxonomy of the Myzini (Homoptera: Aphididae) of the Eastern United States

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Title:
The taxonomy of the Myzini (Homoptera: Aphididae) of the Eastern United States
Creator:
Nielsson, Richard Joseph, 1943-
Publication Date:
Language:
English
Physical Description:
178 leaves : ill. ; 28 cm.

Subjects

Subjects / Keywords:
Abdomen ( jstor )
Animal vivipary ( jstor )
Body length ( jstor )
Cherries ( jstor )
Colors ( jstor )
Genitalia ( jstor )
Head ( jstor )
Insect antennae ( jstor )
Peppers ( jstor )
Species ( jstor )
Aphids ( lcsh )
Dissertations, Academic -- Entomology and Nematology -- UF
Entomology and Nematology thesis Ph. D
Homoptera ( lcsh )
Genre:
bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1971.
Bibliography:
Includes bibliographical references (leaves 166-176).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Richard Joseph Nielsson.

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Th- Tay::c n-, of th.? .j--.-i (Ho.opt..--.: A *
of the E"st.e:n UTnited States


















































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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

Pathology.


iii












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


iv







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


Figure Page
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

By

Richard Joseph Nielsson

June, 1971

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

designated.


vii












INTRODUCTION


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.


Terminology


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

alatae.

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

















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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

segment.

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

primary host.

In southern regions virginogeniae frequently live par-

theno&enLctically throughout the year without the formation

of sexuales.

Literature Review


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

the body.


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,




9

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

Geographical Scope

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.

Mounting Technique

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

Canda balsam.

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.








Organization

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,

Holland HRL.

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
setae........................................ 2
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


I )l







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
1946:197.

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

American authors.

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.
Macrosipn)um Oestlund."

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-

logical relationships.

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

(hemerocallis).

Type. Aphis cerasi Fabricius.
Distribution. Worldwide.


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
subtriangular ..............................lyhri

Myzus cerasi (Fabricus)
Figure 2

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-

tae.

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

edge.

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

setae.

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


D iE
0N

B F
M 0




G I
H

J


Figure 2







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

plate.

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

Station, Taiwan.
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,
cauda.














R


C


H J K E




L

IVF

I vO P
M N
N


Figure 3







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

known.

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
Palmer 1934:203-4.
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
1929:153.

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

light amber.

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

viviparae.

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,

CFS.
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

hosts.
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
latter species.

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 -
'lb~ot^:"}^]








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

sclerotic patch.

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

plates brown.

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

amber.

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,
HGW, JOP.
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

apteriform alatae.
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.










DA L



B F


EG H
E
M L


N P
N P


Figure 4


S


wf




















































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.










1z
I>
E z ,- o


-I


A








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

Pass.

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
1960: 841.
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

genus zyzus.

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

plate.
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

the segment.

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

growth.

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




70

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

anholocyclism.
Distinguishing characteristics. Color yellow to green.

Second hind tarsal segment with 2 setae. Cornicles swollen
(virginogeniae).

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

States.

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. --
i;aier 1954:.361.
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-

laterally.

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

1954).
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.











C


M



B D

E F O





R Q

K H

L S T


Figure 6






'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
196OT~83, 838.
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




80

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




81

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

edge.

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
many pseudosensoria.

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

November-December.

Meier (1954) reported that the species may also live

parthenogenetically throughout the year inside rolled

peach leaves.

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




85

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
Distribution: Holarctic.
One species occurs in the Eastern United States, Phoro-

don humuli (Schrank).

Phorodon humuli (Schrank)
Figure 7
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
posterior edge.

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.
















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Figure 7


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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.
Distribution. Nearctlc.


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




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