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Calliduloidea, Callidulidae, Callidulinae. Biota Taiwanica: Hexapoda,

Lepidoptera

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 台灣生物誌 六足總綱
鱗翅目 錨紋蛾總科
錨紋蛾科 錨紋蛾亞科

BIOTA TAIWANICA
Hexapoda: Lepidoptera, Calliduloidea,
Callidulidae (Callidulinae)

顏聖紘、吳士緯
YEN Shen-Horn, WU Shipher

國立中山大學 行政 院 國家 科 學 委 員會
計 畫 編 號：NSC 96-2621-B-110-006-MY 2
 台灣生物誌 六足總綱
鱗翅目 錨紋蛾總科
錨紋蛾科 錨紋蛾亞科

BIOTA TAIWANICA
Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

顏聖紘、吳士緯
YEN Shen-Horn, WU Shipher

國立中山大學 行政院國家科 學委 員會
National Sun Yat-Sen University & National Science Council
 Contents

Preface 3
The family Callidulidae 4
A key to the Callidulidae species distributed in Taiwan 7
The genus Tetragonus 7
Tetragonus catamitus 9
The genus Callidula 10
Callidula attenuata 12
The genus Pterodecta 13
Pterodecta flederi 14
Appendix 1. Acronyms of the depository 18
Legends of Figure Plates 19
Fig. 1. The Callidulidae species occurring in Taiwan 20
Fig. 2. Male abdominal structures and genitalia of Callidulidae 21
Fig. 3. Female abdominal structures and genitalia of Callidulidae 22
Fig. 4. Live adult and immature stages of Callidula attenuate 23
 BIOTA TAIWANICA
2 3 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

Preface

The project “Biota Taiwanica” was first launched in 2007 under the sponsorship of the National
Science Council (NSC). The aim of the whole series is to present a contemporary account of
Taiwan’s native fauna and flora for various purposes, such as scientific research, education,
professional training, resource management and interest of the general public. Although our
knowledge of the island’s Lepidoptera fauna remains far from complete, the immediate and
potential value of the series is beyond doubt. This series is aimed to provide a synopsis of the native
Lepidoptera species of Taiwan, and to serve as a concise manual for identification. This series also
incorporates colour pictures of each taxon, dichotomous keys, diagnoses of all taxa, valid names
and synonymys, followed by references, geographical distributions and general biology and citation
of representative museum collections.

The arrangement of the whole series does not follow any taxonomic system of high category
taxa because the backbone phylogeny of the Lepidoptera has been undergoing a dramatic change
during the last 10 years. We therefore decided to take the taxa of which the monophyletic status is
confirmed or less suspicious with higher priority. In the whole series, all the synonyms that have
ever appeared in literature are included. Reference citation is generally limited to the original
publication. Chinese names of all taxa are given in traditional Chinese characters. A selected
number of museum collections examined by the author(s) are cited for each specific taxon to
represent the distribution pattern of a taxon in different areas and altitudes of Taiwan. We also
include some specimens collected outside Taiwan to show that the Taiwanese material has been
compared with other material or closely related species. The most importantly, examination of
type specimen of each taxon is included when the type material is accessible. Because the chaotic
and inconsistent anglicized spelling of locality names of Taiwan by various collectors of different
nationalities, we follow the spelling appearing on the collection labels, but give annotations or
corrections whenever necessary. The acronym for museum or private collection where specimens
are deposited is cited. The order of geographical distribution in Taiwan is given from west to east
and from north to south. The names of several areas follow current usage; for example, Sri Lanka
for Ceylon, Sulawesi for Celebes, and Myanmar for Burma. Pictures of adult specimen, genitalia,
larva, pupa, hostplant(s) and habitat are provided whenever available. The scale of the figures is not
always indicated. Some figures contain a scale bar or indicate the scale within the legend.

We acknowledge the generous and continuous financial support provided by the National Science
Council of the Taiwan (ROC). We also thank the following persons who offered great help in
various aspects during our preparation of this volume: Geoff Martin, Martin Honey, Jeremy D.
Holloway (The Natural History Museum, London), Mei-Lin Chang, Cheng-Shing Lin, Yen-Lin
Chen (National Museum of Natural Science, Taichung), Enya Chang (National Taiwan University,
Taipei), Chia-Husna Wei, Ying-Ling Tsai, Chi-Ling Su (National Sun Yat-Sen University), Jung-Tai
Chao, Shen-Shan Lu (Taiwan Forestry Research Institute, Taipei), Joël Minet (Muséum National
d'Histoire Naturelle, Paris).

Shen-Horn Yen
National Sun Yat-Sen University
July 2009
CALLIDULINAE (CALLIDULOIDEA: CALLIDULIDAE)
錨紋蛾亞科 (錨紋蛾總科：錨紋蛾科)

Shen-Horn YEN1, Shipher WU2

顏聖紘1、吳士緯2

1Department of Biological Sciences, National Sun Yat-Sen University, Kaohsiung, Taiwan

2Department of Entomology, National Taiwan University, Taipei, Taiwan

1國立中山大學生物科學系，高雄，台灣

2國立台灣大學昆蟲學系，台北，台灣

Abstract
The Callidulidae of Taiwan is revised. In total 3 species in 3 genera are recognized.
Herimba atkinsoni Moore, 1879, which was listed in Callidulidae in the checklist of
Lepidoptera of Taiwan is removed because this species has already been transferred to
Thyrididae since 1989. The present work provides a morphological diagnosis, illustrations
of adult wings and genitalia as well as the biological information of the species occurring
in Taiwan.

Keywords: Lepidoptera, Ditrysia, Obtectomera, Macrolepidoptera, Calliduloidea

The family Callidulidae Moore, 1877 (錨紋蛾科)

Callidulidae Moore, 1877, Proceedings of the Zoological Society, London: 599. (Type
genus: Callidula Hübner, [1825])

Taxonomic history of Callidulidae

The family Callidulidae has amazed lepidopterists for long because the moths of this
family posses a bizarre combination of characters, such as the “butterfly-like” appearance
with conspicuous colouration, active diurnal flight, peculiar association with the true
ferns and leaf-rolling behaviour of larvae, all making the phylogenetic affinity of this
family puzzled and intriguing.

