of the ENTOMOLOGICAL SOCIETY WASHINGTONfavret.aphidnet.org/pubs/Favret_et-al_2010.pdf · 2017. 4....

PROCEEDINGSof the

ENTOMOLOGICAL SOCIETY of WASHINGTON

ACTUAL AND INFERRED CHECKLIST OF THE APHIDS (HEMIPTERA:APHIDIDAE) OF THE GREAT SMOKY MOUNTAINS NATIONAL PARK,

WITH ATTENDANT ANT AND HOST PLANT ASSOCIATIONS

COLIN FAVRET, JOSHUA J. DUGGAN, NATHAN J. SANDERS, AND L. RICHARD PHILLIPPE

(CF) AphidNet LLC, 18901 Tributary Ln, Gaithersburg, MD 20879, U.S.A.;Systematic Entomology Laboratory, Agricultural Research Service, United StatesDepartment of Agriculture, Bldg. 005, BARC-W, Beltsville, MD 20705, U.S.A.(e-mail: [email protected]); (JJD) 619 Hawthorne St NE, Grand Rapids,MI 49503, U.S.A.; (NJS) University of Tennessee, Department of Ecology andEvolutionary Biology, 569 Dabney Hall, Knoxville, TN 37996, U.S.A.; (LRP)Illinois Natural History Survey, 1816 S Oak St, Champaign, IL 61820, U.S.A.

PROC. ENTOMOL. SOC. WASH.

112(3), 2010, pp. 381–403

ACTUAL AND INFERRED CHECKLIST OF THE APHIDS (HEMIPTERA:APHIDIDAE) OF THE GREAT SMOKY MOUNTAINS NATIONAL PARK,

WITH ATTENDANT ANT AND HOST PLANT ASSOCIATIONS

COLIN FAVRET, JOSHUA J. DUGGAN, NATHAN J. SANDERS, AND L. RICHARD PHILLIPPE

(CF) AphidNet LLC, 18901 Tributary Ln, Gaithersburg, MD 20879, U.S.A.;Systematic Entomology Laboratory, Agricultural Research Service, United StatesDepartment of Agriculture, Bldg. 005, BARC-W, Beltsville, MD 20705, U.S.A.(e-mail: [email protected]); (JJD) 619 Hawthorne St NE, Grand Rapids,MI 49503, U.S.A.; (NJS) University of Tennessee, Department of Ecology andEvolutionary Biology, 569 Dabney Hall, Knoxville, TN 37996, U.S.A.; (LRP)Illinois Natural History Survey, 1816 S Oak St, Champaign, IL 61820, U.S.A.

Abstract.—During 2003–2006, a general aphid survey was conducted in theGreat Smoky Mountains National Park in the American states of Tennessee andNorth Carolina. The project was undertaken within the context of the All TaxaBiodiversity Inventory and funded by Discover Life in America. In all, 121 aphidspecies were documented. When present, attendant ants were also collected andidentified, as were the aphid host plants. The aphids, their attendant ants, and hostplants are listed here. In addition, a checklist of the plants of the park and a catalogof the aphids of North America were cross-referenced to create a list of aphids notactually found but likely present in the Great Smoky Mountains National Park,increasing the number to 206. Finally, we used Chao1 statistical techniques toestimate the total number of aphid species based on our sampling to date. Theseproduced estimates of 201–214 species.

Key Words: biodiversity, plant lice, survey, Formicidae

DOI: 10.4289/0013-8797.112.3.381

The All Taxa Biodiversity Inventory(ATBI) of the Great Smoky MountainsNational Park (GRSM) was initiated in1997 with the aim of cataloging thediversity of all life within the park(Nichols and Langdon 2007). Insectsand other arthropods have constituted asignificant portion of the ATBI (Sanderset al. 2010) due to their tremendousspecies diversity. To date, however, no

studies have reported on aphids (He-miptera: Aphididae).

Aphids are phloem-sucking insectswith over 4,500 species worldwide(Remaudiere and Remaudiere 1997).Most species are strongly host-specific(Dixon 1987) and many are tended byants: the aphids provide nutrient-richhoneydew in return for protection frompredators and parasitoids (Way 1963).Given the high plant diversity in GRSM(over 1,300 species; Jenkins 2007) andthe otherwise biologically rich nature of* Accepted by Michael W. Gates

the park (Nichols and Langdon 2007), alarge number of aphid species are likelypresent. Historically, few aphid collec-tions have been made in the park, andthese were restricted to a few localities.The study presented here is the firstsystematic survey of the aphids ofGRSM.

MATERIALS AND METHODS

A general survey of the aphids of thepark was initiated in late spring 2003and continued with sampling in thespring and summer of 2004, fall of2005, and spring 2006. A cumulativetotal of 46 days were spent collecting at115 sites. Because host identity isimportant in identifying aphids, sam-pling was mostly done by visualinspection of plants, and the aphidswere removed by hand. In order tocollect grass-feeding aphids, which maybe hidden within the thatch of a field, amodified leaf-blower was used to aspi-rate insects into a net. In some cases,the aphids were then allowed to self-sort onto their preferred host in a cagecontaining multiple potential hosts, butin other cases, no clear host associationwas made. A beating sheet was usedoccasionally to sample aphids on treesand shrubs, especially conifers.

The aphids, and when tended, theirattendant ants, were collected into 95%ethanol. A cutting of the host plant waspressed and dried. Geoposition coordi-nates were recorded with a hand-heldGPS unit. Aphids, ants, and plants werebrought back to the laboratory forprocessing and identification. Approxi-mately 1,100 aphids were mounted tomicroscope slides and 144 ants werepinned.

Aphids were identified by CF using alarge volume of literature sources. Mosthelpful were the host-based keys byBlackman and Eastop (1994 and 2006).Other important resources were thoseby: Boudreaux and Tissot (1962),

Corpuz-Raros and Cook (1974), Eastop(1971), Heie (1979), MacGillivray(1958), Pepper and Tissot (1973),Richards (1968a, b, 1972), Robinson(1985, 1986, 1987), and Quednau(1999, 2003). Ants were identified byNJS using standard resources (e.g.,Creighton 1950, Fisher and Cover2007, MacGown et al. 2007).

