Table Of ContentPROC. ENTOMOL. SOC. WASH.
108(3), 2006, pp. 611-618
REVIEW OF THE NEOTROPICAL LEAFHOPPER GENUS
CHLOROGONALIA (HEMIPTERA: CICADELLIDAE: CICADELLINAE),
WITH NOTES ON THE GENUS CALDWELLIOLA
STUART H. MCKAMEY
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S.
Department of Agriculture, c/o National Museum of Natural History, Smithsonian
Institution, P.O. Box 37012, MRC 168, Washington, DC 20013-7012, U.S.A. (e-mail:
smckamey(@sel.barc.usda.gov)
Abstract.4Chlorogonalia Young, 1977, and Caldwelliola Young, 1977, are
superficially very similar genera that may not be closely related genera within the
tribe Cicadellini (Hemiptera: Cicadellidae: Cicadellinae). Two species, Chlorogonalia
ultima Young and Caldwelliola reservata (Fowler), are particularly similar
phenotypically and are sympatric in Ecuador (new country record for C. reservata).
New diagnostic features are given for both genera; two of these features can be
assessed in undissected specimens. Caldwelliola tharma (Yo ung) is proposed as a new
combination (from Chlorogonalia). Species of both genera are possible vectors of the
bacterium YXylella fastidiosa in coffee; Caldwelliola caucana has been directly
implicated as a vector of the crespera disease in Colombia. An illustrated key is
included to distinguish the four known species of Chlorogonalia.
Key Words: Cicadellidae, Xylella fastidiosa, vector, crespera, coffee
There are 328 genera and over 2,200 Brazil, causing citrus variegated chloro-
species of sharpshooters (Hemiptera: sis. Most New World sharpshooters were
Cicadellidae: Cicadellinae) in the New included in two revisions by Young
World (Leafhoppers of the World Data- (1968, 1977), the earlier revision covering
base; McKamey, in preparation). The the tribe Proconiini and the more recent
group is unusual in the Cicadellidae for dealing with Cicadellini. Two of the 163
feeding on the host plant9s xylem rather New World genera of Cicadellini, Ch/lor-
than phloem or parenchyma. The xylem ogonalia and Caldwelliola, are extremely
is deficient in nutrients and is therefore similar in both external and internal
imbibed and excreted in large amounts, features but may not even be closely
the latter activity giving these insects the related, their similarities being superfi-
common name of sharpshooters. Some cial. In the process of identifying a puta-
members of this group are also well tive pest species, it was discovered that
known for vectoring the bacter1um Xy- some specimens (USNM) of Caldwelliola
lella fastidiosa. For example, species of had been misidentified by Young as
the genus Homalodisca Stal vector xX. Chlorogonalia, indicating that a clearer
fastidiosa in grape in the United States, diagnosis of the two genera was needed.
causing Pierce9s Disease, and genera such No host records have been reported in
as Acrogonia Stal vector it in citrus in the literature for either, but both genera
612 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
contain species commonly associated
with coffee, which suffers from a Xy/ella
fastidiosa-induced disease.
REVIEW OF LITERATURE
Among the many new _ leafhopper
genera described by Young (1977) were
Chlorogonalia and Caldwelliola, both
replete with newly described species.
Young considered the closest relatives
of Chlorogonalia to be Cicadella Latreille
and Draeculacephala Spinola, whereas he
suggested that Borogonalia Young is the
closest relative to Caldwelliola.
Young (1977) distinguished Chlorogo-
nalia from Caldwelliola based on the
aedeagal paraphyses, the former having
their two arms (paraphyses) symmetrical
in form but asymmetrical in position,
and the latter having the paraphyses
asymmetrical in form and _ position.
Another feature was the rudimetary
abdominal apodemes in Chlorogonalia,
compared to strongly developed apo-
demes in Caldwelliola. Neither feature
can be assessed without dissection.
Nevertheless, Young used some un-
Figs. 1-2. Oblique aspect of anterior portion.
dissected specimens to report species 1, Chlorogonalia ultima. 2, Caldwelliola reservata.
distributions, leading to incomplete
country records for Caldwelliola. In
particular, Young misidentified some
in Ecuador (Fig. 5), lacking the pronotal
Ecuadorian specimens (USNM) of Cald-
welliola reservata (Fowler), new country markings found in the Central American
record, as Chlorogonalia ultima Young. populations (Figs. 6-7). Neither Chloro-
Caldwelliola reservata had been reported gonalia nor Caldwelliola have received
to occur only in Central America. further taxonomic attention.
