University of Nebraska - Lincoln

DigitalCommons@University of Nebraska - Lincoln Faculty Publications: Department of Entomology

Entomology, Department of

1-1-1994

Patterns among Host Plants of Potato Leafhopper, Empoasca fabae (Homoptera: Cicadellidae) William O. Lamp Dartmouth College

Gary R. Nielsen Dartmouth College

Stephen D. Danielson University of Nebraska - Lincoln, [email protected]

Follow this and additional works at: http://digitalcommons.unl.edu/entomologyfacpub Part of the Entomology Commons Lamp, William O.; Nielsen, Gary R.; and Danielson, Stephen D., "Patterns among Host Plants of Potato Leafhopper, Empoasca fabae (Homoptera: Cicadellidae)" (1994). Faculty Publications: Department of Entomology. Paper 283. http://digitalcommons.unl.edu/entomologyfacpub/283

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JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 67(4),

1994,

pp.

354-368

Patterns

among Host Plants of Potato Leafhopper, Empoasca fabae (Homoptera: Cicadellidae)

and Stephen D. Danielson2 O. Lamp, Gary R. Nielsen,1 of Entomology, of Maryland, Department University 20742 College Park, Maryland

William

abstract:

characteristics of potato Ecological leafhopper, Empoasca (Harris), fabae that non-economic in the landscape be plant may polyphagy, suggest species to its population a host The objectives of this study were to compile important ecology. taxonomic and ecological the list, and to determine host list, to ascertain patterns within including

in non-crop Fabaceae

in 100 genera and The host plant list included 220 species habitats. 47% of the genera and 62% of the species. Yet, the list represented includes a diversity of taxonomic the class Magnoliopsida, groups within representing highly on such and morphological based types. Ecological classifications, chemistry divergent as habitat, characteristics diverse. the di form, and origin, were Thus, growth similarly utilization

26 families.

of plant species suitable for reproduction that non-crop habitats may be a suggests source of potato after spring migration into northern significant leafhopper populations states. Also, the ability of leafhopper adults to utilize additional species (e.g., grasses, pines) as refugia provides a secondary our limited role to non-crop habitats. data suggest Yet, that utilization of non-crop for reproduction is restricted to a relatively habitats few nat

versity

uralized habitat

hosts level,

in non-crop

(e.g., deciduous as well as abiotic

at a landscape or behavior, trees). Host finding operating and biotic factors within limit host utilization habitats, may

habitats.

The potato leafhopper, Empoasca Cicadellidae), fabae (Harris) (Homoptera: as an economic has long been recognized crops including pest of many alfalfa, in the midwestern and eastern United States and potato, bean, and soybean on dynamics Canada has concentrated of this pest in (DeLong, 1938). Research characteristics of the species suggest that non crop systems, yet three ecological economic in its population plant species outside of crop systems may be important on a wide it is highly polyphagous, range of plant First, reproducing ecology. are highly and and families adults Wheeler, 1943, 1949). Second, species (Poos both and short (interhabitat) distances mobile, long (interregional) traversing and Flanders and Poston 1957; 1975; Radcliffe, (Medler, 1989). Third, Pedigo, it is multivoltine, with three to five overlapping per year, and indi generations have been viduals have a life span of up to six months 1938). Adults (DeLong, collected in several habitats outside of crop systems, forest, including deciduous forest understory, habitats old field, and newly-disturbed (Lamp et al., 1989 and to con habitats are believed unpubl. data; Taylor, 1993). As a result, non-crop to potato leafhopper population tribute significantly (Poos, 1935; Hogg dynamics and Hoffman, 1989). As a group, leafhoppers host ranges (Putnam, have restrictive 1941; DeLong, lie in leafhopper may 1965). Part of the reason for the wide host range of potato its ability

1 Department 2 Department Accepted

to vary

feeding

behaviors

in response

to its host

of Biological Dartmouth New Sciences, Hanover, College, of Entomology, of Nebraska, Nebraska Lincoln, University 16 April for publication 1994.

plant.

Hampshire 68583.

For

example,

03755.

