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SYSTEMATICS
Siamusotima aranea, a New Stem-Boring Musotimine
(Lepidoptera: Crambidae) from Thailand Feeding on
Lygodium flexuosum (Schizaeaceae)
M. ALMA SOLIS,
1
SHEN-HORN YEN,
2
JOHN H. GOOLSBY,
3
TONY WRIGHT,
3
ROBERT PEMBERTON,
4
AMPOM WINOTAI,
5
USANEE CHATTRUKUL,
6
AMARA THAGONG,
6
AND SURIONT RIMBUT
6
Ann. Entomol. Soc. Am. 98(6): 887Ð895 (2005)
ABSTRACT Siamusotima aranea Solis & Yen, is a new stem-boring musotimine species from Thai-
land. It was discovered in the stems of Lygodium flexuosum (L.) Sw. (Schizaeaceae) during exploration
for biological control agents of Lygodium microphyllum (Cav.) R. Br., the Old World climbing fern.
This is the Þrst report in the Pyraloidea of a stem-boring larva with unique modiÞcations of the anal
segment resembling that of tenebrionid beetle immatures and with observations of possible mimicry
between the adult moth and spiders.
KEY WORDS Pyraloidea, Old World climbing fern, pteridophagy, spiders, tenebrionid beetles
EXPLORATION FOR BIOLOGICAL control agents of Lygo-
dium microphyllum (L.) Sw., the Old World climbing
fern, has been conducted in Australia and southeast-
ern Asia since 1998 by the Agricultural Research Ser-
viceÐAustralian Biological Control Laboratory in Bris-
bane, Queensland (Goolsby et al. 2003a, b). This fern
is an invasive weed in southern Florida where it
threatens many wetland communities, including the
Everglades (Pemberton and Ferrier 1998; Pemberton
et al. 2002; Goolsby et al. 2003a, b). Among the insect
species discovered on Lygodium species and proposed
for use as biological control agents, those of the Mu-
sotiminae (Crambidae) are especially signiÞcant be-
cause a number of species in different genera have
been found to exclusively consume Lygodium. Muso-
timines discovered previously were external fern feed-
ers (Goolsby et al. 2003a, b; Solis et al. 2004; Yen et al.
2004), but recently one species was found boring into
the stems of L. flexuosum in Thailand. We describe all
life stages of this new genus and species, and discuss
the placement of the genus. We also discuss the pos-
sible mimicry of the adult wing pattern to Thomisidae
spiders and the resemblance of the anal segment of the
lepidopteran larva to anal segment of tenebrionid im-
matures.
Materials and Methods
Biological Exploration Methods. The following
countries were visited between September 1998 and
March 2002: Australia, China, Japan, Indonesia, Ma-
laysia, New Caledonia, Singapore, Taiwan, and Thai-
land. SpeciÞc search areas were selected by climate
matching, habitat similarity, herbarium locality data,
and information provided by knowledgeable special-
ists. Methods for collection included hand searches,
sweeping, beating of foliage, and dissection of under-
ground plant parts (Goolsby et al. 2003a).
Taxa Studied. Specimens were studied from The
Natural History Museum, London, United Kingdom
(BMNH). Type material is deposited in those institu-
tions as designated within the text. To determine the
identity and/or placement of the taxa feeding on Ly-
godium, the type species, and other species if available,
of all described genera currently in the Musotiminae,
including several newly published and unpublished
genera by Solis et al. (2004), Yen et al. (2004), and
S.-H.Y. (unpublished data), were dissected, studied,
and compared. The diagnosis below is composed of
synapomorphies.
Pinned specimens were examined and dissected
after abdomens were soaked in 10% potassium hy-
droxide and wings were soaked in bleach. Dissections
were stained in chlorozal black for genitalia, and Eo-
sin-Y for wings. Genitalia were placed in vials with
1
SEL, USDA, Smithsonian Institution, P.O. Box 37012, National
Museum Natural History, E-517, MRC 168, Washington, DC 20013-
7012.
2
Department of Biological Sciences, Imperial College at Silwood
Park, Ascot, Berkshire SL5 7YP, United Kingdom, and Department of
Biological Sciences, National Sun Yat-Sen University, Kaohsiung 804,
Taiwan.