Historically the composition and systematic position of Callidulidae/Calliduloidea have

been inconsistent in the literature since the 1920s. Before Fredrick Moore proposed the
family in 1877, the species of Callidulidae were placed in either Castniidae or Arctiidae
(Lithosiinae). Hering (1925) first suggested that Ratardidae (now as Ratardinae of
Cosiidae) could be related to Callidulidae based on several plesiomorphic or homoplasic
characters, and this association was followed by several following works, e.g. Heppner
 BIOTA TAIWANICA
4 5 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

& Wang (1987), Minet (1987a, 1989), and Heppner (1993[6]), but rejected by Holloway
(1986a), and Bradley & Carter (1987), who placed Ratardidae to be related to Cossidae.
Another alternative hypothesis proposed by Heppner (1993[6]) suggested an affinity
between the Afrotropical Hibrildidae and Calliduloidea, but the group Hibrildidae
is placed in Bombycoidea by Minet (1989, 1991), and recently Nässig & Oberpieler
(2007) treated Hibrildidae as a junior synonym of the Eupterotidae subfamily Janinae
Aurivillius, 1892.

At the present three subfamilies of Callidulidae, viz. Callidulinae, Griveaudiinae and

Pterothysaninae, are recognized (Minet, 1989[90], 1991). Minet proposed the following
seven putative apomophies to support the monophyletic status of the family: (1) antenna
bent at subapical part and turned up at apex; (2) a pair of apical foreleg spines stronger
on the fourth tarsal segment, but without well-developed spines on the fifth; (3) forewing
vein R4 reaching the termen; (4) in both wings M2 arising closer to M3 than to M1; (5)
subcostal retinaculum lacking in male of some species; (6) a sclerotised bridge uniting
valvae present and connecting ventrally to juxta; (7) female ovipositor lobes flat and
each distinctively bilobed. Minet (1991) also suggested three potential syapomorphies
to place this family closely related to the butterflies (Papilionoidea, Hesperioidea,
Hedyloidea), Drepanoidea, Geometroidea and Cimelioidea (=Axioidea): pupal labial
palpus concealed, fenestrae laterals reduced, and adult thoracic metafurca modified.
However, Tschistjakov & Belyaev’s (1987) study stated that some immature character
states, such as hypognathous larval head, frons united with clypeus, shape of prothoracic
shield, formula of crochets, and presence of maxillary palpus in pupa, are shared between
Callidulidae and Pyraloidea (Pyralidae + Crambidae).

The most recent phylogenetic hypothesis of the Lepidoptera based on five protein-
coding nuclear genes under maximum likelihood analysis (Reigier et al., 2009) suggests
a new picture of the relationships of the groups involving Calliduloidea. None of the
trees obtained from the analyses suggests a close relationship between Callidulidae
and Cimeliidae (=Axiidae) or Ratardinae of Cossidae. The placement of Callidulidae
varies with different analysis but it was never grouped with core macrolepidopterans and
sometimes grouped with Papilionoidea butterflies (Reigier et al., 2009).

Diagnosis

Adult – Head with proboscis developed in Callidulinae, but very short in Pterothysaninae;
compound eyes not hairy; antennae filiform, reaching to half of costal margin of
forewing; labial palpus developed, porrect or bent upright, acute or with a clavate third
joint, 1st segment densely scaled; ocelli small, absent in Pterothysaninae; chaetosemata
present in all subfamilies. Forewnig usually triangular with protruded or rounded apex,
termen convex, sometimes glossy without dense and flattened scales; hindwing with
costal margin straight and termen rounded or with a tip along termen; vein R3 stalked
with R4; both wings held vertically at rest in Callidulinae, but wing holding posture roof-
like in Griveaudiinae and Pterothysaninae; frenulum short; discoidal cell open in both
wings; legs scaled, hind-tibia with long spurs. Abdomen not extending beyond tornus of
hindwing, sometimes with hair brushes along spiracular like or situated between male
8th tergite and genitalic ring. Male genitalia with vinculum and saccus developed, valvae
joined mesally, gnathos incomplete, uncus narrow near apex and more or less hook-like;
aedeagus with a coecum. Female genitalia with corpus bursae variously ornamented,
ostium bursae reaching to anterior edge of 8th sternite, apophyses well developed,
ovipositor flattened and quadrilobate Minet (1989[1990)

Egg – Eggs flattened dorsoventrally, ovoid in Helicomitra (Minet, 1990), elliptical and
flattened in Griveaudiinae (D.C. Lees, pers. comm..) and Callidulinae (Nagano, 1916;
Holloway et al., 1987, Chistyakov & Belyaev, 1987, Christyakov et al., 1994).

Larva – Hypognathous; last instar larva with primary setae only; L group bisetose on
T1; A1 with a pair of lateral gland with unidentified function; prologs short with crochets
arranged in biordinal circles.

Pupa – Maxillary palpus sometimes exposed; labial palpus concealed; proboscis short
in Helicomitra or long in Callidulinae (Minet, [1998]); fore-femora concealed; midlegs
extending beyond apex of antennal sheath; abdomen with two moveable segments in
Callidulinae (A5-A6) (Minet, 1987); cremaster present in Callidulinae, but absent in
Pterothysaninae.

Biological information

Callidulidae adults may be nocturnal or diurnal (Minet, [1998]1999; Holloway, 1998).

Most early stage information for the grouap is from the Callidulinae, though a pupa
for one of the Madagascan Helicomitra Butler (Pterothysaninae) was described by
Minet (1987b). The hostplant of Helicomitra is belonging to the family Bignoniaceae
(D.C. Lees, pers. comm.). Tschistjakov & Belyaev (1987) described the early stages of
Pterodecta felderi Bremer. Callidulinae eggs are laid singly on the edge of host-plant
foliage, and the larvae roall leaflets or groups of leaflets together with silk, living and
pupating within the roll (Barlow, 1982; Holloway, 1998). The recorded host-plants of
Callidulinae are exclusively ferns, with the genera Matteucia and Osmundastrum noted
for Pterodecta felderi by Tschistjakov & Belyaev (1987). The hostplant specificity of
Callidulidae is yet to be assessed.