A complete set of aphid specimensfrom every collection were deposited inthe insect collection of the IllinoisNatural History Survey, Champaign,IL (INHS). Some duplicates wereselected to form synoptic collectionsdeposited in the insect collection of theGreat Smoky Mountains National Park,Gatlinburg, TN and the Aphid Collec-tion of the National Museum of NaturalHistory, Beltsville, MD. The ants andsome of the plants were deposited at theINHS. The insects were databased andtheir data made available on the Dis-cover Life in America (DLIA) ATBIand INHS insect databases (www.dlia.org, www.inhs.uiuc.edu).

During the course of the project, itbecame clear that many species actuallypresent within GRSM would not befound. In order to form an estimate ofthe number of likely but unconfirmedspecies, we cross-referenced a list ofNorth American aphids (Smith andParron 1978) with a checklist of theplants of GRSM (Great Smoky Moun-tains Natural History Association2004). If the typical host of an aphidspecies, listed by Smith and Parron(1978) as occurring in either NorthCarolina or Tennessee, was known tobe present in GRSM, that species wasadded to the inferred checklist of theaphids of GRSM. Aphid names occur-ring in the literature were updated withreference to Aphid Species File (Favret2009), ant names were checked withAntbase (Agosti and Johnson 2005),and host names were checked with the

382 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

USDA Plants database (USDA, NRCS2009).

We estimated the asymptotic numberof aphid species (the total number hadsampling gone to completion) in GRSMby calculating the Chao1 estimate ofspecies richness (Chao 1984) as SChao1

¼ SObs þ F12/2F2, where SObs is the

number of species observed, F1 is thenumber of singletons, and F2 is thenumber of doubletons. We calculatedthe Chao1 estimate in two ways. First,we used the number of sites as samples.So a singleton would be a species thatoccurred at only one site, and adoubleton a species that occurred attwo sites. Second, we used the numberof days as samples. In this case, asingleton would be a species that wascollected on only one day, and adoubleton would be a species that wascollected on only two days. Finally, weestimated the number of sites or daysthat would be required to fully sampleaphid diversity in GRSM following theprocedure outlined in Chao et al.(2009).

RESULTS AND DISCUSSION

Over the course of four field seasons,121 aphid species were documented inGRSM (Table 1). The full complementof collection data, including geoposi-tion coordinates, locality and date ofcollection, collector and host, are avail-able on the searchable databases(www.dlia.org, www.inhs.illinois.edu).All but two species are represented byspecimens deposited in museums. Theremaining species, Grylloprociphilusimbricator (Fitch) and Prociphilus tes-sellatus (Fitch), were sighted and pho-tographed by Jason Love of the GreatSmoky Mountains Institute at Tremont.These two species are distinct andaccurate identifications were straight-forward. The observed aphid data arecomparable to other surveys of homop-terous insects in GRSM. Fifty-three

species of treehopper and 97 speciesof planthopper were observed andrecorded (Wallace et al. 2003, Gonzonet al. 2006, respectively).

The list of 121 species includes oneCapitophorus, two Drepanaphis, andone Tuberculaphis species that were notidentifiable. Listed in Table 1, theyrepresent species clearly not listed byname, and are possibly new to science.Not listed in Table 1 are specimensfrom one collection of Hyperomyzus,nine of Macrosiphum, one of Monel-liopsis, one of Myzus, one of Nasono-via, five of Rhopalosiphum, and three ofUroleucon. Specimens of these latterseven genera remain unidentified for avariety of reasons: a lack of host plantdata, specimen damage, or an inade-quate resolution of their taxonomy andcryptic morphology. Furthermore, itwas not possible to confirm that theselatter species were not otherwise al-ready recorded in the list.

Beyond the 121 listed and confirmedspecies, an additional 85 are likelypresent in the park, as inferred bycross-referencing aphid and plant lists(Table 1). However, well over half ofthe observed aphid species were not onthe inferred list, suggesting that theestimate of 206 species is conservative.The Chao1 estimate of species richnessbased on the number of sites visitedsuggests that the total number of aphidspecies in GRSM might be 214. Tocomprehensively sample GRSM aphidspecies would require sampling at anadditional 1,123 sites. The Chao1estimate of species richness based onthe number of sample days suggest thatthere might be 201 aphid species inGRSM, and sampling all of the specieswould require an additional 470 days.The three methods of estimating thetotal number of species in GRSMproduced remarkably similar estimates(201, 206, and 214), and both estimatesof required additional sampling are

VOLUME 112, NUMBER 3 383

Table

1.

Actu

al

(in

pare

nth

ese

s)and

infe

rred

list

of

aphid

specie

s,host

pla

nts

,and

att

endant

ants

for

GR

SM

.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

1A

bst

ruso

myzu

s

reti

cula

tus

(Heie

)

Oxali

s

2A

cuti

cauda

soli

dagin

ifoli

ae

(Wil

liam

s)

Soli

dago

canadensi

sL

.

3(1

)A

cyrt

hosi

phon

pis

um

(Harr

is)

Tri

foli

um

pra

tense

L.

4A

mphoro

phora

am

pull

ata

Buckto

nO

nocle

ase

nsi

bil

isL

.

5(2

)A

mphoro

phora

senso

riata

Maso

nR

ubus

Rubus

occid

enta

lis

L.

6(3

)A

noecia

corn

i(F

abri

ciu

s)unknow

nhost

7A

noecia

oenoth

era

eW

ilso

nO

enoth

era

8A

phis

cali

gin

osa

Hott

es

and

Fri

son

Corn

us

9(4

)A

phis

card

uell

aW

als

hH

eli

anth

us

Cry

pto

taenia

canadensi

s(L

.)D

C.,

Taenid

iain

tegerr

ima

(L.)

Dru

de

10

Aphis

cly

desm

ithi

Str

oyan

Sanic

ula

canadensi

sL

.