VOLUME 67, NUMBER 4 355 Backus and Hunter tissues, (1989) found that adults on alfalfa fed on vascular on beans they fed on mesophyll whereas adults settle on tissues. Furthermore, different portions of the two hosts (Backus et al., 1990). Artificial plant breeding the host range and plant introductions into North America may have enhanced of the leafhopper. workers have For example, resistance plant long noted an to this leafhopper association of susceptibility with lack of species pubescence in original germplasm and other physical characteristics lines of soybean, potato, et al., 1979; Tingey, and alfalfa (Broersma et al., 1972; Robbins 1985; Brewer et In this resistance is trichomes because al., 1986a, b). part, impede the physical to the plant surface (Lee et al., 1986). A chemical normal attachment of individuals to feeding is less frequently cited compared to a physical basis. basis for resistance For example, of Solanum comparisons species suggest that tri closely-related content is associated with resistance chome presence and not glycoalkaloid (Tingey et al. (1979) found interrupted and Sinden, Raman 1982). Alternatively, feeding in response to feeding on glycoalkaloid behavior within artificial diet in the absence content was as of physical resistance factors, and that increased glycoalkaloid sociated with greater resistance among potato cultivars. A larger study of 100 found potato resistance associated with both the species of Solanum leafhopper and glandular and that artificial led tomatine selection trichomes, glycoalkaloid to increased susceptibility et al., 1992). by potato leafhopper (Flanders These studies suggest that potato is able to feed as an adult on a leafhopper wide range of plant species, although certain morphological and chemical factors for reproduction. Fewer plants serve as hosts for female limit their host utilization as well as survival of small nymphs in comparison to those that serve oviposition as hosts for adult feeding (Lamp et al., 1984b). As a consequence, nymphs develop on fewer species compared to species suitable for adult survivorship. Furthermore, variable plant potato leafhopper response to genetically diverse and phenotypically within variation. The list of plants suitable populations species shows considerable for reproduction (hosts), other plant taxa not suitable for reproduction (non-hosts), and actual utilization of hosts in nature may suggest patterns in host plant selection by potato leafhopper. the host plant lists published by Poos and Wheeler Although (1943, 1949) are are not to the conducive of because the species extensive, patterns they analysis were not grouped by taxonomic or ecological classification characteristics. Also, host plants have been discovered additional A more since their publications. toward the goal of greater under complete listing and its analysis was desired of the role of non-economic plant species in the population standing ecology of between plant characteristics and accept potato leafhopper and the relationship a The objectives to compile of this study were leafhopper. ability by potato to determine host list for this leafhopper, and ecological taxonomic complete the host list, and to determine major groups of non-host patterns within plants. we compared the compiled host list to actual utilization of hosts at Furthermore, two disparate sites. Methods Host plant species were determined by literature search on the basis of suitability for nymphal if or collected from the development, i.e., nymphs were observed in the to field reared and adult. Research of Solanum plant species has received

356

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

considerable

in the search for resistant germplasm. Rather than list all emphasis are as to exotic North not and America used cultivated species (which primarily we as these in cited the table. A in the genus plants), species only major problem is species determination. first study of this leafhopper DeLong (1931) recognized that many Empoasca identical and cannot be separated species are superficially without careful examination of male genitalia. Therefore, only studies published were included. and specimens identified after this discovery common names Plant and authorities) and families were species (including et al. (1964), Fernald determined by consulting Gleason (1952), Radford (1970), and Cronquist Hitchcock classification (1973), and Bailey (1976). Higher (order, was and class) was based on Smith classification subclass, (1977). Ecological the ecological determined and morphological found in information by consulting the taxonomic of ecological references listed above. Analysis trends among the some genera were host plants was based on genera instead of species because as discussed over-represented by species (e.g., Solanum above), and also because the genus is generally of ecological indicative Five classes were type. ecological used. First, habitats of host genera were categorized the level human dis of by to turbance either as occupants of habitats disturbance subjected by humans or that of caused "natural" habitats (including by grazing livestock) (i.e., rarely or never subjected to human disturbance). Two, habitats associated with genera were further categorized Three habitat by the level and frequency of disturbance. at least types were recognized: cropland annually-tilled (land that is cultivated once a year); fields, pastures, on average and waste places (land that is disturbed once every five years and may be grazed by livestock); and woodlands, shrublands, and park-like perennial plants (e.g., trees and shrubs) settings where long-lived or planted. Third, genera were categorized have been naturalized by growth form versus or peren and if herbaceous then annual, biennial, herbaceous (i.e., woody were as or nial life history). Fourth, to the native introduced genera categorized a the included both coincident with range. Occasionally, genus leafhopper's region In native and introduced such the of cases, species (e.g., Medicago). majority was were as to hosts used the genus. Fifth, species "compet classify categorized or "stress tolerant" according to the autecological scheme by itive," "ruderal," Grime (1979). Briefly, this scheme is defined by the role of stress and disturbance are plant species that factors on limiting the distribution of plants. Competitors low stress ruderals exploit of low stress and low disturbance, exploit conditions and high disturbance, and stress tolerants exploit high stress and low disturbance. To determine sites were identified during 1989 actual host utilization, Nebraska a number and 1990 in Lancaster Co. that provided of plant species that were considered (based on an early draft of Table potential hosts for potato leafhopper to established alfalfa fields, and were sampled once 1). The sites were adjacent 30 in 1989 and from each week during the growing season from June 6 to August 15 in 1990. The non-crop June 11 to August species that were sampled at the elm [Ulmus americana 1989 site were American [Morus rubra L.], red mulberry [Leonu (Michx.) MacM.], motherwort L.], wild four o'clock [Mirabilis nyctaginea rus cardiaca L.], catnip [Nepeta cataria L.], common milkweed [Asclepias syriaca dock [Rumex crispus L.], English L.], narrowleaf L.], Siberian elm [Ulmuspumila walnut L.], and sunflower [He [Celtis occidentalis [Juglans regia L.], hackberry sumac [Rhus lianthus annuus L.]. Species sampled at the 1990 site were smooth

