List of the Adoliadini (Lepidoptera, Nymphalidae) of the ... · 6 Takashi Yokochi and Fidel...

INTRODUCTION

Tanaidaceans are typically benthic, free-living crusta-ceans; most are small, up to a few millimeters long. They are highly abundant in shallow-water and deep-sea bottom habi-tats, and play an important role in ecosystems (Larsen et al., 2015). Species in various animal groups have been reported as their consumers, including crustaceans, polychaetes, sea anem-ones, wading birds, and fishes (Shaffer, 1979; Oliver et al., 1982; Larsen, 2005).

Planktonic arrow worms (phylum Chaetognatha) are strictly carnivorous and occupy an important position in the marine food web (Feigenbaum and Maris, 1984). As predators, they consume mainly copepods but are not copepod specialists, and various other organisms have also been reported from their gut contents, including chaetognaths, polychaete larvae, roti-fers, cladocerans, cirripedian nauplii, ostracods, amphipods, decapod larvae, euphausiids, appendicularians, echinoderm larvae, ciliates, diatoms, dinoflagellates, infusorians, and fish larvae and postlarvae (e.g., Feigenbaum and Maris, 1984; Kehayias et al., 1996; Tonnesson and Tiselius, 2005; Nomura et al., 2007).

During a cruise of the TR/V Seisui-maru (Mie University, Japan) in 2019, one chaetognath individual containing a tanaidacean in its gut was collected. As tanaidaceans had not been previously noted among chaetognath prey items, I report this observation here.

MATERIALS AND METHODS

The chaetognath was collected on 23 April 2019 during the cruise 1903 by TR/V Seisui-maru, at Station 3B in Tanabe Bay (33°32.66'N 135°7.2'E to 33°33.67'N 135°6.07'E; 646–752 m depth), in one of two small plankton nets attached inside a larger beam trawl; the sample from Station 3B contained many other chaetognaths and tanaidaceans. The specimen was photographed and then fixed and preserved in 80% ethanol. The tanaidacean was extracted from the gut of the chaetognath by using sharpened needles, mounted on a glass slide in glycerin, and observed with an Olympus BX51 microscope. The chaetognath and tanaidacean were subsequently deposited in the Kitakyushu Museum of Natural History & Human History (KMNH IvR 600,001 and 600,002, respectively).

The tanaidacean was in too poor condition to obtain tissue for DNA extraction. Total DNA was extracted from a piece of the chaetognath’s body muscle by using a NucleoSpin Tissue XS kit (TaKaRa Bio, Japan). Attempts were made to amplify part of the cytochrome c oxidase subunit I (COI) and 18S rRNA (18S) genes by PCR using the primers listed in Table 1. PCR amplification conditions for COI with TaKaRa Ex Taq DNA polymerase (TaKaRa Bio) were 94°C for 1 min; 35 cycles of 98°C for 10 sec, 50°C for 30 sec, and 72°C for 50 sec; and 72°C for 2 min. Conditions for 18S amplification with KOD FX Neo polymerase (Toyobo, Japan) were 94°C for 2 min; 45 cycles of 98°C for 10 sec, 65°C for 30 sec, and 68°C for 1 min; and 68°C for 2 min. Methods for sequencing and sequence assembly

were as described by Tomioka et al. (2016). The nucleotide sequence obtained was deposited in the International Nucle-otide Sequence Database (INSD) through the DNA Data Bank

of Japan (DDBJ). BLAST searches (Altschul et al., 1990) were used to determine the nucleotide sequences in the INSD most similar to the sequence obtained.

RESULTS AND DISCUSSION

The tanaidacean in the chaetognath gut (Fig. 1) was very fragile, and most of its appendages were broken or lost during dissection, but I observed that it has 1) a ring-shaped pereonite 1 (p1 in Fig. 1); 2) the maxillipedal endites partly fused; 3) a ventrodistal blade-like seta on a pereopod (not pereopod 1, but one of pereopods 2–6); 4) a chelipedal fixed finger with one ventral simple seta; and 5) a developing oostegite (part of the brood pouch). Although identification to species was not possible, these features allowed identification of the female individual as Pseudotanais sp., in the family Pseudotanaidae. I morphologically identified the chaetognath specimen as belonging to the family Sagittidae, one of planktonic families (Müller et al., 2018); this was supported by the BLAST result of its 18S sequence (1620 nt long; INSD accession number LC494513) which determined a sagittid sequence in INSD as the most similar to the present sequence (the COI sequence could not be amplified).

The chaetognath I observed was collected in one of two small plankton nets attached inside a larger beam trawl. Both nets had picked up bottom sediments (Fig. 2). As female pseudotanaids are typically benthic tube-dwellers (Heard and Anderson, 2009), whereas sagittid chaetognaths are planktonic, this case of chaetognath predation on a tanaidacean may have resulted from cod-end feeding in the net.

Although chaetognaths are probably uncommon preda-tors on tanaidaceans as they usually occupy different oceanic zones, species in several tanaidacean families produce non-benthic, swimming males (Błażewicz-Paszkowycz et al., 2014), and these males would be exposed to chaetognath pre-

dation in the water column.

ACKNOWLEDGMENTS

I thank Taeko Kimura for providing the specimen used in this study; Captain Yoichi Maekawa and the crew of TR/V Seisui-maru, and researchers aboard, for support during the cruise in 2019; Matthew H. Dick for reviewing the manuscript and editing the English; and Taichiro Goto and one anonymous reviewer for critical comments on the manuscript. This study was supported in part by a KAKENHI grant (JP19K06800) from the Japan Society for the Promotion of Science (JSPS).

REFERENCES

Altschul, S. F., Gish, W., Miller, W., Myers, E. W. and Lipman, D. J. 1990. Basic local alignment search tool. Journal of Molecular Biology, 215: 403–410.

Błażewicz-Paszkowycz, M., Jenning, R. M., Jeskulke, K. and Brix, S. 2014. Discovery of swimming males of Paratanaoidea (Tanaidacea). Polish Polar Research, 35: 415–453.

Feigenbaum, D. L. and Maris, R. C. 1984. Feeding in the Chaetognatha. Oceanography and Marine Biology, An Annual Review, 22: 343–392.

Folmer, O., Black, M., Hoeh, W., Lutz, R. and Vrijenhoek, R. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3: 294–299.

Heard, R. W. and Anderson, G. 2009. Tanaidacea (Crustacea) of the Gulf of Mexico. In: Felder, D. L. and Camp, D. K. (eds.), Gulf of Mexico Origin, Waters, and Biota, Volume 1, Biodiversity, Texas A&M University Press, College Station, pp. 987–1000.

Kehayias, G., Lykakis, J. and Fragopoulu, N. 1996. The diets of the chaetognaths Sagitta enflata, S. serratodentata atlantica and S. bipunctata at different seasons in Eastern Mediterranean coastal waters. ICES Journal of Marine Science, 53: 837–846.

Larsen, K. 2005. Deep-sea Tanaidacea (Peracarida) from the Gulf of Mexico. Brill, Leiden.

Larsen, K., Guţu, M. and Sieg, J. 2015. Order Tanaidacea Dana, 1849. In: Vaupel Klein, J. C. von, Charmantier-Daures, M. and Schram, F. R. (eds.), The Crustacea. Revised and Updated, as well as Extended from the Traité de Zoologie 5, Brill, Leiden, pp. 249–329.

Maraun, M., Erdmann, G., Schulz, G., Norton, R. A., Scheu, S. and Domes, K. 2009. Multiple convergent evolution of

arboreal life in oribatid mites indicates the primacy of ecology. Proceedings of the Royal Society B, 276: 3219–3227.

Müller, C. H. G., Harzsch, S. and Perez, Y. 2018. Chaetognatha. In: Schmidt-Rhaesa, A. (ed.), Miscellaneous Invertebrates, De Gruyter, Berlin, pp. 163–282.

Nakayama, T., Watanabe, S., Mitsui, K., Uchida, H. and Inoue, I. 1996. The phylogenetic relationship between the Chlamydomonadales and Chlorococcales inferred from 18SrDNA sequence data. Phycological Research, 44: 47–55.

Nomura, H., Aihara, K. and Ishimaru, T. 2007. Feeding of the chaetognath Sagitta crassa Tokioka in heavily eutrophicated Tokyo Bay, Japan. Plankton & Benthos Research, 2: 120–127.

Oliver, J. S., Oakden, J. M. and Slattery, P. N. 1982. Phoxocephalid amphipod crustaceans as predators on

larvae and juveniles in marine soft-bottom communities. Marine Ecology Progress Series, 7: 179–184.

Puitika, T., Kasahara, Y., Miyoshi, N., Sato, Y. and Shimano, S. 2007. A taxon-specific oligonucleotide primer set for PCR-based detection of soil ciliate. Microbes and Environments, 22: 78–81.