3
Australian Biological Control Laboratory, USDAÐARS, 120 Meiers
Rd., Indooroopilly, Queensland, Australia 4068.
4
Invasive Plants Research Laboratory, USDAÐARS, 3205 College
Ave. Ft. Lauderdale, FL 33314.
5
Department of Agriculture, Plant Protection Research and De-
velopment OfÞce, Bangkok, Thailand.
6
Royal Project Foundation, Chiang Mai, Thailand.
glycerin and/or slide mounted in Canada balsam or
Euparol (Clarke 1941, Holloway et al. 1987). Mea-
surements were made with an ocular micrometer.
Forewing length was measured from the center of the
axillar area to the apex of the forewing. Terminology
follows Hinton (1946), Klots (1970), Maes (1985, 1995,
1997), Yoshiyasu (1985), Phillips and Solis (1996),
Munroe and Solis (1999), and Solis and Maes (2002).
Results and Discussion
Siamusotima Solis & Yen, New Genus
(Figs. 1A and B, 2A, 4AÐC)
Diagnosis. Forewing with three brown bands curv-
ing anteriorly to apex and hindwing with four longi-
tudinal lines radiating from median line to outer mar-
gin on an otherwise white ground color; uncus base as
broad as vinculum; valva with sclerotized lobe directly
over saccular lobe pointing anteriorly; sacculus me-
dially with a small membranous saccular lobe pointing
posteriorly. Larva elongate with abdominal prolegs
and crochets reduced, and the caudal segment a scle-
rotized cup-like structure.
Etymology. The preÞx of the generic epithet is de-
rived from the Latin “siam” referring to the historical
name of Thailand where this taxon was Þrst discov-
ered, and the generic stem “musotima” referring to its
relationship to this genus in the subfamily. The gender
of the genus is feminine.
Type Species. Siamusotima aranea Solis & Yen
Comparison and Phylogenetic Relationships of Sia-
musotima to Other Genera in the Musotiminae. Sia-
musotima has strikingly distinct larval morphology
from all other musotimine genera. Compared with the
larva of “Cataclysta” angulata Moore, a leaf miner on
A
B
Fig. 1. (A) Male adult of S. aranea, Thailand, Chiang Mai, Ban Pong, Mae Jo. (B) Live adult of S. aranea, Australian
Biological Control Laboratory.
888 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 98, no. 6
Nephrolepis cordifolia (L.) Presl. (Oleandraceae)
(Yen et al. 2004), the larva of Siamusotima is more
highly adapted to an endophagous environment with
its elongated body, reduced prolegs and crochets of
A3-A6, and a cup-like caudal abdominal segment. Al-
though both larvae are internal feeders, we do not
consider that they are closely related because the
pupal shape, genitalic characters, and host association
of “Cataclysta” angulata reveal its phylogenetic afÞnity
with Eugauria Snellen. The pupal shape of Siamuso-
tima is similar to that of Musotima dryopterisivora
Yoshiyasu (as Musotima acclaralis Walker in Naka-
mura 1977) and Neurophyseta Hampson (S.-H.Y., per-
sonal observation), but the latter have reduced fron-
toclypeal setae. The pupal shape of these three genera
is also similar to those of Lygomusotima Solis & Yen,
Neomusotima Yoshiyasu, and Austromusotima Yen &
Solis, but the latter three genera lack the protruded
conical spiracles.
In the adult, the forewing of Siamusotima is basally
brown and distally white with three brown bands
curving anteriorly to the apex. In Lygomusotima, an-
other genus that occurs in Thailand, the entire wing is
light brown except the area between the postmedial
and subterminal line that is distinctively patterned
with a bright, white triangle that spans R
5
and M
1
. The
entire uncus of Siamusotima is broad, and in Lygomu-
sotima the apex is about half as wide as the base.
Lygomusotima has a simple valva without lobes, with
a slightly indented sacculus, but in Siamusotima the
valva has a sclerotized lobe directly over the saccular
lobe pointing anteriorly and the sacculus has a medial,
small membranous saccular lobe pointing posteriorly.
