No. 78（2018） E-discovery of final-instar larvae and pupa of Tacola larymna (Nymphalidae, Limenitidinae) in Bornean Malaysia and central Thailand Keith V. Wolfe 616 Alumrock Drive, Antioch, California 94509, USA : bflyearlystages@comcast.net Abstract The early stages of Tacola larymna are now known thanks to the power of social media, with the vegetative hostplant in Thailand determined to be probably Olea (Oleaceae). Keywords Athyma, Auzakia, Moduza, Olea, Tacola larymna elisa, Tacola larymna siamensis. The limenitidine genus Tacola Moore is comprised of how the Internet and social media can be scientifically three species that range across most of insular Southeast employed in the absence of traditional fieldwork. N.B.: Asia – Tacola larymna Doubleday: Malay Peninsula, After completion of the manuscript, I saw private pictures Sumatra, Java, Borneo, Palawan, and Western Visayas; of the equally atypical larva and pupa of Auzakia danava Tacola eulimene Godart: Sulawesi and Moluccas; and Moore from Godawari, Nepal (ms in prep.). Tacola magindana Semper: Philippines (Mindanao Region sensu Vane-Wright, 1990) (Tsukada, 1991; Treadaway & Schroeder, 2012). The type species is T. larymna, which was originally described as Limenitis Fabricius (Doubleday & Westwood, [1848] - 1852), but thereafter assigned to Athyma Westwood (Moore, 1858). Forty years later, apparently citing distinctive wing characters, Moore erected the genus Tacola (Moore, 1898) for T. larymna and its congeners, a proposal that numerous modern authors (D'Abrera, 1985; Otsuka, 1988; Corbet & Pendlebury, 1992; etc.) elected not to follow, preferring instead his earlier Athyma placement. However, an examination of their wing venation, male genitalia (e.g., "phallus thin and long [versus] thick and short"), and island distribution convinced Tsukada (1991) that 1 a separate genus was indeed justified, thus resurrecting Tacola. Additionally, as part of a cladistic analysis of Neotropical Adelpha Hübner, Willmott (2003b) dissected a male T. larymna from Malaysia, stating that similar to Auzakia Moore, Tacola lacks a clunicula and the aedeagus is distinct from all other Limenitidini. To this conversation I now add the first record of Tacola preadults – the extraordinary last-instar caterpillar and chrysalis of T. larymna, both of which appear dramatically different from any related taxa. It is important to note that the following is yet another example (see Wolfe, 2017) of 52 2 Figs. 1 - 2. Final-instar Tacola larymna elisa. 1. dorsal view. 2. dorsolateral view. Gunung Mulu National Park, Sarawak, Borneo. 18. ⅲ. 2009. © Paul Bertner, used with permission. No. 78（2018） While perusing the online images of longtime F LICKR approximately 580 km from Paul's earlier spotting, and (https://www.flickr.com/) friend Paul Bertner, I discovered uploaded to the G OPENG N ATURAL TRIP (https://www. an amazing caterpillar from Gunung Mulu National Park facebook.com/groups/gopenggnt/) Facebook group. It that was festooned with elaborate fleshy and seemingly too was described as really big and "looking like birds' sclerotized processes plus a befitting head capsule to droppings at first glance." From the beginning, I shared match (Figs. 1 - 2), clearly a never-before-seen limenitid my information and thoughts with Motomu Teshirogi, a or a remarkable lookalike immature moth. Paul replied to dear friend and well-traveled colleague who is an expert my email that because the encounter occurred years ago, on nymphalid butterflies (Teshirogi, 2016, etc.). His he could not recall anything concerning the