Shaffer, P. L. 1979. The feeding biology of Podarke pugettensis (Polychaeta: Hesionidae). Biological Bulletin, 156: 343–355.

Tomioka, S., Kondoh, T., Sato-Okoshi, W., Ito, K., Kakui, K. and Kajihara, H. 2016. Cosmopolitan or cryptic species? A case study of Capitella teleta (Annelida: Capitellidae). Zoological Science, 33: 545–554.

Tonnesson, K. and Tiselius, P. 2005. Diet of the chaetognaths Sagitta setosa and S. elegans in relation to prey abundance and vertical distribution. Marine Ecology Progress Series, 289: 177–190.

4 Keiichi Kakui Bull. Kitakyushu Mus. Nat. Hist. Hum. Hist., Ser. A, 18: 5–42, March 31, 2020

List of the Adoliadini (Lepidoptera, Nymphalidae) of the Philippine Islands (Part 1)

ABSTRACT − The type-specimens of one-hundred and twenty taxa of the tribe Adoliadini from Philippines are examined and the classification of species-subspecies relations is revised. With information of the type locality and the current repository of the type or syntypes all of the taxonomic names are shown alphabetically in part 1. The annotated synonymic list is present in part 2.

KEY WORDS: Lepidoptera, Rhopalocera, Nymphalidae, Adoliadini, Philippines, Bassarona, Cynthia, Dophla, Euthalia, Lexias, Tanaecia, aconthea, adonia, albusequus, alpheda, amlana, anosia, antiquea, aquamarina, aruna, balabacana, balabacensis, boholensis, bongaoensis, borromeoi, cabigoni, calliphorus, candida, canescens, cavarna, chloe, circe, cocytina, cocytus, cusama, dacasini, damalis, darani, dhayma, dinagatensis, dinorah, dirtea, djata, dodong, dohertyi, dumaranensis, dunya, ellora, eson, eva, evelina, exul, felicitacionae, gahiti, galoa, garuda, godartii, goertzi, hikarugenzi, hinunanganensis, hiwaga, howarthi, imperator, ingae, irauana, jama, joloana, kayumanggia, kulaya, laetitiae, leopardina, leucotaenia, leytana, leytensis, liaoi, lubentina, ludonia, lupina, lusiada, macer, mahadeva, malissia, marinduque, medaga, merta, mindanaensis, mindorana, mindorensis, miscus, monara, monina, nadenya, negrosiana, nirodha, obatrata, ormocana, ornata, palawana, pallida, panayana, panopus, pardalis, pelea, phelada, philippensis, phlegethon, pinamalayana, piratica, placida, princesa, proditrix, producta, ramada, rhamases, rodriguezi, romeo, rudraca, saidja, samarensis, sarmana, satrapes, schoenigi, semperi, senilis, sibutana, sibuyana, smaragdifera, soloni, soregina, subpiratica, suluana, susoni, tanagra, tawitawia, tethys, teuta, trapesa, treadawayi, tsukada, tyawena, uposatha, virginalis, visayana, vistrica, volupia, waltraudae, yoshiroui, yui, holotype, paratype, syntype, lectotype, homonym, synonym, taxonomy.

INDEX

(PART 1)1. INTRODUCTION2. ABBREVIATIONS

3. LIST OF DESCRIBED SPECIES AND SUBSPECIES NAME

(PART 2)4. DESCRIPTIONS OF SPECIES AND SUBSPECIES

5. DISCUSSION6. ACKNOWLEDGMENTS

7. REFERENCES

Takashi Yokochi1 and Fidel Bendanillo2

1 Asian Insects Research Society (AIRS). 1-10-26, Shonan, Owariasahi, Aichi, 488-0823, Japan. E-mail: [email protected]

2 University of San Carlos Biological Museum. Learning Resource Center, Josef Baumgartner Bldg. Talamban Campus, Cebu, Republic of Philippines.

E-mail: [email protected]

(Received June 30, 2019; accepted September 6, 2019)

5

6 Takashi Yokochi and Fidel Bendanillo Adoliadini of the Philippine Is. 7

1. INTRODUCTION

The Philippine Islands are situated in the northern end of the Malay Archipelago and are composed of more than 7,000 islands, large and small, extending from north to south about 1,800 km and from east to west about 1,100 km. The insect fauna is rich in diversity and the tribe Adoliadini, Nymphalidae, Lepidoptera, the main group of this study, is no exception. The important works concerned with Philippines were, back in history, “Diagnosen einiger neuer Tagfalter von den Philippinen und Bemerkungen über die Verbreitung einiger derselben” (G. Semper, 1886–1892), and then “Die Gross-Schmetterlinge der Erde, 9” (1908–1927) edited by A. Seitz. Recently, a Japanese entomologist, E. Tsukada, had compiled a great series of “Butterflies of the South East Asian Islands” (1980–1991).

The late Prof. Julian N. Jumalon (1909–2000) (Fig. 1, left), the Lepidoptera Research Associate and Lecturer of Department of Biology, University of San Carlos, published a paper of “New Philippine butterflies” in 1975, and many new taxonomic names concerned with various families were described there. He played a leading part of the study of Philippine butterflies and contributed greatly to entomology.

In 1995, the late Mr. Colin G. Treadaway (Fig. 1, right, died in 2019) published “Checklist of the butterflies of the Philippine Islands (Lepidoptera: Rhopalocera)”. Although, 25 years have passed since his paper presented, several new findings are still remained unpublished — it can be followed by his own handwriting on the glass surface of his specimen cases —. The first author visited Senckenberg Museum in Frankfurt to research on the Treadaway collection on March 2019, and just then was informed that he had died several weeks ago at his home in England. In the middle 1990’s, every time the first author visited to Senckenberg, he came bringing his described type specimens, with his wife driving from

Limbach, about 250 km from Frankfurt. It is an undoubted fact that Mr. Treadaway and Dr. H. Schröder of Senckenberg thoroughly deserve credit for working further on systematic classification in Philippines Rhopalocera.

Here we revised the group of the tribe Adoliadini distributed in the Philippines, based on the checklist by Treadaway (1995). One hundred-twenty taxa of type specimens founded in the Philippines are examined and revised the classification of species-subspecies relation. This paper is published in two parts. All the taxa described before are examined and figured as much as possible in part 1. In the second part, synonymic lists and some important specimens will be shown, affixing several discussions.

2. ABBREVIATIONS

Public Institution (Museum, University). BMNH: The Natural History Museum, London, United Kingdom; FASU: Exhibition Hall of Food, Agricultural and Environmental Sciences, Faculty of Agriculture, Shinshu University, Nagano, Japan; KMNH: Kitakyushu Museum of Natural History & Human History, Fukuoka, Japan; MNHN: Museum National d'Histoire Naturelle Entomologie, Paris, France; SMFL: Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Frankfurt-am-Main, Germany; USC: University of San Carlos, Cebu, Philippines; ZMHU: Zoologisches Museum, Humboldt Universität, Berlin, Germany.

Private collections. ET: Etsuzo Tsukada, Yamanashi, Japan; HH: Hiroto Hanafusa, Tottori, Japan; JF: Jumalon Family, Cebu, Philippines; MKN: Misao Kaneko, Tokyo, Japan; SK: Shun-ichi Kawamura, Tokyo, Japan; STA: Shinji Taga, Tokyo, Japan; TY: Takashi Yokochi, Aichi, Japan.

Specimens concerned. FW: Length of Forewing; HT: Holotype; LT: Lectotype; PLT: Paralectotype; PT: Paratype; TL: Type Locality; TP: Type Preservation; TS: Type Series; ST: Syntype.


The following list contains the taxonomic names concerned with Philippines. The names which are described from the other areas are also included.

albusequus Nihira & Kawamura, 1986 (Fig. 2)TL: Lakit lakit, Sanga Sanga, Tawitawi.TS (TP): HT ♀ (SK).

amlana Jumalon, 1970 (Fig. 3)TL: Amlan, Negros Oriental.TS (TP): HT ♂ (USC), PT10 ♂1♀ (USC, JF).

antiquea Schröder & Treadaway, 1980 (Fig. 4)TL: Culasi, 500 m, Antique, Panay.TS (TP): HT ♂ (SMFL).

aquamarina Fruhstorfer, 1913 (Fig. 5)TL: Mindanao.TS (TP): LT ♂ (SMFL).Note: Description is based on some specimens. The lecto-

type designation was carried out by Yokochi (1998).

balabacana (Bassarona) Tsukada, 1991 (Fig. 6)TL: Balabac.TS (TP): HT ♂ (ET), PT 8♂6♀.

balabacana (Dophla) Tsukada, 1991 (secondary homonym) (Fig. 7)

Note: See below balabacensis.

balabacensis nom. nov. (Fig. 7)TL: Balabac.TS (TP): HT ♂ (ET), PT 2♂2♀.Note: The taxon balabacana was described under the

genus Dophla. As Yokochi (2010) assigned both Dophla and Bassarona to the genus Euthalia, balabacana (Dophla) is a junior secondary homonym of Bassarona teuta balabacana Tsukada, 1991. Here a new name balabacensis is replaced for Dophla evelina balabacana Tsukada, 1991.

