taxonID	type	description	language	source
194CAF38FF89B05F70A5FEA2FE1AF8E2.taxon	description	almost no morphological differences between the two species or the intermediate forms in the male genitalia, although the two species showed small geographical and individual variations in the distal portion of the valva (Figs 15 – 17). Variation of the male genitalia amongst races or subspecies in the Nymphalidae is often found in nonfunctional parts such as the dorsal and distal margins of the valvae, which do not make contact with the female genitalia during copulation (Goulson, 1993). Additionally, there was no variation of the male genitalia between the two species in overlapping areas of their distributions (Figs 16, 17). The morphological and molecular evidence therefore indicates that E. phemius and E. ipona have not become reproductively isolated and that gene exchange is still taking place. Thus it would be reasonable to treat E. ipona as a form of E. phemius. The monophyly of E. euphemia was strongly supported, with relatively high values (Figs 11 – 14; BV 78 – 100 %, PP 0.96 – 1.00). Although E. euphemia was regarded as the sister group to the remaining E. phemius complex in the COI and EF- 1 a trees (Figs 11, 13), the ND 5 and Tpi trees showed that this species is internal within the clade composed of E. phemius and E. ipona (Figs 12, 14), so there was little phylogenetic resolution between E. phemius (and E. ipona) and E. euphemia. Although there are some cases in which recently diverged species fail to form reciprocal monophyly in a locus of mtDNA (ex. Kandul et al., 2004; Oliver & Shapiro, 2007), a single mitochondrial locus generally becomes reciprocally monophyletic much faster than does a single nuclear locus (Hudson & Coyne, 2002). Moreover, the time for reproductively isolated lineages to display reciprocal monophyly in a majority of nuclear loci can be considerably long (Hudson & Coyne, 2002), so different nuclear loci will often conflict, especially in comparisons of closely related taxa (Maddison, 2008). Therefore, it is not easy to decide clearly whether E. euphemia should be a species or a subspecies based on our molecular analyses. However, the genetic distances between E. euphemia and the remaining E. phemius complex species were rather small in all analysed genes (COI: 1.20 – 2.10 %, ND 5: 0.60 – 1.32 %, EF- 1 a: 0.35 – 0.53 %, Tpi: 0.82 – 2.06 %), as compared to the genetic distances between the other Euthalia species (COI: 3.30 – 9.00 %, ND 5: 3.49 – 8.65 %, EF- 1 a: 0.71 – 5.13 %, Tpi: 3.30 – 13.58 %). Nevertheless, as speciation some- times can be caused with low pairwise sequence divergence of mtDNA in Lycaenidae (Kandul et al., 2004; Oliver & Shapiro, 2007), we can only say from our molecular analyses that the specific status of E. euphemia is vague. The results suggest that reproductive isolation may not be complete, or that barriers may have recently been established between E. euphemia and the remaining E. phemius complex. Defining species is problematic, because there is no concrete concept free from any ambiguities. As de Queiroz (1998, 2005, 2007) hypothesizes, during the course of speciation, the two separating lineages acquire different properties relative to each other (species criteria), such as being phenetically distinguishable, reciprocal monophyly, and pre- and postzygotic reproductive isolation. The more evidence of lineage separation is available, the better one can say that different species exist. Therefore, in our case, more data, specifically morphological data, which we will show below, would be required to draw our conclusion on the specific status of E. euphemia.	en	Yago, Masaya, Yokochi, Takashi, Kondo, Mariko, Braby, Michael F., Yahya, Bakhtiar, Peggie, Djunijanti, Wang, Min, Williams, Mark, Morita, Sadayuki, Ueshima, Rei (2012): Revision of the Euthalia phemius complex (Lepidoptera: Nymphalidae) based on morphology and molecular analyses. Zoological Journal of the Linnean Society 164 (2): 304-327, DOI: 10.1111/j.1096-3642.2011.00772.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2011.00772.x
194CAF38FF89B05F70A5FEA2FE1AF8E2.taxon	description	Venation system follows Tsukada (1991). The important point to note is that Euthalia species generally show specific differences in the male genitalia, especially the shape of the valva (Tsukada, 1991). Tsukada did not report the male genital structures of E. phemius, and did not compare this to E. euphemia. We have for the first time compared in detail the genital structures for both E. phemius and E. euphemia. In particular, the shapes of valvae were analysed based on the EFDs (Table 3; Fig. 19; see also Material and methods). As a result, there were no differences in not only the male genital valvae but also the other genitalic parts between E. phemius (and E. ipona) and E. euphemia (Figs 15 – 19; Table 2). Based