Leafhoppers (Homoptera: Cicadellidae) that probe human skin: A

© Koninklijke Brill NV, Leiden, 2013 DOI 10.1163/18749836-06001064

Terrestrial Arthropod Reviews 6 (2013) 201–225 brill.com/tarT A R

Leafhoppers (Homoptera: Cicadellidae) that probe human skin: A review of the world literature and nineteen

new records, from Panama

Edwin Domínguez Núñez and Annette Aiello

Smithsonian Tropical Research Institute, Apartado 0843-03092 Balboa, Ancon, Republic of Panama

e-mails: [email protected]; [email protected] Received on January 16, 2013, Accepted on 17 March 2013.

Final version received on April 17, 2013

SummaryWe review the world literature on leafhoppers (Cicadellidae) that have probed human skin, update the names presented in those published reports, and sort out which are original accounts and which are citations of previous reports (Table 1). We add 19 new records, representing five subfamilies, and at least 13 genera, all from Panama (Table 2). We speculate as to why these normally phytophagous insects sometimes probe human skin, and provide insight into the possible evolutionary consequences of this behavior.

KeywordsInsecta; Homoptera; Cicadellidae; leafhoppers; aberrant feeding; behavior; bite; blood; entomology; evolution; human skin; probe; medical entomology; new records; Panama

Introduction

Members of the insect orders Homoptera and Heteroptera, sometimes treated collec-tively as the Hemiptera, have distinctive mouth parts specialized for piercing and suck-ing that have enabled them to radiate into a variety of niches unavailable to members of most other insect groups. Concomitant with this specialization, essentially all Homoptera and Heteroptera subsist on liquid food alone.

Though the diets of Heteroptera vary from plant juices, to insect prey, to vertebrate blood, the Homoptera are restricted to plant fluids, mainly the xylem and/or phloem of vascular plants. With their high reproductivity and exceptional capacity to transmit plant diseases, the Homoptera include some of the worst agricultural pests. Among the most important in this respect are the leafhoppers (family Cicadellidae), which include highly efficient vectors of plant viruses (Nault and Rodriguez, 1985; Krinsky, 2002).

202 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225

Despite their present limitation to feeding on plants, anecdotal evidence suggests that a number of Homoptera, as well as certain plant-feeding Heteroptera, have the potential to diversify onto other food sources. That the switch from herbivory to preda-tion already has occurred within the Heteroptera, e.g., Reduviidae, lends credence to that idea, as does the fact that species representing at least 15 families of herbivorous Heteroptera and five families of Homoptera are known to have bitten humans. These accounts were reviewed by Bequaert (1926), Myers (1929), Usinger (1934), Ryckman (1979), Ryckman and Bentley (1979), and Alexander (1984).

Homoptera known to annoy humans by probing with their mouthparts are found among the leafhoppers (Cicadellidae), treehoppers (Membracidae), spittle bugs (Cercop-idae), plant hoppers (Fulgoroidea), and cicadas (Cicadidae).

Over the years, a variety of non-blood-feeding insects, e.g., lacewing larvae (Neuroptera: Chrysopidae), has “bitten” each of us (EDN and AA) and, independently, we began to collect the leafhoppers involved. A chance conversation led to the joint preparation of this paper, with the objectives of investigating the history of this phenomenon in cicadellids and reporting our own experiences.

Methods

We tracked down every published report of leafhoppers biting humans that we could find, updated the names, and organized the original reports chronologically, each fol-lowed by the publications that cite them. Thus, the entire history of these reports is laid out in Table 1.

Our own observations (Table 2) were made in three general areas of the Republic of Panama: Panama City (immediately east of the Panama Canal); the town of Arraiján (just west of the Canal watershed); and Cocle Province, which is farther west. Attackers were collected manually and preserved in 70% ethanol or were frozen; later, all speci-mens were mounted on points and the male genitalia were preserved in vials in glyc-erin. Vouchers are deposited in MIUP (Museo de Invertebrados G. B. Fairchild de la Universidad de Panamá) and at STRI (Smithsonian Tropical Research Institute) as noted in Table 2.

Nomenclature of the Cicadellidae was updated Cwikla, 1985a; Dietrich and Dmitriev, 2006; Dietrich and Dmitriev, 2008; Dlabola, 1958c; Emeljanov, 1999c; Ghauri, 1971; Ghauri, 1974a; Kramer,1971; Metcalf, 1967a; Metcalf,1967c; Metcalf, 1968a; Ross, 1968; Takiya et al. 2006; Young, 1977.

Results

In all, we found 174 records, published from 1911-2006, by 27 authors (authors and co-authors being counted as one), in 33 publications. These reports represent three cicadellid subfamilies, 19 genera, and at least 27 species from various regions of the world, including China, Cuba, England, French Polynesia, India, Japan, North Africa, the Philippines, Trinidad, and the United States (Table 1). Of the 174 records,

E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225 203Ta

ble

1. C

icad

ellid

ae re

port

ed to

hav

e pr

obed

hum

an sk

in w

orld

wid

e. O

rigin

al re

port

s are

hig

hlig

hted

and

arr

ange

d ch

rono

logi

cally

, eac

h fo

llow

ed b

y th

e pu

blic

atio

ns th

at c

ite th

em. I

nfor

mat

ion

in sq

uare

bra

cket

s was

add

ed b

y th

e au

thor

s.

Empo

asca

faba

e (H

arris

, 184

1)Ty

phlo

cybi

nae

Dal

las,

Texa

s [U

SA]

Tuck

er (1

911,

p.2

9) (a

s Em

poas

ca m

ali),

spec

imen

s det

erm

ined

by

O

. Hei

dem

ann.

Inge

sted

bloo

d.

Empo

asca

faba

e (H

arris

, 184

1)Ty

phlo

cybi

nae

Rile

y &

Joha

nnse

n (1

915,

p.3

3) (a

s Em

poas

ca m

ali),

sour

ce n

ot

give

n bu

t mos

t lik

ely

Tuck

er (1

911)

.Em

poas

ca fa

bae (

Har

ris, 1

841)

Typh

locy

bina

eLa

wso

n (1

926,

p.7

3), c

iting

Rile

y &

Joha

nnso

n (1

915)

.Em

poas

ca fa

bae (

Har

ris, 1

841)

Typh

locy

bina

eN

orth

Am

eric

aM

yers

(192

9, p

.475

), ci

ting

Tuck

er (1

911)

, Rile

y an

d Jo

hann

sen

(191

5).

Empo

asca

faba

e (H

arris

, 184

1)Ty

phlo

cybi

nae

Ryck

man

& B

entle

y (1

979,

p.3

7-38

) (as

Em

poas

ca m

ali),

citi

ng

Rile

y &

Joha

nnse

n (1

915)

.Em

poas

ca fa

bae (

Har

ris, 1

841)

Typh

locy

bina

eRy

ckm

an &

Ben

tley

(197

9, p

.42)

(as E

mpo

asca

mal

i), c

iting

Tu

cker

(191

1).

Scap

hyto

pius

acu

tus (

Say,

1830

)D

elto

ceph

alin

aeR

iley

& Jo

hann

sen

(191

5, p

.33)

(as P

laty

met

opiu

s acu

tus),

sour

ce

not g

iven

.

Scap

hyto

pius

acu

tus (

Say,

1830

)D

elto

ceph

alin

aeLa

wso

n (1

926,

p.7

3) (a

s Pla

tym

etop

ius a

cutu

s), c

iting

Rile

y &

Jo

hann

sen

(pre

sum

ably

191

5).

Scap

hyto

pius

acu

tus (

Say,

1830

)D

elto

ceph

alin

aeN

orth

Am

eric

aM

yers

(192

9, p

.475

) (as

Pla

tym

etop

ius a

cutu

s), c

iting

Rile

y &

Jo

hann

sen

(191

5).

Scap

hyto

pius

acu

tus (

Say,

1830

)D

elto

ceph

alin

aeR

iley

& Jo

hann

sen

(193

8, p

.143

) (as

Pla

tym

etop

ius a

cutu

s), so

urce

no

t giv

en, b

ut n

ot d

oubt

thei

r ow

n 19

15 re

cord

.Sc

aphy

topi

us a

cutu

s (Sa

y, 18

30)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

3) (a

s Pla

tym

etop

ius a

cutu

s), c

iting

M

yers

(192

9).

Scap

hyto

pius

acu

tus (

Say,

1830

)D

elto

ceph

alin

aeRy

ckm

an &

Ben

tley

(197

9, p

.37-

38) (

as P

laty

met

opiu

s acu

tus),

ci

ting

Rile

y &

Joha

nnse

n (1

915)

.

Scap

hyto

pius

acu

tus (

Say,

1830

)D

elto

ceph

alin

aeRy

ckm

an &

Ben

tley

(197

9, p

.38)

(as P

laty

met

opiu

s acu

tus),

citi

ng

Rile

y &

Joha

nnse

n (1

938)

.

(Con

tinue

d )


Empo

asca

faba

e (H

arris

, 184

1)Ty

phlo

cybi

nae

Faye

ttevi

lle,

Arka

nsas

[USA

]Be

cker

(191

8, p

.101

) (as

Em

poas

ca m

ali),

spec

imen

s det

erm

ined

by

W.D

. Gib

son.

Empo

asca

faba

e (H

arris

, 184

1)Ty

phlo

cybi

nae

Law

son

(192

0, p

.20)

(as E

mpo

asca

mal

i), c

iting

Bec

ker (

1918

).Em

poas

ca fa

bae (

Har

ris, 1

841)

Typh

locy

bina

eN

orth

Am

eric

aM

yers

(192

9, p

.475

), ci

ting

Beck

er (1

918)

.Em

poas

ca fa

bae (

Har

ris, 1

841)

Typh

locy

bina

eRy

ckm

an (1

979,

p.1

1) (a

s Em

poas

ca m

ali),

citi

ng B

ecke

r (19

18).