11

(5)

Aphis

core

opsi

dis

(Thom

as)

Care

xbru

nnnesc

ens

Thunb.,

Eupato

rium

rugosu

mH

outt

.,N

yss

asy

lvati

ca

Mars

h.,

Pla

tanus

occid

enta

lis

L.,

Pre

nanth

es

Cre

mato

gast

er

lineola

ta(S

ay),

Form

ica

subse

ricea

Say,

Para

trechin

alo

ngic

orn

is

(Latr

eil

le),

Para

trechin

aparv

ula

(Mayr)

12

(6)

Aphis

corn

ifoli

ae

Fit

ch

Corn

us

flori

da

L.

Pre

nole

pis

impari

s(S

ay)

13

(7)

Aphis

cra

cciv

ora

Koch

unknow

nhost

14

(8)

Aphis

fabae

Scopoli

Card

uus

nuta

ns

L.,

Cir

sium

alt

issi

mum

(L.)

Hil

lC

rem

ato

gast

er

lineola

taS

ay),

Pre

nole

pis

impari

s(S

ay)

15

(9)

Aphis

goss

ypii

Glo

ver

Pycnanth

em

um

Cre

mato

gast

er

lineola

ta(S

ay)


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

16

(10)

Aphis

illi

nois

ensi

sS

him

er

Aure

ola

ria

laevig

ata

(Raf.

)R

af.

,V

itis

aest

ivali

sM

ichx.

Myrm

ica

puncti

ventr

isR

oger

17

(11)

Aphis

lugenti

sW

illi

am

sunknow

nhost

Myrm

ica

puncti

ventr

isR

oger,

Tapin

om

ase

ssil

e(S

ay)

18

(12)

Aphis

mid

dle

tonii

Thom

as

unknow

nhost

19

Aphis

nast

urt

iiK

alt

enbach

Capse

lla

burs

a-p

ast

ori

s(L

.)M

edik

.

20

Aphis

oest

lundi

Gil

lett

eO

enoth

era

bie

nnis

L.

21

(13)

Aphis

pulc

hell

aH

ott

es

and

Fri

son

Euphorb

iaunknow

nhost

22

Aphis

rubic

ola

Oest

lund

Rubus

idaeus

L.

stri

gosu

s

(Mic

hx.)

Focke

23

(14)

Aphis

rubif

oli

i

(Thom

as)

Rubus

occid

enta

lis

L.

24

(15)

Aphis

rum

icis

Lin

naeus

Rum

ex

Rum

ex

obtu

sifo

lius

L.

25

(16)

Aphis

spir

aecola

Patc

hC

rata

egus

macro

sperm

aA

she,

Pru

nus

sero

tinus

Ehrh

.,P

seudognaphali

um

obtu

sifo

lium

(L.)

Hil

liard

&B

.L.

Burt

t,P

yru

s,R

anunculu

sbulb

osu

sL

.,T

ilia

am

eri

cana

L.

var.

hete

rophyll

a(V

ent.

)L

ouden

26

Aphis

vib

urn

iphil

aP

atc

hV

iburn

um

27

(17)

Aula

cort

hum

sola

ni

(Kalt

enbach)

Aconit

um

uncin

atu

mL

.,A

mbro

sia

trif

ida

L.,

Am

ela

nchie

rla

evis

Wie

gand,

Ara

lia

racem

osa

L.,

Ast

er

punic

eus

L.,

Conopholi

sam

eri

cana

(L.)

Wall

r.,

Core

opsi

sm

ajo

rW

alt

er,

Cry

pto

taenia

canadensi

s(L

.)D

C.,

Eri

gero

nphil

adelp

hic

us

L.,

Eupato

rium

,

Form

ica

subse

ricea

Say


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

Hie

raciu

mcaesp

itosu

mD

um

ort

.,L

iliu

msu

perb

um

L.,

Osm

orh

iza

cla

yto

nii

(Mic

hx)

C.B

.C

lark

e,

Oxydendru

marb

ore

um

(L.)

DC

.,

Pedic

ula

ris

canadensi

sL

.,P

lata

nus

occid

enta

lis

L.,

Pru

nus,

Pyru

lari

a

pubera

Mic

hx.,

Querc

us

rubra

L.,

Ranunculu

sbulb

osu

sL

.,R

hododendro

n

maxim

um

L.,

Rubus

all

eghenie

nsi

sP

ort

er,

Rubus

canadensi

sL

.,R

ugeli

a

nudic

auli

sS

hutt

lw.

ex

Chapm

.,R

um

ex

cri

spus

L.,

Rum

ex

obtu

sifo

lius

L.,

Salv

ialy

rata

L.,

Sam

bucus

racem

osa

L.,

Saxif

raga

care

yana

A.

Gra

y,

Tia

rell

acord

ifoli

aL

.,T

ilia

am

eri

cana

L.,

Vera

trum

vir

ide

Ait

on

28

Bra

chycaudus

card

ui

(Lin

naeus)

Card

uus

29

(18)

Bra

chycaudus

heli

chry

si

(Kalt

enbach)

Ere

chti

tes

hie

racii

foli

a(L

.)R

af.

ex

DC

.,E

rigero

n,

Eupato

rium

30

Bre

vic

ory

ne

bra

ssic

ae

(Lin

naeus)

Bra

ssic

a

31

Cachry

phora

canadensi

sH

ille

Ris

Lam

bers

Soli

dago

canadensi

sL

.

32

(19)

Cala

phis

betu

laecole

ns

(Fit

ch)

Am

ela

nchie

rarb

ore

a(M

ichx.

f.)

Fern

ald

33

Cala

phis

betu

lell

a

Wals

h

Betu

lanig

raL

.


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

34

(20)

Capit

ophoru

sela

eagni

(Del

Guerc

io)

Ela

eagnus

Cir

sium

alt

issi

mum

(L.)

Hil

l,C

irsi

um

dis

colo

r(M

uhl

ex

Wil

d.)

Spre

ng.

Cre

mato

gast

er

lineola

ta(S

ay),

Pre

nole

pis

impari

s(S

ay)

35

Capit

ophoru

sjo

pepperi

Corp

uz-R

aro

sand

Cook

Am

bro

sia

art

em

isii

foli

aL

.