VOLUME 67, NUMBER 4 357 rose [Rosa arkansana], bird's foot trefoil glabra L.], willow [Salix sp.], Arkansas [Lotus corniculatus L.], Siberian elm, hemp [Cannabis sativa L.], box elder [Acer red bud [Cercis canadensis [Medicago hispida Gaertn.], negundo L.], bur clover in 1989 Five individual plants of each species were sampled L.], and hackberry. in 1990. and three plants per species were sampled and Education Center, Wash Research sites at theWestern Maryland Similarly, were host for utilization leafhopper. sampled by potato ington Co., Maryland, Sites were sampled each week from June 7 in 1989, June 11 in 1990, and May three man-hours consisted of approximately 28 in 1991 until mid-July. Sampling each week of inspecting leaves of various plants in non-crop habitats near alfalfa. species sampled were black locust [Robinia pseudoacacia Primary L.], chestnut oak [Quercus prinus L.], hackberry, redbud, slippery elm [Ulmus rubra Muhl.], black nightshade [Solanum nigrum L.], bitternut [Carya cordiformis hickory (Wang.) K. Koch.], blackberry [Acer rubrum L.]. Other [Rubus sp.], and red maple less common reared species were also inspected, but no nymphs were successfully to adulthood. In both states, large nymphs found on plants were collected in vials and returned to the laboratory. The nymphs were reared in one dram vials. A second screw cap vial, filled with 5% sucrose in water and covered with Parafilm 'M,' was set on were to feed the vial the allowed top-down containing nymph. Large nymphs on the solution until molting to the adult stage. Adults were frozen and later to species. identified Results

and Discussion

list of known species of plants that were suitable for potato leafhopper was composed and of 220 species, in 100 genera and reproduction development 26 families the majority of host species (Table 1). In spite of the species diversity, are represented 61.8% of the species, by the family Fabaceae (=Leguminosae, 47% of the genera). This fact, plus the fact that the leafhopper was originally on 'Windsor' bean, Vicia faba L. (Harris, from a specimen collected described the use of the common name, potato leafhopper. The following 1841), contradicts over 75% of the species on the list: Fabaceae five families represent (61.8%), Asteraceae (5.0%), Fagaceae (4.5%), Rosaceae (3.6%), and Cucurbitaceae (3.6%). The families of known potato leafhopper hosts include 16 orders which represent all six North American subclasses of the class Magnoliopsida (=dicotyledonous a Table and include plants, range of relatively ancestral families 2), (e.g., Berber more to derived families the families idaceae) Furthermore, (e.g., Asteraceae). and represent species with highly divergent types. chemistry morphological The diversity in the taxonomic classification is reflected in the diversity within the ecological the leafhopper's classification range, 73% of its (Table 3). Within host genera occur exclusively in manmade habitats. Although the remaining 27% of the host genera are members of natural communities, they are often used as or pasture/forage can be ornamentals these plants occupy crops. The habitats on the level and frequency of anthropomorphic divided into three types depending disturbance. About one-third associated with annually tilled (32%) are commonly The

and waste places, and the crop land, another third (32%) inhabit fields, pastures, final third (36%) reside in woodlands, and park-like shrubland, settings. Herbs represent almost two-thirds (64%) of the host genera, most of which are perennial.

358 1.

Table field

Plant

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

suitable

species

for potato

leafhopper

as discovered

development

nymphal

studies. Plant family, species

name Source'

Common

Aceraceae Acer

sp.