Empo

asca

faba

e (H

arris

, 184

1)Ty

phlo

cybi

nae

Ryck

man

& B

entle

y (1

979,

p.3

3), c

iting

Mye

rs (1

929)

.Em

poas

ca fa

bae (

Har

ris, 1

841)

Typh

locy

bina

eAl

exan

der (

1984

, p.5

8) (a

s Em

poas

ca m

ali),

citi

ng B

ecke

r (19

18).

Nep

hote

ttix

nigr

opict

us (S

tål,

1870

)D

elto

ceph

alin

aeM

anila

[Phi

lippi

nes]

Bank

s (19

19, p

.181

) (as

Nep

hote

ttix

apica

lis).

Nep

hote

ttix

nigr

opict

us (S

tål,

1870

)D

elto

ceph

alin

aePh

ilipp

ines

Ryck

man

(197

9, p

.11)

(as N

epho

tetti

x ap

icalis

), ci

ting

Bank

s (1

919)

.N

epho

tetti

x ni

grop

ictus

(Stå

l, 18

70)

Del

toce

phal

inae

Phili

ppin

esAl

exan

der (

1984

, p.5

8) (a

s Nep

hote

ttix

apica

lis),

citin

g Ba

nks

(191

9).

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aeM

anila

[Phi

lippi

nes]

Bank

s (19

19, p

.181

) (as

Nep

hote

ttix

bipu

ncta

tus).

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aePh

ilipp

ines

Brum

pt (1

922,

p.8

15) (

as N

epho

tetti

x bi

punc

tatu

s), c

iting

Ban

ks

(pre

sum

ably

191

9).

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aePh

ilipp

ines

Bequ

aert

(192

6, p

.189

) (as

Nep

hote

ttix

bipu

ncta

tus),

citi

ng B

anks

as

repo

rted

by

Brum

pt (1

922)

.N

epho

tetti

x vi

resce

ns (D

istan

t, 19

08)

Del

toce

phal

inae

Phili

ppin

esBr

umpt

(192

7, p

.952

-953

) (as

Nep

hote

ttix

bipu

ncta

tus),

citi

ng

Bank

s, as

in th

e 19

22 e

ditio

n.N

epho

tetti

x vi

resce

ns (D

istan

t, 19

08)

Del

toce

phal

inae

Phili

ppin

esM

yers

(192

9, p

.474

) (as

Nep

hote

ttix

bipu

ncta

tus),

citi

ng B

rum

pt

(192

2).

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aeM

artin

i (19

29, p

.385

) (as

Nep

hote

ttix)

, sou

rce

not g

ive,

but

pr

esum

ably

orig

inat

ing

from

Ban

ks (1

919)

or o

ne o

f tho

se c

iting

hi

m.

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aePh

ilipp

ines

Brum

pt (1

936,

p.1

305)

(as N

epho

tetti

x bi

punc

tatu

s), c

iting

Ban

ks,

as in

his

prev

ious

two

editi

ons.

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aePh

ilipp

ines

Brum

pt (1

949,

p.1

276)

(as N

epho

tetti

x bi

punc

tatu

s), c

iting

Ban

ks,

as in

his

prev

ious

thre

e ed

ition

s.

Tabl

e 1.

(C

ont.)

(Con

tinue

d )

E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225 205

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aePh

ilipp

ines

Ryck

man

(197

9, p

.11)

(as N

epho

tetti

x bi

punc

tatu

s), c

iting

Ban

ks

(191

9).

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aeRy

ckm

an &

Ben

tley

(197

9, p

.31)

(as N

epho

tetti

x), c

iting

Mar

tini

(192

9).

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aeRy

ckm

an &

Ben

tley

(197

9, p

.33)

(as N

epho

tetti

x bi

punc

tatu

s),

citin

g M

yers

(192

9).

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aePh

ilipp

ines

Alex

ande

r (19

84, p

.58)

(as N

epho

tetti

x bi

punc

tatu

s, m

isspe

lled

as

bipu

ncta

ta),

citin

g Ba

nks (

1919

).

Empo

asca

faba

e (H

arris

, 184

1)Ty

phlo

cybi

nae

Kan

sas [

USA

]La

wso

n (1

920,

p.2

0) (a

s Em

poas

ca m

ali).

Empo

asca

faba

e (H

arris

, 184

1)Ty

phlo

cybi

nae

Law

son

(192

6, p

.73)

(as E

mpo

asca

mal

i), a

repe

at o

f his

own

repo

rt

of 1

920.

Empo

asca

faba

e (H

arris

, 184

1)Ty

phlo

cybi

nae

Nor

th A

mer

ica

Mye

rs (1

929,

p.4

75),

citin

g La

wso

n (1

926)

.Em

poas

ca fa

bae (

Har

ris, 1

841)

Typh

locy

bina

eR

iley

& Jo

hann

sen

(193

8, p

.143

) (as

Em

poas

ca m

ali),

sour

ce n

ot

give

n.Em

poas

ca fa

bae (

Har

ris, 1

841)

Typh

locy

bina

eK

ansa

s [U

SA]

Ryck

man

& B

entle

y (1

979,

p.2

8), c

iting

Law

son

(192

6).

Empo

asca

faba

e (H

arris

, 184

1)Ty

phlo

cybi

nae

Ryck

man

& B

entle

y (1

979,

p.3

8) (a

s Em

poas

ca m

ali),

citi

ng R

iley

& Jo

hann

sen

(193

8).

Exiti

anus

indi

cus (

Dist

ant,

1908

)D

elto

ceph

alin

aeM

adra

s [In

dia]

Don

ovan

(192

0a, p

.220

; 192

0b, p

.212

) (as

Phr

ynom

orph

us

indi

cus),

bas

ed o

n sp

ecim

ens c

olle

cted

by

him

, tak

en to

Gah

an,

and

dete

rmin

ed b

y D

istan

t.

Exiti

anus

indi

cus (

Dist

ant,

1908

)D

elto

ceph

alin

ael’I

nde

Brum

pt (1

922,

p.8

14) (

as P

hryn

omor

phus

indi

cus),

citi

ng D

onov

an

(pre

sum

ably

192

0).

Exiti

anus

indi

cus (

Dist

ant,

1908

)D

elto

ceph

alin

aeM

adra

s (In

dia)

Berg

evin

(192

5a, p

. 40)

(as A

thys

anus

indi

cus),

citi

ng p

erso

nal

corr

espo

nden

ce fr

om M

arsh

all,

who

repo

rted

that

M.E

. Chi

na

foun

d tw

o sp

ecim

ens i

n th

e Br

itish

Mus

eum

that

wer

e co

llect

ed b

y D

onov

an in

Nov

. 191

9 an

d an

nota

ted

as “

bloo

dsuc

kers

.” U

nkno

wn

to h

im, D

onov

an (1

920)

alre

ady

had

publ

ished

his

acco

unt o

f the

ir bl

ood-

suck

ing

beha

vior

.

(Con

tinue

d )

Tabl

e 1.

(C

ont.)


Exiti

anus

indi

cus (

Dist

ant,

1908

)D

elto

ceph

alin

aeM

adra

s, In

dia

Bequ

aert

(192

6, p

.189

) (as

Phr

ynom

orph

us in

dicu

s), c

iting

D

onov

an (1

920)

.Ex

itian

us in

dicu

s (D

istan

t, 19

08)

Del

toce

phal

inae

Indi

aBr

umpt

(192

7, p

.952

) (as

Phr

ynom

orph

us in

dicu

s), c

iting

Don

ovan

(p

resu

mab

ly 1

920)

.Ex

itian

us in

dicu

s (D

istan

t, 19

08)

Del

toce

phal

inae

Indi

aM

yers

(192

9, p

.474

) [as

Ath

ysan

us (P

hryn

omor

phus

) ind

icus],

citi

ng

Don

ovan

(192

0).

Exiti

anus

indi

cus (

Dist

ant,

1908

)D

elto

ceph

alin

aeM

adra

s (In

dia)

Chi

na (1

929,

p.9

7) (a

s Ath

ysan

us in

dicu

s), in

not

e re

ad b

y E.

B.

Poul

ton

at 1

6 O

ct. 1

929

mee

ting

of th

e En

tom

olog

ical

Soc

iety

of

Lond

on, i

n w

hich

he

men

tions

that

he

“dre

w B

erge

vin’s

atte

ntio

n”

to D

onov

an sp

ecim

ens i

n th

e Br

itish

Mus

eum

.Ex

itian

us in

dicu

s (D

istan

t, 19

08)

Del

toce

phal

inae

Indi

aBr

umpt

(193

6, p

.130

5) (a

s Phr

ynom

orph

us in

dicu

s), c

iting

D

onov

an (p

resu

mab

ly 1

920)

, as i

n hi

s 192

7 ed

ition

.Ex

itian

us in

dicu

s (D

istan

t, 19

08)

Del

toce

phal

inae

Mad

ras,

Inde

an

glai

seFo

ley

& A

udou

ze (1

938,

p.5

26) (

as A

thys

anus

indi

cus),

repo

rtin

g ob

serv

atio

ns b

y D

onov

an in

Nov

. 191

9.Ex

itian

us in

dicu

s (D

istan

t, 19

08)

Del

toce

phal

inae

Indi

aBr

umpt

(194

9, p

.127

5) (a

s Ath

ysan

us in

dicu

s), c

iting

Don

ovan

(p

resu

mab

ly 1

920)

, as i

n hi

s pre

viou

s tw

o ed

ition

s.Ex

itian

us in

dicu

s (D

istan

t, 19

08)

Del

toce

phal

inae

Mad

ras,

Indi

aRy

ckm

an (1

979,

p.1

6) (a

s Phr

ynom

orph

us in

dicu

s), c

iting

Don

ovan

(1

920a

).Ex

itian

us in

dicu

s (D

istan

t, 19

08)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

3) (a

s Ath

ysan

us in

dicu

s), c

iting

M

yers

(192

9).