36

(21)

Capit

ophoru

ssp

.unknow

nhost

37

(22)

Caro

linaia

cari

cis

Wil

son

Care

xunknow

nhost

38

(23)

Caro

linaia

rhois

(Monell

)A

mela

nchie

rarb

ore

a(M

ichx.

f.)

Fern

ald

,E

lym

us

hyst

rix

L.,

Fest

uca

pra

tensi

s

Huds.

,P

hle

um

pra

tense

L.,

Sphenopholi

snit

ida

(Bie

hle

r)S

cri

bn.

39

Cata

merg

us

kic

kapoo

(Hott

es

and

Fri

son)

Poly

gonatu

m

40

(24)

Cavari

ell

aaeg

opodii

(Scopoli

)A

rali

ara

cem

osa

L.,

Osm

orh

izae

cla

yto

nii

(Mic

hx.)

C.B

.C

lark

e,

Verb

esi

na

occid

enta

lis

(L.)

Walt

er

41

Cavari

ell

ahenders

oni

(Know

lton

and

Sm

ith)

Sali

x

42

Chait

ophoru

snig

rae

Oest

lund

Sali

xnig

raM

ars

h.

43

Chait

ophoru

spopuli

foli

i(E

ssig

)P

opulu

s

44

Chait

ophoru

svim

inali

sM

onell

Sali

x

45

(25)

Chait

ophoru

svim

inic

ola

Hil

leR

is

Lam

bers

Sali

xSali

xse

ricea

Mars

h.

Cre

mato

gast

er

lineola

ta(S

ay)


Table

1.

Conti

nued.

Nu

mb

er

(obse

rved)

Aphid

specie

s

Ty

pic

al

host

of

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

46

(26)

Cin

ara

atl

anti

ca

(Wil

son)

Pin

us

Pin

us

rigid

aM

ill.

,P

inus

stro

bus

L.,

Pin

us

vir

gin

iana

Mil

l.

Form

ica

subse

ricea

Say,

Lasi

us

ali

enus

(Fo

rste

r),

Tapin

om

ase

ssil

e(S

ay)

47

(27)

Cin

ara

bra

ggii

(Gil

lett

e)

Pic

ea

rubens

Sarg

.

48

(28)

Cin

ara

confi

nis

(Koch)

Abie

sfr

ase

ri(P

urs

h)

Poir

.,P

icea

rubens

Sarg

.49

(29)

Cin

ara

cupre

ssi

(Buckto

n)

Junip

eru

svir

gin

iana

L.

50

(30)

Cin

ara

engelm

annie

nsi

s

(Gil

lett

eand

Palm

er)

Pic

ea

rubens

Sarg

.

51

(31)

Cin

ara

gra

cil

is

(Wil

son)

Pin

us

vir

gin

iana

Mil

l.

52

(32)

Cin

ara

mela

ina

Boudre

aux

Pin

us

rigid

aM

ill.

53

Cin

ara

new

ell

iT

isso

tP

inus

palu

stri

sM

ill.

54

(33)

Cin

ara

perg

andei

(Wil

son)

Pin

us

stro

bus

L.,

Pin

us

vir

gin

iana

Mil

l.

55

(34)

Cin

ara

pin

ivora

(Wil

son)

Pin

us

Pin

us

rigid

aM

ill.

,P

inus

vir

gin

iana

Mil

l.F

orm

ica

subse

ricea

Say

56

(35)

Cin

ara

stro

bi

(Fit

ch)

Pin

us

stro

bus

L.

Cam

ponotu

spennsy

lvanic

us

(DeG

eer)

57

Cin

ara

taedae

(Tis

sot)

Pin

us

taeda

L.

58

(36)

Cin

ara

vandykei

(Wil

son)

Abie

sfr

ase

ri(P

urs

h)

Poir

.,P

icea

rubens

Sarg

.59

(37)

Cin

ara

wats

oni

(Tis

sot)

Pin

us

taeda

L.

Pin

us

rigid

aM

ill.

,P

inus

vir

gin

iana

Mil

l.

60

Dip

hyll

aphis

mic

rotr

em

aQ

uednau

Querc

us

rubra

L.

61

(38)

Dre

panaphis

aceri

foli

ae

(Thom

as)

Acer

rubru

mL

.


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

62

Dre

panaphis

caro

linensi

sS

mit

hA

cer

saccharu

mM

ars

h.

63

(39)

Dre

panaphis

choanotr

icha

Sm

ith

and

Dil

lery

Acer

saccharu

mM

ars

h.

Acer

saccharu

mM

ars

h.

64

(40)

Dre

panaphis

know

ltoni

Sm

ith

and

Dil

lery

Acer

rubru

mL

.

65

(41)

Dre

panaphis

nig

ricans

Sm

ith

Acer

rubru

mL

.A

cer

rubru

mL

.C

am

ponotu

ssu

bbarb

atu

sE

mery

,

Myrm

ica

puncti

ventr

isR

oger

66

(42)

Dre

panaphis

parv

a

Sm

ith

Acer

rubru

mL

.A

mela

nchie

rarb

ore

a(M

ichx.

f.)

Fern

ald

67

Dre

panaphis

sabri

nae

Mil

ler

Acer

sacchari

num

L.

68

Dre

panaphis

sim

pso

ni

Sm

ith

Acer

saccharu

mM

ars

h.

69

(43)

Dre

panaphis

sp.

1unknow

nhost

70

(44)

Dre

panaphis

sp.

2unknow

nhost

71

(45)

Eri

caphis

wakib

ae

(Hott

es)

Pedic

ula

ris

canadensi

sL

.R

ubus

all

eghenie

nsi

sP

ort

er

72

(46)

Eri

oso

ma

am

eri

canum

(Ril

ey)

Ulm

us

am

eri

cana

L.

unknow

nhost

73

Eri

oso

ma

cra

taeg

i

(Oest

lund)

Cra

taeg

us

puncta

taJa

cq.