Acer

negundo saccharum

Acer

Marshall

d

maple boxelder

L. sugar

1 d, maple

j

Amaranthaceae Amaranthus

L.

retroflexus

d

pigweed

Anacardiaceae Rhus

chinensis

Rhus

copallina

Rhus

glabra

Rhus

Mill.

sumac d

shining L. L.

typhina

Asteraceae

e tree

nutgall L.

sumac d

smooth

sumac d

staghorn

(=Compositae)

Artemisia

L.

absinthium tinctorius

Carthamus

wormwood

L.

d e

safflower Dahlia

Guizotia

(L.f.) Cass. L.

abyssinica annuus

Helianthus Helianthus

ramtilla

L.

inula

artichoke d

strawflower

d

d

erecta

Tagetes Xanthium

d

Jerusalem

sp.

d

a,

d

sunflower L.

tuberosus

Helichrysum Inula helenium

dahlia

sp.

L.

African

marigold d d

cocklebur

sp.

Zinnia sp.

zinnia

d

Berberidaceae Berberis

Pursh

aquifolium

d

barberry

Betulaceae Betula

sp.

Betula

nigra

birch

d L.

e birch

river

Cannabinaceae L.

sativa

Cannabis

d

hemp

Caricaceae Carica

L.

papaya

papaya

d

Chenopodiaceae Beta

vulgaris

L.

d beet

sugar

Convolvulaceae Ipomoea

batatas

sweet

Poir.

d potato

Cucurbitaceae

Cucurbita ficifolia Bouche Cucurbita

lundelliana

Cucurbita

maxima

Cucurbita

mixta

Cucurbita Cucurbita Cucurbita Cucumis

sativus

h

pumpkin winter

h squash h squash h squash

butternut acorn

L. Bailey L.

h squash

wild

Pangalo Duchesne moschata pepo sororia

h gourd

leaf

fig

Bailey Duchesne

wild cucumber

squash d

h

during

VOLUME 67, NUMBER 4 359 1. Continued.

Table Plant family, species

Common name

Source1

Euphorbiaceae Croton

capitatus Michx. L. communis

Ricinus Fabaceae

glabra Poir. herbacea Walt. Amorpha L. Arachis hypogaea L. canadensis Astragalus

Astragalus

glycyphylla

Astragalus

uliginosus arborescens

Cassia

milk

L. L.

var.

Clitoria

vetch

d

vetch

e

Colutea Coronilla

varia

Coronilla

cretica

partridge (Pollard) Macb.

Crotalaria

lanceolata

Crotalaria

usaramoensis

Desmodium

paniculatum strictum DC lablab

Galactia

texana

Galactia

wrightii

Indigofera Indigofera Indigofera Indigofera Indigofera Indigofera Indigofera

indigo

e trefoil

tick

e trefoil bean d e

pea

e

pea

e

pea

e

goat's d

soybean licorice

(Nutt.) Fisch.

d

Pursh

indigo E. G.Baker farichildii kirilowi Maxim. indigo Scheele lindheimeriana

licorice

d e

licorice e e

indigo e

e

indigo flax-leaved

Retz. Vent.

rue e locust

honey

L.

macrostachys

e trefoil

bonavist milk

d sp. indigo arrecta Benth.

linifolia

e bush

indigo tick

milk

Galega officinalis L. Gleditsia triacanthos max Glycine (Merrill)

Glycyrrhiza

e dalea edalea e dalea

black

milk

A. Gray L.

lepidota uralensis

d

foxtail

tick

Galactia

glabra

d

rattlebox

A. Gray (L.) DC

A. Gray volubilis (L.) Britton

Glycyrrhiza

d

rattlebox f.

L.

Glycyrrhiza

d

rattlebox

Baker

A. Gray batacaulon

Dolichos

e senna 1d, vetch

rattlebox Kotschy E. Mey.

Willd. alopecuroides A. Gray frutescens Robins lumholtzii and Fern.

Desmodium

e

crown

L.

intermedia

Desmodium

d

yellow-wood butterfly-pea bladder

e

L.

Crotalaria

ordiae

d

butterfly-pea md,

K. Koch

L.

incana

Dalea

wild

sp. redbud

L.

Crotalaria

Dalea

e pea tree e pea e pea partridge sensitive plant e

Siberian

robusta

lutea (Michx.) laurifolia Poir. L. arborescens

d

milk

L.

canadensis

Cladrastis

vetch

milk

Centrosema Cercis

e

vetch

Lam.

indigo

indigo

e

vetch

milk

Michx.

fasciculata nictitans

false 1 d,

peanut

Lam.

falcatus

Cassia

herbaceous

milk

Astragalus

Caragana Cassia fasciculata

d indigo

false

cicer L.