foodplant, best guess was a deformed Moduza procris Cramer that adding "the bird dropping [larva was] about 4 - 5 inches failed to molt properly, an excellent contender given their and had the curl-up-into-a-ball defensive display." significant morphological resemblance (Fig. 4). This Notwithstanding that field estimates of length can be second Sarawak caterpillar, however, effectively removed problematic, thus often unreliable even under the best all such bodily doubts, but not the probability of it being a circumstances, the remembered size of four to five inches moth; only a resulting chrysalis or confirmed adult would suggested that the caterpillar was in fact a moth, or it prove its lepidopteran kinship. represented the overstated length of a nonetheless very big butterfly. So if the latter, which genus and species? That much-anticipated opportunity finally arose on Having lived in Asia for nine years and personally reared another Facebook group, this time C ATERPILLARS or observed a good many of the T HAILAND หนอนผี เ สื ้ อ ในประเทศไทย (https://www.facebook. OF region's Limenitidini early stages then and since, I was eventually able to confidently eliminate all widely accepted possibilities based upon either larval appearance (Adelpha [Neotropical], Athyma, Lebadea C. Felder, Limenitis, Moduza Moore [but see below], Pandita Moore, Parasarpa Moore, Patsuia Moore, S u m a l i a M o o r e , a n d Ta r a t t i a Moore) or reported geographic range (Auzakia [not Borneo], Litinga Moore [China], and Lamasia Moore [Sulawesi]) – all, that is, except Tacola. Specifically T. larymna, the so-called Great Sergeant, the 3 Fig. 3. Final-instar Tacola larymna elisa, dorsal view. Kuching, Sarawak, Borneo. ⅴ. 2014. © Clement Sim, used with permission. largest Athyma, where in Borneo flies Tacola larymna elisa Fruhstorfer, an apparently uncommon species that D'Abrera (1985) calls "a giant race, especially in the ♀." Hope aside, the odds of it being a moth caterpillar were still greater than 50 percent. The mystery would lie dormant for three years until an identical larva (Fig. 3) was photographed by Clement Sim in Kuching, 4 Fig. 4. Comparison of final-instar Moduza procris (left) on Mussaenda sp. (Rubiaceae): Mumbai, Maharashtra, India, and final-instar Tacola larymna (right) on unknown hostplant: Kuching, Sarawak, Borneo. 53 No. 78（2018） com/groups/390324371076210/), with the posting of narrowly acuminate, base acute, matt in appearance, no an unknown larva found by Paradorn Dokchan, an aromatic smell (pellucid glands) when crushed; venation entomological researcher at Kasetsart University's prominent with intramarginal veins absent; no stipules; Kamphaeng Saen Campus. Four last-instar caterpillars petioles distinct; stems pale. Other families considered said to each measure around 4.5 cm were collected from only to be readily disqualified were the Euphorbiaceae neighboring small trees of the same species, their basic and Phyllanthaceae, as eventually were Memecylon colors being seashell white, peach (Fig. 5), chestnut, and Linnaeus (Melastomataceae) and Syzygium P. Browne ex blackish taupe, surprisingly variable compared to the Gaertner (Myrtaceae), two tricky genera that contain trees above two independent encounters in Borneo. Cuttings superficially similar to Olea. sans flowers or fruits were taken to a local herbarium for determination, but the specimen was allegedly lost. The Oleaceae is an ancient family in the order Lamiales Despite Paradorn's numerous return visits to the same and consists of 600+ species in 25 extant genera with a marked