Global diversity and distribution

At the present about 60 species placed in 8 genera are recognized in Callidulidae. The
Pterothysaninae contains two genera, Helicomitra in Madagascar and Pterothysanus in
the Himalayan region. The Griveaudiinae contains a single genus, Griveaudia, confined
to Madagascar (Minet, [1998]1999). The Callidulinae range from India and southern
Siberia through the Indo-Australian tropics to the Solomons but not Australia (Common,
 BIOTA TAIWANICA
6 7 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

1990; Nielsen, Edwards & Rangsi, 1996; Holloway, 1998; Minet, [1998]1999). New
Guinea may harbor the highest diversity of this family (Holloway, 1998). The formation
of the present distribution pattern of Callidulidae remains debatable since Minet
(1989[1990]) suggested a double vicariant pattern in the Callidulidae. However, Holloway
(1998) considered that this interpretation contrasts strikingly with the situation in the
Castniidae, and is not consistent with the current consensus on separation of fragments
of Gondwanaland. A comprehensive phylogenetic study using appropriate molecular
markers and divergence time estimate methods may help to resolve this puzzle. The
wings of both Pterothysaninae and Callidulinae are brightly coloured, but to date there is
no evidence suggesting that their colouration is relevant to aposematism corresponding
to any secondary defense mechanism. In S.E. Asia and some south Pacific islands, some
Callidulinae species may participate in several mimicry complexes which are dominated
by unpalatable Geometridae (Ennominae) and Arctiidae (Lithosiinae) (Yen et al., 2005).

Diversity in Taiwan

In Taiwan only 3 Callidulidae species in 3 genera are recorded. In the checklist of the
Lepidoptera of Taiwan, Herimba atkinsoni Moore, 1879 was included in Callidulidae
(Inoue, 1992). This species, however, was already transferred to Thyrididae by Minet
(1989) (see also Yen & Kishida, 1997). We therefore exclude this species from the
Callidulidae list of Taiwan.

A key to the Callidulidae species distributed in Taiwan

1. Forewing upperside without a conspicuous antemedial stripe, hindwing with a short

tail formed by M3 Tetragonus catamitus
-. Forewing uppsdier with a conspicuous antemedial stripe, hindwing without a short tail
formed by M3 2
nd rd
2. 2 labial palpal segment extending anteriorly with 3 one bending downward,
forewing termen protruded at M2 Pterodecta felderi
nd rd
-. 2 labial palpal segment short, 3 segment porrect, forewing termen smooth
Callidula attenuata

The genus Tetragonus Gryer, 1832 (隱錨紋蛾屬)

Tetragonus Geyer, 1832, Zuträge zur Sammlung exotischer Schmettlinge 4: 17. (Type
species: Tetragonus catamitus Geyer)
Cleosiris Boisduval, 1836, Histoire Naturelle des Insectes. Species Général des
Lépidoptères 1: pl. 23, fig. 3, Expl. Planches 6. (Type species: Tetragonus catamitus
Geyer)
Agonis Felder, 1874, in Felder & Rogenhoder, Reise der Oesterreichischen Fregatte
Novara um die Erde in den Jahren 1857, 1858, 1859. Zoologischer Theil, 2
(Lepidoptera): pl. 107, fig. 21. (Type species: Agonis lycaenoides Felder, 1874).
Diagnosis

Adult external features – Head with compound eyes, antennae, labial palpus and
proboscis developed; antennae length about 1/3 of forewing costa; labial palpus long with
2nd segment as long as 1.5 x width of compound eye, 3rd palpal segment slender, porrect
anteriorly; costal margin of forewing smooth; apex sharp; termen slightly incised below
apex; R2 separated from R3+R4; R5 derived from discoidal cell; cross veinlet between
M1 and M2 vestigial; hindwing Sc+R1 very close to dorsal margin of cell near wing base;
discoidal cell open; M3 stalked with CuA1; 1A+2A and 3A present completely; hindwing
margin slightly protruded at M3 forming a short tail; no androconial scale patch present
on both fore- and hindwings.

Pregenital abdomen – Tergites 4-7 much broader than other segments; anterior margin
of 8th tergite sclerotized, forming an arm-like extension, 8th sternite rather membranous
and narrow with two slender arms formed by lateral margin of sternite; two pairs of
androconial hair tufts, or culcita (sensu Kobes, 1990), extending on broad patches
laterally between 8th and 7th tergites with intersegmental membrane expanded and
slightly protruded laterally.

Male genitalia – Uncus single and pointed at apex, fused with tegumen; tegument not
densely scaled dorsally; vinculum slender with dorsal part slightly fused with ventral
part of tegument, vinculum arms arched from lateral view; saccus bifid and extending
ventrally; valve broad with costal margin nearly straight, sacculus not sclerotized; juxta
(or aedeagus supporting apparatus) present as an elongated structure extending antero-
dorsally with lateral part slightly folded inwardly; manica and transtilla absent; aedeagus
stout and short, phallobase extending anteriorly, ductus ejaculatorix arising from
phallobase ventrally, vesica with a row of sharp cornuti when protruded.

Female genitalia – Papillae analis flattened cephalocaudally and slightly bilobed medially;
apophysis posterioris extending to posterior part of 7th tergite; apophysis anterioris short;
8th tergite expanding ventrally; antrum located at hind margin of 7th sternite; ostium
bursae sclerotized and swollen; ductus bursae with a corrugated sclerotization; corpus
bursae with a large ovate signum.

Global distribution and diversity

The genus Tetragonus is widely distributed from the E. Himalayan region to S.E. Asian
islands. At the present only two species are recognized (Pagenstecher, 1902).

Diversity in Taiwan

The genus has only one species distributed in Taiwan.