Astragalus

Dalea

d

(=Leguminosae)

Amorpha

Dalea

e

hogwort castor-bean

e indigo e

e

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

360

Table

1. Continued.

Plant family, species

Indigofera

Mats. pseudotinctoria Franch. reticulata

Indigofera

sphaerocarpa

Indigofera Laburnum

suffruticosa

Indigofera

e

indigo

A. Gray Mill. indigo sp.

latifolius ornatus Nutt.

wild

Sylvester L. venosus Muhl.

perennial wild

sericea

L.

Lespedeza Masam.

e pea

bush

bush

Lespedeza

maximowiczi

Lespedeza Lespedeza

Michx. procumbens seiboldi Miq.

Lespedeza

stuvei Nutt.

Lespedeza

thunbergii

C. K.

(Benth.) Greene (Nutt.)

hemicycla

Medicago

hispida

Medicago

lupulina

indica

Melilotus

e clover

and Clements

arenaria

Onobrychis Petalostemun

sativa

vetch e

deer e

vetch

e

vetch

deer d

lupine

1 medic

spotted toothed

e bur clover e

clover

bur bur

e

clover

1e, medic

black L. L.

e

clover

sweet

white sour

yellow Lam.

sweet s

onobrychis and Standi.

cloverd,f d

nissoliae d

onobrychis Woot.

clover d, f, 1 1 clover

yellow mimosa

Schotf

Lam.

prostratum

1a, d,

alfalfa bur

(L.) All.

(L.)

1

clover

bur

A. Gray DC.

Onobrychis

trefoil e

large-flowered e

(L.) All.

officinalis

trefoil e

bird's-foot

Desr.

sp. schottii

e clover

bush

L.

polymorpha sativa

Melilotus

e

bush

Willd.

Gaertn.

e

clover clover

Grossheim

scutellata

e clover

bush bush

Ottley

Medicago

e clover

bush

deer

denticulata

alba

Schneider

Clements

Medicago

e

clover

trefoil

Medicago

Medicago Melilotus

bush

Greene

Dougl. arabica (L.) Huds.

Medicago

e clover

bush

L.

Lotus

Medicago

e clover

bush

(DC.) Nakai (L.) Britt.

(Benth.) Greene

scoparius nanus

e clover

bush

bush

grandiflorus humistratus

Lotus

e clover

bush

L. H.

Lespedeza

salsuginosus

clovere

bush

Bailey juncea (L.f.) Pers. latissima Nakai

purshianus

e clover

(Masam.)

japonica

Lotus

clover d

bush

Lespedeza

Lotus

e clover

bush

Lespedeza

Lespedeza virginica Lotus corniculatus

e e

clover

and Hosak

Lespedeza

clover d

clover

G. Don

(Dumont)

(L.) Hornem. frutescens hirta (L.) Hornem.

Lespedeza

Nissolia

e

bush

Miq. cyrtobotrya v. shimadae daurica

Lespedeza

Mimosa

pea

bush

Michx.

capitata cuneata

Lespedeza

Lupinus

e

sp.

Lespedeza

Lotus

everlasting pea

e

Thunb.

Ell. angustifolia bicolor Turez.

Lespedeza

1pea e pea

singletary

Lespedeza Lespedeza

chain d

golden L.

Lathyrus

Lathyrus Lebeckia

e

indigo e

hirsutus

Lathyrus

e

indigo

Lathyrus Lathyrus

name Source1

Common

e prairie

clover e

VOLUME 67, NUMBER 4 361 1. Continued.

Table

Phaseolus

aconitifolius

Phaseolus Phaseolus

v. acutifolius aureus Roxb.

Phaseolus

calcaratus

Phaseolus

lunatus

Phaseolus

metcalfd

Phaseolus

vulgaris

Piptanthus Pueraria

mung

thunbergiana

Rhynchosia

rariflora texana Torr.

Rhynchosia Robinia pseudoacacia exaltata vesicaria

Sophora

Tephrosia Tephrosia Trifolium

Trifolium

pratense

Trifolium

repens

Trifolium

resupinatum corniculata

rhynchosia e m locust d, e river hemp

black Colorado

Japanese bean

wild and Gray

e rue e

d

alsike

clover d clover d

zigzag red L.

m d,

clover

1 clover d, 1 clover d,

white

L.

Persian e

L. trigonella L. Trigonella foenum-grecum trigonella v. oblongifolia e vetch Vicia amoena Regel L. Vicia angustifolia narrow-leaved Trigonella

Vicia

cracca

Vicia

dasycarpa Vicia faba Vicia ludoviciana

cow

L. Ten.

e vetch1

vetch

d 1vetch

winter L.

d

broadbean

Nutt.