74

(47)

Ess

igell

apin

iW

ilso

nP

icea

rubens

Sarg

.,P

inus

rigid

aM

ill.

,

Pin

us

vir

gin

iana

Mil

l.75

(48)

Eucall

ipte

rus

tili

ae

(Lin

naeus)

Til

iaam

eri

cana

L.

76

(49)

Eucera

phis

mucid

a

(Fit

ch)

Am

ela

nchie

rarb

ore

a(M

ichx.

f.)

Fern

ald

,

Betu

lale

nta

L.


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

77

Eucera

phis

puncti

pennis

(Zett

ers

tedt)

Betu

la

78

(50)

Gry

llopro

cip

hil

us

imbri

cato

r(F

itch)*

Fagus

Fagus

gra

ndif

oli

aE

hrh

.

79

Ham

am

eli

stes

spin

osu

sS

him

er

Ham

am

eli

svir

gin

iana

L.

80

Hyalo

myzu

scoll

inso

nia

e(P

epper)

Coll

inso

nia

canadensi

sL

.

81

(51)

Hyalo

pte

roid

es

hum

ilis

(Walk

er)

unknow

nhost

82

Hypero

myzu

sin

flatu

s(R

ichard

s)Soli

dago

83

(52)

Hypero

myzu

snabali

(Oest

lund)

Pre

nanth

es

alt

issi

ma

L.

Form

ica

subse

ricea

Say

84

(53)

Hypero

myzu

spic

ridis

(Born

er)

unknow

nhost

85

(54)

Hyst

ero

neura

seta

riae

(Thom

as)

Tri

dens

flavus

(L.)

Hit

chc.

86

(55)

Illi

noia

gold

mary

ae

(Know

lton)

Eri

gero

nphil

adelp

hic

us

L.

87

(56)

Illi

noia

liri

odendri

(Monell

)

Lir

iodendro

ntu

lipif

era

L.

Lir

iodendro

ntu

lipif

era

L.,

Magnoli

a

trip

eta

la(L

.)L

.88

Illi

noia

pepperi

(MacG

illi

vra

y)

Vaccin

ium

89

(57)

Illi

noia

rhokala

za

(Tis

sot

and

Pepper)

Rhododendro

nm

axim

um

L.

Rhododendro

nm

axim

um

L.

90

(58)

Illi

noia

spir

aecola

(Patc

h)

Magnoli

afr

ase

riW

alt

er,

Poly

stic

hum

acro

stic

hoid

es

(Mic

hx.)

Schott

.,P

renanth

es,

Til

iaam

eri

cana

L.


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

91

(59)

Izip

hya

flabell

a(S

anborn

)C

are

xbro

moid

es

Schkuhr

ex

Wil

d.,

Care

xpensy

lvanic

aL

am

,

92

(60)

Macro

siphonie

lla

mil

lefo

lii

(De

Geer)

unknow

nhost

93

Macro

siphonie

lla

pennsy

lvanic

a

(Pepper)

Achil

lea

mil

lefo

lium

L.

94

(61)

Macro

siphonie

lla

tapusk

ae

(Hott

es

and

Fri

son)

Achil

lea

mil

lefo

lium

L.

unknow

nhost

95

Macro

siphum

carp

inic

ole

ns

Patc

h

Carp

inus

caro

linia

na

Walt

er

96

Macro

siphum

cory

li

Davis

Cory

lus

am

eri

cana

Walt

er

97

Macro

siphum

gera

nii

(Oest

lund)

Gera

niu

mm

acula

tum

L.

98

Macro

siphum

pse

udocory

liP

atc

h

Ost

rya

vir

gin

iana

(Mil

l.)

K.

Koch

99

(62)

Macro

siphum

rosa

e

(Lin

naeus)

Rosa

caro

lina

L.

100

Mic

ropars

us

desm

odio

rum

Sm

ith

and

Tuata

y

Desm

odiu

mpanic

ula

tum

(L.)

DC

.

101

(63)

Mic

ropars

us

singula

ris

(Hott

es

and

Fri

son)

unknow

nhost

102

Mic

ropars

us

tephro

siae

(Sm

ith)

Tephro

sia

vir

gin

iana

(L.)

Pers

.

103

(64)

Min

daru

spin

icola

(Thom

as)

Abie

sfr

ase

ri(P

urs

h)

Poir

.


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

host

of

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

104

Monell

iahis

pid

a

Quednau

Cary

agla

bra

(Mil

l.)

Sw

eet.

105

Monell

iam

icro

seto

sa

Ric

hard

s

Cary

a

106

(65)

Monell

iopsi

sbis

sell

i

Quednau

unknow

nhost

107

(66)

Monell

iopsi

scary

ae

(Monell

)

Cary

agla

bra

(Mil

l.)

Sw

eet

108

(67)

Monell

iopsi

s

nig

ropuncta

ta(G

ranovsk

y)

Jugla

ns

Cary

agla

bra

(Mil

l.)

Sw

eet,

Cary

a

pall

ida

(Ash

e)

Engl.

&G

raebn.

109

(68)

Monell

iopsi

specanis

Bis

sell

Jugla

ns

cin

ere

aL

.

110

(69)

Myzo

call

iscast

aneae

(Fit

ch)

Cast

anea

denta

ta(M

ars

h.)

Bork

h.

111

Myzo

call

iscast

aneoid

es

(Baker)

Cast

anea

112

(70)

Myzo

call

isdis

colo

r(M

onell

)Q

uerc

us

alb

aL

.

113

Myzo

call

isfr

isoni

Boudre

aux

and

Tis

sot

Querc

us

114

(71)

Myzo

call

isgra

novsk

yi

Boudre

aux

and

Tis

sot

Kalm

iala

tifo

lia

L.

115

(72)

Myzo

call

islo

ngir

ost

ris

Ric

hard

sQ

uerc

us

Querc

us

116

Myzo

call

islo

ngiu

nguis

Boudre

aux

and

Tis

sot

Querc

us

117

Myzo

call

ism

ela

nocera

Boudre

aux

and

Tis

sot

Querc

us

118

Myzo

call

ism

ult

iseti

s

Boudre

aux

and

Tis

sot

Querc

us


Table

1.