plants (Figs. 6 - 7) in Khao Yai National Park, reproductive parts remain unseen. Realizing that field pictures of strictly vegetative material are always a challenge to identify, I ultimately contacted 11 botanists in seven different countries, many of whom are authorities on the individual plant families proffered and employed at such respected institutions as the Naturalis Biodiversity Center (Netherlands), Royal Botanic Garden Edinburgh (Scotland), Department of National Parks' Office of the Forest Herbarium (Thailand), and National University of Singapore. After multiple emails, photos, and followup discussions, the subject immature/sterile hostplant is strongly suspected to be one of Thailand's five species of Olea Linnaeus (Oleaceae) – specifically Olea brachiata 6 (Loureiro) Merrill or Olea salicifolia Wallich ex G. Don – based upon these characteristics: leaves simple, opposite (few subopposite), mostly entire with scattered, somewhat serrated margins on distal half; leaf apex 7 5 Fig. 5. Final-instar Tacola larymna siamensis, dorsal view. Ex Khao Yai National Park, Nakhon Ratchasima, Thailand. 3. ⅸ. 2014. © Paradorn Dokchan (cell phone photo), used with permission. 54 Figs. 6 - 7. Immature/sterile hostplant (see text) of Tacola larymna siamensis. 6. Branch. 7. Close-up. Evergreen hill, 700 m above sea level, Khao Yai National Park, Nakhon Ratchasima, Thailand. 5. ⅺ. 2015. © Paradorn Dokchan (cell phone photos), used with permission. No. 78（2018） 2010). The sole exception is Athyma ranga Moore, which feeds exclusively on the Oleaceae, namely Chionanthus Linnaeus, Fraxinus Linnaeus, Ligustrum Linnaeus, Olea, and Osmanthus Loureiro (Robinson et al., 2001; Robinson et al., 2017). For a comprehensive review of the Oleaceae and iridoid glucosides (a subgroup of glycosides), bitter chemical compounds with presumed antifeedant 8 Fig. 8. Pupa of Tacola larymna siamensis, dorsal view. Ex Khao Ya i N a t i o n a l P a r k , Nakhon Ratchasima, Thailand. 8. ⅸ. 2014. © Paradorn Dokchan (cell phone photo), used with permission. and growth inhibiting properties that were present in the leaves of virtually every oleaceous taxon tested, sometimes in high concentrations, see Jensen et al., 2002. As an aside, the ±90 recognized species of New World Adelpha limenitids are currently reported to eat 89 genera in 27 different and diverse plant families, including four in the Lamiales: Acanthaceae, Bignoniaceae, Lamiaceae, and Verbenaceae, yet not a single Neotropical Oleaceae (Constantino, 2002; Willmott, 2003a; Beccaloni et al., 2008; Oliveira-Neto et al., 2016; Janzen & Hallwachs, worldwide distribution (Wallander & Albert, 2000), while 2018). the 33 species of Olea (Green, 2002; Kiew, 2016) are native to warm temperate and tropical areas of the Old Wild-collected larvae are often parasitized or otherwise World. Interestingly, a frequently overlooked article by die of disease, particularly under artificial laboratory Morrell (1954) records O. brachiata (as Olea "maritima" conditions in tropical climates, so it was with immense Wallich ex G. Don) as a foodplant for Moduza procris relief that the four Thai caterpillars successfully pupated in Singapore, a widespread butterfly said to be confined five days later. And what an incredible chrysalis it was mainly to the Rubiaceae. Altogether, just a handful of – completely unlike any known butterfly or moth (Fig. Nymphalidae* are reported to use the Oleaceae, with 8)! Motomu and I spent the next week exchanging the original unsourced ithomiine listing almost certainly various published and unpublished