Vicia minutiflora Dietr. Vicia variabilis Freyn

pea

d

cracca

L.

1 vetch

deer-pea

vetch1

pygmy-flowered and

Sint.

vetch

d

pagoda-tree e

goat's

hoary cracca

(L.) Pers. a

L. hybridum L. medium

e Texas

and Gray

L.

Trifolium

Trifolium

e

rhynchosia rhynchosia

(Raf.) Cory e Ell.

vivginiana clover

e

e cowpea Vigna capensis Walp. d cowpea (L.) Endl. Vigna sinensis Wisteria f. rosea (Bean) Rehd. and Wils. floribunda Wisteria sinensis Chinese (Sims) Sweet

American

wisteria

d

wisteria d

Fagaceae Castanea Castanea

dentata

Castanea

pumila alba L.

Quercus Quercus Quercus Quercus Quercus Quercus Quercus

(Marsh.) (L.) Mill.

Borkh.

Engelm. muhlenbergii palustris Muench. L. phellos rubra (Marsh.)

sp.

Ashe

Chinese

chestnut

American d

chinquapin d oak

scrub

doak chestnut

yellow oak j d northern

oak

d

blackjack pin willow

d

chestnut d

white

ilicifolia Wang. marilandica Muench.

a, d

d

L.

sp.

bean

kidney

e

piptanthus

kudzu

spicata tenella A. Gray toxicaria (L.) Pers.

Tephrosia

s bean e bean,

garden

Sweet

(L.) Britt. (Walt.) Torr.

Tephrosia

lima bean d Metcalf

japonica helvola

Strophostyles

d

bean

Benth.

(Ell.) DC. Standi.

difformis

Sesbania

rice

L.

Rhynchosia

e bean

tepary

d

Roxb.

(Hook.) Benth.

e

bean

Freeman

var. macrocarpus Woot. Standi.

nepalensis

Sesbania

moth

Jacq. latifolius

name Source1

Common

Plant family, species

oak red doak

oak d

362

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 1. Continued.

Table Plant family, species stellata

Quercus

d

oak

post

Wang.

name Source1

Common

Junglandaceae sp.

Carya L.

regia

Juglans Lamiaceae

d, j walnut d

hickory

English

(=Labiatae)

Lallemantia

Fisch.

iberica

Leonurus

hortensis

L.

officinalis

L.

Satureja

d

& Mey.

L.

cardiaca

i

motherwort e

savory

Malvaceae Althaea Althaea

roseus

Hibiscus

d e

hibiscus

L.

sabdariffa

Malva

da,

rose-mallow

Thore

d

hollyhock d

cotton

sp. L.

Gossypium cannabinus Hibiscus Hibiscus

sp. marshmallow

d

roselle

d

mallow

sp.

Moraceae Humulus

lupulus

L. americanus

Nutt.

e

hop

Phytolaccaceae sp.

Phytolacca

d

pokeweed

Polygonaceae pensylvanicum Polygonum L. Rheum rhaponticum Rumex

crispus

Rumex

obtusifolius

d

smartweed

L.

da,

rhubarb

leaf dock d, m d dock

narrow

L. broadleaf

L.

Rhamnaceae Rhamnus

buckthorn d

alder

L.

Frangula

Rosaceae paradoxa

Fallugia

americana

Prunus

spp.

rose

Endl.

Marsh.

cerasus

Prunus Rosa

(Don)

plum L.

Fragaria Malus

sp.

e plume d strawberry a, d apple

Prunus

sp.

cherry

Apache

d,

j

d sour

d

cherry

d Rubus

Sanguisorba

sp.

minor

salad

Scop.

sp.

m

d, blackberry e burnet

Salicaceae Salix Salix Salix

sp. willow exigua Nutt. Barratt hookeriana

d

willow e e willow

Hooker's

Solanaceae L. belladonna Atropa Solanum spp. c, g, k L. carolinense Solanum Solanum

melongenaL.

Solanum

onigerum

d

horsenettle eggplant

b

d

belladonna

a,d b,

VOLUME 67, NUMBER 4 363 Table

1. Continued.

Solanum

torvum

Solanum

tuberosum

name Source1

Common

Plant family, species Mexican

Swartz L.

solanum d c a,

potato

Tiliaceae L.

americana

Tilia

American

basswood j

Ulmaceae Celt

is occidentalis

Celtis

laevigata

L. Willd.

Ulmus

pumila rubra Muhl.