Conti

nued.

Nu

mb

er

(obse

rved)

Aphid

specie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

119

(73)

Myzo

call

ispuncta

tus

(Monell

)Q

uerc

us

monta

na

Wil

d.

120

Myzo

call

issp

inosu

sB

oudre

aux

and

Tis

sot

Querc

us

121

(74)

Myzo

call

istu

berc

ula

tus

Ric

hard

sunknow

nhost

122

(75)

Myzo

call

isw

als

hii

(Monell

)Q

uerc

us

rubra

L.

Querc

us

rubra

L.

123

Myzu

scera

si(F

abri

ciu

s)P

runus

cera

sus

L.

124

Myzu

shem

ero

call

isT

akahash

iH

em

ero

call

isfu

lva

(L.)

L.

125

Myzu

sorn

atu

sL

ain

gV

iola

126

(76)

Naso

novia

caro

linensi

s

Heie

Saxif

raga

care

yana

A.

Gra

y

127

(77)

Naso

novia

heuchera

e

(Thom

as)

Heuchera

longif

lora

Rydb.

128

(78)

Naso

novia

purp

ura

scens

(Oesl

tund)

Thali

ctr

um

cori

aceum

(Bri

tton)

Sm

all

,T

hali

ctr

um

pubesc

ens

Purs

h

129

(79)

Naso

novia

ribis

nig

ri

(Mosl

ey)

Hie

raciu

mcaesp

itosu

mD

um

ort

.

130

(80)

Naso

novia

tiare

llae

Heie

Tia

rell

acord

ifoli

aL

.

131

(81)

Nearc

taphis

bakeri

(Cow

en)

Tri

foli

um

pra

tense

L.

Hie

raciu

mvenosu

mL

.,T

rifo

lium

pra

tense

L.

Pre

nole

pis

impari

s(S

ay)

132

Nearc

taphis

cly

desm

ithi

Hil

leR

isL

am

bers

Cra

taeg

us

133

Nearc

taphis

cra

taeg

ifoli

ae

(Fit

ch)

Cra

taeg

us

puncta

taJa

cq.


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

134

(82)

Neom

yzu

scir

cum

flexum

(Buckto

n)

Cacali

aatr

ipli

cif

oli

aL

.F

orm

ica

subse

ricea

Say

135

Papula

phis

sleesm

ani

(Pepper)

Adia

ntu

mpedatu

mL

.

136

Pem

phig

us

burs

ari

us

(Lin

naeus)

Populu

snig

raL

.

137

Pem

phig

us

ephem

era

tus

Hott

es

and

Fri

son

Betu

lanig

raL

.

138

Pem

phig

us

monophagus

Maxso

n

Populu

sbals

am

ifera

L.

139

Pem

phig

us

nort

onii

Maxso

n

Populu

sdelt

oid

es

Bart

ram

ex

Mars

h.

140

Pem

phig

us

tart

are

us

Hott

es

and

Fri

son

Populu

sdelt

oid

es

Bart

ram

ex

Mars

h.

141

(83)

Ple

otr

ichophoru

s

ast

eri

foli

ae

(Str

om

)

unknow

nhost

142

Ple

otr

ichophoru

s

gla

ndulo

sus

(Kalt

enbach)

Art

em

isia

vulg

ari

sL

.

143

Ple

otr

ichophoru

shott

esi

Hil

leR

is

Lam

bers

Achil

lea

144

Ple

otr

ichophoru

s

pato

nkus

(Hott

es

and

Fri

son)

Achil

lea

mil

lefo

lium

L.

145

Ple

otr

ichophoru

s

pse

udopato

nkus

Corp

uz-R

aro

sand

Cook

Achil

lea

mil

lefo

lium

L

146

Pro

cip

hil

us

fraxin

ifoli

i

(Ril

ey)

Fra

xin

us

am

eri

cana

L.


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

host

of

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

147

Pro

cip

hil

us

pic

eaeru

bensi

s(S

mit

h)

Pic

ea

rubens

Sarg

.

148

(84)

Pro

cip

hil

us

tess

all

atu

s(F

itch)*

Aln

us

serr

ula

ta(A

iton)

Wil

d.

149

Pte

rocom

ma

smit

hia

e

(Monell

)

Sali

xalb

aL

.

150

(85)

Rhodobiu

mporo

sum

(Sanders

on)

unknow

nhost

151

(86)

Rhopalo

myzu

s

lonic

era

e(S

iebold

)

unknow

nhost

152

Rhopalo

myzu

spoae

(Gil

lett

e)

Poa

pra

tensi

sL

.

153

(87)

Rhopalo

siphum

cera

sifo

liae

(Fit

ch)

Care

xpla

nta

gin

ea

Lam

.L

asi

us

ali

enus

(Fo

rste

r)

154

(88)

Rhopalo

siphum

maid

is

Fit

ch

Pla

tanus

occid

enta

lis

L.

155

(89)

Rhopalo

siphum

oxyacanth

ae

(Schra

nk)

unknow

nhost

156

(90)

Rhopalo

siphum

padi

(Lin

naeus)

Care

xbru

nnesc

ens

(Pers

.)P

oir

.,C

are

xlu

rida

Wahle

nb.,

Holc

us

lanatu

sL

.

157

(91)

Rhopalo

siphum

rufi

abdom

inali

s

(Sasa

ki)

unknow

nhost

158

(92)

Schiz

aphis

gra

min

um

(Rondani)

Care

xlu

rida

Wahle

nb.

159

(93)

Schiz

ola

chnus

flocculo

sus

(Wil

liam

s)

Pin

us

rigid

aM

ill.

,P

inus

vir

gin

iana

Mil

l.


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

160

Schiz

ola

chnus

parv

us

(Wil

son)

Pin

us

vir

gin

iana

Mil

l.

161

Shenahw

eum

min

utu

m

(Davis

)

Acer

saccharu

mM

ars

h.