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aeC

alcu

tta [I

ndia

]Br

umpt

(192

2, p

.815

) (as

Nep

hote

ttix

bipu

ncta

tus),

citi

ng A

. Al

cock

, with

no

date

or p

lace

of p

ublic

atio

n. W

e ha

ve b

een

unab

le

to lo

cate

any

Alc

ock

publ

icat

ion

that

men

tions

Cic

adel

lidae

as

bitin

g hu

man

s.

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aeC

alcu

tta [I

ndia

]Be

quae

rt (1

926,

p.1

89) (

as N

epho

tetti

x bi

punc

tatu

s), c

iting

Alc

ock

as

repo

rted

in B

rum

pt (1

922)

.N

epho

tetti

x vi

resce

ns (D

istan

t, 19

08)

Del

toce

phal

inae

Cal

cutta

(Ind

ia)

Brum

pt (1

927,

p.9

52-9

53) (

as N

epho

tetti

x bi

punc

tatu

s), c

iting

A.

Alc

ock,

as i

n th

e 19

22 e

ditio

n.N

epho

tetti

x vi

resce

ns (D

istan

t, 19

08)

Del

toce

phal

inae

Indi

aM

yers

(192

9, p

.474

) (as

Nep

hotet

tix b

ipun

ctatu

s), c

iting

Bru

mpt

(1

922)

.

Tabl

e 1.

(C

ont.)

(Con

tinue

d )


Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aeC

alcu

tta (I

ndia

)Br

umpt

(193

6, p

.130

5) (a

s Nep

hote

ttix

bipu

ncta

tus),

citi

ng A

. Al

cock

, as i

n hi

s pre

viou

s 2 e

ditio

ns.

Nep

hote

ttix

vire

scens

(Dist

ant,

1908

)D

elto

ceph

alin

aeC

alcu

tta (I

ndia

)Br

umpt

(194

9, p

.127

6) (a

s Nep

hote

ttix

bipu

ncta

tus),

citi

ng A

. Al

cock

(miss

pelle

d as

Alc

ok).

Exiti

anus

fasci

olat

us (M

elic

har,

1911

)D

elto

ceph

alin

aeIn

Sal

ah [s

outh

ern

Alge

ria]

Berg

evin

(192

5a, p

.38-

39) (

as A

thys

anus

und

escr

ibed

sp.);

de

scrib

ed b

y hi

m in

a fo

llow

ing

pape

r (19

25b,

p.4

2) a

s Ath

ysan

us

vuln

eran

s, fro

m si

x sp

ecim

ens (

4 m

ales

, 2 fe

mal

es) s

ent t

o hi

m b

y D

r. Fo

ley,

who

repo

rted

thei

r biti

ng b

ehav

ior a

s com

mun

icat

ed in

a

lette

r fro

m D

r. Fo

uque

, dat

ed 1

2 Se

pt. 1

924.

Exiti

anus

fasci

olat

us (M

elic

har,

1911

)D

elto

ceph

alin

aeIn

Sal

ah (s

outh

ern

Alge

ria)

Berg

evin

(192

5b, p

p. 4

2) (a

s Ath

ysan

us v

ulne

rans

sp. n

ov.),

from

sp

ecim

ens c

olle

cted

by

Dr.

Fouq

ue, a

nd se

nt to

him

by

Dr.

Fole

y.Ex

itian

us fa

sciol

atus

(Mel

icha

r, 19

11)

Del

toce

phal

inae

“Sah

ara”

in e

rror

Bequ

aert

(192

6, p

.189

) (as

Ath

ysan

us v

ulne

rans

), ci

ting

Berg

evin

(1

925)

, but

he

misr

ead

Sala

h as

“Sa

hara

.”Ex

itian

us fa

sciol

atus

(Mel

icha

r, 19

11)

Del

toce

phal

inae

“Sah

ara”

in e

rror

Mye

rs (1

929,

p.4

74) (

as A

thys

anus

vul

nera

ns),

citin

g Be

rgev

in

(192

5b),

thou

gh p

erpe

tuat

ing

Bequ

aert’

s err

or c

once

rnin

g th

e lo

catio

n; p

erha

ps h

e di

d no

t hav

e ac

cess

to B

erge

vin’s

pub

licat

ion.

Exiti

anus

fasci

olat

us (M

elic

har,

1911

)D

elto

ceph

alin

aeIn

Sal

ah (s

outh

ern

Alge

ria)

Chi

na (1

929,

p.9

7) (a

s Ath

ysan

us v

ulne

rans

), ci

ting

Berg

evin

(1

925a

& 1

925b

), in

lette

r rea

d by

E.B

. Pou

lton

at 1

6 O

ct. 1

929

mee

ting

of th

e En

tom

olog

ical

Soc

iety

of L

ondo

n.Ex

itian

usD

elto

ceph

alin

aeM

artin

i (19

29, p

.385

) (as

Ath

ysanu

s), so

urce

not

giv

en, b

ut b

ased

on

the m

atch

ing

list o

f gen

era,

pre

sum

ably

orig

inat

ing

from

Be

rgev

in (1

925a

).Ex

itian

us fa

sciol

atus

(Mel

icha

r, 19

11)

Del

toce

phal

inae

Alge

riaRy

ckm

an (1

979,

p.1

2) (a

s Ath

ysan

us sp

.), c

iting

Ber

gevi

n (1

925)

.Ex

itian

usD

elto

ceph

alin

aeRy

ckm

an &

Ben

tley

(197

9, p

.31)

(as A

thys

anus

sp.),

citi

ng M

artin

i (1

929)

.Ex

itian

us fa

sciol

atus

(Mel

icha

r, 19

11)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

3) (a

s Ath

ysan

us v

ulne

rans

), ci

ting

Mye

rs (1

929)

.

(Con

tinue

d )

Tabl

e 1.

(C

ont.)


Eusce

lis cu

rtice

ps L

indb

erg,

192

7D

elto

ceph

alin

aeK

hart

oum

(Sou

dan

Egyp

tien)

Berg

evin

(192

5a, p

p. 3

9-40

) (as

Ath

ysan

us sp

.), re

gard

ing

a sp

ecim

en c

olle

cted

by

H.B

. Joh

nsto

n, 1

7 O

ct. 1

924,

and

sent

by

Mar

shal

l to

Berg

evin

, who

pas

sed

it on

to L

indb

erg

who

then

de

scrib

ed it

(192

7, p

.92-

94) a

s Eus

celis

curt

iceps

.

Eusce

lis cu

rtice

ps L

indb

erg,

192

7D

elto

ceph

alin

aeK

hart

oum

, Ang

lo-

Egyp

tian

Suda

nBe

quae

rt (1

926,

p. 1

89) (

as A

thys

anus

), ci

ting

Berg

evin

(192

5).

Eusce

lis cu

rtice

ps L

indb

erg,

192

7D

elto

ceph

alin

aeK

hart

oum

(Gou

ver-

nem

ent d

u So

udan

)Li

ndbe

rg (1

927c

, p. 9

2), d

escr

ibin

g sp

ecim

ens s

ent t

o hi

m b

y Be

rgev

in.

Eusce

lis cu

rtice

ps L

indb

erg,

192

7D

elto

ceph

alin

aeSu

dan

Mye

rs (1

929,

p.4

75),

citin

g Li

ndbe

rg (1

927)

.Eu

scelis

curt

iceps

Lin

dber

g, 1

927

Del

toce

phal

inae

Suda

nC

hina

(192

9, p

.97)

, citi

ng B

erge

vin

(192

5a),

in le

tter r

ead

by

Poul

ton

at 1

6 O

ct. 1

929

mee

ting

of th

e En

tom

olog

ical

Soc

iety

of

Lond

on.

Eusce

lis cu

rtice

ps L

indb

erg,

192

7D

elto

ceph

alin

aeK

hart

oum

(Sud

an)

Fole

y &

Aud

ouze

(193

8, p

.526

) (as

Ath

ysan

us),

men

tioni

ng th

at

Mar

shal

l sen

t 5 u

ndes

crib

ed sp

ecie

s of b

lood

-suc

king

“Ja

ssid

ae”

to

Berg

evin

in 1

925.

Eusce

lis cu

rtice

ps L

indb

erg,

192

7D

elto

ceph

alin

aeSu

dan

Ryck

man

& B

entle

y (1

979,

p.2

8), c

iting

Lin

dber

g (1

927)

.Eu

scelis

curt

iceps

Lin

dber

g, 1

927

Del

toce

phal

inae

Kha

rtou

m (S

udan

)Le

wis

(195

8, p

.45)

(miss

pelle

d as

curt

ipes)

, citi

ng L

indb

erg

(192

7).

Eusce

lis cu

rtice

ps L

indb

erg,

192

7D

elto

ceph

alin

aeSu

dan

Ryck

man

& B

entle

y (1

979,

p.2

8), c

iting

Lew

is (1

958)

.Eu

scelis

curt

iceps

Lin

dber

g, 1

927

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

3), c

iting

Mye

rs (1

929)

.Eu

scelis

curt

iceps

Lin

dber

g, 1

927

Del

toce

phal

inae

Nor

th A

frica

Alex

ande

r (19

84, p

.58)

(miss

pelle

d as

Ath

ysau

rus s

p.),

citin

g Be

rgev

in (1

925)

.

Acon

urell

a ne

ar p

rolix

a (L

ethi

erry

, 188

5)D

elto

ceph

alin

aeK

hart

oum

(Sou

dan

Egyp

tien)

Berg

evin

(192

5a, p

p. 3

9-40

) (as

Tham

note

ttix,

miss

pelle

d as

Th

amot

ettiy

), re

gard

ing

spec

imen

s col

lect

ed b

y H

.B. J

ohns

ton,

17

Oct

. 192

4, a

nd se

nt b

y M

arsh

all t

o Be

rgev

in, w

ho p

asse

d th

em o

n to

Lin

dber

g w

ho th

en d

escr

ibed

them

(192

7, p

.88-

90) a

s 5

spec

imen

s of Th

amno

tetti

x sa

ngui

suga

. Lat

er, M

yers

(192

9, p

.474

) re

port

ed th

at a

ccor

ding

to W

. E. C

hina

, the

y co

rrec

tly b

elon

g to

Ac

onur

a.