Euthalia (Dophla) evelina balabacensis nom. nov.pro Dophla evelina balabacana Tsukada, 1991, non

Bassarona teuta balabacana Tsukada, 1991

boholensis (Lexias) M. & T. Okano, 1988 (Fig. 8)TL: Bilar, Bohol.TS (TP): HT ♂ (FASU), PT 6♂2♀.

boholensis (Euthalia) M. & T. Okano, 1990 (Fig. 9)TL: Bilar, Bohol.TS (TP): HT ♀ (FASU), PT 4♀.

bongaoensis Schröder & Treadaway, 1990 (Fig. 10)TL: Bongao Peak, Bongao, Tawitawi Group.TS (TP): HT ♀ (SMFL), PT 18♂20♀.

borromeoi Schröder, 1977 (Fig. 11)TL: Espana, 2000 ft, Sibuyan.TS (TP): HT ♂ (SMFL), PT 1♀.

cabigoni Schröder, Treadaway, A. Mohagan & N. Mohagan, 2009 (Fig. 12)

TL: Balut Is, Sarangani Group.TS (TP): HT ♂ (SMFL), PT 2♂1♀.

calliphorus C. & R. Felder, 1861 (Fig. 13)TL: Luzon.TS (TP): ST ♂ (BMNH).

candida Schröder, Treadaway & Nuyda, 1990 (Fig. 14)TL: 30 m NN, Calamian Group.TS (TP): HT ♂ (SMFL), PT10 ♂.

cavarna Fruhstorfer, 1913 (Fig. 15)TL: Balabas [Balabac].TS (TP): LT ♀ (ZMHU).

Note: Description is based on a few females, and one spec-imen is preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

chloe Schröder & Treadaway, 1990 (Fig. 16)TL: Jao Asan, 200 m, Sibuyan, Romblon Group.TS (TP): HT ♂ (SMFL), PT 11♂9♀.

circe Schröder & Treadaway, 1990 (Fig. 17)TL: Cavan Cavan, Sibutu, Tawitawi Group.TS (TP): HT ♂ (SMFL), PT 9♂10♀.

cusama Fruhstorfer, 1913 (Fig. 18)TL: Mindanao, Camiguin de Mindanao.TS (TP): ST ♂ (MNHN), ♀ (SMFL).Note: Figured specimen is a syntype female in SMFL.

dacasini Hanafusa, 1990 (Fig. 19)TL: Balabac.TS (TP): HT ♂ (HH), PT 1♀.

damalis Erichson, 1834 (Fig. 117)TL: Luzon.TS (TP): ST ♂ [untraced, not examined].Note: The type series should be preserved in ZMHU, but

the specimens that corresponded with the types could not be found. The Fig. 117 is reproduced from the original description (pl. 50, fig. 4, ♂).

darani Fruhstorfer, 1913 (Fig. 20)TL: Sulu Archipelago.TS (TP): HT ♀ (BMNH).Note: Though the original description does not mention

the number of type series, one female would be the holotype.Fig. 1. Younger age of Julian N. Jumalon (left) and Colin G. Treadaway (right).

dhayma Fruhstorfer, 1913 (Fig. 21)TL: Sulu Archipelago.TS (TP): HT ♀ (BMNH).Note: Though the original description does not mention the

number of type series, one female would be the holotype.

dinagatensis Tsukada, 1991 (Fig. 22)TL: Dinagat.TS (TP): HT ♂ (ET), PT 2♂1♀.

dinorah Fruhstorfer, 1899 (Fig. 23)TL: Basilan.TS (TP): ST 2♂2♀ (MNHN).

dodong Schröder & Treadaway, 1978 (Fig. 24)TL: Mobo, 1000 ft, Masbate.TS (TP): HT ♂ (SMFL), PT 3♂7♀.

dohertyi Butler, 1901 (Fig. 25)TL: Sulu Archipelago.TS (TP): HT ♂ (BMNH).

dumaranensis Hanafusa, 1992 (Fig. 26)TL: Dumaran.TS (TP): HT ♂ (HH).

dunya Doubleday, [1848] (Fig. 27)TL: Assam, India.TS (TP): ST ♂ (BMNH).

ellora Fruhstorfer, 1898 (Fig. 28)TL: Mindoro.TS (TP): ST ♂ (MNHN).Note: Though the original description does not mention

the number and sex, a syntype specimen in MNHN is a male.

eson de Nicéville, 1894 (Fig. 29)TL: Palawan.TS (TP): LT ♂ (SMFL), PLT 1♀.Note: Description is based on a single pair, and all

materials are preserved in SMFL. The lectotype designation was carried out by Yokochi (1998).

eva C. & R. Felder, [1867] (Fig. 30)TL: Luzon.TS (TP): ST ♂ (BMNH).Note: According to the original description, eva is distrib-

uted in Assam, Luzon, and Celebes. The details of type series are not mentioned. Staudinger (1886) is the first revisory of which Luzon is recognized as a locality of eva.

exul Tsukada & Nishiyama, 1980 (Fig. 31)TL: Bo. Melgar, Dinagat.TS (TP): HT ♂ (ET), PT 3♂6♀.

felicitacionae Jumalon, 1989 (Fig. 32)TL: Kantakpal Creek, Awihao, Ormoc, Western Leyte.TS (TP): HT ♂ (JF).

gahiti M. & T. Okano, 1990 (unavailable name) (Fig. 33)TL: Bilar, Bohol.TS (TP): HT ♀ (FASU).Note: As this taxon gahiti was described as a form of

Lexias panopus boholensis (infrasubspecific name), it should be treated as an unavailable name.

galoa Fruhstorfer, 1913 (Fig. 34)TL: Mindoro.TS (TP): ST (BMNH).Note: Though the original description does not mention

the number and sex, syntype specimen in BMNH is a male.

goertzi Jumalon, 1975 (Fig. 35)TL: Amlan, Negros Oriental.TS (TP): HT ♀ (ET).Note: Though the original description mentions the

holotype is preserved in USC, it is deposited in ET.

hikarugenzi Tsukada & Nishiyama, 1980 (Fig. 36)TL: near Baguio, N. Luzon.TS (TP): HT ♂ (MKN), PT 1♀ (MKN).

hinunanganensis M. & T. Okano, 1986 (Fig. 37)TL: Hinunangan, Leyte.TS (TP): HT ♂ (FASU), PT 1♂.

hiwaga Nuyda & Kawamura, 1989 (Fig. 38)TL: Camiguin, Babuyan.TS (TP): HT ♂ (SK), PT 7♂4♀ (SK, ET).

howarthi Jumalon, 1975 (Fig. 39)TL: Amlan, Negros Oriental.TS (TP): HT ♂ (USC), PT 4♀ (USC, JF).

imperator Hewitson, 1863 (Fig. 40)TL: Luzon.TS (TP): HT ♀ (BMNH).

ingae (Lexias) Schröder & Treadaway, 1987 (Fig. 41)TL: Negros Or. [Oriental], 800 m.TS (TP): HT ♂ (SMFL), PT 5♂5♀.

ingae (Euthalia) Schröder & Treadaway, 1990 (Fig. 42)TL: Bongao Peak, 200 m, Bongao, Tawitawi Group.TS (TP): HT ♀ (SMFL), PT 10♂12♀.

irauana Jumalon, 1975 (Fig. 43)TL: Irauan [Irawan], Puerto Princesa, Palawan.TS (TP): HT ♀ (JF), PT 1♂1♀ (USC).Note: The holotype is deposited in Jumalon family collec-

tion.

joloana Staudinger, 1889 (Fig. 44)TL: Sulu.TS (TP): LT ♂ (ZMHU).Note: Description is based on two males and one female,

and all of them are preserved in ZMHU. The lectotype desig-nation was carried out by Yokochi (1999).

kayumanggia Treadaway & Nuyda, 1994 (Fig. 45)TL: Mt. Bayog, S. Palawan.TS (TP): HT ♂ (SMFL), PT 13♂4♀.

kulaya Treadaway & Nuyda, 1994 (Fig. 46)TL: Magellanes, Homonhon.TS (TP): HT ♂ (SMFL), PT 14♂10♀.Note: The name kulaya is spelt “kúlaya” in the original

description.