Both Lygomusotima and Siamusotima have a basally
quadrate, bifurcate juxta, but in Siamusotima the juxta
is only three-quarters as long as the vinculum with
indistinct juxtal arms and in Lygomusotima the juxta is
as long as the vinculum with distinct juxtal arms. The
female genitalia of Siamusotima, has a membranous
corpus bursae without a signum and the ductus bursae
is slightly constricted with an internal circle of scle-
rotized, minute, teeth. The corpus bursae of Lygomu-
sotima is rugose with a signum.
The male genitalia of Siamusotima is similar to those
of Ambia ptolycusalis Walker, Musotima aduncalis
Felder & Rogenhofer, Neurophyseta, Melanochroa Yo-
shiyasu, Undulambia Lange, and Uthinia Snellen in
having a ventral valval process. However, the valval
process of Siamusotima is more robust and cornate.
Also, the broad uncus of Siamusotima indicates that
this new genus is not closely related to those genera
that have a slender uncus, e.g., Aeolopetra Meyrick,
Cilaus Joannis, Cymoriza Guene´e, Drosophantis Mey-
rick, and Parthenodes Guene´ e. Siamusotima may be
most closely related to Musotima because both genera
have a medial ventral valval process and very similar
Fig. 2. (A) Male genitalia of S. aranea. (B) Male aedoeagus of S. aranea. (C) Female genitalia and caudal segments of
S. aranea.
November 2005 SOLIS ET AL.: FERN-BORING MUSOTIMINE PYRALOIDS 889
pupal shape, but Siamusotima is still distinguished
from Musotima by having much narrower valva,
broader uncus, and internally feeding larvae.
Challenges in the Definition and Placement of
Musotimine Genera
Because the classiÞcation of Hampson (1897) clas-
siÞcation of the Hydrocampinae based on adults, the
majority of musotimine species have been placed in
two large and systematically chaotic genera, Ambia
Walker (type species Ambia ptolycusalis Walker) and
Musotima Meyrick (type species Diathrausta adunca-
lis Felder & Rogenhofer). All other musotimine gen-
era, except for the Neotropical Neurophyseta (Munroe
et al. 1995, Phillips and Solis 1996), are either mono-
typic or comprise two to 10 species. However, it is still
difÞcult to determine whether we can apply these two
generic names to at least 100 musotimine species other
than their type-species groups. The morphology of A.
ptolycusalis is very “simple” and “conservative,” and
similar character states also can be seen in some spe-
cies presently placed in Musotima. With very little
information, we are unable to determine whether
these two genera are monophyletic, closely related,
congeneric, or applicable to some undescribed spe-
cies-groups. In this article and other studies of muso-
timine systematics, we have avoided creating new
genera that would lead existing genera, such as Ambia
and Musotima, to be para- or polyphyletic, and we
have recognized all other existing genera that dem-
onstrate clear morphological gaps from the A. ptoly-
cusalis and the M. aduncalis species-groups based on
both adults and immatures. Yen et al. (2004) illus-
trated that there is low phylogenetic correlation be-
tween adult and immature morphology of Musotimi-
nae. Thus, using only adult or immature characters
may not successfully justify generic status and phylo-
genetic afÞnity of a musotimine species.
Siamusotima aranea Solis & Yen
(Figs. 1A and B, 2AÐC, 3A and B, 4AÐC, 5AÐF,
6AÐC)
Adult. Head (Fig. 1A and B). Frons brown, white
laterally, and vertex brown; antennae white, prismatic,
laterally compressed; labial palpi three-segmented,
white medially, brown laterally, second segment with
a fan of scales; maxillary palpi white medially, brown
laterally. Proboscis scaled white basally. Chaetose-
mata and ocelli absent.
Thorax (Fig. 1A and B). Thorax, patagium, and te-
gulae white, with dark brown-tipped scales. Legs
white, tibiae brown laterally.