162

(94)

Sip

ha

flava

(Forb

es)

Care

x

163

(95)

Sit

obio

navenae

(Fabri

ciu

s)A

nth

oxanth

um

odora

tum

L.,

Care

xpra

sina

Wahle

nb.,

Dacty

lis

glo

mera

ta

L.,

Fest

uca

pra

tensi

sH

uds.

,H

olc

us

lanatu

sL

.,Ju

ncus,

Phle

um

pra

tense

L.,

Pla

nta

go

,P

oa

als

odes

A.

Gra

y,

Salv

ialy

rata

L.

164

Sit

obio

npte

rinig

rum

(Ric

hard

s)

165

Subsa

ltusa

phis

vir

gin

ica

(Baker)

Care

x

166

Takecall

isaru

ndin

ari

ae

(Ess

ig)

Aru

ndo,

Aru

ndin

ari

a

167

(96)

Takecall

ista

iwanus

(Takahash

i)A

undin

ari

agig

ante

a(W

alt

er)

Muhl.

168

(97)

Tetr

aneura

ulm

i(L

innaeus)

unknow

nhost

169

Thecabiu

saff

inis

(Kalt

enbach)

Populu

snig

raL

.

170

(98)

Thecabiu

sgra

vic

orn

is(P

atc

h)

unknow

nhost

171

(99)

Theri

oaphis

trif

oli

i(M

onell

)unknow

nhost

172

Til

iphagus

lycoposu

gus

Sm

ith

Til

iaam

eri

cana

L.

173

Tin

ocall

isulm

ifoli

i

(Monell

)

Ulm

us

am

eri

cana

L.


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

174 (1

00)

Tuberc

ula

tus

sp.

unknow

nhost

175

Tubero

lachnus

sali

gnus

(Gm

eli

n)

Sali

x

176

Uro

leucon

anom

ala

e

(Hott

es

and

Fri

son)

Ast

er

novae-a

ngli

ae

L.

177

Uro

leucon

bra

chychaetu

m(O

live)

Kri

gia

monta

na

(Mic

hx.)

Nutt

.

178

Uro

leucon

bra

dbury

i(O

live)

Ast

er

179 (1

01)

Uro

leucon

chry

santh

em

i

(Oest

lund)

Leucanth

em

um

vulg

are

Lam

.

180

Uro

leucon

chry

sopsi

dic

ola

(Oli

ve)

Chry

sopsi

sm

ari

ana

(L.)

Ell

iot

181

Uro

leucon

cie

fi(O

live)

Eupato

rium

capil

lifo

lium

(Lam

.)S

mall

182 (1

02)

Uro

leucon

cre

pusi

siphon

(Oli

ve)

Ast

er

Ast

er

cord

ifoli

us

L.,

Ast

er

pate

ns

Ait

on,

Sym

phyotr

ichum

dum

osu

m(L

.)G

.L.

Neso

m,

Sym

phyotr

ichum

late

rifl

oru

m(L

.)A

.L

ove

&D

.L

ove

183 (1

03)

Uro

leucon

eupato

ricole

ns

(Patc

h)

Eupato

rium

,H

iera

ciu

mvenosu

mL

.,Senecio

aure

a(L

.)A

.L

ove

&D

.L

ove,

Verb

esi

na

alt

ern

ifoli

a(L

.)B

ritt

on

ex

Kearn

ey

184 (1

04)

Uro

leucon

eupato

rifo

liae

(Tis

sot)

Agera

tina

alt

issi

ma

(L.)

Kin

g&

H.

Rob.


Table

1.

Conti

nued.

Nu

mb

er

(obse

rved)

Aphid

specie

s

Ty

pic

al

host

of

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

185 (1

05)

Uro

leucon

gra

vic

orn

e(P

atc

h)

Soli

dago

Eri

gero

nannuus

(L.)

Pers

.,E

rigero

npulc

hell

us

Mic

hx.,

Lysi

machia

quadri

foli

aL

.186 (1

06)

Uro

leucon

heli

anth

icola

(Oli

ve)

Heli

anth

us

mic

rocephalu

s

Torr

.&

A.

Gra

y

Heli

anth

us

atr

oru

bens

187 (1

07)

Uro

leucon

hie

racic

ola

(Hil

leR

isL

am

bers

)

unknow

nhost

188

Uro

leucon

illi

ni

(Hott

es

and

Fri

son)

Heli

anth

us

189 (1

08)

Uro

leucon

impati

ensi

cole

ns

(Patc

h)

unknow

nhost

190 (1

09)

Uro

leucon

leonard

i(O

live)

Rudbeckia

lacin

iata

L.

191 (1

10)

Uro

leucon

lute

olu

m(W

illi

am

s)Soli

dago

gig

ante

aA

iton

192

Uro

leucon

nig

roti

biu

m(O

live)

Soli

dago

193 (1

11)

Uro

leucon

nig

rotu

berc

ula

tum

(Oli

ve)

Soli

dago

Soli

dago

canadensi

sL

.

194 (1

12)

Uro

leucon

parv

otu

berc

ula

tum

(Oli

ve)

Heli

anth

us

atr

oru

bens

L.

Ere

chti

tes

hie

racif

oli

a(L

.)R

af.

ex

DC

.,N

yss

asy

lvati

ca

Mars

h.,

Osm

orh

iza

cla

yto

nii

(Mic

hx.)

C.B

.C

lark

e,

Verb

esi

na

occid

enta

lis

(L.)

Walt

er

195

Uro

leucon

paucose

nso

riatu

m

(Hil

leR

isL

am

bers

)

Ast

er

196

Uro

leucon

pepperi

(Oli

ve)

Cir

sium


Table

1.

Conti

nued.

Nu

mb

er

(ob

serv

ed

)A

ph

idsp

ecie

s

Ty

pic

al

ho

stof

aphid

record

ed

fro

mN

Co

rT

N(S

mit

hand

Parr

on

19

76

)fo

un

din

park

(Gre

at

Sm

oky

Mo

un

tain

sN

atu

ral

His

tory

Ass

ocia

tion

2004)

Obse

rved

host

Att

endant

ants

197 (1

13)

Uro

leucon

pie

loui

(Ric

hard

s)Soli

dago

canadensi

sL

.F

orm

ica

subse

ricea

Say

198 (1

14)

Uro

leucon

pse

udam

bro

siae

(Oli

ve)

Lactu

ca

Ere

chti

tes

hie

racif

oli

a(L

.)R

af.

ex

DC

.,L

actu

ca

canadensi

sL

.,P

ackera

aure

a

(L.)