Tabl

e 1.

(C

ont.)

(Con

tinue

d )


Acon

urell

a ne

ar p

rolix

a (L

ethi

erry

, 188

5)D

elto

ceph

alin

aeK

hart

oum

, Ang

lo-

Egyp

tian

Suda

nBe

quae

rt (1

926,

p. 1

89) (

as Th

amno

tetti

x), c

iting

Ber

gevi

n (1

925)

.

Acon

urell

a ne

ar p

rolix

a (L

ethi

erry

, 188

5)D

elto

ceph

alin

aeK

hart

oum

(Gou

ver-

nem

ent d

u So

udan

)Li

ndbe

rg (1

927c

, p. 8

8), d

escr

ibin

g (a

s Tham

note

ttix

sang

uisu

ga)

spec

imen

s sen

t to

him

by

Berg

evin

. Lat

er, W

. E. C

hina

said

they

co

rrec

tly b

elon

g to

Aco

nura

.Ac

onur

ella

near

pro

lixa

(Let

hier

ry, 1

885)

Del

toce

phal

inae

Suda

nM

yers

(192

9, p

.474

) (as

Tham

note

ttix

sang

uisu

ga),

citin

g Li

ndbe

rg

(192

7), w

ho d

escr

ibed

it a

s a n

ew sp

ecie

s, Th

amno

tetti

x sa

ngui

suga

, bu

t in

foot

note

, Mye

rs sa

ys W

.E. C

hina

ass

ured

him

it b

elon

gs to

Ac

onur

a ve

ry n

ear A

. pro

lixa.

Acon

urell

a ne

ar p

rolix

a (L

ethi

erry

, 188

5)D

elto

ceph

alin

aeSu

dan

Chi

na (1

929,

p.9

7) (a

s Tham

note

ttix

sang

uisu

ga),

citin

g Be

rgev

in

(192

5a),

in le

tter r

ead

by E

.B. P

oulto

n at

16

Oct

ober

192

9 m

eetin

g of

the

Ento

mol

gica

l Soc

iety

of L

ondo

n.Ac

onur

ella

near

pro

lixa

(Let

hier

ry, 1

885)

Del

toce

phal

inae

Mar

tini (

1929

, p.3

85) (

as Th

amno

tetti

x), s

ourc

e no

t giv

en, b

ut

base

d on

the

mat

chin

g lis

t of g

ener

a, p

resu

mab

ly o

rigin

atin

g fro

m

Berg

evin

(192

5a).

Acon

urell

a ne

ar p

rolix

a (L

ethi

erry

, 188

5)D

elto

ceph

alin

aeK

hart

oum

(Sud

an)

Fole

y &

Aud

ouze

(193

8, p

.526

) (as

Ath

ysan

us),

men

tioni

ng th

at

Mar

shal

l sen

t 5 u

ndes

crib

ed sp

ecie

s of b

lood

-suc

king

“Ja

ssid

ae”

to

Berg

evin

in 1

925.

Acon

urell

a ne

ar p

rolix

a (L

ethi

erry

, 188

5)D

elto

ceph

alin

aeBr

umpt

(194

9, p

.127

6) (a

s Tham

note

ttix,

miss

pelle

d as

Ta

mno

tetti

x), c

iting

Ber

gevi

n (1

925)

.Ac

onur

ella

near

pro

lixa

(Let

hier

ry, 1

885)

Del

toce

phal

inae

Kha

rtou

m (S

udan

)Le

wis

(195

8, p

.45)

(as A

conu

ra n

r. pr

olix

a: a

s “a

bug

whi

ch m

ay b

e an

Aco

nura

”), c

iting

Lin

dber

g (1

927)

and

Mye

rs (1

929)

.Ac

onur

ella

near

pro

lixa

(Let

hier

ry, 1

885)

Del

toce

phal

inae

Suda

nRy

ckm

an &

Ben

tley

(197

9, p

.28)

(as Th

amno

tetti

x sa

ngui

suga

), ci

ting

Lind

berg

(192

7).

Acon

urell

a ne

ar p

rolix

a (L

ethi

erry

, 188

5)D

elto

ceph

alin

aeRy

ckm

an &

Ben

tley

(197

9, p

.33)

(as Th

amno

tetti

x sa

ngui

suga

), ci

ting

Mye

rs (1

929)

.Ac

onur

ella

near

pro

lixa

(Let

hier

ry, 1

885)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

1) (a

s Tham

note

ttix)

, citi

ng M

artin

i (1

929)

.

(Con

tinue

d )

Tabl

e 1.

(C

ont.)


Oro

sius c

ellul

osus

(Lin

dber

g, 1

927)

Del

toce

phal

inae

Kha

rtou

m (S

ouda

n Eg

yptie

n)Be

rgev

in (1

925a

, pp.

39-

40) (

as D

eltoc

epha

lus),

rega

rdin

g sp

eci-

men

s col

lect

ed b

y H

.B. J

ohns

ton,

17

Oct

. 192

4, a

nd se

nt b

y M

arsh

all t

o Be

rgev

in, w

ho p

asse

d th

em o

n to

Lin

dber

g, w

ho

desc

ribed

them

(192

7, p

.90-

92) a

s thr

ee sp

ecim

ens o

f Tham

note

ttix

cellu

losu

s (m

isspe

lled

as ce

llulo

sa).

Oro

sius c

ellul

osus

(Lin

dber

g, 1

927)

Del

toce

phal

inae

Kha

rtou

m, A

nglo

- Eg

yptia

n Su

dan

Bequ

aert

(192

6, p

. 189

) (as

Delt

ocep

halu

s), c

iting

Ber

gevi

n (1

925)

.

Oro

sius c

ellul

osus

(Lin

dber

g, 1

927)

Del

toce

phal

inae

Kha

rtou

m (G

ouve

r-ne

men

t du

Soud

an)

Lind

berg

(192

7c, p

. 90)

, des

crib

ing

(as Th

amno

tetti

x ce

llulo

sa) a

sp

ecim

en se

nt to

him

by

Berg

evin

.O

rosiu

s cell

ulos

us (L

indb

erg,

192

7)D

elto

ceph

alin

aeSu

dan

Mye

rs (1

929,

p.4

75) (

as Th

amno

tetti

x ce

llulo

sa),

citin

g Li

ndbe

rg

(192

7).

Oro

sius c

ellul

osus

(Lin

dber

g, 1

927)

Del

toce

phal

inae

Suda

nC

hina

(192

9, p

.97)

(as Th

amno

tetti

x ce

llulo

sa),

citin

g Be

rgev

in

(192

5a),

in le

tter r

ead

by E

.B. P

oulto

n at

16

Oct

. 192

9 m

eetin

g of

th

e En

tom

olog

ical

Soc

iety

of L

ondo

n.O

rosiu

s cell

ulos

us (L

indb

erg,

192

7)D

elto

ceph

alin

aeM

artin

i (19

29, p

.385

) (as

Delt

ocep

halu

s), so

urce

not

giv

en, b

ut

base

d on

the

mat

chin

g lis

t of g

ener

a, p

resu

mab

ly o

rigin

atin

g fro

m

Berg

evin

(192

5a).

Oro

sius c

ellul

osus

(Lin

dber

g, 1

927)

Del

toce

phal

inae

Brum

pt (1

949,

p.1

276)

(as D

eltoc

epha

lus),

citi

ng B

erge

vin

(192

5).

Oro

sius c

ellul

osus

(Lin

dber

g, 1

927)

Del

toce

phal

inae

Kha

rtou

m (S

udan

)Le

wis

(195

8, p

.45-

46) (

as Th

amno

tetti

x ce

llulo

sa),

citin

g Li

ndbe

rg

(192

7).

Oro

sius c

ellul

osus

(Lin

dber

g, 1

927)

Del

toce

phal

inae

Suda

nRy

ckm

an &

Ben

tley

(197

9, p

.28)

(as Th

amno

tetti

x ce

llulo

sa),

citin

g Li

ndbe

rg (1

927)

.O

rosiu

s cell

ulos

us (L

indb

erg,

192

7)D

elto

ceph

alin

aeRy

ckm

an &

Ben

tley

(197

9, p

.28)

(as Th

amno

tetti

x ce

llulo

sa),

citin

g Le

wis

(195

8).

Oro

sius c

ellul

osus

(Lin

dber

g, 1

927)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

1) (a

s Delt

ocep

halu

s), c

iting

Mar

tini

(192

9).

Oro

sius c

ellul

osus

(Lin

dber

g, 1

927)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

3) (a

s Tham

note

ttix

cellu

losa

), ci

ting

Mye

rs (1

929)

.

Erat

oneu

ra b

asila

ris (S

ay, 1

825)

Typh

locy

bina

eLa

wso

n (1

926,

p.7

3) (a

s Ery

thro

neur

a ba

silar

is), c

iting

unp

ublis

hed

expe

rienc

e of

Ray

mon

d H

. Bea

mer

.

Tabl

e 1.

(C

ont.)

(Con

tinue

d )


Erat

oneu

ra b

asila

ris (S

ay, 1

825)

Typh

locy

bina

eM

yers

(192

9, p

.475

) (as

Ery

thro

neur

a ba

silar

is), c

iting

Law

son

(192

6).

Erat

oneu

ra b

asila

ris (S

ay, 1

825)

Typh

locy

bina

eR

iley

& Jo

hann

sen

(193

8, p

.143

) (Er

ythr

oneu

ra b

asila

ris),

citin

g La

wso

n (1

926)

.Er

aton

eura

bas

ilaris

(Say

, 182

5)Ty

phlo

cybi

nae

Kan

sas [

USA

]Ry

ckm

an &

Ben

tley

(197

9, p

.28)

(as E

ryth

rone

ura

basil

aris)

, citi

ng

Law

son

(192

6).