 BIOTA TAIWANICA
8 9 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

Biological information

The adults of Tetragonus are active day-flying moths, but occasionally attracted to light
at night. They mostly fly around edge of shady forest, where the host fern species are
abundant. Larvae are leaf-rollers and pupate inside the shelter.

Tetragonus catamitus Geyer, 1832 (隱錨紋蛾)

(Figs. 1A-B, 2A-C, 3A-B)

Tetragonus catamitus Geyer, 1832, Zuträge zur Sammlung exotischer Schmettlinge 4: 17.
(Type locality: Java. Type specimen not examined)
Cleosiris fasciata Moore, 1883, Proceedings of the Zoological Society of London 1883:
15. (Type locality: Borneo. Type specimen not examined)

Specimen examined

TAIWAN: Taoyuan County, 1♀, Fuhsing Lodge, 20-VIII-1990, B. S. Chang leg.

(NMNS); 2♀♀, Fuhsing, Paling, 24-VIII-1969, B. S. Chang leg. (NMNS); Taichung
County, 1♀, Sijiaolin, 21-X-1991, Y. B. Fan leg. (TFRI); Nantou County, 1♀, Lianhuachi,
30-V-1960, Y. C. Chang leg. (TFRI), 2♂♂, same locality, 30-V-1960, Y. C. Chang leg.
(TFRI); 1♀, same locality, 17-V-1985, Y. C. Sen leg. (TFRI); 2♂♂, same locality, 14-II-
1990, Y. B. Fan leg. (TFRI); 1♂, same locality, 23-V-1990, Y. B. Fan leg. (TFRI); 1♂3♀♀,
same locality, 4-VII-1991, Y. B. Fan leg. (TFRI); 1♂, same locality, 5-VIII-1991, Y. B. Fan
leg. (TFRI); 3♂♂, same locality, 16-VII-1992, Y. B. Fan leg. (TFRI); 1♂, Jenai Wushe,
23-10-1991, C. S. Lin leg. (NMNS); 1♀, Jenai Sungkang, 6-V-1965, B. S. Chang leg.
9NMNS); 1♀, Jenai Kaofeng, 7-V-2003, C. C. Lo leg. (NMNS); 1♂, Puli, Shihtzutou, 10-
V-2002, C. C. Lo leg. (NMNS); same locality, 1♀, 29-IX-2002, C. C. Lo leg. (NMNS);
Chiayi County, 1♂, Alishan, 17-V-1968, B. S. Chang leg. (NMNS); 1♂, Fenchifu, 1-IV-
1969, B. S. Chang leg. (NMNS); Pintung County, 1♀, Kending (=Kenting), 27-II-1992,
Y. B. Fan leg. (TFRI); Hualien County, 2♂♂1♀, locality unknown, 15-X-1991, Y. B. Fan
leg. (TFRI); 1♂, Halun, 7-VIII-1963, B. S. Chang leg. (NMNS); Ilan County, 1♂, Fushan
Botanical Garden, 27-VII-1991, Y. B. Fan leg. (TFRI).

Diagnosis

Adult – Forewing length 18-19 mm (n = 12). Wing background yellowish brown,

forewing upperside without conspicuous antemedial band running from costa to tornus
instead of an ochreous light brown zone laying between Sc+R1 and CuA1 and proximal
half of discoidal cell; underside of both fore- and hindwings having several dark brown
dots centered with silver white spot.; postmedial line irregular..

Global distribution

This species ranges throughout the E. and S.E. Asian lowland of seasonal or tropical
forests where the host fern species are abundant. Kobes (1990) recognized three
subspecies, but Holloway (1998) treated all of them as the junior synonyms of the
nominotypical subspecies and indicated there is great variation in size and colouration.

Distribution in Taiwan

This species occurs in lowland forests of Taiwan.

Biological information

According to Holloway (1998), the manuscript made by Bell provides the first description
of the immature stages of this species from S. India. Barlow (1982) also described the
larva as translucent greenish, with a dark black semi-prognathous head and a prothorax
with a well-developed prothoracic shield. Young instar larvae live in a rolled leaf, and the
mature larvae live between leaves drawn together with silk (Holloway, 1998). In Taiwn,
this species was reared from Pteridium aquilinum (L.) Kuhn subsp. latiusculum (Desv.)
Shieh, 1973 (Dennstaedtiaceae) (Duo-na, Kaohsiung, S. Wu, unpublished rearing data).

The genus Callidula Hübner, [1819]

Callidula Hübner, [1819], Verzeichniss bekannter Schmettlinge 5: 66. (Type species:

Papilio petavius Stoll,1781)
Cleis Guérin-Méneville, [1831], Voyage Coquille 2 (2): pl. 18, f. 5. (Type species: Cleis
posticalis Guérin-Méneville, [1831]); ditto: 286 [1838].
Datanga Moore, 1879, in Hewitson & Moore, Descriptions of new Indian lepidopterous
insects from the collection of the late Mr. W. S. Atkinson 1: 21. (Type species:
Datanga minor Moore, 1879)
Petavia Horsfield, [1828], Descriptions and catalogue of the lepidopterous insects of the
Museum of East India Company 1: 59, pl. 2, f. 1, 1a-f. (Type species: Petavia sakuni
Horsfield, [1828))

Diagnosis

Adult external features – Head with compound eyes, antennae, labial palpus and
proboscis developed; antennae length about 1/3 of forewing costa; labial palpus long with
2nd segment as long as 1.2 x width of compound eye, 3rd palpal segment slender, porrect
anteriorly; costal margin of forewing smooth; apex sharp; termen slightly incised below
apex; R2 stalked from R3+R4; R5 derived from discoidal cell; cross veinlet between
M1 and M2 vestigial; hindwing Sc+R1 very close to dorsal margin of cell near wing
base; discoidal cell open; M3 stalked with CuA1; 1A+2A and 3A present completely;
hindwing margin without a short tail; androconial scale patch not present on both fore-
and hindwings.
 BIOTA TAIWANICA
10 11 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

Pregenital abdomen – Tergites 4-7 much broader than other segments; anterior margin
of 8th tergite sclerotized, forming an arm-like extension, 8th sternite rather membranous
and narrow with two slender arms formed by lateral margin of sternite; two pairs of
androconial hair tufts, or culcita (sensu Kobes, 1990), extending on broad patches
laterally between 8th and 7th tergites with intersegmental membrane expanded and
slightly protruded laterally; androconial hair tufts much longer and developed than those
of Tetragonus.

