Table Of ContentPROC. ENTOMOL. SOC. WASH.
109(4), 2007, pp. 938-947
A NEW SPECIES OF DICHRORAMPHA (LEPIDOPTERA: TORTRICIDAE:
GRAPHOLITINI) FROM JAMAICA: A POTENTIAL BIOCONTROL AGENT
AGAINST CHROMOLAENA ODORATA (ASTERACEAE)
John W. Brown and Costas Zachariades
(JWB) 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: [email protected]); (CZ) ARC-Plant Protection Research
Institute, Weeds Research Division, Private Bag X6006, Hilton 3245, South Africa
(e-mail: [email protected])
—
Abstract. Dichrorampha odorata Brown and Zachariades, new species, is described
and illustrated from Jamaica. It is most similar to D. sapodilla Heppner among
described species, both superficially and in the male genitalia. However, the two are
easily separated by the long costal fold of the male forewing of D. odorata, which is
absent in D. sapodilla. The shapes of the valva and cucullus also distinguish the two.
The related D. aztecaWalsingham, revised status, which shares adistinctmaleforewing
costal foldwithD. odorata, is returnedto Dichrorampha. Dichroramphaodoratainduces
galls in the shoot tips ofthe invasive weed Chromolaena odorata (L.) R. M. King & H.
Robinson (Asteraceae), commonly known as triffid, Jack-in-the-bush, bitter bush,
Christmas bush, and Siam weed. The new species appears to have considerable
potential as a biological control agent against this weedy shrub in South Africa.
Key Words: Systematics, Tortricidae, Dichrorampha, new species, biological control,
triffid, Jack-in-the-bush, Chromolaena odorata, Jamaica, South Africa
With over a dozen common names, to its efficient short- and long-distance
including triffid, Jack-in-the-bush, Siam dispersal abilities (Muniappan 2005). In
weed, bitter bush, and Christmas bush. South Africa it is a problem primarily for
Chromolaena odorata (L.) R. M. King & biodiversity conservation, grazing, and
H. Robinson (Asteraceae) (formerly forestry (Goodall and Erasmus 1996).
known as Eupatorium odoratum L.) is Invasive populations ofC odorata in Asia
a perennial shrub native to South and and southern Africa both originated from
Central America (Holm et al. 1977). In the Caribbean, although the southern
C
recent decadesit has become a serious pest African form (biotype) of odorata is
in the humid tropics ofsoutheastern Asia, distinct from that invading southeastern
Africa, and the Pacific Islands. It has been Asia and western Africa (Zachariades et
nominated as among the top 100 "world's al. 2004). Plants identical to the South
worst" invaders (Wilson 2006). It spreads African biotype have been found only on
rapidly in lands used for forestry, pasture, islands in the northern Caribbean, most
and plantation crops such as rubber, notably Jamaica and Cuba (Zachariades
coffee, coconut, cocoa, andcashew, owing et al. 2004).
—
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109, 4 939
During efforts to identify biological America, 10 in the Neotropics, and one
control agents against this weed, a new in South Africa. The new species is
species of tortricid moth was discovered assigned to the genus on the basis of
in Jamaica that induces galls in the the following character states: male fore-
terminal shoots of the plant. The pur- wing with a well developed costal fold
poses of this paper are to describe and (present or absent in Dichrorampha, but
illustrate this new species and present absent in all other New World Grapho-
information on its biology. litini) (Heinrich 1926); a distinct row of
dark dots along the termen of the fore-
Materials and Methods
wing (Komai 1999) (Fig. 1); female
Specimens oflarvae were discovered in genitalia with the sterigma, seventh
galls of Chrornolaena odorata in three sternite, and sclerotized posterior por-
general areas in Jamaica by personnel tion of ductus bursae fused (Komai
from the Plant Protection Research In- 1999) (Fig. 3); and female genitalia with
stitute, South African Agricultural Re- a single thornlike signum (usually with
search Council: (1) near Mandeville, (2) two thornlike signa in other genera of
in the Blue Mountains, and (3) along the Grapholitini) (Heinrich 1926) (Fig. 3).
north coast from Port Maria to south- Most species of Dichrorampha for which
west of Montego Bay. The galls were host plants are known (i.e., about 25
imported into quarantine in South species) feed on Asteraceae (e.g.,
Africa and successfully cultured (see McDunnough 1946; Swatschek 1958;
Biology below). Representative larvae MacKay 1959; Danilevsky and Kuznet-
and pupae were placed in boiling water zov 1968; Bradley et al. 1979; Jensen and
for approximately 2 minutes and trans- Palm 1981; Park 1983; Godfrey et al.