The most obvious feature of both
MATERIALS AND METHODS
species is a pair of marks on the head,
one dorsally and one anteriorly (Figs. 14 The pygofer and genitalia were pre-
2), which vary intraspecifically in size and pared by placing the abdomen, separated
shape. Unfortunately these are not di- from the body at its base, in a weak
agnostic, being found in other cicadelline solution of potassium hydroxide (KOH)
genera and species4e.g., Macunolla ven- overnight. KOH-treated parts were pre-
tralis (Signoret), which, like most species served in glycerin in polypropylene
of Chlorogonalia and Caldwelliola, has microvials beneath the specimens. Digi-
green wings. Young9s confusion of the tal images were captured at 400 dpi
two species is understandable given that resolution using a Sony DKC5000®
Chlorogonalia ultima (Figs. 3-4) resem- digital camera mounted on a _ Leica
bles populations of Caldwelliola reservata MZ-APO# stereoscopic microscope.
VOLUME 108, NUMBER 3 613
Figs. 347. 3-4, Ecuadorian Chlorogonalia ultima. 5, Ecuadorian Caldwelliola reservata. 6-7, Central
American populations of C. reservata have additional macula on the pronotum that distinguish them
more easily from C. u/tima.
Specimens examined are deposited in subgenital plates triangular9 except
the National Museum of Natural Histo- abruptly narrowed in distal 1/5 (Fig. 8);
ry, Smithsonian Institution (USNM). paired, symmetrical aedeagal paraphyses
Literature citations designated with located at its base; male genital style with
key letters are consistent with the bibli- apex acute; and female with abdominal
graphy by Metcalf (1964). sternum VII acute to weakly emarginate
and approximately 3 length of exposed
RESULTS pygofer in ventral view (Fig. 9).
Chlorogonalia Young Discussion.4Species of Chlorogonalia
usually can be recognized by their green
Type species: Tettigonia coeruleovittata
body and forewings, except for hyaline
Signoret 1855, by original designation.
forewing apical cells (Figs. 2, 4), and
Diagnosis.4The genus Chlorogonalia a pair of apical marks on the head
differs from other New World Cicadel- (Figs. 3, 12-13), which in some individ-
lini in having the following combination uals are confluent but constricted at mid
of characters: head vertex with small point. As noted above, however, the
lenticular impressions between the ocelli; species Caldwelliola reservata (Fig. 5)
forewing lacking numerous crossveins may be confused with Chlorogonalia
distally (Fig. 4); abdominal segment XI ultima (Fig. 3). The features Young
rudimentary (less than or equal to 0.2 (1977) used to distinguish Chlorogonalia
length of entire pygofer); male with from Caldwelliola, namely the asymmet-
614 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 8-11. External features distinguishing Chlorogonalia species (8, 9) from Caldwelliola species (10,
11) in males (8, 10) and females (9, 11). SGP, subgenital plate; StV//, abdominal sternum VII.
rical aedeagal paraphyses and enlarged but also present in the female (Fig. 19),
abdominal sternum II apodemes and in the length of sternum VII relative
(Figs./ 15.) 17))an' -the: latter, | pane; not to the exposed pygofer (Figs. 9, 11) were
consistent. The paired paraphyses in neither mentioned nor illustrated by
Caldwelliola are sometimes symmetrical Young (1977). As a result of this re-
and also often flattened, the combination definition of the genus and examination
not found in Chlorogonalia. The sternum of the holotype (USNM), one new
II apodemes in some Caldwelliola indi- combination is proposed: Caldwelliola
viduals (e.g., C. cunahua Young holo- tharma (Young) (from Chlorogonalia;
type) are sometimes diminutive. Four Figs. 14-15). The genera can now be
consistent features were found that distinguished from undissected speci-
mens by examining the form of the
distinguish the genera: the form of the
subgenital plates in the male (Figs. 8,
subgenital plates (Fig. 8), the apical form
of the styles (Fig. 17), the relative length 10) and sternum VII in the female
(Figs, 9.1L):
of abdominal segment9X I in the male
(Figs. 18-19), and the relative size of
Kry TO SPECIES OF CHLOROGONALIA
abdominal sternum VII in the female
1. Forewing subhyaline with brown veins
(Figs 9... 1)» The} differences. snayjthe
ME Sat ot, ernest 8 Rb, A, delongi Young
relative length of abdominal segment
4 Forewing mostly green, with green veins
XI, most notable in the male (Fig. 18) except apical cells hyaline (Hissin ee 2
VOLUME 108, NUMBER 3 615
tN Head vertex with black mark with length Notes.4Chlorogonalia delongi differs
more than 2X its median width (Fig. 12)...
from other Chlorogonalia in having
2 & ase Cee CeCe losoplanensis (Schréder)
nearly translucent forewings, in contrast
4 Head vertex with black mark with length
equal to or less than its median width 3 to the green forewing basal and discal
3. Pronotum and forewing sometimes marked cells of other species, and an angulate
with blue (Fig. 13); aedeagus in lateral view head with the central dorsal macula
convex at mid length (Young 1977: Fig.
distinctly longer than wide. The species
A GHD iee6 eees, 3 coeruleovittata (Signoret)
4 Pronotum and forewing not marked with is known only from Mexico (Cuerna-
blue; aedeagus in lateral view gradually vaca, Distrito Federal, Michoacan, and
narrowing to apex, not convex at mid length Puebla) (Young 1977).