Male genitalia – Uncus single and pointed at apex, fused with tegumen; tegument not
densely scaled dorsally; vinculum slender with dorsal part slightly fused with ventral
part of tegument, vinculum arms arched from lateral view; saccus not as pronounced as
in Tetragonus; valve broad with costal margin nearly straight, sacculus not sclerotized;
juxta (or aedeagus supporting apparatus) present as an elongated structure extending
antero-dorsally with lateral part slightly folded inwardly; manica and transtilla absent;
aedeagus stout and short, phallobase extending anteriorly, ductus ejaculatorix arising
from phallobase ventrally, vesica with a row of sharp cornuti when protruded.

Female genitalia – Papillae analis flattened cephalocaudally and slightly bilobed medially;
apophysis posterioris extending to posterior part of 7th tergite; apophysis anterioris very
long, extending anteriorly beyond anterior margin of 7th tergite; 8th tergite expanding
ventrally; antrum located at hind margin of 7th sternite; ostium bursae sclerotized and
swollen; ductus bursae with a corrugated sclerotization; corpus bursae without a signum.

Global distribution and diversity

The genus is diverse with more than 10 species ranging throughout the Indo-Australian
tropics east to the Solomons, but is not known from Australia (Holloway, 1998).

Diversity in Taiwan

The genus is represented by only one species in Taiwan.

Biological information

The adults are active day-fliers along the edge of secondary or primary forests. The
larvae are fern-feeders and lead-rollers (Holloway, 1998)

Callidula attenuata (Moore, 1879) (帶錨紋蛾)

(Figs. 1C-D, 2D-F, 3C-D, 4)

Petavia attenuata Moore, 1879, Descriptions of Indian Lepidoptera Heterocera from

the collection of the late Mr. WS Atkinson: 21 (Type locality: N. India. Type not
examined)
Callidula attenuata, Kirby, 1892, A synonymic catalogue of Lepidoptera Heterocera
(Moths) 1: 396; Hampson, 1892, Fauna of British India, Moths 1: 324.
Callidula formosana Wileman, 1910, The Entomologist 43: 290. (Type locality: Formosa.
Type in NHM)

Specimens examined: TAIWAN: Taipei County, 1♀, Shenkeng, II-VI-1992, Y.

B. Fan leg. (TFRI); 2♂♂, Shenkeng, II-VI-1992, Y. B. Fan leg. (TFRI); 1♂, Sindian
Feitsuishuiku, 29-VII-1984, B. S. Chang leg. (NMNS); Taoyuan County, 1♂, Fushin
Hsiapaling, 15-VIII-1981, B. S. Chang leg. (NMNS); 1♂, Fushin Kuanghua, 6-XI-1981,
B. S. Chang leg. (NMNS); Nantou County, 1♀, Yuchih Lienhuachih, 9-IX-2004, H. H.
Liang & H. Y. Chang leg. (NMNS); Chiayi County, 1♀, Chuchi Fenchifu, 9-10-I-2003,
C. S. Lin & W. T. Yang leg. (NMNS); Kaohsiung County, 2♂♂, Liugui, 29-XI-1966, Y.
C. Chang leg. (TFRI); 1♂, Shanping, 29-I-1992, Y. B. Fan leg. (TFRI); 1♂, Shanping, 16-
IV-1992, Y. B. Fan leg. (TFRI); 1♂2♀, Shanping, 30-3-XI-XII-1993, C. S. Lin & M. L.
Chang leg. (NMNS); Ilan County, 1♂, Fushan Botanical Garden, 10-VI-1992, Y. B. Fan
leg. (TFRI); 1♂, same locality, 27-IX-1995, Y. B. Fan leg. (TFRI); 1♀, same locality, 28-
XII-1992, Y. B. Fan leg. (TFRI); 1♀, Romtong, 15-II-1990, C. C. Chang leg. (NMNS).

Diagnosis

Adult – Forewing length 17-18 mm (n = 6), ground colouration darkish brown; forewing
with a reddish orange band starting from costal margin and ending at 1A+2A; underside
of both wings yellow tinged with blackish margin; forewing discoidal spots 3, silver
white centrally and surrounded by black margin; postmedial band light yellow; hindwing
discoidal spots 3 with similar arrangement as those in forewing; an ochreous area present
between antemedial to postmedial zone and bordered by 1A+2A and M1.

Global distribution

This species ranges throughout the E. Himalayan region, E. China, Taiwan and Japan.

Distribution in Taiwan
This species is distributed in the low and mid-elevation forests of the main island of
Taiwan.

Biological information:

The adults are active day-fliers in both secondary and primary forests. In Taiwan,
the immature stages were reared out from Angiopteris lygodiifolia Rosenst., 1917
(Maratiiaceae) (Wulai, Taipei, S. Wu, unpublished data, 2007).
 BIOTA TAIWANICA
12 13 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

The genus Pterodecta Butler, 1877 (錨紋蛾屬)

Pterodecta Butler, 1877, Annals & Magazine of Natural History (4) 20: 399. (Type
species: Pterodecta gloriosa Butler, 1877)

Diagnosis

Adult external features – Head with compound eyes, antennae, labial palpus and
proboscis developed; antennae length about 1/3 of forewing costa; labial palpus long
with 2nd segment as long as 2.5 x width of compound eye, 3rd palpal segment bending
downward, forming a geniculate appearance with 2nd one; costal margin slightly incised
near apex; termen pronounced with an excavate area near M2; R2 stalked with R3+R4;
R5 derived from discoidal cell; cross veinlet between M1 and M2 absent, making an
open discoidal cell; hindwing Sc+R1 very close to dorsal margin of cell near wing base;
discoidal cell open; M3 stalked with CuA1; 1A+2A and 3A present completely; no
androconial scale patch present on both fore- and hindwings.