ferred to 90% EtOH for preservation. 1987; Sterling 1991; Corley 1992) with
Dissection methodology follows that a few notable exceptions: Dichrorampha
presented in J. Brown and Powell okui Komai has been reported from
(1991, 2000). The image of the aduh acorns of Quercus sp. (Fagaceae) (Oh et
was captured using a Microptics® digital al. 2001); Dichrorampha petiverella (Lin-
camera system. Preliminary illustrations naeus) (reported as dorsana) from
of the genitalia were drawn from slide- Lathyrus sp. and Pisum sp. (Fabaceae)
mounted preparations using a Ken-a- (Disque 1908; Bradley et al. 1979); and
Vision microprojector; final illustrations Dichrorampha manilkara Heppner and
were inked using dissecting and com- D. sapodilla Heppner from Mcmilkara sp.
pound microscopes. Terminology for
(Sapotaceae) (Heppner 1981). Five ofthe
genitalia structures and wing venation six species reported from Europe by
follows Horak (1984); terminology for
Swatschek (1958) feed in the roots of
larval chaetotaxy follows R. Brown
their asteraceous hosts, with the sixth
(1987); terminology for forewing pattern feeding on shoots.
follows R. Brown and Powell (1991), as
modified by Baixeras (2002); and plant Dichvorampha odorata Brown and
taxonomy follows GRIN (2006) and Zachariades, new species
METAFRO
(2006).
(Figs. 1-9)
Systematics Diagnosis. Dichrorampha odorata is
Dichrorampha, with 1 12 described most similar to D. sapodilla Heppner
species worldwide (Brown 2005), is (TL: USA, Florida), both superficially
primarily Palaearctic (ca. 80 species), and in the male genitalia. However, the
with 20 described species in North two are easily separated by the long
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
940
mm-
•5
Figs. 1-2 Dichrorampha odorata. 1, Adult male (with scale bar), 2, Male genitalia.
costal fold of the male forewing of D. venter (Fig. 2). Also, D. sapodilla has
odorata, which is absent in D. sapodilla. been reared only from Manilkara zapota
The shape of the cucullus is also distinc- (L.) P. Royen (Sapotaceae) (Heppner
tive between the two: in D. sapodilla it 1981), whereas the new species is known
has distinct dorsal and basal lobes and is only from Chromolaena odorata (Aster-
well separated from the basal part ofthe aceae).
valva by a rounded excavation along the Dichrorampha odorata is superficially
venter of the valva; in D. odorata the similar to D. azteca Walsingham, revised
cucullus lacks the dorsal lobe and is status (TL: Mexico, Guerrero), particu-
separated from the basal part of the larly in the shared possession of a well-
valva by a narrow incision along the developed costal fold in the male fore-
VOLUME NUMBER
109, 4 941
Schaffer and AndriopJecta Obraztsov,
but it is considerably more widespread
and variable within the tribe (Brown and
Baixeras 2006).
—
Description. Adult: Head: Vertex
rough-scaled, cream and pale tan, scales
extending forward nearly to middle of
labial palpus; frons pale tan; outer
surface of labial palpus cream in basal
half with small dark spot at base,
blackish gray in distal half, with scaling
broadly expanded, inner surface cream;
antenna pale gray; ocellus present. Tho-
rax: Dorsum brown, mixed with pale
scales posteriorly, without posterior
crest; legs striped cream and gray-brown,
male without conspicuous secondary sex
scales. Forewing (Fig. 1) ground color
mottled gray-brown, brown, and ochre-
ous; termen with distinct subapical in-
dentation between R5 and Mi at base of
apical-most, white, terminal strigula;
a pair of parallel, narrow, brown, ob-
lique fasciae, arising between costal
strigulae 7-8 and 9 (i.e., subterminal
fascia) and 5-6 and 7-8 (i.e., postmedian
fascia), narrowly bordered by pale or-
ange and separated by narrow black line,
originating at costa ca. 0.5 and 0.7
distance from base to apex, extending
toward termen; costa with 7 pairs ofpale
strigulae, basal pairs indistinct, distal
pairs frequently coalesced; basal half of
wing with ill-defmed, dark, oblique
Fig. 3. Dichrorampha odorata, female genitalia.
band, originating at dorsal margin ca.