laetitiae Schröder & Treadaway, 2000 (Fig. 47)TL: Cavan Cavan, 5 m, Sibutu, Tawitawi Group.TS (TP): HT ♂ (SMFL), PT 4♂.

leopardina Fruhstorfer, 1913 (Fig. 48)TL: Sulu Islands.TS (TP): HT ♀ (BMNH).

leucotaenia G. Semper, 1878 (Fig. 49)TL: Camotes.TS (TP): LT ♂ (SMFL).Note: This taxon was described based on sixty-nine

materials from Camotes, Bohol, Leyte, Panaon, E. Mindanao, and N. Mindanao. Yokochi (1998) mentioned the type series of leucotaenia was untraced so designated the lectotype for the figure of the original description, but the type series specimen was found after the third investigation in SMFL. We designate here again the lectotype for a male specimen shown in Fig. 49 (labelled “Typus (red) / Camotes / Bohol / 113 / 211 / Coll. C. Semper”).

leytana Schröder & Treadaway, 1982 (Fig. 50)TL: St. Bernard, Calpio, 280 m, S. Leyte.TS (TP): HT ♂ (SMFL), PT 18♂10♀.

leytensis Jumalon, 1975 (Fig. 51)TL: Lake Danao, Ormoc City, Leyte.TS (TP): HT ♀ (USC).

liaoi Schröder & Treadaway, 1982 (Fig. 52)TL: Mt. Madja-an [Mt. Madja-as], 500 m, Antique, Panay.TS (TP): HT ♂ (SMFL), PT 1♂2♀.

ludonia Staudinger, 1889 (Fig. 53)TL: Palawan.TS (TP): LT ♂ (ZMHU).Note: Description is based on four males and six females,

and two pairs are preserved in ZMHU. The lectotype designa-tion was carried out by Yokochi (1999).

lupina Druce, 1874 (Fig. 54)TL: Philippine Islands [Sulu].TS (TP): HT ♀ (BMNH).

lusiada C. & R. Felder, 1863 (Fig. 55)TL: Luzon.TS (TP): ST ♂ (SMFL).Note: Though the original description does not mention

the number of types, a male syntype is preserved in SMFL.

macer Tsukada & Nishiyama, 1980 (Fig. 56)TL: Bo. Melgar, Dinagat.TS (TP): HT ♂ (ET), PT 3♂2♀.

malissia Fruhstorfer, 1913 (Fig. 57)TL: Basilan.TS (TP): ST♂ (MNHN).Note: Though the original description does not mention

the number of types, a male syntype is preserved in MNHN.

marinduque K. Okano, 1992 (Fig. 58)TL: Marinduque.TS (TP): HT ♀ (KMNH), PT 1♂ (KMNH).

medaga Fruhstorfer, 1913 (Fig. 59)TL: Camiguin de Mindanao.TS (TP): ST ♂ (MNHN).Note: Though the original description does not mention


mindanaensis Schröder & Treadaway, 1978 (Fig. 60)TL: Surigao del Sur, Mindanao.TS (TP): HT ♂ (SMFL).

mindorana (Tanaecia) Fruhstorfer, 1904 (Fig. 61)TL: Mindoro.

TS (TP): ST 2♂1♀ (BMNH).Note: The figured female (Fig. 61) is preserved in the

Rothschild collection (Ro 98/7), BMNH. It might be one of the types, although the attached label is only “Mindoro”. No other specimen is found that corresponded with the type series.

mindorana (Euthalia) Fruhstorfer, 1899 (Fig. 62)TL: Mindoro.TS (TP): ST (MNHN).

mindorana (Euthalia) Tsukada, 1991 (permanently invalid)Note: The taxon mindorana described under the genus

Euthalia is a junior primary homonym of Euthalia lusiada mindorana Fruhstorfer, 1899 (permanently invalid name). Now it is replaced by tsukada Koçak (1996).

mindorensis Schröder & Treadaway, 1982 (Fig. 63)TL: Mindoro.TS (TP): HT ♂ (SMFL), PT 6♂7♀.

miscus Fruhstorfer, 1913 (Fig. 64)TL: Mindanao.TS (TP): ST (MNHN, BMNH).Note: According to the original description, types

(syntypes) are preserved in Fruhstorfer collection (now in MNHN) and BMNH.

monara Fruhstorfer, 1913 (Fig. 65)TL: N. Borneo.TS (TP): ST (MNHN).

nadenya Fruhstorfer, 1913TL: Luzon.TS (TP): HT ♀ [untraced, not examined].Note: Though the original description indicates that the

type female should be preserved in ZMHU, the specimen corresponding with nadenya could not be found there. No figure is presented in the original paper.

negrosiana Schröder & Treadaway, 1987 (Fig. 66)TL: Negros Or. [Oriental].TS (TP): HT ♂ (SMFL), PT 1♂1♀.

nirodha Fruhstorfer, 1913 (Fig. 67)TL: Basilan.TS (TP): ST ♂♀ (MNHN).

obatrata Yokochi, 1994 (Fig. 68)TL: Dumaran.TS (TP): HT ♀ (HH), PT 1♂1♀.

ormocana Jumalon, 1970 (Fig. 69)TL: Lake Danao, Ormoc City, Leyte.TS (TP): HT ♂ (USC), PT 1♀.

ornata Schröder & Treadaway, 1979 (Fig. 70)TL: Romblon.TS (TP): HT ♂ (SMFL), PT 1♂2♀.

palawana (Lexias) Moore, 1897 (Fig. 71)TL: Palawan.TS (TP): HT ♂ (BMNH).

palawana (Tanaecia) Staudinger, 1889 (Fig. 72)TL: Palawan.TS (TP): LT ♂ (ZMHU).Note: Description is based on males and females, and six

males and three females are preserved in ZMHU. The lecto-type designation was carried out by Yokochi (1999).

palawana (Euthalia) Staudinger, 1889 (Fig. 73)TL: Palawan.TS (TP): LT ♂ (ZMHU).Note: Description is based on one male and six females,

and one male and four females are preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

pallida Schröder, Treadaway & Nuyda, 1990 (Fig. 74)TL: 30 m NN, Calamian Group.TS (TP): HT ♂ (SMFL), PT 11♂16♀.

panayana Schröder & Treadaway, 1980 (Fig. 75)TL: Culasi, 800 ft, Antique, Panay.TS (TP): HT ♂ (SMFL), PT 4♂4♀.

panopus C. & R. Felder, 1861 (Fig. 76)TL: Luzon.TS (TP): ST ♂ (BMNH).Note: The original description does not mention the number of type series.

phelada G. Semper, [1887] (Fig. 77)TL: Luzon.TS (TP): LT ♀ (SMFL).Note: This taxon was described based on five materials

from Luzon, Camiguin de Mindanao, and SE. Mindanao. Yokochi (1998) mentioned the type series of phelada was untraced so designated the lectotype for the figure of the original description, but the target specimen was found after the third investigation in SMFL. We designate here again the lectotype for a female specimen shown in Fig. 77 (labelled “Typus (red) / Euth. phelada typ. ♀ Semper / Marivcles (?)

11.11.81 / 110 / Coll. C. Semper / Lectotype ♀ Euthalia phelada Semper [87]:1888 Designated by T. Yokochi, 1997”).

philippensis Fruhstorfer, 1899 (Fig. 78)TL: Basilan.TS (TP): LT ♂ (MNHN).Note: Description is based on one male from Basilan

(MNHN) and one female from Luzon (ZMHU). As the Luzon-group should be classified as nadenya. We select a male (Fig. 78) deposited in MNHN as the lectotype, labelled, “Type (red) / lubentina philippensis Fruhst. / Philippinen Bazilan II-III, 98 Doherty ex. coll. H. Fruhstorfer / MUSEUM PARIS 1934 COLL H Fruhstorfer”.

phlegethon G. Semper, [1887] (Fig. 79)TL: Mindanao.TS (TP): LT ♀ (SMFL).Note: Description is based on twenty-one males and seven

females from Samar, Camiguin de Mindanao, N. Mindanao, E. Mindanao, SE. Mindanao, and several materials are preserved in SMFL. The lectotype designation was carried out for the Mindanao specimen by Yokochi (1998).

pinamalayana Yokochi, 1997 (Fig. 80)Note: The new name pinamalayana was proposed by

Yokochi (1997), because mindorana Tsukada, 1991 was a permanently invalid name. However, pinamalayana is a junior synonym of tsukada Koçak (1996).

piratica G. Semper, [1887] (Fig. 81)TL: Camiguin de Mindanao.TS (TP): LT ♂ (SMFL).Note: Description is based on six materials from Luzon,

Mindoro, Camiguin de Mindanao, N. Mindanao. The lectotype designation was carried out for the Camiguin de Mindanao specimen by Schröder & Treadaway (1987).

princesa Fruhstorfer, 1899 (Fig. 82)TL: Palawan.TS (TP): HT ♀ (MNHN).

proditrix Fruhstorfer, 1913 (Fig. 83)TL: Basilan.TS (TP): ST ♂ (MNHN).

producta Butler, 1901 (Fig. 84)TL: Philippines [recte W. Malaysia].TS (TP): ST ♀ (BMNH).

rhamases Staudinger, 1889 (Fig. 85)TL: Palawan.