Pregenital abdomen – Tergites 4-7 much broader than other segments; anterior margin of
8th tergite sclerotized, forming an arm-like extension, concave medially, 8th sternite rather
membranous with two slender arms formed by lateral margin of sternite; androconial
hair tufts present between 7th and 8th intersegmental menbrane.

Male genitalia – Uncus single and pointed at apex, fused with tegumen; tegument not
densely scaled dorsally; vinculum slender with dorsal part slightly fused with ventral part
of tegument; valve broad with costal margin slightly protruded, sacculus not strongly
sclerotized; juxta (or aedeagus supporting apparatus) present as an elongated structure
extending antero-dorsally with lateral part slightly folded inwardly; manica and transtilla
absent; aedeagus stout and short, phallobase not prominent, ductus ejaculatorix extending
anteriorly, a cornate process present dorsally near base of vesica; cornutus present as a
single sclerotized clavate structure.

Female genitalia – Papillae analis flattened cephalocaudally and slightly bilobed medially;
apophysis posterioris extending to middle of 7th tergite; apophysis anterioris longer than
posterial ones; antrum located at hind margin of 7th sternite; ostium bursae membranous
without sclerotized modification around; ductus bursae not obviously differentiated from
corpus bursae, a sclerotized ring present near base; corpus bursae somewhat spherical
with an appendix bursae derived posteroventrally; signum and other scobination on
corpus bursae absent.

Global diversity and distribution

According to Pagenstecher (1902), there are two species, namely Pterodecta felderi and
P. anchora Moore, 1887, being recognized in this genus. The distribution ranges from E.
Himalayan region and Russian Far East to the mountain areas of E. Asia.

Diversity in Taiwan

Only one species of Pterodecta is distributed in Taiwan.

Biological information

Adults of this genus are actively fliers in daytime. The hostplant are ferns (Felicopsida),
but species identifications are not available in most previous literature.

Pterodecta felderi (Bremer, 1864) (錨紋蛾)

(Figs. 1E-F, 2G-I, 3D-F)

Callidula felderi Bremer, 1864, Mémoires de l’Academie Imperialedes Sciences de Saint

Pétersbourg 8(1): 38, pl. 4, f. 3 (Type locality: Amurland. Type not examined)
Pterodecta gloriosa Butler, 1877, Annals & Magazine of Natural History (4)20: 399.
(Type locality: Japan, Yokohama. Type in NHM)

Specimens examined

TAIWAN: Taichung County, 1♂, Wuling, 17-IX-1992, Y. B. Fan leg. (TFRI); Taoyuan
County, 1♂, Fushin Shangpaling (=Upper Paling), 14-VIII-1981, B. S. Chang leg.
(NMNS); Kaohsiung County, 1♀, Taoyuan, Tengchih, 20-VIII-1996, W. T. Wu leg.
(NMNS); Ilan County, 1♂, Taipingshan, 26-VIII-1994, C. H. Mu leg. (TFRI); Hualien
County, 2♀♀, Hepinglindao, 16-VI-1992, Y. B. Fan leg. (TFRI)

Diagnosis

Adult – Wing length 17-18 mm (n = 5); ground colouration of both wings dark brown;
forewing upperside with a broad postmedial band, extending from costa to hind margin,
orange band broader between M2 and CuA2 with a protrusion laying between M3 and
CuA2; underside patterned with numerous cross-veined find lines, forewing yellowish
orange from wing base to submarginal zone, discoidal cell with 3 laterally compressed
spots; hindwing underside colouration less bright as that in forewing, basal and subbasal
white spots rather small compared with forewing ones, middle spot white with very fine
dark brown broader; both fore- and hindwing tips tinged white.

Global distribution:

This species has a wide distribution ranging from the Pacific coast of Russian Far East,
Japan, Korean Peninsula, palaearctic China and Taiwan.
 BIOTA TAIWANICA
14 15 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

Distribution in Taiwan

In Taiwan, this species is distributed in the mid- and high altitude mountain areas with
indigenous forests.

Biological information:

The species is actively day-flying in the primary forests in mid- and high elevation of
Taiwan. The hostplant utilized by the larvae in Taiwan is yet to be investigated.

Remarks

Chang (1989) adopted formosana Wileman as a valid subspecific name for the Taiwanese
population. However, having examined a long series of the species from Japan, Russian
and N.E. China, we failed to identify any diagnostic character to distinguish any
morphological variations corresponding to geographical regions. We therefore follow
the concept of Inoue (1992) to treat the Taiwanese population as the nominotypical
subspecies.