images of comparable erroneous (however, see Forbes, 1958: 316) and the few limenitidine pupae, resulting in a composite (Fig. 9 others probably reflecting isolated instances or secondary excluding Auzakia danava, which was discovered only utilization of primarily two ornamental genera (e.g., recently and is added here for reference) that he then took Braby, 2000; Igarashi & Fukuda, 2000; Robinson et al., to 東京大学総合研究博物館 (The University Museum, 9 Fig. 9. Representative similar-appearing Asian Limenitidinae pupae, those of other closely related genera Athyma, Lebadea, Limenitis, Pandita, Parasarpa, Sumalia, and Tarattia being significantly different. Left to right: Neptis alwina Bremer & Grey (China), Neptis sankara Kollar (China), Patsuia sinensium Oberthür (China), Moduza procris (Malaysia), Auzakia danava (Nepal), and Tacola larymna (Thailand). 55 No. 78（2018） for numerous hours of interesting and valuable debate regarding not just the morphology and taxonomic position of Tacola, but also many other nymphalid genera over the years. A sincere "khop-khun-mak" to Paradorn Dokchan (Kamphaeng Saen, Thailand) for his patience in addressing my myriad questions and requests concerning the T. l. siamensis. Blaise Pereira (Mumbai, India) kindly 10 Fig. 10. Resulting adult female of Tacola larymna siamensis, dorsal view. Ex Khao Yai National Park, Nakhon Ratchasima, Thailand. 21.ⅸ. 2014. © Paradorn Dokchan (cell phone photo), used with permission. provided the larval photo of M. procris used in Fig. 4, while Motohiro Harada (Kamakura, Japan) graciously supplied the pupal pictures of Neptis Fabricius and Patsuia, as did L. C. Goh (Shah Alam, Malaysia) of M. procris and Huang Hang-Chi (Taipei, Taiwan) of A. danava, for Fig. 9. The University of Tokyo) for comments from several References Japanese lepidopterists who were working there on a Beccaloni, G. W., Viloria, Á. L., Hall, S. K. & Robinson, special project. The consensus was indeed a limenitid, G. S., 2008. Catalogue of the hostplants of the NOT a moth, the proof coming in 13 days when a very Neotropical butterflies/Catálogo de las plantas large female Tacola larymna siamensis Fruhstorfer huésped de las mariposas neotropicales. Monografías eclosed (Fig. 10). One voucher specimen is awaiting Tercer Milenio, Vol. 8. Sociedad Entomológica deposition at Kasetsart University's Department of Aragonesa, Zaragoza. Entomology in Kamphaeng Saen, whereas the other three Bell, T. R., 1910. The common butterflies of the plains of raised adults were returned to Khao Yai National Park. India (including those met with in the hill stations of I now defer to future, more knowledgeable researchers Hist. Soc. 19(4): 846-879. the Bombay Presidency), Part VII. J. Bombay Nat. to explicate the classification of Tacola and its subtribal Braby, M. F., 2000. Butterflies of Australia: Their relationship with Moduza, Auzakia, and other possible Identification, Biology and Distribution; Volume Two. basal members of the Limenitidini. CSIRO Publishing, Collingwood. Constantino, L. M., 2002. Utilización de Trichanthera * Methona themisto Hübner – Jasminum Linnaeus gigantea como alimento para cría de seis especies Charaxes sempronius Fabricius – Jasminum de mariposas diurnas. Pp. 195-203. In: Ospina, S. D. Athyma asura Moore – Chionanthus & Murgueitio R., E. (Eds.). Tres Especies Vegetales Nymphalis antiopa Linnaeus – Fraxinus Promisorias: Nacedero Trichanthera gigantea (H. & Vanessa annabella Field – Ligustrum B.) Nees, Botón de oro Tithonia diversifolia (Hemsl.) Euphydryas maturna Linnaeus – Fraxinus, Ligustrum, Gray y Bore