TS (TP): LT ♂ (ZMHU).Note: Description is based on males, and four males are

preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

rodriguezi Schröder & Treadaway, 1982 (Fig. 86)TL: 28 km W of Puerto Princesa, Palawan.TS (TP): HT ♂ (SMFL), PT 2♂2♀.

romeo Schröder & Treadaway, 1987 (Fig. 87)TL: Mt. Halcon, N. Mindoro.TS (TP): HT ♀ (SMFL), PT 1♀.

rudraca Fruhstorfer, 1913 (Fig. 88)TL: Balabac.TS (TP): LT ♂ (ZMHU).Note: Description is based on some specimens, and two

males and two females are preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

samarensis Tsukada, 1991 (Fig. 89)TL: San Juan, Samar.TS (TP): HT ♂ (ET), PT 3♂2♀.

sarmana Fruhstorfer, 1913 (Fig. 90)TL: Basilan.TS (TP): ST (MNHN).Note: Though the original description does not mention


satrapes C. & R. Felder, 1861 (Fig. 91)TL: Luzon.TS (TP): ST ♂ (BMNH).Note: According to the original description, the type series

(number is unknown) are deposited in Semper and Felder collections.

schoenigi Schröder & Treadaway, 1978 (Fig. 92)TL: Amlan, 600 m, Negros.TS (TP): HT ♀ (SMFL).

semperi Staudinger, 1889 (Fig. 93)TL: Palawan.TS (TP): LT ♂ (ZMHU).Note: Description is based on males and females, and four

males and four females are preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

senilis Hanafusa, 1992 (Fig. 94)TL: Dumaran.TS (TP): HT ♂ (HH), PT 1♂.

sibutana Schröder & Treadaway, 1990 (Fig. 95)TL: Cavan Cavan, 5 m, Sibutu, Tawitawi Group.TS (TP): HT ♂ (SMFL), PT 17♂15♀.

sibuyana Schröder & Treadaway, 1982 (Fig. 96)TL: Sibuyan, Romblon Group.TS (TP): HT ♂ (SMFL), PT 2♂5♀.

smaragdifera Fruhstorfer, 1913 (Fig. 97)TL: Locality unknown, presumably Mindoro [Mindoro].TS (TP): ST (MNHN).Note: Though the original description does not mention


soloni M. & T. Okano, 1990 (Fig. 98)TL: Bilar, Bohol.TS (TP): HT ♂ (FASU), PT 2♂3♀.

soregina Fruhstorfer, 1913 (Fig. 99)TL: Sulu Archipelago.TS (TP): ST ♀ (BMNH).Note: According to the original description, the type series

are exactly female, but the number is unknown.

subpiratica Schröder & Treadaway, 1987 (Fig. 100)TL: Nueva Ecija, Bongabon, Centr. Luzon.TS (TP): HT ♂ (SMFL), PT1♀.

suluana Fruhstorfer, 1902 (Fig. 101)TL: Sulu Archipelago.TS (TP): HT ♀ (BMNH).

susoni Jumalon, 1975 (Fig. 102)TL: Camp 7, Minglanilla, Cebu.TS (TP): HT ♂ (USC), PT 9♂9♀ (USC, JF, SMFL, TY).

tanagra Staudinger, 1889 (Fig. 103)TL: Palawan.TS (TP): LT ♂ (ZMHU).Note: Description is based on seven males and eleven

females, and two males and three females are preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

tawitawia Treadaway & Nuyda, 1994 (Fig. 104)TL: Magsaggao, Tawitawi.TS (TP): HT ♂ (SMFL).

tethys Tsukada, 1991 (Fig. 105)TL: N. Palawan.TS (TP): HT ♂ (ET), PT 10♂12♀.

Note: Attached label to the holotype specimen shows “Balabac”, but the correct type locality is Palawan.

trapesa G. Semper, [1887] (Fig. 106)TL: Camiguin de Mindanao.TS (TP): LT ♂ (SMFL).Note: Description is based on seven materials from E.

Mindanao. The lectotype designation was carried out for the Camiguin de Mindanao specimen by Yokochi (1998).

treadawayi Tsukada, 1991 (Fig. 107)TL: Romblon, Sibuyanes.TS (TP): HT ♂ (SMFL).

tsukada Koçak, 1996 (Fig. 80)TL: Mindoro.TS (TP): HT ♂ (ET), PT 10♂11♀.Note: Euthalia lubentina mindorana Tsukada, 1991 is a

permanently invalid name (nom. nov. pro mindorana Tsukada, 1991, non Fruhstorfer,1899). New name tsukada was proposed by Koçak (1996).

tyawena Fruhstorfer, 1913 (Fig. 108)TL: unknown [Palawan].TS (TP): LT ♀ (ZMHU).Note: Description is based on some specimens, and one

male and one female are preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

uposatha Fruhstorfer, 1913 (Fig. 109)TL: Banggi (Banguey), E. Malaysia.TS (TP): LT ♂ (ZMHU).Note: According to the original description, the type series

(male and female, but the number is unknown) are deposited in Staudinger collection (ZMHU). The lectotype designation was carried out by Yokochi (1999).

virginalis Fruhstorfer, 1913TL: locality not given.TS (TP): ST ♀ (BMNH) [not examined].Note: This is a female form of panopus. The type locality

suspected is Mindanao, though the specific location is not shown. The type series are deposited in the Rothschild collection (BMNH, London), but we have not located them at present.

visayana (Lexias) Schröder & Treadaway, 1987 (Fig. 110)TL: Catmon, S. Leyte.TS (TP): HT ♂ (SMFL), PT 6♂4♀.

visayana (Cynitia) Schröder & Treadaway, 1981 (Fig. 111)TL: St. Bernard, Catmon, S. Leyte.

TS (TP): HT ♂ (SMFL), PT 10♂2♀.

vistrica Fruhstorfer, 1913 (Fig. 112)TL: locality unknown, presumably one of the Central

Philippine islands [recte Dinagat].TS (TP): ST ♂ (BMNH).Note: Tsukada (1991) suspected this taxon is distributed

in Basilan, but it presumably comes from Dinagat.

volupia Tsukada & Nishiyama, 1981 (Fig. 113)TL: Camiguin.TS (TP): HT ♂ (ET), PT 2♂5♀.

waltraudae Schröder & Treadaway, 1990 (Fig. 114)TL: Indalawan, Balabac.TS (TP): HT ♀ (SMFL), PT 2♂2♀.

yoshiroui K. Okano, 1992 (Fig. 115)TL: Babuyan.TS (TP): HT ♂ (KMNH).

yui Yokochi, 1994 (Fig. 116)TL: Dumaran. TS (TP): HT ♂ (TY), PT 11♂4♀.

(to be continued)

1. INTRODUCTION






2. ABBREVIATIONS

Public Institution (Museum, University). BMNH: The Natural History Museum, London, United Kingdom; FASU: Exhibition Hall of Food, Agricultural and Environmental Sciences, Faculty of Agriculture, Shinshu University, Nagano, Japan; KMNH: Kitakyushu Museum of Natural History & Human History, Fukuoka, Japan; MNHN: Museum National d'Histoire Naturelle Entomologie, Paris, France; SMFL: Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Frankfurt-am-Main, Germany; USC: University of San Carlos, Cebu, Philippines; ZMHU: Zoologisches Museum, Humboldt Universität, Berlin, Germany.

Private collections. ET: Etsuzo Tsukada, Yamanashi, Japan; HH: Hiroto Hanafusa, Tottori, Japan; JF: Jumalon Family, Cebu, Philippines; MKN: Misao Kaneko, Tokyo, Japan; SK: Shun-ichi Kawamura, Tokyo, Japan; STA: Shinji Taga, Tokyo, Japan; TY: Takashi Yokochi, Aichi, Japan.

Specimens concerned. FW: Length of Forewing; HT: Holotype; LT: Lectotype; PLT: Paralectotype; PT: Paratype; TL: Type Locality; TP: Type Preservation; TS: Type Series; ST: Syntype.