(Young 1977: Fig. 472p) .... ultima Young
C. losoplanensis (Schréder)
SPECIES OF CHLOROGONALIA
Tettigella losoplanensis Schréder 1959:
C. coeruleovittata (Signoret 1855) 67 [n.sp.].
Tettigonia lineata Signoret 1854a: 21 Chlorogonalia losoplanensis: Young
[n.sp., preoccupied]. 19772 S75; Fie. 470 (ntcomb:|:
Tettigonia coeruleovittata Signoret Notes.4This uncommon species (10
s55d: 813 [nom. nov. for 7. lineata specimens known) is recorded from
Signoret]. Mexico and Panama. The two medial
Tettigonia coeruleovittata delineata macula (Fig. 12) on the head are some-
Fowler 1900c: 277 [n. subsp.]. times confluent.
Chlorogonalia coeruleovittata: Young
C. ultima Young
1977-575, Fig. 469 [n.comb.].
Notes.4Chlorogonalia coeruleovittata Chlorogonalia ultima Young 1977: 577,
is a widespread species, known from Fig. 472 [n.sp.].
southern Mexico, Guatemala, El Salva- Notes.4 Chlorogonalia ultima (Figs. 1,
dor, Costa Rica, Venezuela, Colombia, 3-4) 1s recorded only from South Amer-
and Brazil (Young 1977 and USNM ica (Colombia, Ecuador, and Peru),
specimens). It recently was found to be where it is common (USNM specimens,
common in a coffee plantation in south- including holotype). Young (1977) went
ern Mexico (McKamey, unpublished). so far as to say that <small green or
Although the coffee was not suffering yellowish Neotropical Cicadellini are
from crespera disease (see below for likely to belong in Scopogonalia, Cald-
characteristics), it is likely that the welliola, new genus, or in Chlorogonalia
species also feeds on coffee elsewhere in ultima, tm: 8sp:~ It <closely wesemibles
its range, including other coffee-produc- Ecuadorian populations of Caldwelliola
ing areas, and might be involved in the reservata (Figs. 2, 5), which had been
spread of crespera (see notes for Cald- misidentified by Young as C. ultima, but
welliola below). Usually having distinc-
differs in the four diagnostic features
tive blue marks on the pronotum and
mentioned earlier. They are sympatric 1n
wings (Fig. 13), C. coeruleovittata is the
Ecuador in the most narrow sense of the
most easily identified species in the
word: in two instances Chlorogonalia
genus. Male individuals without the blue
ultima and Caldwelliola reservata have
markings can be identified based on the
been collected in the same collection
aedeagal structure (Young 1977, fig. 469).
lot4in <Victoria/Aranillas,= 150 m, 18-
C. delongi Young 19 Aug 1977; deus Es Pena; sel Bore
Chlorogonalia delongi Young 1977: Province (USNM), and <8Los Rios=9 [city
575, Fig. 473 [n.sp.]. and province], 2 March 1938, Wm. Cl-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
13
Figs. 12-15. Features of Chlorogonalia and Caldwelliola. 12, Chlorogonalia losoplanensis. 13, C.
coeruleovittata. 14-15, Dorsal habitus and anterior portion of cleared abdomen of Ca/dwelliola tharma.
VOLUME 108, NUMBER 3 617
Figs. 16-19. Features of Cal/dwelliola. 16, Dorsal habitus of C. caucana, an implicated vector of coffee
crespera disease in Colombia. 17, C. reservata, cleared male abdomen, showing the largedsternum II
apodemes (APO) often, but not always present in Ca/dwelliola, and the blunt genital style (STY) . 18-19,
Dorsal and lateral aspects of large abdominal segment XI. 18, C. caucana, male. 19, C. reservata, female.
Macintyre (USNM). This is a new coun- South America. Caldwelliola reservata
try record for C. reservata. previously was known only from Central
America, where populations have black
Caldwelliola Young marks on the pronotum and scutellum
(Fig. 647) similar to those of C. caucana
Type species: Tefttigonia reservata Fowler
Young in Colombia (Fig. 16).