Alocasia macrorrhiza (Linneo.) Schott. Syringa Linnaeus Fundación CIPAV, Cali. Euphydryas phaeton Drury – Fraxinus Bell's (1910) record of the Oleaceae for Neptis hylas Linnaeus (as Neptis "eurynome" Westwood), which was repeated by Wynter-Blyth (1957 page 192, but not page 495) and others, has always been an error for the Olacaceae, with "Mappia foetida" (Wight) Miers presently treated as Nothapodytes nimmoniana (Graham) Mabberley in the Icacinaceae. Corbet, A. S. & Pendlebury, H. M., 4th Edition revised by Eliot, J. N., 1992. The Butterflies of the Malay Peninsula. Malayan Nature Society, Kuala Lumpur. D'Abrera, B., 1985. Butterflies of the Oriental Region, Part II: Nymphalidae, Satyridae & Amathusidae. Hill House, Melbourne. Doubleday, E. & Westwood, J. O., [1848]-1852. The Genera of Diurnal Lepidoptera, Vol. II. Longman, Brown, Green, and Longmans, London. Acknowledgements My hearty thanks to Motomu Teshirogi (Tokyo, Japan) 56 Forbes, W. T. M., 1958. Caterpillars as Botanists. Pp. 313317. In: Becker, E. C. (Ed.). Proceedings of the Tenth No. 78（2018） International Congress of Entomology, Volume 1. Montreal. Green, P. S., 2002. A revision of Olea L. (Oleaceae). Kew Bull. 57(1): 91-140. Igarashi, S. & Fukuda, H., 2000. The Life Histories of Asian Butterflies Vol. 2. Tokai University Press, Tokyo. lidae&PFamily=Oleaceae accessed 31 March 2018. Robinson, G. S., Ackery, P. R., Kitching, I. J., Beccaloni, G. W. & Hernández, L. M., 2017. HOSTS. http:// hosts.myspecies.info/taxonomy/term/42449 accessed 31 March 2018. Teshirogi, M., 2016. Nymphalid Butterflies of the World: Eggs, Larvae, Pupae, Adults and Host Plants. database for an inventory of the macro caterpillar Hokkaido University Press, Sapporo (in Japanese). Treadaway, C. G. & Schroeder, H. G., 2012. Revised fauna of Área de Conservación Guanacaste, checklist of the butterflies of the Philippine Islands northwestern Costa Rica. http://janzen.sas.upenn. (Lepidoptera: Rhopalocera). Nachr. Entomol. Ver. Janzen, D. H. & Hallwachs, W., 2018. Dynamic edu/caterpillars/dblinks/searchplaycat4.lasso?Search=GCAcaterpillars337&herbivore%20 species=Adelpha accessed 31 March 2018. Jensen, S. R., Franzyk, H. & Wallander, E., 2002. Chemotaxonomy of the Oleaceae: iridoids as taxonomic markers. Phytochemistry 60(3): 213-231. Apollo, Suppl. 20: 1-64. Tsukada, E., 1991. Butterflies Of The South East Asian Islands V, Nymphalidae (II). Azumino Butterflies Research Institute, Matsumoto (in Japanese). Vane-Wright, R. I., 1990. The Philippines – Key to the biogeography of Wallacea? Pp. 19-34. In: Knight, Kiew, R., 2016. Olea luzonica (Oleaceae), a new name W. J. & Holloway, J. D. (Eds.). Insects and the Rain for O. obovata from the Philippines. Gard. Bull. Forests of South East Asia (Wallacea). The Royal Singapore 68(1): 97-98. Moore, F., 1858. Monograph of the Asiatic species of Entomological Society of London, London. Wallander, E. & Albert, V. A., 2000. Phylogeny and Neptis and Athyma, two genera of diurnal Lepidoptera classification of Oleaceae based on rps16 and trnL-F belonging to the family Nymphalidae. Proc. Zool. sequence data. Am. J. Bot. 87(12): 1827-1841. Soc. Lond. 26: 3-20. Moore, F., 1898. Lepidoptera Indica, Vol. III, Parts 32-34. Lovell Reeve & Co., Ltd., London. Morrell, R., 1954. Notes on the larval habits of a group of nymphalid butterflies. Malay. Nat. J. 8(4): 157-164. Willmott, K. R., 2003a. The Genus Adelpha: Its Systematics, Biology and Biogeography (Lepidoptera: Nymphalidae: Limenitidini). Scientific Publishers, Gainesville. Willmott, K. R., 2003b. Cladistic analysis of the Oliveira-Neto, J. F., Dias, F. M. S., Pawoski, E. L., Neotropical butterfly genus Adelpha (Lepidoptera: Oliveira, P., Casagrande, M. M. & Mielke, O. H. H., Nymphalidae), with comments on the subtribal 2016. Immature Stages of Six Species of Adelpha classification of Limenitidini. Syst. Entomol. 