References

Barlow, H.S. 1982. An Introduction to the Moths of South East Asia. Kuala Lumpur: the
author.
Boisduval, [J.B.A.] 1836. Histoire Naturelle des Insectes. Species Général des
Lépidoptères. Tome Premier. Paris: Librairie Encyclopédique de Roret, 690 pp. +
24 pls.
Bremer, O. 1864. Lepidopteren Ost-sibiriens, insbesondere des Amur-landes, gesammelt
von den Herren G. Radde, R.Maack und P. Wulffius. Mémoires de l’ Academie
Imperialedes Sciences de Saint Pétersbourg (7) 8 (1): 1–103, pls. 1-8.
Butler, A.G. 1877. Descriptions of new species of Heterocera from Japan. Part I. Annals
& Magazine of Natural History Series 4, 20 (119): 393-404.
Chang, B.S. 1989. Illustrated moths of Taiwan 1. Taipei: Taiwan Provincial Museum,
242 pp.
Geyer, 1832. Zuträge zur Sammlung exotischer Schmettlinge 4: pls. [104-131], f. 601-764
(1826), pls. [132-137], figs. 765-800 (1827 - 1831), [1-3], 4-6, [7], 8-48 (1832).
Guérin-Méneville, F.E. 1831. Zoologie Atlas; Insects. pls. 1–21. In: Duperrey, L.I. (ed).
Voyage Autour du Monde, exécuté par ordre du Roi, sur la corvette de sa Majesté,
La Coquille, pendant les années 1822, 1823, 1824 et 1825, sous le minestère et
conformément aux instructions de S.E.M. le Marquis de Clermont-Tonnerre,
Ministre de la Marine; et publié sous les auspices de son Excellence Mgr le Cte de
Chabrol, Ministre de la Marine et des Colonies. Paris: A. Bertrand .
Guérin-Méneville, F.E. 1838. Insectes. pp. 57–302. In : Duperrey, L.I. (ed). Voyage
Autour du Monde, exécuté par ordre du Roi, sur la corvette de sa Majesté, La
 Coquille, pendant les années 1822, 1823, 1824 et 1825, sous le minestère et
conformément aux instructions de S.E.M. le Marquis de Clermont-Tonnerre,
Ministre de la Marine; et publié sous les auspices de son Excellence Mgr le Cte de
Chabrol, Ministre de la Marine et des Colonies. Paris: A. Bertrand.
Heppner, J.B. 1993[6]. Keys to families of Lepidoptera. Tropical Lepidoptera 4.
Supplement 3. Gainesville: Association of Tropical Lepidoptera.
Heppner, J.B., Wang, H.Y. 1987. A rare moth, Ratarda tertia Strand (Lepidoptera:
Ratardidae), from Palin. Taiwan Journal of Taiwan Museum 40: 91-94.
Hering, E.M. 1925. Die Familie der Ratardidae (Lep.), mit Beschreibung 5 neuer Arten.
Mitteilungen aus dem Zoologischen Museum in Berlin 12: 149-158.
Holloway, J.D., Bradley, J.D., Carter, D.J. 1987. Lepidoptera. CIE Guides to Insects of
Importance to Man, Volume 1 (C.R. Betts ed.). Wallingford: CAB International.
Holloway JD. 1998. The moths of Borneo: families Castniidae, Callidulidae, Drepanidae
and Uraniidae. Malayan Nature Journal 52: 1-155.
Hübner, J. 1816-1819. Verzeichniss bekannter Schmettlinge. Augsburg: Jacob Hübner.
[1], pp. 1-16 (1816), (2-8), pp. 17-128 [1819].
Hübner, J. 1818. Zuträge zur Sammlung exotischer Schmetterlinge, bestehend in
Bekundigung einzelner Fliegmuster neuer oder rarer nichteuropäischer Gattungen
3: 35 pls.
Inoue, H., Sugi, S., Kuroko, H., Moriuti, S. & Kawabe, A. 1982. Moths of Japan. 2
Volumes. Tokyo: Kodansha.
Inoue, H. 1992. Callidulidae. p. 110. In: Heppner, J.B. & Inoue, H. (eds.), Lepidoptera of
Taiwan 1 (2): Checklist. Gainesville: Association of Tropical Lepidoptera. xlix +
276 pp.
Kobes, L.W.R. 1990. The Callidulidae of Sumatra. Heterocera Sumatrana 6: 101-116.
Minet, J. 1986. Ébauche d'une classification moderne de l'ordre des Lépidoptères.
Alexanor 14: 291-313.
Minet, J. 1987a. Les Ratardidae proposés comme groupe-frère des Callidulidae (Lep.
Ditrysia). Bulletin de la Société entomologique de France 92: 39-44.
Minet, J. 1987b. Description d'une chrysalide de Pterothysaninae (Lep. Callidulidae).
Nouvelle revue d’Entomologie (N.S.) 4: 312.
Minet, J. 1989 [1990]. Nouvelles frontières, géographiques et taxonomiques, pour la
famille des Callidulidae (Lepidoptera, Calliduloidea). Nouvelle revue d’Entomologie
(N.S.) 6: 351-368.
Minet, J. 1991. Tentative reconstruction of the ditrysian phylogeny (Lepidoptera:
Glossata). Entomologica scandinavica 22: 69-95.
Moore, F. 1858. In: Horsfield, T. & Moore, F. (eds.), Catalogue of the Lepidopterous
Insects in the Museum of the Honourable East India Company 1. London: 298 pp. +
18 pls.
Moore, F. 1883. Descriptions of new genera and species of Asiatic Lepidoptera
Heterocera. Proceedings of the Scientific Meetings of the Zoological Society of
London 1883: 15-18. Pls. V-VI.
Moore, F. 1879. Descriptions of Indian Lepidoptera Heterocera from the collection of
 BIOTA TAIWANICA
16 17 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

the late Mr. WS Atkinson. London: 299 pp. + 8 pls.

Pagenstecher, A. 1887. Beiträge zur Lepidopteren-Fauna des malayischen Archipels
(IV). Ueber die Calliduliden. Jahrbücher des Nassauischen Vereins für Naturkunde
40: 205-244.
Pagenstecher, A. 1902. Callidulidae. Das Tierreich 17: ix + 1-24.
Regier, J.C., Zwick, A., Cummings, M.P., Kawahara, A.K., Cho, S., Weller, S., Roe,
A., Baixeras, J., Brown, J.W., Parr, C., Davis, D.R., Epstein, M., Hallwachs,
W., Hausmann, A., Janzen, D.H., Kitching I.J., Solis, A., Yen, S.H., Bazinet,
A.L., Mitter C. 2009. Toward reconstructing the evolution of advanced moths and
butterflies (Lepidoptera: Ditrysia): an initial molecular study. BMC Evolutionary
Biology (in press).
Tschistjakov, Yu.A., Belyaev, E.A. 1987. The immature stages of Pterodecta felderi
(Bremer) and systematic position of the family Callidulidae (Lepidoptera). Tinea 12
(Suppl): 285-289.
Wileman, A.E. 1910. New Lepidoptera-Heterocera from Formosa. The Entomologist 43:
285-291.
Yen, S.H. & Kishida, Y. 1997. Revision on Herimba atkinsoni Moore, 1879 (Thyrididae:
Striglininae) and descriptions of three new subspecies from China and Taiwan.
Transactions of the Lepidopterological Society of Japan 48 (2): 97-108.
Yen, S.H., Robinson, G.S., Quicke, D.L.J. 2005. The Phylogenetic relationships of
Chalcosiinae (Lepidoptera, Zygaenoidea, Zygaenidae), Zoological Journal of the
Linnean Society 143: 161-341.
Appendix 1. Acronyms of the depository