1 Sources

are as follows:

and Wheeler, 1967; h, Howe 1992;

Table 1. All

1

americana

Ulmus Ulmus

1943; e, Poos and Rhodes,

1,Taylor,

1993; m,

md, j,

hackberry

sugarberry L.

L.

elm d, j, 1, m

American

dwarf

d elm melm

slippery

a, DeLong, 1931; c, Sleesman, 1940; d, Poos 1931; b, Poos and Haenseler, and Lauer, and Wheeler, and Jarvis, 1949; f, Manglitz 1966; g, Radcliffe and Radcliffe, 1976; i, Lamp et al., 1984; j, Lamp et al., 1989; k, Flanders this first

report.

2. of higher based classification, Summary are within subclasses the class Magnoliopsida.

on Smith

of hose

(1977),

in Table

listed

plants

Number of Subclass

Magnoliidae Hamamelidae

Order

Juglandales Caryophyllales

Berberidaceae 1 Betulaceae 1 Fagaceae 2 Moraceae 1 Cannabaceae 1 Ulmaceae 2 Juglandaceae 2 Amaranthaceae 1

Polygonales Violales

Chenopodiaceae 1 Polygonaceae 3 Caricaceae 1

Ranunculales Fagales Uticales

Caryophyllidae

Genera Family

1

Phytolaccaceae

Dilleniidae

Cucurbitaceae

Malvales

Rosidae

Salicales Rosales Euphorbiales Rhamnales Sapindales

Asteridae

Polemoniales

Asteraceae Lamiaceae Total

1 1 10 1 1 5 2 1 1

2

1 4 1 8

Malvaceae 4 Tiliaceae 1 Salicaceae 1 Rosaceae 7 Fabaceae 47 Euphorbiaceae 2 Rhamaceae 1 Aceraceae 1 Anacardiaceae 1 Solanaceae 2 Convolvulaceae

Asterales Lamiales

Species

1

10 3 100

6 1 2 8 136 2 1 2 4 6 1

11 3 220

364 3.

Table Table

Ecological

classification

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY of plant

suitable

genera

for potato

leafhopper

reproduction,

from

1. Genera within class

Class Category (%) Source

of disturbance

Human

73

Natural Habitat

disturbance

27

Field,

pastures,

Woodlands,

Growth form

32 cropland waste places

Annually-tilled

shrublands,

Herbaceous

36

40

perennial biennial

Herbaceous Herbaceous

annual

Plant origin

Native

Plant autecology

Competitive

2 22

38

Introduced

Stress

32

36

Woody

Ruderal

parks

62

64

36 tolerant

0

22% of the host genera and woody perennials 36%. Moreover, into the leafhopper's represent species that have been introduced to Grime's range. According system, 36% are ruderals while (1979) classification are the remaining These genera genera represent her competitive. competitive or woody baceous with old fields, hay associated species commonly perennial and below that little disturbance. The lack of pastures fields, experience ground stress tolerant hosts is likely a function of a number of factors, few including over use our the stressful environments limited of such range, leafhopper's highly plants for crop and ornamental plants, and limited observations. The breadth of plant hosts alone, and especially when classified taxonomically and ecologically, that this species has readily adopted hosts from demonstrated most of the major and disparate dicotyledonous taxa within its range. Wherever utilizes native and introduced dicot crops, forages, it occurs, potato leafhopper its propensity for openings weeds, shrubs, and trees. From (fields, meadows, created by and shared with humans, it is not surprising that most of its pastures) hosts are found in anthropogenic and the majority habitats, most are herbaceous, in So have been introduced. A recent analysis of potato resistance leafhopper have clones resistant while that ancestral lanum characteristics, suggest types human lack intermediate from selection resistance derived types (Flanders et al., use occur not does the use of its without of human-derived plants 1992). Yet,

Annuals comprise 62% of the genera

trees and native plants in natural landscapes, legumes (Lamp et including and data This observation discussed al., 1989; Taylor, 1993, suggests that below). the has enhanced the human modification of leafhopper's landscape although the leafhopper utilization of hosts, likely was able to utilize native hosts and the in North America. habitats before the presence of agriculture Furthermore, use of plant monocultures has likely led to for agriculture and land management at overwintering densities both during spring generations increased leafhopper sites (e.g., the use of legumes for erosion control, Taylor, 1993) as well as during

native

VOLUME 67, NUMBER 4 365 Table

4.

Common

hosts non-crop determined

Site Year NE:

Lancaster

at two disparate sites within potato nymphs. by rearing of collected

10 1990 MD:

Washington

Co.