Forewing (Fig. 1A and B). Average forewing length
6.0 mm (n ⫽ 7). Costal swelling absent. Costa brown
to apex or white from basal area to apex. Margin
incised or slightly crenulate with brown scales as a
continuous line from the apex along the outer margin
around the anal angle and along posterior margin to
base of 1A⫹2A. Vein R
1
and R
2
from discal cell, R
3
and
R
4
stalked near apex of wing, arising from apex of
discal cell, R
5
close at base; M
1
straight; M
2
connected
to M
3
by perpendicular vein at base just beyond where
M
3
and CuA
1
are more or less approximate at base;
1A⫹2A strongly sclerotized. Basal area dark brown,
light brown distally. Median line dark brown. Two
brown bands extending distally from median line on
white wing, one along the length of Cu and CuA
2
curving anteriorly to apex of wing extending to fringe,
one a dark brown line along M
1
curving anteriorly to
Fig. 3. (A) Frontal view of cranial chaetotaxy of S. aranea. (B) Lateral view of cranial chaetotaxy of S. aranea. A, anterior
setae; Aa, pore near seta A2; S, stemmatal setae; SS, submental setae; F, frontal setae; AF, adfrontal setae; C, clypeal setae;
L, lateral setae; La, pore near L1; P, posteriodorsal setae; MG, genal setae; M, mandibular setae; AL, labral setae.
890 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 98, no. 6
costa. Short brown band from median line along R vein
to half the length of wing. Fringe white. Underside
white with dark brown bands highly visible; color and
pattern as on upperside of wing. Male retinaculum a
short set of setae pointing toward costa located basally
on Cu.
Hindwing (Fig. 1A and B). Costa straight; apex
rounded; termen slightly sinuous to tornus. Veins Sc
and R separate at base, vein Sc⫹R
1
then anastomosed
with Rs for short distance, M
1
straight with a perpen
-
dicular line toward M
2
;M
2
to CuA
1
as in forewing;
CuA
2
,1A⫹2A, and 3A complete. Margin crenulate.
Basal area dark brown, light brown distally. Median
line dark brown. Four longitudinal lines radiating from
median line to outer margin along Rs, M
2
, CuA
2
, and
1A⫹2A. Underside white with dark brown basal area
and longitudinal bands highly visible. Male frenulum
with one seta, female with three setae.
Abdomen. Tergites one and two white with some
brown-tipped scales, tergites 3 and 4 completely
brown, remaining tergites white. Tympanal organs:
Tympanal cases (⫽bulla tympani, caisses tympa-
niques) reduced, tympanic frame (⫽fornix tympani,
cadre tympanique) highly sclerotized, processes tym-
pani (⫽sailles tympaniques) absent, ramus tympani
absent.
Fig. 4. Last instar of S. aranea. (A) Lateral view S. aranea. (B) Ventral view of cup-like structure of S. aranea. (C) Lateral
view of cup-like structure of S. aranea. (D) Lateral view of tenebrionid immature, P. nigrocyaneus Motschulsky, from Hayashi
(1966). (E) Dorsal view of cup-like structure of P. nigrocyaneus from Hayashi (1966). (F) Lateral view of cup-like structure
P. nigrocyaneus from Hayashi (1966). A, abdominal segments.
November 2005 SOLIS ET AL.: FERN-BORING MUSOTIMINE PYRALOIDS 891
Male genitalia (Fig. 2A and B). Uncus basally as
broad as vinculum, slightly tapering to a membranous
round apex; gnathos (⫽pseudognathos of Maes 1997,
Solis and Maes 2002) mostly membranous with scle-
rotized spinules dorsally, arms tapered to tegumen;
tegumen continuous, forming an inverted “U” with
ventral sclerotized extension to costa of valva; juxta
bifurcated, three-quarters as long as vinculum, basally
quadrate; transtilla medially membranous; valva sim-
ple, apical margin rounded, sclerotized lobe directly
over saccular lobe pointing anteriorly; basally sacculus
overlaps juxta, sacculus medially with a small mem-
branous saccular lobe pointing posteriorly; vinculum
broadly sclerotized, two times as long as tegumen.
Aedoeagus simple without cornuti, distal half lightly
sclerotized; coecum one-third length of aedoeagus.