Erat

oneu

ra b

asila

ris (S

ay, 1

825)

Typh

locy

bina

eRy

ckm

an &

Ben

tley

(197

9, p

.33)

(as E

ryth

rone

ura

basil

aris)

, citi

ng

Mye

rs (1

929)

.Er

aton

eura

bas

ilaris

(Say

, 182

5)Ty

phlo

cybi

nae

Ryck

man

& B

entle

y (1

979,

p.3

8) (a

s Ery

thro

neur

a ba

silar

is), c

iting

R

iley

& Jo

hann

sen

(193

8).

Para

phlep

sius i

rror

atus

(Say

, 183

0)D

elto

ceph

alin

aeLa

wso

n (1

926,

p.7

4) (a

s Phl

epsiu

s irr

orat

us),

citin

g un

publ

ished

ex

perie

nce

of E

.M. B

ecto

n.

Para

phlep

sius i

rror

atus

(Say

, 183

0)D

elto

ceph

alin

aeN

orth

Am

eric

aM

yers

(192

9, p

.475

) (as

Phl

epsiu

s irr

orat

us),

citin

g La

wso

n (1

926)

.Pa

raph

lepsiu

s irr

orat

us (S

ay, 1

830)

Del

toce

phal

inae

Rile

y &

Joha

nnse

n (1

938,

p.1

43) (

as P

hlep

sius i

rror

atus

), ci

ting

Law

son

(192

6).

Para

phlep

sius i

rror

atus

(Say

, 183

0)D

elto

ceph

alin

aeK

ansa

s [U

SA]

Ryck

man

& B

entle

y (1

979,

p.2

8) (a

s Phl

epsiu

s irr

orat

us),

citin

g La

wso

n (1

926)

.Pa

raph

lepsiu

s irr

orat

us (S

ay, 1

830)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

3) (a

s Phl

epsiu

s irr

orat

us),

citin

g M

yers

(192

9).

Para

phlep

sius i

rror

atus

(Say

, 183

0)D

elto

ceph

alin

aeRy

ckm

an &

Ben

tley

(197

9, p

.38)

(as P

hlep

sius i

rror

atus

), ci

ting

Rile

y &

Joha

nnse

n (1

938)

.

Endr

ia in

imica

(Say

, 183

0)D

elto

ceph

alin

aeLa

wso

n (1

926,

p.7

4) (a

s Delt

ocep

halu

s ini

micu

s).

Endr

ia in

imica

(Say

, 183

0)D

elto

ceph

alin

aeN

orth

Am

eric

aM

yers

(192

9, p

.475

) (as

Delt

ocep

halu

s ini

micu

s), c

iting

Law

son

(192

6).

Endr

ia in

imica

(Say

, 183

0)D

elto

ceph

alin

aeR

iley

& Jo

hann

sen

(193

8, p

.143

) (as

Delt

ocep

halu

s ini

micu

s),

citin

g La

wso

n (1

926)

.En

dria

inim

ica (S

ay, 1

830)

Del

toce

phal

inae

Kan

sas [

USA

]Ry

ckm

an &

Ben

tley

(197

9, p

.28)

(as D

eltoc

epha

lus i

nim

icus),

ci

ting

Law

son

(192

6).

(Con

tinue

d )

Tabl

e 1.

(C

ont.)


Endr

ia in

imica

(Say

, 183

0)D

elto

ceph

alin

aeRy

ckm

an &

Ben

tley

(197

9, p

.33)

(as D

eltoc

epha

lus i

nim

icus),

ci

ting

Mye

rs (1

929)

.En

dria

inim

ica (S

ay, 1

830)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

8) (a

s Delt

ocep

halu

s ini

micu

s),

citin

g R

iley

& Jo

hann

sen

(193

8).

Dra

ecul

acep

hala

mol

lipes

(Say

, 184

0)C

icad

ellin

aeLa

wso

n (1

926,

p.7

4).

Dra

ecul

acep

hala

mol

lipes

(Say

, 184

0)C

icad

ellin

aeN

orth

Am

eric

aM

yers

(192

9, p

.474

) (m

isspe

lled

as D

racu

laec

epha

la m

ollip

es),

citin

g La

wso

n (1

926)

.D

raec

ulac

epha

la m

ollip

es (S

ay, 1

840)

Cic

adel

linae

Rile

y &

Joha

nnse

n (1

938,

p.1

43),

citin

g La

wso

n (1

926)

.D

raec

ulac

epha

la m

ollip

es (S

ay, 1

840)

Cic

adel

linae

Kan

sas [

USA

]Ry

ckm

an &

Ben

tley

(197

9, p

.28)

, citi

ng L

awso

n (1

926)

.D

raec

ulac

epha

la m

ollip

es (S

ay, 1

840)

Cic

adel

linae

Ryck

man

& B

entle

y (1

979,

p.3

3), c

iting

Mye

rs (1

929)

.D

raec

ulac

epha

la m

ollip

es (S

ay, 1

840)

Cic

adel

linae

Ryck

man

& B

entle

y (1

979,

p.3

8), c

iting

Rile

y &

Joha

nnse

n (1

938)

.

Exiti

anus

obs

curin

ervi

s (St

ål, 1

859)

Del

toce

phal

inae

Law

son

(192

6, p

.74)

(as E

usce

lis o

bscu

riner

vis).

Exiti

anus

obs

curin

ervi

s (St

ål, 1

859)

Del

toce

phal

inae

Nor

th A

mer

ica

Mye

rs (1

929,

p.4

75) (

as E

usce

lis o

bscu

riner

vis),

citi

ng L

awso

n (1

926)

.Ex

itian

us o

bscu

riner

vis (

Stål

, 185

9)D

elto

ceph

alin

aeR

iley

& Jo

hann

sen

(193

8, p

.143

) (as

Eus

celis

obs

curin

ervi

s), c

iting

La

wso

n (1

926)

.Ex

itian

us o

bscu

riner

vis (

Stål

, 185

9)D

elto

ceph

alin

aeK

ansa

s [U

SA]

Ryck

man

& B

entle

y (1

979,

p.2

8) (E

usce

lis o

bscu

riner

vis),

citi

ng

Law

son

(192

6).

Exiti

anus

obs

curin

ervi

s (St

ål, 1

859)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

3) (a

s Eus

celis

obs

curin

ervi

s), c

iting

M

yers

(192

9).

Exiti

anus

obs

curin

ervi

s (St

ål, 1

859)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

8) (a

s Eus

celis

obs

curin

ervi

s), c

iting

R

iley

& Jo

hann

sen

(193

8).

Psam

mot

ettix

livi

dellu

s (Ze

tters

tedt

, 18

40) o

r nea

rD

elto

ceph

alin

aeH

such

owfu

, K

iang

su, C

hina

Law

son

(192

6, p

.74)

(as D

eltoc

epha

lus s

triat

us o

r nea

r), c

iting

co

rres

pond

ence

from

Mar

shal

Her

tig.

Psam

mot

ettix

livi

dellu

s (Ze

tters

tedt

, 18

40) o

r nea

rD

elto

ceph

alin

aeK

iang

su, C

hina

Mye

rs (1

929,

p.4

75) (

as D

eltoc

epha

lus),

citi

ng L

awso

n (1

926)

.

Tabl

e 1.

(C

ont.)

(Con

tinue

d )


Psam

mot

ettix

livi

dellu

s (Ze

tters

tedt

, 18

40) o

r nea

rD

elto

ceph

alin

aeK

iang

su, C

hina

Ryck

man

& B

entle

y (1

979,

p.2

8) (a

s Delt

ocep

halu

s), c

iting

Law

son

(192

6).

Psam

mot

ettix

livi

dellu

s (Ze

tters

tedt

, 18

40) o

r nea

rD

elto

ceph

alin

aeRy

ckm

an &

Ben

tley

(197

9, p

.33)

(as D

eltoc

epha

lus),

citi

ng M

yers

(1

929)

.

Exiti

anus

exiti

osus

(Uhl

er, 1

880)

Del

toce

phal

inae

St. L

ouis,

Miss

ouri

[USA

]C

rosb

y (1

926,

p.8

12) (

as E

usce

lis ex

itios

us),

dete

rmin

ed b

y W

.L.

McA

tee.

Exiti

anus

exiti

osus

(Uhl

er, 1

880)

Del

toce

phal

inae

Nor

th A

mer

ica

Mye

rs (1

929,

p.4

75) (

as E

usce

lis ex

itios

us),

citin

g C

rosb

y (1

926)

.Ex

itian

us ex

itios

us (U

hler

, 188

0)D

elto

ceph

alin

aeSt

. Lou

is, M

issou

ri [U

SA]

Ryck

man

(197

9, p

.15)

(as E

usce

lis ex

itios

us),

citin

g C

rosb

y (1

926)

.

Exiti

anus

exiti

osus

(Uhl

er, 1

880)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.3

3) (a

s Eus

celis

exiti

osus

), ci

ting

Mye

rs (1

929)

.Ex

itian

us ex

itios

us (U

hler

, 188

0)D

elto

ceph

alin

aeU

SAAl

exan

der (

1984

, p.5

8) (a

s Eus

celis

exiti

osus

), ci

ting

Cro

sby

(192

6).

Empo

asca

faba

e (H

arris

, 184

1), a

false

re

cord

Typh

locy

bina

eRe

adio

(192

7) p

ublis

hed

“Stu

dies

on

the

biol

ogy

of th

e Re

duvi

idae

of

Am

eric

a N

orth

of M

exic

o,”

but a

s far

as w

e ca

n de

term

ine

didn

’t m

entio

n C

icad

ellid

ae.