1988

summer

range,

Minimum number of host species sampled Host(s) 198911

Co.

leafhopper

Rumex

crispus,

Cercis

canadensis,

Ulmus

9

Robinia

psuedoacacia

91989

Robinia

pseudoacacia, rubra

Ulmus 91990

Celtis

occidentalis,

americana,

Celtis

Trifolium

pratense

occidentalis sp., Celtis

Rubus Ulmus

occidentalis,

rubra

landscapes (e.g., the use of alfalfa monocul agricultural et tures, Lamp, al., 1984a). 1991; Lamp can only speculate why some plant taxa are not suitable hosts for this We insect. One of the more groups is all of the species polyphagous striking non-host as grasses and sedges). in the class Liliopsida such plants (i.e., monocotyledonous of this class may be a function Its inability to produce offspring on any member to access smaller vascular bundles or the inability of its nymphs of the relatively are frequently collected on grasses (e.g., corn, them (Lamp et al., 1984b). Adults Poos and Wheeler, 1943), and in no choice tests adults will survive but will not on weed and crop grasses (Lamp et al., 1984b, and unpubl. data). various oviposit on the list is gymnosperms taxon not represented Another (Division Pinophyta). on eastern white Potato has been collected strobus L., pine, Pinus leafhopper taeda L., and longleaf pine, Pinus palustris Mill loblolly pine, Pinus (Poos and Wheeler, 1943; Taylor, 1993). Recent evidence using elemental markers has dem onstrated that adults do ingest fluids from loblolly pine, a common site of over the summer

in northern

in southern U.S. potato leafhopper (Taylor et al., 1993). wintering in the class Magnoliopsida, encountered Among eight major orders frequently the leafhopper's Ge range have not been found as hosts: Ericales, Gentianales, Some and Umbellales. raniales, Myrtales, Rubiales, Papaverales, Scrophulariales, of these orders are well known for possessing defenses chemical prominent (e.g., some have both chemical and and Umbellales), Papaverales, Scrophulariales, some we Geraniales and and know little about defenses for physical Ericales), (e.g., their defensive and Myrtales) and Turner, strategies (e.g., Rubiales (Harborne 1984). In spite of its inability to reproduce on these groups, non-host plants for potato to its population leafhopper may contribute ecology during times when hosts are For example, unavailable. studies during the winter have shown that the leaf hopper uses non-host plant species as refugia (Decker and Cunningham, 1968; increases in population densities in corn have been Taylor, 1993). Furthermore, associated with the harvesting of adjacent alfalfa fields during the summer (Lamp, these non-hosts may allow adults to survive during times unpubl. data). Thus, or are difficult when host plants are unavailable to locate. The nymph successful with a laboratory culture although rearing technique, success had with field-collected (Lamp, unpubl. data), varying nymphs. Approx and 48% of the Maryland imately 20% of the Nebraska nymphs collected, nymphs sex to was skewed to females: 72% survived adulthood. the ratio collected, Also,

366

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

of the Nebraska nymphs and 80% of the Maryland nymphs surviving to adulthood were female. This may have been the result of the rearing procedure. All males that survived were identified as E. fabae. Of the 26 potential hosts sampled, potato leafhopper nymphs were verified on only one to four plant species per year and site (Table 4). Thus, in spite of the large list, host utilization under field conditions in that at one location may the actual host fact for suggests range reproduction consist of only a few species. DeLong further (1965) suggested that host utilization at the time of first varies between years as a result of varying host phenology spring migration. In conclusion, the diversity of host plants suitable for reproduction suggests that non-crop habitats may be a significant source of potato leafhopper populations states. Also, the ability of leafhopper after spring migration into northern adults to utilize at least some non-host a secondary role species as refugia contributes to non-crop habitats. Yet, our limited data suggest that utilization of non-crop is restricted to a relatively few naturalized habitats hosts. Additional information on the role of these habitats to better understand is needed potato leafhopper population

dynamics.

Acknowledgments We thank Marilyn Morris in the early development for her assistance of a host we In list for potato thank David Clara addition, Fuentes, Liewehr, leafhopper. on earlier manuscript and Bob Wright Robert Peterson, for their comments drafts. came in for this from No. USDA-CSRS MD-H-176 part Support study Project and (NC-193) Article Number

the Maryland Station. This is Scientific Experiment Agriculture of Contribution Number 8764 the A6553, Maryland Agricultural and Journal Series Number of Entomology, Station, Department the Agricultural Research Division of the University of Nebraska

Experiment 10375 of Lincoln.

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