Fig. 5. (A) SEM of ventral view of cranial setae and structures of S. aranea. (B) Scanning electron micrograph (SEM)
of reduced proleg of A3 of S. aranea. (C) SEM of lateral view of cranial setae of S. aranea. (D) SEM of ventral view of
metathoracic legs and the associated setae of S. aranea. (E) SEM of dorsal view of A3 of S. aranea. (F) SEM of ventral view
of caudal prolegs of S. aranea. BS, basistipes setae; DS, dististipes setae; LA, labio-hypopharyngeal setae; D, dorsal setae; L,
lateral setae; SV, subventral setae; V, ventral setae; CX, coxal setae; FE, femoral setae.
892 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 98, no. 6
Female Genitalia (Fig. 2C). Corpus bursae mem-
branous, as long as seventh segment; signum absent;
appendix bursae absent; ductus seminalis originating
from ventroposterior area of corpus bursae; ductus
bursae membranous posteriorly, slightly sclerotized
from ostium bursae to half the length of ductus where
it is slightly constricted with an internal circle of
minute sclerotized teeth; posterior apophyses two-
thirds as long as anterior apophyses; papillae analis
anteroposteriorly ßattened, indented mediodorsally.
Larva (Figs. 3A and B, 4AÐC, and 5AÐF). Head
hypognathous, epicranial suture very short; height of
frontoclypeus 1.15 times width; six stemmata, C1 and
C2 approximate, C3 and C4 approximate, C5 anterio-
ventrad to C6; S2 longer than S1, SS2 longer than SS1,
MG1 as long as S3, F1 present near middle of fronto-
clypeus, AF1 and AF2 on adfrontal area extremely
short, La and Aa absent, D short, P2 shorter than P1;
labrum with Þve short setae on each side externally;
mandible with one row of serrations, M1 as long as M2.
T1Ð3 and A1Ð10 integument smooth, segments with-
out chalazae, pinacula only very weakly present dor-
sally. Prothoracic shield weakly sclerotized, D1
shorter than D2, XD1 shorter than XD2, SD1 and SD2
absent, L and SV setae absent. T2ÐT3 with D setae
bisetose; SD setae bisetose; L setae unisetose; SV
present, as long as L setae; V setae very short. A1-A2
with D setae bisetose; SD group bisetose, SD1 as long
as L2 of T2Ð3; L, SV and V groups unisetose. Setal map
of A3Ð8 in general similar to A1Ð2. A9 with D, SD, L,
SV and V groups unisetose, and with spiracle switched
to posterodorsal part. A10 with anal shield modiÞed as
a cup-like structure with margin surrounded by D, SD,
PP and L group setae; L1 situated in front of cup-like
structure. Thoracic legs present, with developed CX
and FE setal groups. Prolegs of A3ÐA6 reduced and
crochets of A3ÐA5 absent; crochets of A6 weakly
present as semicircular and uniserial form. Caudal
prolegs of A10 extending posteriorly and situated be-
neath cup-like structure, crochets absent.
Pupa (Fig. 6AÐC). Ventral view: Frontoclypeus of
head slightly concave; labrum narrow; maxillary pal-
pus visible near base of proboscis case; proboscis ex-
tending to middle of A2; fore-femur visible, protho-
racic legs (foreleg) two-thirds length of forewing case,
reaching to end of A2; mesothoracic leg (midleg)
extending to middle of A4, longer than antennal
sheath; hindtarsus visible; abdominal segments 5Ð 6
without vestigial prolegs; abdominal segments 8 Ð10
with genital oriÞce anteriorly; anus posteriorly with-
out lateral conical depressions. Dorsal view: prothorax
smooth, with two very short dorsolateral horn-like
structures; mesothorax without setae; surface of ab-
dominal tergites and wing sheath smooth; cremaster
somewhat ßattened dorsoventrally with a truncate
apex and three pairs of curved setae.
Type Material. HOLOTYPE: Male, Thailand,
Chiang Mai, Ban Pong, Mae Jo, May 2003, A. Winotai,
Ex: Lygodium flexuosum. Deposited at the Rafßes Mu-
seum of Biodiversity Research, National University of
Singapore, Singapore (NUS).