Empo

asca

faba

e (H

arris

, 184

1), a

false

re

cord

Typh

locy

bina

eRy

ckm

an &

Ben

tley

(197

9, p

.36)

(as E

mpo

asca

mal

i), c

iting

Re

adio

(192

7) in

err

or; R

eadi

o ne

ver p

ublis

hed

on b

iting

ci

cade

llids

.Em

poas

ca fa

bae (

Har

ris, 1

841)

, a fa

lse

reco

rdTy

phlo

cybi

nae

Alex

ande

r (19

84, p

.58)

(as E

mpo

asca

mal

i), c

iting

Rea

dio

(192

7),

a fa

lse re

cord

per

haps

bas

ed o

n m

isint

erpr

etat

on o

f con

fusin

g te

xt

in R

yckm

an &

Ben

tley

(197

9, p

. 36)

.

Hor

tens

ia si

mili

s (W

alke

r, 18

51)

Cic

adel

linae

Trin

idad

Mye

rs (1

929,

p.4

74) (

as K

olla

sim

ilis)

repo

rtin

g co

mm

unic

atio

n w

ith H

.A. B

allo

u.

Hor

tens

ia si

mili

s (W

alke

r, 18

51)

Cic

adel

linae

Ryck

man

& B

entle

y (1

979,

p.3

3) (a

s Kol

la si

mili

s), c

iting

Mye

rs

(192

9).

(Con

tinue

d )

Tabl

e 1.

(C

ont.)


Typh

locy

ba q

uerc

us (F

abric

ius,

1777

), bu

t M

yers

had

dou

bts a

bout

the

iden

tifica

tion

Typh

locy

bina

eJa

pan

Mye

rs (1

929,

p.4

75) [

He

had

doub

ts ab

out t

he id

entifi

catio

n],

repo

rtin

g un

publ

ished

obs

erva

tion

by T

. Esa

ki.

Typh

locy

ba q

uerc

us (F

abric

ius,

1777

)Ty

phlo

cybi

nae

Ryck

man

& B

entle

y (1

979,

p.3

3), c

iting

Mye

rs (1

929)

, but

Mye

rs

had

doub

ts ab

out t

he id

entifi

catio

n.

Erat

oneu

ra m

acul

ata

(Gill

ette

, 189

8)Ty

phlo

cybi

nae

Japa

nM

yers

(192

9, p

.475

) [as

Ery

thro

neur

a (Z

ygin

a) m

acul

ata?

], re

port

ing

unpu

blish

ed o

bser

vatio

n by

T. E

saki

.

Erat

oneu

ra m

acul

ata

(Gill

ette

, 189

8)Ty

phlo

cybi

nae

Ryck

man

& B

entle

y (1

979,

p.3

3) (a

s Ery

thro

neur

a m

acul

ata)

, ci

ting

Mye

rs (1

929)

.

Empo

asca

?Ty

phlo

cybi

nae

no lo

calit

y gi

ven,

bu

t lik

ely

to h

ave

been

Cin

nam

ara,

As

sam

, Ind

ia

Andr

ews (

1929

, p.9

6), p

erso

nal e

xper

ienc

e re

port

ed in

lette

r rea

d by

E.B

. Pou

lton

at 1

6 O

ct. 1

929

mee

ting

of th

e En

tom

olog

ical

So

ciet

y of

Lon

don.

Kyb

os sm

arag

dula

(Dla

bola

, 195

8c:5

1)Ty

phlo

cybi

nae

Engl

and

Mye

rs (1

929,

p.4

75 &

477

), re

port

ing

the

as y

et u

npub

lishe

d ex

perie

nce

of W

.E. C

hina

.

Kyb

os sm

arag

dula

(Dla

bola

, 195

8c:5

1)Ty

phlo

cybi

nae

Esth

er, S

urre

y [E

ngla

nd]

Chi

na (1

929,

p.9

6-97

) (as

Em

poas

ca sm

arag

dula

), hi

s per

sona

l ex

perie

nce

repo

rted

in n

ote

read

by

E.B.

Pou

lton

at 1

6 O

ct. 1

929

mee

ting

of th

e En

tom

olog

ical

Soc

iety

of L

ondo

n, w

here

in h

e m

entio

ns th

at “

Mye

rs h

as re

cent

ly p

ublis

hed

a no

te o

n th

is su

bjec

t, bu

t I h

ave

had

no ti

me

to tr

ace

it.”

Kyb

os sm

arag

dula

(Dla

bola

, 195

8c:5

1)Ty

phlo

cybi

nae

Ryck

man

& B

entle

y (1

979,

p.3

3), c

iting

Mye

rs (1

929)

.

Neo

alitu

rus t

enell

us (E

mel

jano

v, 19

99c:

56)

Del

toce

phal

inae

Uni

vers

ity o

f C

alifo

rnia

[USA

]U

singe

r (19

34, p

.97)

(as E

utet

tix te

nellu

s), re

port

ing

unpu

blish

ed

expe

rienc

e of

Juliu

s Fre

itag.

Tabl

e 1.

(C

ont.)

(Con

tinue

d )


Neo

alitu

rus t

enell

us (E

mel

jano

v, 19

99c:

56)

Del

toce

phal

inae

Her

ms (

1950

, p.5

74) (

as E

utet

tix te

nellu

s), c

iting

Usin

ger (

1934

).

Neo

alitu

rus t

enell

us (E

mel

jano

v, 19

99c:

56)

Del

toce

phal

inae

Ryck

man

& B

entle

y (1

979,

p.4

2) (a

s Eut

ettix

tene

llus),

citi

ng

Usin

ger (

1934

).N

eoal

ituru

s ten

ellus

(Em

elja

nov,

1999

c:56

)D

elto

ceph

alin

aeC

alifo

rnia

[USA

]Al

exan

der (

1984

, p.5

8) (a

s Eut

ettix

tene

llus),

citi

ng U

singe

r (19

34).

Eryt

hron

eura

com

es (S

ay, 1

825)

Typh

locy

bina

eFr

esno

, Cal

iforn

ia

[USA

]U

singe

r (19

34, p

.97-

98),

repo

rtin

g un

publ

ished

exp

erie

nces

of

Cal

vert

E. N

orla

nd a

nd G

ordo

n L.

Sm

ith.

Eryt

hron

eura

com

es (S

ay, 1

825)

Typh

locy

bina

eH

erm

s (19

50, p

.574

), ci

ting

Usin

ger (

1934

).Er

ythr

oneu

ra co

mes

(Say

, 182

5)Ty

phlo

cybi

nae

Ryck

man

& B

entle

y (1

979,

p.4

2) (a

s Ery

thro

neur

a co

mes

), ci

ting

Usin

ger (

1934

).Er

ythr

oneu

ra co

mes

(Say

, 182

5)Ty

phlo

cybi

nae

Cal

iforn

ia [U

SA]

Alex

ande

r (19

84, p

.58)

(miss

pelle

d as

Ery

thro

nem

a co

mes)

, citi

ng

Usin

ger (

1984

).

Exiti

anus

fasci

olat

us (M

elic

har,

1911

)D

elto

ceph

alin

aeIn

Sal

ah [s

outh

ern

Alge

ria]

Fole

y &

Aud

ouze

(193

8, p

.526

) (as

Ath

ysan

us v

ulne

rans

), re

port

ing

obse

rvat

ions

by

Fonq

ue in

Sep

t. 19

24.

Exiti

anus

fasci

olat

us (M

elic

har,

1911

)D

elto

ceph

alin

aesu

d al

gerie

nBr

umpt

(194

9, p

.127

6) (a

s Ath

ysan

us v

ulne

rans

), re

port

ing

obse

rvat

ion

mad

e by

Dr.

Fouq

ue (m

isspe

lled

as F

onqu

e).

Neo

alitu

rus t

enell

us (E

mel

jano

v, 19

99c:

56)

Del

toce

phal

inae

Loga

n, U

tah

[USA

]K

now

lton

(195

1, p

.112

) (as

Circ

ulife

r ten

ellus

), ob

serv

atio

ns b

y hi

m a

nd M

.F. B

owen

.

Eryt

hron

eura

Typh

locy

bina

eW

ad M

edan

i [Su

dan]

Lew

is (1

958,

p.4

5), a

ppar

ently

an

inde

pend

ent r

epor

t.

Eryt

hron

eura

Typh

locy

bina

eSu

dan

Ryck

man

& B

entle

y (1

979,

p.2

8), c

iting

Lew

is (1

958)

.

Delt

ocep

halu

sD

elto

ceph

alin

aeW

ad M

edan

i [S

udan

]Le

wis

(195

8, p

.45)

, app

aren

tly a

n in

depe

nden

t rep

ort.

Delt

ocep

halu

sD

elto

ceph

alin

aeSu

dan

Ryck

man

& B

entle

y (1

979,

p.2

8), c

iting

Lew

is (1

958)

.

(Con

tinue

d )

Tabl

e 1.

(C

ont.)


Exiti

anus

fasci

olat

us (M

elic

har,

1911

)D

elto

ceph

alin

aeK

hart

oum

[Sud

an]

Lew

is (1

958,

p.4

5) (a

s E. v

ulne

rans

Ber

gevi

n). Th

ere

is co

nfus

ion

here

: the

gen

us n

ame

is no

t spe

lled

out a

nd th

e pr

eced

ing

genu

s is

Eusce

lis, a

lthou

gh v

ulne

rans

Ber

gevi

n w

as d

escr

ibed

in A

thys

anus

. Re

gard

less

, app

aren

tly th

is is

an in

depe

nden

t rep

ort b

ecau

se a

ll pr

evio

us re

cord

s are

for S

alah

.

Exiti

anus

fasci

olat

us (M

elic

har,

1911

)D

elto

ceph

alin

aeSu

dan

Ryck

man

& B

entle

y (1

979,

p.2

8) (a

s E. v

ulne

rans

), ci

ting

Lew

is (1

958)

.

Exiti

anus

?D

elto

ceph

alin

aeW

ad M

edan

i [S

udan

]Le

wis

(195

8, p

.45)

, app

aren

tly a

n in

depe

nden

t rep

ort.

Tham

note

ttix

2 sp

p.D

elto

ceph

alin

aeW

ad M

edan

i [S

udan

]Le

wis

(195

8, p

.46)

, app

aren

tly a

n in

depe

nden

t rep

ort.