0.33 distance from base to tornus,
wing. However, the male genitalia of D. attenuating before reaching mid-costa;
azteca have a relatively broad, deep, U- ocellus (patch) not developed; 2>-^ small
shaped excavation immediately basad of black spots forming line parallel to
the cucullus as in D. sapodilla and unlike termen in yellow-speckled terminal re-
that of D. odorata. gion; interfacial spot ill-defined, mostly
The female of D. odorata has a frenu- obscured with gray-brown. Fringe gray-
lum comprised of two bristles. Although ish brown. Ventral surface lustrous
somewhat unusual, this character state is bronze. Hindwing uniform dark brown
found in several other Grapholitini (e.g., above, except broad area ofoverlap with
a few species in each of the genera forewing whitish. Fringe grayish brown.
Pammene, Cydia, and Grapholita). Ko- Ventral surface lustrous bronze. Male
mai (1999) proposed that it represented frenulum with one bristle, female frenu-
a synapomorphy for Strophedra Herrich- lum with two. Abdomen: Scaling dark
942 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
brown, without conspicuous secondary and SV-group 2:1:1 on Tl-3. Abdomen
sex scaling in male. Male genitalia with spiracles small and circular, only
(Fig. 2; based on 2 preparations) with slightly smaller than that of Tl; spiracle
tegumen rather broad, rounded-triangu- on A8 slightly posterior to middle of
lar dorsally, with small mid-dorsal segment; segments 1-8 with SD2 in-
hump; gnathos a narrow, ill-defmed conspicuous, SDl dorsad of spiracle on
band dorsad of aedeagus; vinculum A1-7, anteriorad of spiracle on A8; LI
weakly sclerotized; valva moderately and L2 on same pinaculum on A1-8;
broad, with large, subrectangular basal D2s on shared dorsal pinaculum on A9;
excavation, costa weakly arched from ca. Dl and SDl on a shared pinaculum on
base to ca. middle of cucullus, apex A9; L-group trisetose on A9; V's about
rounded; sacculus not developed; cucul- the same distance apart on A7-9, or
lus broad with large reflexed basal lobe, slightly further apart on A9; SV Group
densely setose, separated from basal on 1, 2, 7, 8, 9 is 2:2:2:2:1. Crochets
portion of valva along ventral edge by uniordinal, 18-20 on prolegs (A3-6), 10-
deep, narrow incision; "neck" of valva 14 on A10. Anal fork present.
inconspicuous owing to narrow incision; According to Swatschek (1958), the
aedeagus relatively long, ca. 0.5 length of
larvae ofDichroranipha are characterized
valva, arched, slightly dilated distally, by the following: seta SDl on A8
with diffuse patch of fixed cornuti.
immediately anterior of the spiracle and
Female genitalia (Fig. 3; 2 preparations) SV group bisetose on A1, 2, 7, and 8, and
with papillae anales slender, slightly
unisetose on A9, except occasionally
broader in distal 0.5; apophyses poster-
trisetose on A2. The only species of
iores ca. 0.75 times length of apophyses Dichrorampha treated by MacKay (1959)
anteriores; sterigma symmetrical, un-
sclerotized except for narrow crescent- (i.e., D. bittana) was reported to lack an
anal fork; Swatschek (1958) did not
shaped patches extending from lateral
edge of rounded, sclerotized ostium; mention this feature is his key or di-
agnoses of larval Dichrorainpha.
ductus bursae comparatively broad, ca.
f8iftthismemsorleonsgcelrerotthiazned,wisdlei,ghtdliysteallontgwaot-e widPtuhpa:ca.Fus1imfomrm,(nlen=gth5);ca.typ5icmamll,y
funnel-shaped, remainder of ductus bur- tortricoid, without cephalic projection
sae membranous; ductus seminalis from or conspicuous sculpturing. Dorsum of
ductus bursae ca. 0.6 distance from segment Al lacking spines; dorsum of
ostium to junction of ductus and corpus A2 8 with two rows of spines, anterior
bursae; corpus bursae oblong, membra- row situated at anterior edge ofsegment,
nous, with a single, slender, thornlike with larger spines, row attenuate well
signum at right lateral side of corpus. before spiracle; posterior row situated at
Larva: General (last instar): Length posterior edge of segment, with smaller
6.0-7.0 mm (n = 3), head somewhat spines but frequently extending further
circular in outline in dorsal view, amber laterad; segment A8 sometimes with
to pale yellow with variably developed posterior row weak or absent. Segments
brown stemmatal patch and spot at genal A9-10 each with short row of larger
A
angle; prothoracic shield pale yellow thorns; 10 with 6 slender, hook-tipped
with faint or no traces of pattern; body setae; cremas—ter absent.