1900, by original designation.
In a study of Cicadellidae as possible
Species of the genus Ca/ldwelliola re- vectors9 (of) the 4 crespera: disease) the
semble Chlorogonalia or Crossogonalia Centro Nacional de Investigacion de
Young but may be recognized by the Café (CENICAFE, Chinchina, Caldas,
features given by Young (1977) in Colombia) sent specimens of six abun-
addition to the new features noted above dant species to the Systematic Entomol-
that distinguish it from Chlorogonalia. ogy Laboratory, U.S. Department of
With regardt o the lengthened abdomi- Agriculture, for identification. In Costa
nal segment XI in Ca/dwelliola, in males Rica, Rodriguez et al. (2001) have in-
the range is 0.340.7X the length of the dicated that the causal agent for crespera
entire pygofer (Fig. 18). This feature is is the bacterium YXy/ella fastidiosa (also
widespread in Cicadellinae but serves to responsible for citrus chlorosis in Brazil
distinguish it from Chlorogonalia and and Pierce9s Disease of grape and peach
was the basis for referring C. tharma, to in9 the United9 <States))" and 1) 1s <vec
Caldwelliola, new combination, as men- tored by two sharpshooter leafhoppers.
tioned above. The nine described species Among the CENICAFE specimens were
of the genus are Neotropical, recorded three sharpshooters capable of trans-
from Mexico and much of Central and mitting X. fastidiosa: Graphocephala n.
618 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
sp., Juliaca scalarum Young, and C. Touchet for assisting with the digital
caucana. Both species previously were photography.
recorded from Colombia but host plants
LITERATURE CITED
were unknown. Young (1977) provided
a diagnosis and illustration for J. sca- Fowler, W. W. 1900c. Order Rhynchota. Suborder
Hemiptera-Homoptera (continued). Biologia
larum (his Fig. 387) and Caldwelliola
Centrali-Americana 2: 2814292.
caucana (his Fig. 845; here an anterior,
Metcalf, Z. P. 1964. General Catalogue of the
dorsal habitus, Fig. 16). The specimens Homoptera. Fascicle VI. Cicadelloidea. Bibli-
were collected in Dept. Risaralda. Ju- ography of the Cicadelloidea (Homoptera
liaca scalarum is known from Cundina- Auchenorrhyncha). United States Department
of Agriculture, Agriculture Research Service,
marca as well as Cochabamba, Peru.
349 pp.
Caldwelliola caucana was known from Rodriguez, C. M., J. J. Obando, W. Villalobos, L.
Popayan and Valle, Colombia. The Moreira, and C. Rivera. 2001. First report of
coffee crespera disease is characterized Xylella fastidiosa infecting coffee in Costa
Rica. Plant Disease 85(9), electronic publica-
by proliferation of vegetative buds,
tion number D-2001-0723-02N (on-line; URL:
which leads to an over-production of
http://www.apsnet.org/pd/+toc/2001/dse01tc.
leaves and shortening of internode dis- htm).
tance, thus producing dwarf trees with Schréder, H. 1959. Taxonomische und tiergeogra-
a <witches broom9 appearance. In ex- phische Studien an Neotropischen Zikaden
(Cicadellidae, Tettigellinae). Abhandlungen
treme cases, only vegetative tissue is
Senckenbergischen Naturforchenden Gesell-
produced, causing a total loss of pro-
schaft 499: 1-93.
ductivity. Chlorogonalia coeruleovittata Signoret, V. 1854a. Revue iconographique des
also feeds on coffee and thus has the Tettigonides. Annales de la Société Entomolo-
potential to spread the disease. gique de France 2(3): 5428.
. 1855d. Revue iconographique des Tettigo-
nides. Annales de la Société Entomologique de
ACKNOWLEDGMENTS France 3(3): 765-836.
Young, D. A. Jr. 1968. Taxonomic study of the
I am grateful to C. A. Galvis Garcia of
Cicadellinae (Homoptera, Cicadellidae). Part
CENICAFE for bringing to my atten-
1. Proconiini. United States National Museum
tion specimens suspected of transmitting Technical Bulletin 261, 287 pp.
coffee crespera disease in Colombia. J. . 1977. Taxonomic study of the Cicadellinae
Brown, L. L. Deitz, R. Robbins, and an (Homoptera: Cicadellidae). Part 2. New World
Cicadellini and the genus Cicadella. North
anonymous reviewer are thanked for
Carolina Agricultural Experiment Station
comments on an earlier draft, and M. Technical Bulletin 239, 1,135 pp.