The following list contains the taxonomic names concerned with Philippines. The names which are described from the other areas are also included.

albusequus Nihira & Kawamura, 1986 (Fig. 2)TL: Lakit lakit, Sanga Sanga, Tawitawi.TS (TP): HT ♀ (SK).

amlana Jumalon, 1970 (Fig. 3)TL: Amlan, Negros Oriental.TS (TP): HT ♂ (USC), PT10 ♂1♀ (USC, JF).

antiquea Schröder & Treadaway, 1980 (Fig. 4)TL: Culasi, 500 m, Antique, Panay.TS (TP): HT ♂ (SMFL).

aquamarina Fruhstorfer, 1913 (Fig. 5)TL: Mindanao.TS (TP): LT ♂ (SMFL).Note: Description is based on some specimens. The lecto-

type designation was carried out by Yokochi (1998).

balabacana (Bassarona) Tsukada, 1991 (Fig. 6)TL: Balabac.TS (TP): HT ♂ (ET), PT 8♂6♀.

balabacana (Dophla) Tsukada, 1991 (secondary homonym) (Fig. 7)

Note: See below balabacensis.

balabacensis nom. nov. (Fig. 7)TL: Balabac.TS (TP): HT ♂ (ET), PT 2♂2♀.Note: The taxon balabacana was described under the

genus Dophla. As Yokochi (2010) assigned both Dophla and Bassarona to the genus Euthalia, balabacana (Dophla) is a junior secondary homonym of Bassarona teuta balabacana Tsukada, 1991. Here a new name balabacensis is replaced for Dophla evelina balabacana Tsukada, 1991.

Euthalia (Dophla) evelina balabacensis nom. nov.pro Dophla evelina balabacana Tsukada, 1991, non

Bassarona teuta balabacana Tsukada, 1991

boholensis (Lexias) M. & T. Okano, 1988 (Fig. 8)TL: Bilar, Bohol.TS (TP): HT ♂ (FASU), PT 6♂2♀.

boholensis (Euthalia) M. & T. Okano, 1990 (Fig. 9)TL: Bilar, Bohol.TS (TP): HT ♀ (FASU), PT 4♀.

bongaoensis Schröder & Treadaway, 1990 (Fig. 10)TL: Bongao Peak, Bongao, Tawitawi Group.TS (TP): HT ♀ (SMFL), PT 18♂20♀.

borromeoi Schröder, 1977 (Fig. 11)TL: Espana, 2000 ft, Sibuyan.TS (TP): HT ♂ (SMFL), PT 1♀.

cabigoni Schröder, Treadaway, A. Mohagan & N. Mohagan, 2009 (Fig. 12)

TL: Balut Is, Sarangani Group.TS (TP): HT ♂ (SMFL), PT 2♂1♀.

calliphorus C. & R. Felder, 1861 (Fig. 13)TL: Luzon.TS (TP): ST ♂ (BMNH).

candida Schröder, Treadaway & Nuyda, 1990 (Fig. 14)TL: 30 m NN, Calamian Group.TS (TP): HT ♂ (SMFL), PT10 ♂.

cavarna Fruhstorfer, 1913 (Fig. 15)TL: Balabas [Balabac].TS (TP): LT ♀ (ZMHU).

Note: Description is based on a few females, and one spec-imen is preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

chloe Schröder & Treadaway, 1990 (Fig. 16)TL: Jao Asan, 200 m, Sibuyan, Romblon Group.TS (TP): HT ♂ (SMFL), PT 11♂9♀.

circe Schröder & Treadaway, 1990 (Fig. 17)TL: Cavan Cavan, Sibutu, Tawitawi Group.TS (TP): HT ♂ (SMFL), PT 9♂10♀.

cusama Fruhstorfer, 1913 (Fig. 18)TL: Mindanao, Camiguin de Mindanao.TS (TP): ST ♂ (MNHN), ♀ (SMFL).Note: Figured specimen is a syntype female in SMFL.

dacasini Hanafusa, 1990 (Fig. 19)TL: Balabac.TS (TP): HT ♂ (HH), PT 1♀.

damalis Erichson, 1834 (Fig. 117)TL: Luzon.TS (TP): ST ♂ [untraced, not examined].Note: The type series should be preserved in ZMHU, but

the specimens that corresponded with the types could not be found. The Fig. 117 is reproduced from the original description (pl. 50, fig. 4, ♂).

darani Fruhstorfer, 1913 (Fig. 20)TL: Sulu Archipelago.TS (TP): HT ♀ (BMNH).Note: Though the original description does not mention

the number of type series, one female would be the holotype.

8 Takashi Yokochi and Fidel Bendanillo Adoliadini of the Philippine Is. 9

dhayma Fruhstorfer, 1913 (Fig. 21)TL: Sulu Archipelago.TS (TP): HT ♀ (BMNH).Note: Though the original description does not mention the

number of type series, one female would be the holotype.

dinagatensis Tsukada, 1991 (Fig. 22)TL: Dinagat.TS (TP): HT ♂ (ET), PT 2♂1♀.

dinorah Fruhstorfer, 1899 (Fig. 23)TL: Basilan.TS (TP): ST 2♂2♀ (MNHN).

dodong Schröder & Treadaway, 1978 (Fig. 24)TL: Mobo, 1000 ft, Masbate.TS (TP): HT ♂ (SMFL), PT 3♂7♀.

dohertyi Butler, 1901 (Fig. 25)TL: Sulu Archipelago.TS (TP): HT ♂ (BMNH).

dumaranensis Hanafusa, 1992 (Fig. 26)TL: Dumaran.TS (TP): HT ♂ (HH).

dunya Doubleday, [1848] (Fig. 27)TL: Assam, India.TS (TP): ST ♂ (BMNH).

ellora Fruhstorfer, 1898 (Fig. 28)TL: Mindoro.TS (TP): ST ♂ (MNHN).Note: Though the original description does not mention


eson de Nicéville, 1894 (Fig. 29)TL: Palawan.TS (TP): LT ♂ (SMFL), PLT 1♀.Note: Description is based on a single pair, and all

materials are preserved in SMFL. The lectotype designation was carried out by Yokochi (1998).

eva C. & R. Felder, [1867] (Fig. 30)TL: Luzon.TS (TP): ST ♂ (BMNH).Note: According to the original description, eva is distrib-

uted in Assam, Luzon, and Celebes. The details of type series are not mentioned. Staudinger (1886) is the first revisory of which Luzon is recognized as a locality of eva.

exul Tsukada & Nishiyama, 1980 (Fig. 31)TL: Bo. Melgar, Dinagat.TS (TP): HT ♂ (ET), PT 3♂6♀.

felicitacionae Jumalon, 1989 (Fig. 32)TL: Kantakpal Creek, Awihao, Ormoc, Western Leyte.TS (TP): HT ♂ (JF).

gahiti M. & T. Okano, 1990 (unavailable name) (Fig. 33)TL: Bilar, Bohol.TS (TP): HT ♀ (FASU).Note: As this taxon gahiti was described as a form of

Lexias panopus boholensis (infrasubspecific name), it should be treated as an unavailable name.

galoa Fruhstorfer, 1913 (Fig. 34)TL: Mindoro.TS (TP): ST (BMNH).Note: Though the original description does not mention

the number and sex, syntype specimen in BMNH is a male.

goertzi Jumalon, 1975 (Fig. 35)TL: Amlan, Negros Oriental.TS (TP): HT ♀ (ET).Note: Though the original description mentions the

holotype is preserved in USC, it is deposited in ET.

hikarugenzi Tsukada & Nishiyama, 1980 (Fig. 36)TL: near Baguio, N. Luzon.TS (TP): HT ♂ (MKN), PT 1♀ (MKN).

hinunanganensis M. & T. Okano, 1986 (Fig. 37)TL: Hinunangan, Leyte.TS (TP): HT ♂ (FASU), PT 1♂.

hiwaga Nuyda & Kawamura, 1989 (Fig. 38)TL: Camiguin, Babuyan.TS (TP): HT ♂ (SK), PT 7♂4♀ (SK, ET).

howarthi Jumalon, 1975 (Fig. 39)TL: Amlan, Negros Oriental.TS (TP): HT ♂ (USC), PT 4♀ (USC, JF).

imperator Hewitson, 1863 (Fig. 40)TL: Luzon.TS (TP): HT ♀ (BMNH).

ingae (Lexias) Schröder & Treadaway, 1987 (Fig. 41)TL: Negros Or. [Oriental], 800 m.TS (TP): HT ♂ (SMFL), PT 5♂5♀.

ingae (Euthalia) Schröder & Treadaway, 1990 (Fig. 42)TL: Bongao Peak, 200 m, Bongao, Tawitawi Group.TS (TP): HT ♀ (SMFL), PT 10♂12♀.