pale, pinacula weakly pigmented, incon- Holotype. Male, Jamaica, Jackson
WP
spicuous; anal shield very pale yellow, Town, 042, 18.25N, 77.29W, 13
unmarked. Thoracic chaetotaxy typically Nov 1999, C. Zachariades. Deposited in
tortricoid, with L-group trisetose on Tl Institute of Jamaica, Kingston, Jamaica.
—
VOLUME NUMBER
109, 4 943
Paratypes. \S, 10$. Jamaica: Blue by Brown (2005), who placed it in
WP
Mountains, 032-5, 047-051, 18.03N, "Grapholitini Unplaced Species." How-
76.45W to 18.01N, 76.40W, 9, 10, 14 Nov ever, an examination ofthe syntype male
1999 (22), C. Zachariades. Blue Moun- (The Natural History Museum, London)
tains; north along coast from Port Maria revealed that it is appropriately assigned
to southwest ofMontego Bay, 18°01.959- to Dichrorainpha based on its similarity
18°27.630N, 78°42.048-78°01.423W, 29- with D. odorata and D. sapodilla.
846 m, 16-24 Nov 2005 (Ic?, 8?), C.
Zachariades & L. D. Khumalo. Deposit- Biology and Discussion
ed in the South African National Collec- During field surveys for biological
tion ofInsects, Pretoria, South Africa; the control agents against Chromolaena
National Museum of Natural History, odorata, Dichrorampha odorata was col-
Smithsonian Institution, Washington, lected in Jamaica and Cuba, but only in
DC, USA; and The Natural History small numbers. However, in November
Museum, Londo—n, England. 2005 large numbers of galled shoot tips
Distribution. Known from Jamaica were discovered on Chromolaena odorata
and Cuba. — in Jamaica, and a large percentage ofthe
Etymology. The specific epithet is galls contained larvae of Dichrorampha
derived from the species name of the odorata. These galls were imported into
larval host plant. quarantine in South Africa and success-
—
Remarks. The paratypes were col- fully cultured.
lected over a range of localities (i.e., The damage caused by Dichrorampha
"waypoints J05-008, 009, 010, Oil, 013, odorata is superficially similar to that
014, 016, 017, 018, 019, 029, 021, 022, caused by Phestinia costella (Hampson)
025, 026"); hence the precise locality of (Lepidoptera: Pyralidae: Phycitinae), al-
each individual is unknown. The male so on C. odorata, but less frass is
holotype, although not the best speci- produced, the eggs probably are laid in
men, was selected because it has more the shoot tip, and the larvae bore intomthme
precise locality data. shoot tip causing the terminal 15-20
of the stem to swell slightly and often
Dichrorainpha azteca Walsingham, turn yellow. The terminal shoot tip dies,
revised status and the stem ceases its growth. The
Dichrorainpha azteca Walsingham 1914: swollen stems (loosely interpreted as
"galls") induced by Dichrorampha odor-
258.
"Dichroramphcr [misplaced] azteca: PoweU ata are smaller and less obvious than
those induced by Phestinia species, but
et al. 1995: 157.
"Grapholitini Unplaced Species" azteca: growth ofthe terminal shoot is effective-
Brown 2005: 365. ly stopped (Figs. 4-5). In the field, galls
of D. odorata were found only on
Based on features ofthe male genitalia terminal shoots whereas those of Phesti-
and forewing pattern, D. azteca is un- nia were found in axillary shoots as well.