NHM The Natural History Museum, London, UNITED KINGDOM

NMNS National Museum of Natural Sciences, Taichung, TAIWAN
NSYSU National Sun Yat-Sen University, Kaohsiung, TAIWAN
TFRI Taiwan Forest Research Institute, Taipei, TAIWAN
 BIOTA TAIWANICA
18 19 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

Legends of Figure Plates

Fig. 1. The Callidulidae species occurring in Taiwan

Fig. 2. Male abdominal structures and genitalia of Callidulidae
Fig. 3. Female abdominal structures and genitalia of Callidulidae
Fig. 4. Live adult and immature stages of Callidula attenuata
A B

C D

E F

Fig. 1. The Callidulidae species occurring in Taiwan. A. Tetragonus catamitus Geyer, 1832; B. Callidula
attenuata (Moore, 1879); C. Pterodecta felderi (Bremer, 1864). Photo: Shipher Wu. Courtesy of
specimens: TFRI.
 BIOTA TAIWANICA
20 21 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

A B

E
D

H
G

Fig. 2. Male abdominal structures and genitalia of Callidulidae. A-C. Tetragonus catamitus; D-F. Callidula
attenuata; G-I. Pterodecta felderi; A, D & G. male genitalia without aedeagus; B, E & H. aedeagus;
C, F & I. pregenital abdomen showing the hair brushes. Photo: Shipher Wu. Courtesy of specimens:
TFRI.
A
B

Fig. 3. Female abdominal structures and genitalia of Callidulidae. A-B. Tetragonus catamitus; C-D.
Callidula attenuata; E-F. Pterodecta felderi (TFRI); A, C & E. genitalia; B, D & F. pregenital
abdomen. Photo: Shipher Wu; Courtesy of specimens: TFRI.
 BIOTA TAIWANICA
22 23 Hexapoda: Lepidoptera,
Calliduloidea, Callidulidae, Callidulinae

A B

C E

Fig. 4. Live adult and immature stages of Callidula attenuata. A. a male displaying hair brushes (Wulai,
Taipei); B. female (Muzha, Taipei); C-E. a larva making a leaf roll shelter on Angiopteris
lygodifolia (Muzha, Taipei). Photo: Enya Chang (A); Shipher Wu (B-E).
Index to taxa
Agonis 7 gloriosa 13, 14
Anchora 13 Griveaudiinae 5, 6
Angiopteris lygodiifolia 12 Hedyloidea 5
Arctiidae 4,7 Hesperioidea 5
attenuata 7,11 Helicomitra 6
Axiidae 5 Herimba atkinsoni 4
Axioidea 5 Hibrildidae 5
Callidula 4, 7, 10, 11, 14 Lithosiinae 4
Callidulidae 4, 7 Lycaenoides 7
Callidulinae 4, 7 Maratiiaceae 12
Calliduloidea 4 Matteucia 6
catamitus 7, 9 minor 10
Castiniidae 4 Osmundastrum 6
Cimeliidae 5
Papilio 10
Cimelioidea 5
Papilionoidea 5
Cleosiris 9
Petavia 10, 11
Cleis 10
Cossidae 4 Petavius 10
Crambidae 5 posticalis 10
Datanga 10 Pteridium aquilinum subsp. latiusculum 10
Dennstaedtiaceae 24 Pterodecta 6, 7, 12, 13, 14
Drepanoidea 5 Pterothysaninae 5
Ennominae 7 Pterothysanus 6
fasciata 9 Pyralidae 5
felderi 6, 7, 13, 14 Pyraloidea 5
Felicopsida 14 Ratardidae 5
formosana 12, 15 sakuni 10
Geometridae 7 Tetragonus 7, 8, 9, 11
Geometroidea 5 Thyrididae 4

台灣生物誌 Biota Taiwanica

六足總綱 鱗翅目 Hexapoda Lepidoptera
錨紋蛾總科 錨紋蛾科 錨紋蛾亞科 Calliduloidea Callidulidae Callidulinae
作者/ 顏聖紘、吳士緯 Authors/ Shen-Horn Yen, Yen-Lin Chen, Shipher Wu
封面攝影/ 吳士緯 Cover photo/ Shipher Wu
總編輯/ 張清風 Editor-in-Chief/ Chang Ching-Fong
編輯委員/ 陳天任、蔣鎮宇 Editorial board/ Tin-Yam Chan, Tzen-Yuh Chiang,
謝長富、林良恭、呂光洋 Chang-Fu Hsieh, Liang-Kong Lin, Kuang-Yang Lue,
彭鏡毅、邵廣昭、曾顯雄 Ching-I Peng, Kwang-Tsao Shao, Shean-Shong Tzean,
王震哲、吳聲華、吳文哲 Jenn-Che Wang, Sheng-Hua Wu, Wen-Jer Wu
出版/ 國立中山大學 Publisher/ National Sun Yat-Sen University
地址/ 高雄市鼓山區蓮海路70號 Address/ 70, Lien-Hai Rd., Kaohsiung 804, Taiwan
電話/ 07 - 5252000 Telephone/ 07 - 5252000

印刷/ 秋雨印刷股份有限公司 Printer/ Choice Lithograph

編排設計/ 天晴文化事業 Designer/ Dawn Cultural Enterprises
出版日期/ 中華民國九十八年七月三十一日 Publishing date/ July 31, 2009
定價/ 新台幣450元整 Price/ 450 NTD
版次/ 初版 Edition/ First edition

ISBN: 978-957-9014-39-7 (平裝)

GPN: 1009801866
ISBN:978-957-9014-39-7

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