PARATYPES. Two males, Thailand, Chiang Mai,
Ban Pong, Mae Jo, May 2003, A. Winotai, Ex: Lygodium
flexuosum; one male, Thailand, Ban Pong, Mae Jo,
Chiang Mai, N 18⬚ 55.61 E 99⬚ 33.01, 15-VI-2001, A.
Winotai, Lygodium stem borer; three females, Thai-
Fig. 6. Pupal cases of S. aranea. (A) Dorsal view. (B) Lateral view. (C) Ventral view. T, thoracic segments; A, abdominal
segments.
November 2005 SOLIS ET AL.: FERN-BORING MUSOTIMINE PYRALOIDS 893
land, Chiang Mai, Ban Pong, Mae Jo, May 2003, A.
Winotai, Ex: Lygodium flexuosum. Paratypes deposited
at The National Museum of Natural History, Smith-
sonian Institution, Washington, DC.
Distribution. Thailand.
Etymology. The speciÞc name is derived from the
Latin aranea and denotes the resemblance of the adult
moth wing pattern to a spider.
Biology. A comprehensive description of laboratory
and Þeld rearings of this species can be found in
Goolsby et al. (2003a) from which the following is
taken. Eggs are laid on the upper surface of the leaf and
remain in this stage 7Ð10 d. Newly eclosed larvae move
to the tip of the stem, and then they move and tunnel
downwards through the stem, periodically making
holes through which frass can be seen outside the
stem; up to 15 larvae have been found in a single stem.
Pupation occurs on the outside of the stem. Adults are
attracted to light from late April to June, and it seems
that there are two generations per year in Thailand. L.
flexuosum is found in upland tropical forests that are
partly deciduous during the dry season in northern
Thailand, in comparison to L. microphyllum that is
mostly a swamp inhabiting species, but it can live on
heavy soils in high rainfall areas. Near its northern
limits in southern China L. flexuosum is sympatric with
L. microphyllum and L. japonicum (Thunb.) Sw. Al-
though S. aranea has been found only in Thailand thus
far, this species may be more widely distributed, be-
cause the host plant, L. flexuosum, occurs widely from
Southeast Asia to Australia.
Mimicry and Morphological Convergence. The
adult and larva exhibit extraordinary morphological
modiÞcations whose functions are unknown but may
presumably be for defense against predators and/or
parasites. The wing pattern of the adult (Figs. 1A and
B) resembles legs held in a laterigrade fashion resem-
bling those of the Thomisidae, the crab spider family
(J. Miller, personal communication). There have been
a few behavioral observations on adult moth species
mimicking spider behavior in defense against spider
predation (Manu 2003, Rota 2003, Aiello and Becker
2004), but it has not been observed in S. aranea. The
larvae S. aranea have a morphological adaptation of
the anal shield that does not occur in any other known
stem-boring pyraloid that may provide protection
from predation and/or parasitism. The anal shield is
modiÞed as a cup-like structure with a margin sur-
rounded by setae (Fig. 4AÐC), very similar to that of
some beetle larvae (Fig. 4DÐF), speciÞcally Amaryg-
mini tenebrionid larvae (Spilman 1969).
Acknowledgments
We thank Kevin Tuck and Michael Shaffer for access to
The Natural History Museum (London) pyraloid collections.
We especially thank Linda Lawrence (SEL, USDA) who
photographed the adults and their internal structures en-
hancing them with Adobe Photoshop and Jon Lewis (SEL,
USDA) for dissections. We thank Jeff Makinson and Ryan
Zonneveld (Commonwealth ScientiÞc and Industrial Re-
search Organization Entomology, Australian Biological Con-
trol Laboratory, Indooroopilly, Queensland) for rearing and
curation. J. Miller provided the spider family identiÞcation.
W. Steiner, Smithsonian Institution, and N. Vandenberg
(SEL, USDA) provided information and citations relevant to
beetles. The following individuals provided comments and
suggestions that improved the manuscript: D. Davis (Smith-
sonian Institution) N. Vandenberg, and M. Pogue (SEL,
USDA).
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