doubtedly congeneric with D. odorata Axillary buds are attacked by Dichro-
and D. sapodilla. It can be separated rampha odorata in the laboratory prob-
from those species by the features ably owing to a shortage of oviposition
discussed above in the diagnosis of D. sites. Larvae (Figs. 6-7) bore down the
mm
odorata. Although described in Dichro- stem for a distance of 15 to 20 (x ± 1
rampha, azteca was treated as "'Dichro- SD = 18.6 ± 3.47 mm for n = 11
rainpha"" [misplaced] by Powell et al. vacated galls or galls containing mature
(1995), and this treatment was followed larvae) and are easily transferred be-
944 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
s^3**^
VOLUME NUMBER
109, 4 945
100 cm). Thus far the species has been leaf-feeding agents, Pareuchaetes insulata
bred easily and quickly (generation time (Walker) (Lepidoptera: Arctiidae) and
of 6-7 weeks) in these cages, showing Calycornyza eupatorivora Spencer (Dip-
a fairly high rate of increase (the highest tera: Agromyzidae), already are estab-
was a six-fold increase from one gener- lished in South Africa. A stem-tip attack-
ation to the next). er has long been considered an important
Larval no-choice tests have been initi- element in any successful biological con-
ated, similar to those conducted by trol program against C. odorata, both to
Cruttwell (1977) on insects identified as decrease the extremely high growth rate
Mescinia nr. parvula (Zeller) from Trini- of the stems and to decrease the number
dad, but which likely are Phestinia of shoot tips available for flowering (in
costella (A. Solis, personal communica- the laboratory, D. odorata larvae also
tion). These tests are considered appro- bored into flower buds). Although several
priate because the larvae are highly other stem-tip attackers are known, many
mobile and are unlikely to bore into have proven difficult to rear under
plants that they find unsuitable as hosts. laboratory conditions. In addition, poor
In initial tests, mid-instar larvae were compatibility between the southern Afri-
placed on marked shoot tips of 10 species can C. odorata biotype and insects or
of Asteraceae as well as C. odorata. pathogens collected from other morpho-
C
Asteraceae were targeted because this logical forms of odorata in South or
family includes the most commonly Central America frequently has been
documented hosts of Dichrorampha. observed or suspected (Zachariades et al.
Larvae caused galls on only two species 2004). This should not be a problem with
{Mikania capensis DC. and Ageratum D. odorata because the biotype of C.
conyzoides L.) other than C. odorata, but odorata invading southern Africa is be-
galling was minimal, and none of the lieved to have originated in Jamaica or
larvae on these plants pupated. None of Cuba. When released in the field in South
the eight other Asteraceae species offered Africa, the ultimate distribution of D.
to larvae (i.e., Adenostemma caffrum odorata may be confined to the coastal
DC, Ageratina adenophora (Sprengel) belt and other areas with a less severe dry
King & Robinson, A. riparia (Regel) season. In certain inland areas (e.g.,
King & Robinson, Synphyotrichwn novi- Zululand) C. odorata dies back during
belgii (L.) G. L. Nesom, Chrysanthemum winter, losing its leaves and sometimes its
X morifolium Ramat., Delairea odorata upper stems; fire is also a common
Lem., Distephanus angulifolius (DC.) occurrence in these areas. The two
&
Robinson B. Khan., Microglossa established leaf-feeding agents, neither
mespilifolia (DC.) Robinson) was fed of which has a prominent diapause
upon to any degree. The insect thus mechanism, currently are restricted to
seems highly host specific. the coastal belt. Dichrorampha odorata
Provided that it has a host range does not have an obvious diapause, but
restricted to C. odorata, Dichrorampha may achieve a somewhat wider distribu-
odorata has several attributes that make tion than P. insulata and C. eupatorivora
it a promising biocontrol agent for this because it does not rely on leaves for its
weed in South Africa. It is bred easily biology to the same degree as these two
undercagedconditions, making it a good species.
species for mass-rearing for release. It
Acknowledgments
has a short life cycle and high fecundity,
and therefore is likely to increase and We thank Kevin Tuck (The Natural
spread quite rapidly in the field. Two History Museum, London) and Jose
946 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Montero (Institute Nacional de Biodiver- Brown, R. L. and J. A. Powell. 1991. Description
sidad, Santo Domingo, Costa Rica) for of a new species of Epiblema (Lepidoptera:
Tortricidae: Olethreutinae) from coastal red-
allowing us to examine specimens in their
wood forests in California with an analysis of
care. Diana Marques provided the draw- the forewing pattern. Pan-Pacific Entomolo-
ings ofthegenitalia, and Karolyn Darrow gist 67: 107-114.
provided the photograph ofthe adult. We Corley, M. F. V. 1992. The summer brood of
thank the following for helpful reviews of Dichrorampha acuminatana (Lienig & Zeller)
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