Empo

asca

dec

ipien

s (Pa

oli,

1930

) or

Empo

asca

sola

ni (C

urtis

, 184

6)Ty

phlo

cybi

nae

Oxf

ord

[Eng

land

]M

cCra

e (1

974,

238

). Sp

ecim

en d

eter

min

ed b

y W

. J. L

e Q

uesn

e as

as

a fe

mal

e of

eith

er E

mpo

asca

dec

ipien

s or E

. pte

ridis,

the

fem

ales

of

whi

ch “c

anno

t be

diffe

rent

iate

d.”

Hor

tens

ia si

mili

s (W

alke

r, 18

51)

Cic

adel

linae

Cub

aN

ovoa

& A

layo

(198

6, p

.80)

.

Hor

tens

ia si

mili

s (W

alke

r, 18

51)

Cic

adel

linae

Repu

blic

of P

anam

aG

uerr

a V.

(200

2, p

.38-

39),

unpu

blish

ed th

esis.

Hom

alod

isca

vitri

penn

is G

erm

ar, 1

821

Cic

adel

linae

Fren

ch P

olyn

esia

Gra

ndgi

rard

et a

l. (2

006,

p.4

35) (

as H

omal

odisc

a co

agul

ata)

.

Tabl

e 1.

(C

ont.)


we surmise that only 37 are original, and the rest are citation upon citation of those 37. Not included in the 174 are three false records that were based upon a misreading of Readio (1927).

We add 19 new records, all from Panama, representing 13 genera, and at least 13 species to the list of Cicadellidae known to have attacked humans (Table 2). These new records represent five subfamilies, two of which, Gyponinae and Xestocephalinae, have not been reported previously. Eleven are first records for their genera: Dilobopterus, Tylozygus, Balclutha, Chlorotettix, Hecalapona, Planicephalus, Plesiommata, Portanus, Xestocephalus, Xyphon, and Zyginama. Two attacks were to the back of the neck, but the other 17 were to the extremities, mostly the forearm, and in each case caused itching or mild stinging lasting several seconds to several minutes.

Fifteen of the attacks took place at night or at dusk, usually on people reading or watching television. One of the four daytime attacks was by an individual of Dilobopterus stolli that alighted on EDN’s leg and probed repeatedly for some three minutes but did not pierce the skin. It was then captured in a vial and placed on the leg of EDN’s wife (LV). The insect immediately pierced her skin, causing pain and itching. A second incident involved a lab researcher who was attacked upon reaching into a brightly lighted chamber containing his study subjects, Hortensia similis (Guerra, 2002). A third daytime attacker was a typhlocybine that alighted on EDN while he was wait-ing at a bus stop. The fourth daytime attack took place in a patio. A fifth daytime attack is not included in our data because the specific insect individual was not cap-tured; it was to a person (LV) who was watering a garden, and thus disturbed a large number of Dilobopterus stolli on their plant, Pseuderanthemum reticulatum (Hooker f.) Radlkofer (Acanthaceae). Including our own contributions, we calculate that now there are 193 published records of Cicadellidae having bitten humans, of which 56 are the original observations, and that they represent 30 genera and at least 39 species in five subfamilies.

In addition to our experiences, Pierre Jolivet has been probed by various species near rice fields and the Awash River in Ethiopia; and by Nephotettix, probably virescens, dur-ing a typhoon in Hong-Kong (personal communication). As well, William Eberhard (in the Neotropics) has been probed occasionally by leafhoppers (personal communi-cation). Neither of them captured their attackers.

Discussion

Several authors doubted that leafhoppers showed interest in humans until they experi-enced it themselves. Donovan (1920a, 1920b) commented “Several people used to complain to me of being stung by these insects, but, not being cognizant at the time of such habits among the Homoptera, I discountenanced these tales of the presumed aggressors. It was only when I was myself a victim of the insects’ action that I was disillusioned.” Lawson (1920) stated “The writer has had several people tell him about being bitten by little green leafhoppers, but not till a short time ago did he have any personal proof of the fact. One night, while collecting under a light, he felt a little prick on his hand, and on looking down saw a little green leafhopper [Empoasca fabae

218 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225Ta

ble

2. C

icad

ellid

ae th

at h

ave

prob

ed h

uman

skin

in th

e Re

publ

ic o

f Pan

ama

Nam

eSu

bfam

ilyG

ende

rD

ay/n

ight

Dat

eLo

calit

y, vi

ctim

, and

par

t of b

ody

atta

cked

Dep

ositi

on

Dilo

bopt

erus

stol

li Si

gnor

et, 1

850

Cic

adel

linae

mal

eD

2011

Nov

. 13

Pana

má:

San

Isid

ro, E

l Val

le, i

n pa

tio: p

robe

d ED

N’s

leg

with

no

succ

ess,

but p

robe

d Li

nette

Veg

a’s (L

V) l

eg

afte

r bei

ng tr

ansfe

rred

ther

e

MIU

P

Hor

tens

ia si

mili

s (W

alke

r, 18

51)

Cic

adel

linae

fem

ale

N20

09 D

ec. 2

3Pa

nam

á: S

an Is

idro

, El V

alle

, in

livin

g ro

om: p

robe

d LV

’s fo

rear

mM

IUP

Hor

tens

ia si

mili

s (W

alke

r, 18

51)

Cic

adel

linae

fem

ale

D20

01Pa

nam

á: U

nive

rsid

ad d

e Pa

nam

á, in

Lab

. de

Estu

dios

Bi

ológ

icos

de

Plag

as A

gríc

olas

, in

lab:

“atta

cked

” Is

mae

l Gue

rra

[Unp

ublis

hed

thes

is]

MIU

P

Tylo

zygu

s geo

met

ricus

(S

igno

ret,

1854

)C

icad

ellin

aefe

mal

eN

2008

Oct

. 23

Pana

má:

Arr

aijá

n, L

oma

del R

ío, 8

.940

7 -7

9.65

68, i

n liv

ing

room

: pro

bed

AA’s

fore

arm

STR

I

Xyph

on re

ticul

atum

(S

igno

ret 1

854)

Cic

adel

linae

mal

edu

sk20

12 F

eb. 4

Pana

má:

Can

al A

rea,

nea

r Can

al A

dmin

istra

tion

Build

ing:

pro

bed

LV’s

fore

arm

MIU

P

Plesi

omm

ata

sp.

Cic

adel

linae

mal

edu

sk20

12 N

ov. 1

5Pa

nam

á: C

hiriq

uí, D

avid

, Dol

egui

ta, 8

th S

treet

: pr

obed

Indi

ra S

imon

’s le

gST

RI

Plesi

omm

ata

sp.

Cic

adel

linae

fem

ale

N20

12 N

ov. 2

9Pa

nam

á: A

rrai

ján,

Pue

rta

de H

ierr

o: p

robe

d Je

rald

ín

Verg

ara’s

left

ring

finge

rM

IUP

Balcl

utha

sp.

Del

toce

phal

inae

fem

ale

N20

09 O

ct. 2

6Pa

nam

á: S

an Is

idro

, El V

alle

, in

livin

g ro

om: p

robe

d LV

’s fo

rear

mM

IUP

Chl

orot

ettix

min

imus

Ba

ker,

1898

Del

toce

phal

inae

mal

eN

2012

Mar

. 14

Pana

má:

Arr

aijá

n, L

oma

del R

ío, 8

.940

7 -7

9.65

68,

unde

r rea

ding

lam

p: p

robe

d th

e ba

ck o

f AA’

s han

d (0

2: 5

2 H

)

STR

I

Plan

iceph

alus

flav

icosta

(S

tål,

1862

)D

elto

ceph

alin

aem

ale

N20

12 M

ar. 5

Pana

má:

San

Isid

ro, E

l Val

le, i

n liv

ing

room

: pro

bed

LV’s

fore

arm

MIU

P

Hec

alap

ona

brev

isens

D

eLon

g an

d Fr

eyta

g,

1975

Gyp

onin

aefe

mal

eN

2008

Aug

. 19

Pana

má:

Arr

aijá

n, L

oma

del R

ío, 8

.940

7 -7

9.65

68,

unde

r rea

ding

lam

p: p

robe

d AA

’s fo

rear

mST

RI

Sola

nsac

a ste

vens

i (G

haur

i, 19

74a)

Typh

locy

bina

em

ale

D20

09 F

eb. 2

5Pa

nam

á: S

anta

Lib

rada

, Pol

iclin

ica,

at b

us st

op:

prob

ed E

DN

’s fo

rear

mM

IUP

(Con

tinue

d )


Sola

nasca

stev

ensi

(Gha

uri,

1974

a)Ty

phlo

cybi

nae

dam

-ag

edN

2009

June

2Pa

nam

á: S

an Is

idro

, El V

alle

, in

livin

g ro

om: p

robe

d th

e ba

ck o

f ED

N’s

neck

MIU

P

Sola

nasca

stev

ensi

(Gha

uri,

1974

a)Ty

phlo

cybi

nae

mal

eD

2012

May

3Pa

nam

á: S

an Is

idro

, El V

alle

, in

livin

g ro

om: p

robe

d th

e ba

ck o

f LV

’s ha

ndM

IUP

Zygi

nam

a sp

.Ty

phlo

cybi

nae

fem

ale

N20

07 A

ug.

Pana

má:

San

Isid

ro, E

l Val

le, i

n liv

ing

room

: pro

bed

LV’s

fore

arm

MIU

P

Uni

dent

ifiab

le to

ge

nus,

but n

one

of th

e ab

ove.

Typh

locy

bina

efe

mal

eN

2009

Oct

. 26

Coc

lé: A

ntón

, El V

alle

, in

hote

l res

taur

ante

: pro

bed

LV’s

fore

arm

MIU

P

Xesto

ceph

alus

dese

rto-

rum

(Ber

g)X

esto

ceph

alin

aefe

mal

eN

2011

Feb

. 4Pa

nam

á: A

rrai

ján,

Lom

a de

l Río

, 8.9

407

-79.