irauana Jumalon, 1975 (Fig. 43)TL: Irauan [Irawan], Puerto Princesa, Palawan.TS (TP): HT ♀ (JF), PT 1♂1♀ (USC).Note: The holotype is deposited in Jumalon family collec-

tion.

joloana Staudinger, 1889 (Fig. 44)TL: Sulu.TS (TP): LT ♂ (ZMHU).Note: Description is based on two males and one female,

and all of them are preserved in ZMHU. The lectotype desig-nation was carried out by Yokochi (1999).

kayumanggia Treadaway & Nuyda, 1994 (Fig. 45)TL: Mt. Bayog, S. Palawan.TS (TP): HT ♂ (SMFL), PT 13♂4♀.

kulaya Treadaway & Nuyda, 1994 (Fig. 46)TL: Magellanes, Homonhon.TS (TP): HT ♂ (SMFL), PT 14♂10♀.Note: The name kulaya is spelt “kúlaya” in the original

description.

laetitiae Schröder & Treadaway, 2000 (Fig. 47)TL: Cavan Cavan, 5 m, Sibutu, Tawitawi Group.TS (TP): HT ♂ (SMFL), PT 4♂.

leopardina Fruhstorfer, 1913 (Fig. 48)TL: Sulu Islands.TS (TP): HT ♀ (BMNH).

leucotaenia G. Semper, 1878 (Fig. 49)TL: Camotes.TS (TP): LT ♂ (SMFL).Note: This taxon was described based on sixty-nine

materials from Camotes, Bohol, Leyte, Panaon, E. Mindanao, and N. Mindanao. Yokochi (1998) mentioned the type series of leucotaenia was untraced so designated the lectotype for the figure of the original description, but the type series specimen was found after the third investigation in SMFL. We designate here again the lectotype for a male specimen shown in Fig. 49 (labelled “Typus (red) / Camotes / Bohol / 113 / 211 / Coll. C. Semper”).

leytana Schröder & Treadaway, 1982 (Fig. 50)TL: St. Bernard, Calpio, 280 m, S. Leyte.TS (TP): HT ♂ (SMFL), PT 18♂10♀.

leytensis Jumalon, 1975 (Fig. 51)TL: Lake Danao, Ormoc City, Leyte.TS (TP): HT ♀ (USC).

liaoi Schröder & Treadaway, 1982 (Fig. 52)TL: Mt. Madja-an [Mt. Madja-as], 500 m, Antique, Panay.TS (TP): HT ♂ (SMFL), PT 1♂2♀.

ludonia Staudinger, 1889 (Fig. 53)TL: Palawan.TS (TP): LT ♂ (ZMHU).Note: Description is based on four males and six females,

and two pairs are preserved in ZMHU. The lectotype designa-tion was carried out by Yokochi (1999).

lupina Druce, 1874 (Fig. 54)TL: Philippine Islands [Sulu].TS (TP): HT ♀ (BMNH).

lusiada C. & R. Felder, 1863 (Fig. 55)TL: Luzon.TS (TP): ST ♂ (SMFL).Note: Though the original description does not mention

the number of types, a male syntype is preserved in SMFL.

macer Tsukada & Nishiyama, 1980 (Fig. 56)TL: Bo. Melgar, Dinagat.TS (TP): HT ♂ (ET), PT 3♂2♀.

malissia Fruhstorfer, 1913 (Fig. 57)TL: Basilan.TS (TP): ST♂ (MNHN).Note: Though the original description does not mention


marinduque K. Okano, 1992 (Fig. 58)TL: Marinduque.TS (TP): HT ♀ (KMNH), PT 1♂ (KMNH).

medaga Fruhstorfer, 1913 (Fig. 59)TL: Camiguin de Mindanao.TS (TP): ST ♂ (MNHN).Note: Though the original description does not mention


mindanaensis Schröder & Treadaway, 1978 (Fig. 60)TL: Surigao del Sur, Mindanao.TS (TP): HT ♂ (SMFL).

mindorana (Tanaecia) Fruhstorfer, 1904 (Fig. 61)TL: Mindoro.

TS (TP): ST 2♂1♀ (BMNH).Note: The figured female (Fig. 61) is preserved in the

Rothschild collection (Ro 98/7), BMNH. It might be one of the types, although the attached label is only “Mindoro”. No other specimen is found that corresponded with the type series.

mindorana (Euthalia) Fruhstorfer, 1899 (Fig. 62)TL: Mindoro.TS (TP): ST (MNHN).

mindorana (Euthalia) Tsukada, 1991 (permanently invalid)Note: The taxon mindorana described under the genus

Euthalia is a junior primary homonym of Euthalia lusiada mindorana Fruhstorfer, 1899 (permanently invalid name). Now it is replaced by tsukada Koçak (1996).

mindorensis Schröder & Treadaway, 1982 (Fig. 63)TL: Mindoro.TS (TP): HT ♂ (SMFL), PT 6♂7♀.

miscus Fruhstorfer, 1913 (Fig. 64)TL: Mindanao.TS (TP): ST (MNHN, BMNH).Note: According to the original description, types

(syntypes) are preserved in Fruhstorfer collection (now in MNHN) and BMNH.

monara Fruhstorfer, 1913 (Fig. 65)TL: N. Borneo.TS (TP): ST (MNHN).

nadenya Fruhstorfer, 1913TL: Luzon.TS (TP): HT ♀ [untraced, not examined].Note: Though the original description indicates that the

type female should be preserved in ZMHU, the specimen corresponding with nadenya could not be found there. No figure is presented in the original paper.

negrosiana Schröder & Treadaway, 1987 (Fig. 66)TL: Negros Or. [Oriental].TS (TP): HT ♂ (SMFL), PT 1♂1♀.

nirodha Fruhstorfer, 1913 (Fig. 67)TL: Basilan.TS (TP): ST ♂♀ (MNHN).

obatrata Yokochi, 1994 (Fig. 68)TL: Dumaran.TS (TP): HT ♀ (HH), PT 1♂1♀.

ormocana Jumalon, 1970 (Fig. 69)TL: Lake Danao, Ormoc City, Leyte.TS (TP): HT ♂ (USC), PT 1♀.

ornata Schröder & Treadaway, 1979 (Fig. 70)TL: Romblon.TS (TP): HT ♂ (SMFL), PT 1♂2♀.

palawana (Lexias) Moore, 1897 (Fig. 71)TL: Palawan.TS (TP): HT ♂ (BMNH).

palawana (Tanaecia) Staudinger, 1889 (Fig. 72)TL: Palawan.TS (TP): LT ♂ (ZMHU).Note: Description is based on males and females, and six

males and three females are preserved in ZMHU. The lecto-type designation was carried out by Yokochi (1999).

palawana (Euthalia) Staudinger, 1889 (Fig. 73)TL: Palawan.TS (TP): LT ♂ (ZMHU).Note: Description is based on one male and six females,

and one male and four females are preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

pallida Schröder, Treadaway & Nuyda, 1990 (Fig. 74)TL: 30 m NN, Calamian Group.TS (TP): HT ♂ (SMFL), PT 11♂16♀.

panayana Schröder & Treadaway, 1980 (Fig. 75)TL: Culasi, 800 ft, Antique, Panay.TS (TP): HT ♂ (SMFL), PT 4♂4♀.

panopus C. & R. Felder, 1861 (Fig. 76)TL: Luzon.TS (TP): ST ♂ (BMNH).Note: The original description does not mention the number of type series.

phelada G. Semper, [1887] (Fig. 77)TL: Luzon.TS (TP): LT ♀ (SMFL).Note: This taxon was described based on five materials

from Luzon, Camiguin de Mindanao, and SE. Mindanao. Yokochi (1998) mentioned the type series of phelada was untraced so designated the lectotype for the figure of the original description, but the target specimen was found after the third investigation in SMFL. We designate here again the lectotype for a female specimen shown in Fig. 77 (labelled “Typus (red) / Euth. phelada typ. ♀ Semper / Marivcles (?)