A.

Love

&D

.L

ove,

Verb

esi

na

occid

enta

lis

(L.)

Walt

er

199 (1

15)

Uro

leucon

reynold

ense

(Oli

ve)

Core

opsi

sm

ajo

rW

alt

er

Core

opsi

dis

majo

rW

alt

er

200 (1

16)

Uro

leucon

rudbeckia

e(F

itch)

Rudbeckia

,Soli

dago

Rudbeckia

lacin

iata

L.

201 (1

17)

Uro

leucon

rura

le(H

ott

es

and

Fri

son)

Verb

esi

na

alt

ern

ifoli

a(L

.)B

ritt

on

ex

Kearn

ey

202 (1

18)

Uro

leucon

sonchell

um

(Monell

)L

actu

ca

flori

dana

(L.)

Gaert

n.

Pre

nole

pis

impari

s(S

ay)

203 (1

19)

Uro

leucon

tard

ae

(Hott

es

and

Fri

son)

Hele

niu

mautu

mnale

L.

unknow

nhost

204 (1

20)

Uro

leucon

tuata

iae

(Oli

ve)

Am

bro

sia

art

em

isii

foli

aL

.A

mbro

sia

art

em

isii

foli

aL

.

205 (1

21)

Uro

leucon

verb

esi

nae

(Boudre

aux)

Verb

esi

na

occid

enta

lis

(L.)

Walt

er

206

Vesi

cula

phis

cari

cis

(Full

aw

ay)

Care

x

*S

igh

tre

co

rdb

yJa

son

Lov

e.


roughly ten times the actual sampling(115 actual versus 1,123 estimatedsites; 46 actual versus 470 estimateddays).

A side-by-side comparison of theinferred and observed checklists sug-gests future study of particular taxa.Chaitophorus, Myzocallis, Pemphigus,and Pleotrichophorus are four genera ofrelatively host-specific aphid species.Only a small number of species of thesegenera were actually collected, but theycontain a large number of specieswhose hosts are known to occur in thepark. Future targeted collecting ofspecies of these four genera shouldfocus on all the GRSM species of Salix,Populus, Quercus, and Achillea (wil-lows, poplars, oaks, and yarrow, respec-tively). The genus Uroleucon needs a

significant amount of revisionary taxo-nomic work. The inferred and observedlists each contain 20 species, and whencombined indicate 30 species at GRSM,making it the most speciose aphidgenus in the park. This diversity maysimply be a result of historical relianceon host plant data to describe species.Host based keys (e.g., Blackman andEastop 2006) permitted the identifica-tion of 20 observed species, but contin-ued preliminary study suggests that anumber of these are synonyms. Incontrast, the observed list for speciesof Cinara is over twice as long as theinferred list, reflecting the first author’staxonomic expertise.

Aphid species exhibit the full rangeof ant tending, with some speciesobligately tended, some never tended,

Fig. 1. Diagram of trilateral associations between aphids, their attendant ants, and their hostplants.


and some facultatively tended (Stadlerand Dixon 2005). Ten species of antswere found tending 20 species of aphids(Figure 1). Unfortunately, ants were notalways systematically collected duringthe project, so the recorded associationsare an undercount. Aphis coreopsidis(Thomas) had the greatest variety ofants tending it, despite it being recordedwith ants on only one host plant species.Of the four ant species found tending A.coreopsidis, two of them were exclusiveto it, not being found tending any otheraphid species. In contrast, Formicasubsericea Say was found tending sevenaphid species, Prenolepis imparis (Say)tended six, and Crematogaster lineolata(Say) tended five.

Several aphid-host associations bearmentioning because of their uniqueness.Aphis illinoisensis Shimer was foundfeeding on Aureolaria laevigata (Raf.)Raf., a member of the Scrophulariaceae(figwort family). This host, a perennialherb, is a new host record for A.illinoisensis and is a far departure fromits typical use of Vitis spp. as host, aperennial vine in the Vitaceae (grapefamily). Aulacorthum solani (Kalten-bach), a polyphagous aphid, was foundon 32 different host plants representing16 families. Of particular note was acolony of A. solani feeding on Con-opholis americana (L.) Wallr., a para-sitic perennial herb found on the rootsof several kinds of oaks and a memberof the Orobanchaceae (broomrape fam-ily).

Finally, five aphid species found inGRSM were previously known onlyfrom western North America or Canada(Blackman and Eastop 1994, 2006):Cinara engelmanniensis (Gillette andPalmer), C. vandykei (Wilson), Schizo-lachnus flocculosus (Williams), Sito-bion pterinigrum (Richards), and Uro-leucon hieracicola (Hille Ris Lambers).These Smoky Mountain records mayrepresent relict populations, or perhaps

sibling species of their western andnorthern counterparts.

Taken together, our work on aphiddiversity in GRSM, and the aphidassociations with both their host plantsand their attendant ants, provides animportant baseline for studying changesin the distribution and abundance ofthese important species as a result ofhabitat modification and climaticchange. Importantly, we have alsosuggested several important lines offuture research.

ACKNOWLEDGMENTS

Barb Fuller helped process some ofthe aphid material in the laboratory andextracted typical host data for NorthCarolina and Tennessee aphids from theList of North American Aphids. JasonLove spotted several aphid species notfound by the authors. Doris Lagosperformed determinations on someaphid material. Ed DeWalt, Ani Katch-ova, and Sue Favret assisted withcollecting efforts. Jeanie Hilten, JohnMorse, and Chuck Parker helped withATBI logistics. Housing in the park wasprovided by Discover Life in America.Favret, Sanders, and Phillippe wereeach separately funded by DLIA. Sand-ers was further supported by a grantfrom the Department of Energy.

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