6568

, w

hile

wat

chin

g T

V: p

robe

d AA

’s ne

ck (2

2: 1

5 H

)ST

RI

Xesto

ceph

alus

dese

rto-

rum

(Ber

g)X

esto

ceph

alin

aem

ale

N20

13 Ja

n. 7

Pana

má:

Arr

aijá

n, L

oma

del R

ío, 8

.940

7 -7

9.65

68,

whi

le w

orki

ng a

t a c

ompu

ter:

prob

ed A

A’s l

eft w

rist

(22:

14

H)

STR

I

Port

anus

sp.

Xes

toce

phal

inae

fem

ale

N20

12 A

ug. 2

5Pa

nam

á: V

illa

de la

s Fue

ntes

Nº2

, ady

acen

te a

l Río

C

urun

dú, i

n be

droo

m: p

robe

d C

arm

en G

alda

mes

’ fo

rear

m

MIU

P

Tabl

e 2.

(C

ont.)


(Harris, 1841)] at work.” Lawson later (1926) published his personal encounters with leafhoppers belonging to three additional species, and reported correspondence from Marshal Hertig, from Hsuchowfu, Kiangsu, China who wrote “Several times I thought I felt a pricking, but believing that Jassids wouldn’t do such a thing, dismissed the idea as imagination. The apparent pricking whenever these insects rested on my arm, how-ever, continued, and so I watched them with my hand lens. In practically every case, (half a dozen or so) after alighting, the proboscis appeared, they braced themselves for the thrust and the prick was felt.”

The fact that leafhoppers do probe human skin is well-documented, however, though a number of writers speak of their cicadellid attackers as “blood suckers,” there is no evidence that the purpose of the probing is to obtain blood, and only Tucker (1911) gives proof that blood actually was obtained—when he crushed a leafhopper that bit him, he produced a bloody smear. In none of our own cases do we know whether the attackers actually obtained blood. Our first priority was to capture and identify the leafhoppers that probed us. To have waited to see whether they engorged would have risked losing the specimens. Perhaps the authors of previous reports made the same decision, or they simply assumed that the insects were seeking blood.

What induces these normally phytophagous insects to seek human fluids? Though no one has answered that question convincingly, we tend to agree with Myers (1929) and Schaefer (2000) who speculate that, in most cases, the insects found themselves in water-stressed situations and, in the absence of plants, sought moisture by probing human skin. Such tiny insects attracted to light and unable to escape from it and the heat it produces might be particularly prone to water-stress. Similarly, Chris Dietrich (personal communication) notes, “Most species spend much of their time feeding, so it’s possible that when they probe human skin they are trying to determine whether they have landed on a suitable host plant.”

Other explanations have been offered including the idea that a virus transmitted by these insects affects their behavior and disorientates them (McCrae, 1974); the possi-bly that the insects are stimulated by substances in solution, such as salt or urea—a speculation based on the observation that often they appear to be attracted to perspira-tion (Lindberg, 1927; Lewis, 1958; Schaefer, 2000); or simple thirst or hunger (Myers, 1929). Rakitov et al. (2005) reviewed the literature on leafhopper aggregations (males in most cases) on moist ground. Their analysis of the excreta produced at a site in Peru indicated “significant sodium retention.” Mammalian blood contains many of the sug-ars, proteins, and salts also found in plants, but it is much richer in nitrogen com-pounds and the protein is rather different from that found in [cell sap] (Usinger, 1934). Whether cicadellids are able to utilize the protein component of mammalian blood is not known, however, Usinger (1934), using mammalian blood serum, carried out experiments on a herbivorous heteropteran, Leptocoris trivittatus (Say) (Rhopalidae ), and learned that at least that plant feeder could survive on it to some degree.

Whatever the explanation for biting behavior in cicadellids, it is common and wide-spread in the group, and one can imagine the phenomenon as an indicator of the potential for dietary diversification within the Homoptera. Though no adventitious biter has been shown to transmit any disease (Schaefer, 2000), the detection of


Filovirus-like particles in a leafhopper (Lundsgaard, 1997) further expands the possi-bilities, as does the intriguing finding that a recently evolved bacterial symbiont harbored by a leafhopper, Euscelidius variegatus (Cicadellidae), is closely related to a bacterial symbiont found in bedbugs (Cimex lectularius), insects whose sole source of nourishment is mammalian blood (Hypsa and Aksoy, 1997; Degnan et al, 2011).

Comments on the literature

The published records of leafhoppers piercing human skin began a century ago with Tucker (1911), who reported having been bitten by Empoasca fabae (as E. mali) in Dallas, Texas, in July of 1909. Two additional cases involving the same species of leaf-hopper occurred elsewhere in the United States: one in Arkansas (Becker, 1918), the other in Kansas (Lawson, 1920).

From then on the literature on biting leafhoppers becomes confusing at times, per-haps thanks to the enthusiasm generated by the novelty of the subject. Authors cited each other frequently, often quite informally as letters read into the minutes (proceed-ings) of society meetings, and they sent specimens back and forth from one museum to another. As a result, in a number of cases several authors published independently, based on the same specimen and biting event. In some cases the leafhoppers were iden-tified only to genus and later described elsewhere, or incorrectly identified and later corrected by someone else, or the source of the record is not given. We have followed the threads of these records to try and determine which are original and which are cita-tions and, assuming that those identifcations were correct, to update the nomencla-ture. An unsolved mystery involves Alfred William Alcock (1859-1933), a British naturalist who worked in India and was Superintendent of the Indian Museum. Though principally a fish specialist he had a broad interest in all things natural. He is cited by Brumpt (1922, 1927, and 1936; and again in 1949, misspelled as Alcok) as the source of a record of Nephotettix virescens (Distant, 1908) (as N. bipunctatus) biting humans in Calcutta, but not in the earlier Brumpt editions (1910, 1913). However, the Alcock publications that we have seen (1900, 1903, 1911, 1920) do not mention biting leafhoppers. In fact, in the insect pests section of the 1900 and 1903 “Indian Museum Notes,” for which he was the editor, leafhoppers are not mentioned at all, and in “Entomology for Medical Officers” (1911, 1920) he states that Homoptera are “of no particular interest to the medical officer.” It seems most likely that Brumpt was referring to a letter or other communication received directly from Alcock between the years 1920 and 1922.

Bergevin (1925a, pp. 40) was a communication hub for observations and speci-mens: e.g., citing personal correspondence from Marshall, he reported that M.E. China found two specimens of Exitianus indicus (Distant, 1908) (as Athysanus indicus) in the British Museum that were collected in Madras, India, by Donovan in November 1919 and annotated as “bloodsuckers.” Unknown to Bergevin, Donovan (1920a, 1920b) already had published his account of the blood-sucking behavior. In another instance, Bergevin (1925a, pp. 39-40), reported that specimens of Eucelis curticeps Lindberg, 1927 (as Athysanus sp.), collected by H.B. Johnston in Khartoum, Sudan on 17 Oct


1924, were sent to him by Marshall, and that he passed them on to Lindberg for description.

Myers (1929) contains a numerical error that caused us confusion when analyzing the list of biting leafhoppers he attributes to various authors. In the second paragraph of his text, he credits eight to Bequaert (1926), instead of three. Another confusion appears to have originated due to a reorganization of his list of published biting inci-dents. He uses l.c. three times: the first time immediately after a citation of Crosby (1926), who never published on the species indicated, and the second and third times after citations of Lawson (1926). At first we thought the three l.c. referred to the three records by Bequaert (1926), who is quoted extensively in the previous section of the paper, but the taxa turned out not to match. Without a doubt the three are citations of Lawson (1926), a conclusion supported by the fact that Lawson is the only other source for the record from Kiangsu, China.

On 16 October 1929, at a meeting of the Entomological Society of London, E. B. Poulton read a letter from E. A. Andrews and a note by W.E. China describing their encounters with biting cicadellids. That information was published on 31 December of the same year in the proceedings of the society as “Jassidae as blood-sucking insects.” However, because Myers (1929) already had published news of the W. E. China incident, shortly before the meeting took place, he is listed as the original source in our Table 1, and Mr. China is listed as having cited his own record.

In Ryckman and Bentley (1979, pp. 36), the entry for Readio (1927) is confusing for lack of a clear break after quoting him, and initially that misled us to believe that Readio reported Empoasca fabae (as E. mali) as biting. In fact Readio wrote only of Heteroptera and did not report biting by any cicadellid. Apparently Alexander (1984) was equally misled and did not consult Readio (1927) directly; he incorrectly credits him for a report of biting by Empoasca fabae (as E. mali).

McCrae (1974) was in error when he stated, “I can find no records of leafhoppers biting or attempting to bite man in Britain, and Dr. W. J. LeQuesne who I have con-sulted also knows of no such instances.” In truth, separate accounts of the experience of W.E. China, in Surrey were published in 1929 by China and by Myers. The claim by Guerra (2002) that his laboratory study subjects, Hortensia similis, attacked him in defense of their eggs has little credibility.

As shown in Table 1, we have attempted to sort out the confusing network of reports of leafhoppers probing human skin. We would welcome additions and corrections to those efforts.

Acknowledgements

Our gratitude goes to Linette Vega, whom leafhoppers find especially attractive; to Angel Aguirre for tirelessly tracking down copies of the literature pertaining to this study; and to Carmen Galdames for not squashing the Portanus that bit her. For assis-tance with identifications we thank Dmitry A. Dmitriev (Illinois Natural History Survey) for Typhlocybinae, Carolina Godoy (Museum of Zoology, School of Biology, University of Costa Rica) for Cicadellinae, and especially Therese A. Catanach (Texas


A&M University) who identified Xyphon reticulatum. Thanks also to Stuart McKamey (Systematic Entomology Laboratory, USDA) and Chris Dietrich (Illinois Natural History Survey) for their comments and advice, and to Pierre Jolivet (Paris) and William Eberhard (STRI, Costa Rica).

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Leafhoppers (Homoptera: Cicadellidae) that probe human skin: A