28(3): Hübner (Lepidoptera: Nymphalidae: Limenitidinae) 279-322. from Southern Brazil Coastal Forests. Neotrop. Entomol. 45(5): 524-536. Otsuka, K., 1988. Butterflies of Borneo, Vol. 1. Tobishima Corporation, Tokyo. Wolfe, K. V., 2017. Probable last-instar larvae of Opoptera staudingeri mexicana (Nymphalidae, Satyrinae, Brassolini). News Lepid. Soc. 59(1): 2829. N.B.: Identity subsequently verified by photos and Robinson, G. S., Ackery, P. R., Kitching, I. J., Beccaloni, pers. comm. with Rolando Cubero regarding his in G. W. & Hernández, L. M., 2001. Hostplants of the prep. comprehensive paper on the life history of Costa moth and butterfly caterpillars of the Oriental Region. The Natural History Museum, London, and Southdene Sdn. Bhd., Kuala Lumpur. Robinson, G. S., Ackery, P. R., Kitching, I. J., Beccaloni, Rican O. s. staudingeri. Wynter-Blyth, M. A., 1957. Butterflies of the Indian Region. The Bombay Natural History Society, Bombay. G. W. & Hernández, L. M., 2010. HOSTS - A Database of the World's Lepidopteran Hostplants. Natural History Museum, London. http://www. nhm.ac.uk/our-science/data/hostplants/search/list. dsml?searchPageURL=index.dsml&Family=Nympha 57 No. 78（2018） ［和文摘要］ し，そのうちタイ産については食草も特定すること ボルネオ及びタイ中部で撮影されたオニミスジの終 ができた。これはモクセイ科オリーブ属 Olea の 1 種 齢幼虫と蛹 と思われる。 オニミスジ Tacola larymna は，ミャンマー，イン タテハチョウ科でモクセイ科を食草とする種は複 ドシナからフィリピン，ボルネオ，ジャワ等の島嶼 数知られているが，その例は少なく，かつ副次的な に分布するタテハチョウ科の大形種である。本種を 利用にとどまっている。ただし，モクセイ科を主食 Athyma 属に含める場合もあるが，本稿では斑紋や交 草とするカワリイチモンジ Athyma ranga のような例 尾器の形質の差異を考慮し，別属 Tacola として扱う 外的な種も存在する。 見解を支持する。 本種の蛹の形態は，近縁他種と比較しても特異で 本種の幼生期はこれまで知られていなかったが， インターネットの画像を活用した調査により，終齢 幼虫及び蛹の形態が判明したので報告する。 あり，種名が判明するまでは「もしかしたら蛾では ないか？」という疑念を抱いたほどであった。 な お，Tacola 属 は Moduza 属，Auzakia 属 等 と 類 本 種 の 終 齢 幼 虫 は チ ャ イ ロ イ チ モ ン ジ Moduza 縁関係にあると思われるが，検証するために必要な procris にやや似ており，約 5 ㎝と大型になる。今回 データが得られていないことから，本稿ではその点 の調査ではボルネオとタイで撮影された画像を確認 についての考察は行わないこととした。 ( 文責：宇野 Butterflies 割付 Butterflies 投稿規定 1. 投稿資格 原則として，会員に限ります。編集委員会が依頼した原 稿は，この限りではありません。 2. 採否および内容の修正 編集委員会にご一任いただくものとします。内容や表記 法について，編集委員会あるいは学術委員会から著者に修正 を求める場合があります。専門用語以外の字句やカナ遣いに ついては，編集委員会の判断で変更することがあります。 3. 言語 和文または英文とします。短報を除き，和文の場合には 原則として英文表記のタイトル・著者名・著者住所をつけて ください。Abstract と Keywords についても，原則として著 者が書いてください。本文が英文の場合には文末に和文摘要 をつけてください。なお当会の会員向けサービスとして英文 作成の代行が可能ですので，もし必要な場合には原稿の当該 箇所にその旨を記入してください。 4. 投稿方法 通常はテキスト （文書ファイル）をワープロソフトで作成 し，学会に電子メールで直接お送りいただくか，CD 等のメ ディアに記録して郵便または宅配便でお送りください。内容 確認のため，プリントアウトも 1 部つけてください。 手書き原稿も受理いたしますが，できるかぎり電子テキ ストでおねがいいたします。ボランティアによる入力作業の ために日数がかかることをご理解ください。 5. 図や写真の扱い ・報文中においては，図表を番号順に引用してください。 ・デジカメ写真は画質を落とさずに高精細のままでお送りくだ さい。数百万〜 1,000 万画素級のデジカメを使って最高精細 度で写したくらいが適当とお考えください。原則として JPEG 58 ファイル形式で，できるだけ圧縮をかけず，CD 等のメディア に記録して送ってください。大体，1 枚の写真が 500 KB から 3MB くらいのファイルサイズになるのが妥当です。 ・ポジフィルム・ネガフィルムの場合は，スキャナーで読み 取り，JPEG ファイル形式の CD を作成してお送りいただく ことを希望します。写真店でも引き受けてくれます。もしプ リントやポジが送られた場合には委員会でスキャンして電子 化しますので，裏面か枠に必ず通し番号等を打ってください （この場合には掲載までの日数が余分にかかることをご諒解 ください）。印刷の仕上がりの点でも，電子ファイルによる ご投稿がお薦めです。 ・標本写真はなるべく灰色ベタの背景でお送りください。電 子ファイルの場合，本誌では通常 R=G=B=240（色座標） に設 定しますので，著者が予め塗りつぶして下さると大変助かり ます。 6. 著作権 掲載された原稿の著作権は日本蝶類学会に帰属するもの とします。但し，著者によるテキスト・図・表・写真等の部 分的な再使用を妨げるものではありません。また，当学会で は掲載された原稿の一部または全部をインターネット上で公 開することがあります。 7. 別刷 原則 30 部を差し上げます。それ以上の部数は有料扱いに なります。必要であればご投稿時にお知らせください。 [ 電子メールの送り先 ] welcome_scripts@yahoo.co.jp [ 郵便または宅配便の送り先 ] 住所は日本蝶類学会，宛先は編集委員会としてください。 彰）