11.11.81 / 110 / Coll. C. Semper / Lectotype ♀ Euthalia phelada Semper [87]:1888 Designated by T. Yokochi, 1997”).

philippensis Fruhstorfer, 1899 (Fig. 78)TL: Basilan.TS (TP): LT ♂ (MNHN).Note: Description is based on one male from Basilan

(MNHN) and one female from Luzon (ZMHU). As the Luzon-group should be classified as nadenya. We select a male (Fig. 78) deposited in MNHN as the lectotype, labelled, “Type (red) / lubentina philippensis Fruhst. / Philippinen Bazilan II-III, 98 Doherty ex. coll. H. Fruhstorfer / MUSEUM PARIS 1934 COLL H Fruhstorfer”.

phlegethon G. Semper, [1887] (Fig. 79)TL: Mindanao.TS (TP): LT ♀ (SMFL).Note: Description is based on twenty-one males and seven

females from Samar, Camiguin de Mindanao, N. Mindanao, E. Mindanao, SE. Mindanao, and several materials are preserved in SMFL. The lectotype designation was carried out for the Mindanao specimen by Yokochi (1998).

pinamalayana Yokochi, 1997 (Fig. 80)Note: The new name pinamalayana was proposed by

Yokochi (1997), because mindorana Tsukada, 1991 was a permanently invalid name. However, pinamalayana is a junior synonym of tsukada Koçak (1996).

piratica G. Semper, [1887] (Fig. 81)TL: Camiguin de Mindanao.TS (TP): LT ♂ (SMFL).Note: Description is based on six materials from Luzon,

Mindoro, Camiguin de Mindanao, N. Mindanao. The lectotype designation was carried out for the Camiguin de Mindanao specimen by Schröder & Treadaway (1987).

princesa Fruhstorfer, 1899 (Fig. 82)TL: Palawan.TS (TP): HT ♀ (MNHN).

proditrix Fruhstorfer, 1913 (Fig. 83)TL: Basilan.TS (TP): ST ♂ (MNHN).

producta Butler, 1901 (Fig. 84)TL: Philippines [recte W. Malaysia].TS (TP): ST ♀ (BMNH).

rhamases Staudinger, 1889 (Fig. 85)TL: Palawan.

TS (TP): LT ♂ (ZMHU).Note: Description is based on males, and four males are

preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

rodriguezi Schröder & Treadaway, 1982 (Fig. 86)TL: 28 km W of Puerto Princesa, Palawan.TS (TP): HT ♂ (SMFL), PT 2♂2♀.

romeo Schröder & Treadaway, 1987 (Fig. 87)TL: Mt. Halcon, N. Mindoro.TS (TP): HT ♀ (SMFL), PT 1♀.

rudraca Fruhstorfer, 1913 (Fig. 88)TL: Balabac.TS (TP): LT ♂ (ZMHU).Note: Description is based on some specimens, and two

males and two females are preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

samarensis Tsukada, 1991 (Fig. 89)TL: San Juan, Samar.TS (TP): HT ♂ (ET), PT 3♂2♀.

sarmana Fruhstorfer, 1913 (Fig. 90)TL: Basilan.TS (TP): ST (MNHN).Note: Though the original description does not mention


satrapes C. & R. Felder, 1861 (Fig. 91)TL: Luzon.TS (TP): ST ♂ (BMNH).Note: According to the original description, the type series

(number is unknown) are deposited in Semper and Felder collections.

schoenigi Schröder & Treadaway, 1978 (Fig. 92)TL: Amlan, 600 m, Negros.TS (TP): HT ♀ (SMFL).

semperi Staudinger, 1889 (Fig. 93)TL: Palawan.TS (TP): LT ♂ (ZMHU).Note: Description is based on males and females, and four

males and four females are preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

senilis Hanafusa, 1992 (Fig. 94)TL: Dumaran.TS (TP): HT ♂ (HH), PT 1♂.

sibutana Schröder & Treadaway, 1990 (Fig. 95)TL: Cavan Cavan, 5 m, Sibutu, Tawitawi Group.TS (TP): HT ♂ (SMFL), PT 17♂15♀.

sibuyana Schröder & Treadaway, 1982 (Fig. 96)TL: Sibuyan, Romblon Group.TS (TP): HT ♂ (SMFL), PT 2♂5♀.

smaragdifera Fruhstorfer, 1913 (Fig. 97)TL: Locality unknown, presumably Mindoro [Mindoro].TS (TP): ST (MNHN).Note: Though the original description does not mention


soloni M. & T. Okano, 1990 (Fig. 98)TL: Bilar, Bohol.TS (TP): HT ♂ (FASU), PT 2♂3♀.

soregina Fruhstorfer, 1913 (Fig. 99)TL: Sulu Archipelago.TS (TP): ST ♀ (BMNH).Note: According to the original description, the type series

are exactly female, but the number is unknown.

subpiratica Schröder & Treadaway, 1987 (Fig. 100)TL: Nueva Ecija, Bongabon, Centr. Luzon.TS (TP): HT ♂ (SMFL), PT1♀.

suluana Fruhstorfer, 1902 (Fig. 101)TL: Sulu Archipelago.TS (TP): HT ♀ (BMNH).

susoni Jumalon, 1975 (Fig. 102)TL: Camp 7, Minglanilla, Cebu.TS (TP): HT ♂ (USC), PT 9♂9♀ (USC, JF, SMFL, TY).

tanagra Staudinger, 1889 (Fig. 103)TL: Palawan.TS (TP): LT ♂ (ZMHU).Note: Description is based on seven males and eleven

females, and two males and three females are preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

tawitawia Treadaway & Nuyda, 1994 (Fig. 104)TL: Magsaggao, Tawitawi.TS (TP): HT ♂ (SMFL).

tethys Tsukada, 1991 (Fig. 105)TL: N. Palawan.TS (TP): HT ♂ (ET), PT 10♂12♀.

Note: Attached label to the holotype specimen shows “Balabac”, but the correct type locality is Palawan.

trapesa G. Semper, [1887] (Fig. 106)TL: Camiguin de Mindanao.TS (TP): LT ♂ (SMFL).Note: Description is based on seven materials from E.

Mindanao. The lectotype designation was carried out for the Camiguin de Mindanao specimen by Yokochi (1998).

treadawayi Tsukada, 1991 (Fig. 107)TL: Romblon, Sibuyanes.TS (TP): HT ♂ (SMFL).

tsukada Koçak, 1996 (Fig. 80)TL: Mindoro.TS (TP): HT ♂ (ET), PT 10♂11♀.Note: Euthalia lubentina mindorana Tsukada, 1991 is a

permanently invalid name (nom. nov. pro mindorana Tsukada, 1991, non Fruhstorfer,1899). New name tsukada was proposed by Koçak (1996).

tyawena Fruhstorfer, 1913 (Fig. 108)TL: unknown [Palawan].TS (TP): LT ♀ (ZMHU).Note: Description is based on some specimens, and one

male and one female are preserved in ZMHU. The lectotype designation was carried out by Yokochi (1999).

uposatha Fruhstorfer, 1913 (Fig. 109)TL: Banggi (Banguey), E. Malaysia.TS (TP): LT ♂ (ZMHU).Note: According to the original description, the type series

(male and female, but the number is unknown) are deposited in Staudinger collection (ZMHU). The lectotype designation was carried out by Yokochi (1999).

virginalis Fruhstorfer, 1913TL: locality not given.TS (TP): ST ♀ (BMNH) [not examined].Note: This is a female form of panopus. The type locality

suspected is Mindanao, though the specific location is not shown. The type series are deposited in the Rothschild collection (BMNH, London), but we have not located them at present.

visayana (Lexias) Schröder & Treadaway, 1987 (Fig. 110)TL: Catmon, S. Leyte.TS (TP): HT ♂ (SMFL), PT 6♂4♀.

visayana (Cynitia) Schröder & Treadaway, 1981 (Fig. 111)TL: St. Bernard, Catmon, S. Leyte.

TS (TP): HT ♂ (SMFL), PT 10♂2♀.

vistrica Fruhstorfer, 1913 (Fig. 112)TL: locality unknown, presumably one of the Central

Philippine islands [recte Dinagat].TS (TP): ST ♂ (BMNH).Note: Tsukada (1991) suspected this taxon is distributed

in Basilan, but it presumably comes from Dinagat.

volupia Tsukada & Nishiyama, 1981 (Fig. 113)TL: Camiguin.TS (TP): HT ♂ (ET), PT 2♂5♀.

waltraudae Schröder & Treadaway, 1990 (Fig. 114)TL: Indalawan, Balabac.TS (TP): HT ♀ (SMFL), PT 2♂2♀.

yoshiroui K. Okano, 1992 (Fig. 115)TL: Babuyan.TS (TP): HT ♂ (KMNH).

yui Yokochi, 1994 (Fig. 116)TL: Dumaran. TS (TP): HT ♂ (TY), PT 11♂4♀.

(to be continued)

1. INTRODUCTION






2. ABBREVIATIONS

Public Institution (Museum, University). BMNH: The Natural History Museum, London, United Kingdom; FASU: Exhibition Hall of Food, Agricultural and Environmental Sciences, Faculty of Agriculture, Shinshu University, Nagano, Japan; KMNH: Kitakyushu Museum of Natural History & Human History, Fukuoka, Japan; MNHN: Museum National d'Histoire Naturelle Entomologie, Paris, France; SMFL: Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Frankfurt-am-Main, Germany; USC: University of San Carlos, Cebu, Philipp

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