on the genetic analyses and genitalic differences, we consider E. euphemia to be a subspecies of E. phemius. Probably, the small genetic variation, and the characteristic wing markings, of E. p. euphemia were caused by a bottleneck effect (founder effect) as a result of promotion of genetic drift, i. e. the ancestral E. p. euphemia had been isolated to a confined area such as Borneo, where a loss of genetic variation rapidly occurred and in a short period the new population was distinctly different, both genetically and phenotypically, with an accumulation of genetic mutations. This taxonomic statement may be clarified through breeding experiments planned in the near future. In this study we also analysed divergence times in the E. phemius complex from combined sequences (1497 bp) of mitochondrial COI and ND 5 genes. There are two reasons why mitochondrial sequences in butterflies are particularly useful for estimating genetic divergence both within and between species (Mallet et al., 2007). First, recombination between mitochondria, because of unisexual inheritance, is unlikely to occur, and thus genetic divergence is considered not to be influenced by occasional introgression. Secondly, in butterflies, hybrid females are often sterile or have low viability according to Haldane’s rule (Coyne, 1985; Presgraves, 2002; Lukhtanov et al., 2005). Haldane’s rule implies that introgression of maternally inherited mitochondria is prevented at an earlier stage of speciation than that for nuclear loci in female-heterogametic species (Sperling, 1990; Jiggins et al., 2001 a, b; Naisbit et al., 2002), which may be transferred between species by backcrossing of male hybrids. Using this combined mitochondrial sequence for divergence time estimation, a linearized NJ tree based on Kimura’s twoparameter model was constructed (Fig. 20). Analysis based on Tajima’s relative-rate test confirmed that a molecular clock could be hypothesized for our data set (P <0.05). A molecular clock of 1.1 – 1.2 ¥ 10 - 8 substitutions per site per year (Brower, 1994) was applied (see Material and Methods). Judging from the linearized tree, the common ancestor of Euthalia was initially divided into six lineages, and this diversification occurred about 2.5 – 3.1 Mya. Beginning around 3 Mya, fluctuations between glacial and interglacial periods gradually occurred throughout the world (Lisiecki & Raymo, 2005; Reymo, Lisiecki & Nisancioglu, 2006). The start of the extreme climatic variation almost corresponds to early divergences within the genus. After this, the common ancestor of the E. phemius complex appeared about 2.1 – 2.3 Mya. In the lineage of the E. phemius complex, the divergence age of the ancestral E. phemius phemius and the ancestral E. phemius euphemia was estimated at about 0.5 – 0.6 Mya. We infer that at this point in time E. phemius was divided into two populations by the formation of the Strait of Malacca between Borneo and the Malay Peninsula, extending from Indochina, perhaps because of a climatic or geographical change. Subsequently, it is considered that one lineage evolved into the extant E. phemius phemius, widely distributed in the Asian continent, whereas the other was isolated in a limited area for a long period and became the extant E. phemius euphemia, endemic to Borneo. Euthalia monina may also have evolved in a similar way, the nominotypical subspecies occurring on the Malay Peninsula and the subspecies E. bipunctata (Snellen van Vollenhoven, 1862) on Borneo (Tsukada, 1991; Eliot, 1992). The division of the populations of E. monina seems to be at almost the same time as those of E. phemius (Fig. 20).	en	Yago, Masaya, Yokochi, Takashi, Kondo, Mariko, Braby, Michael F., Yahya, Bakhtiar, Peggie, Djunijanti, Wang, Min, Williams, Mark, Morita, Sadayuki, Ueshima, Rei (2012): Revision of the Euthalia phemius complex (Lepidoptera: Nymphalidae) based on morphology and molecular analyses. Zoological Journal of the Linnean Society 164 (2): 304-327, DOI: 10.1111/j.1096-3642.2011.00772.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2011.00772.x
194CAF38FF90B04370FCFCFDFB63FEF9.taxon	distribution	Distribution: Bangladesh, Nepal, north India (including Sikkim, Assam), Bhutan, Myanmar, west Malaysia, Langkawi, Thailand [including Tarutao Island and Samui Island (new record)], Laos, Vietnam, south and south-west China (including Hong Kong and Hainan Island).	en	Yago, Masaya, Yokochi, Takashi, Kondo, Mariko, Braby, Michael F., Yahya, Bakhtiar, Peggie, Djunijanti, Wang, Min, Williams, Mark, Morita, Sadayuki, Ueshima, Rei (2012): Revision of the Euthalia phemius complex (Lepidoptera: Nymphalidae) based on morphology and molecular analyses. Zoological Journal of the Linnean Society 164 (2): 304-327, DOI: 10.1111/j.1096-3642.2011.00772.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2011.00772.x
194CAF38FF90B04370FCFCFDFB63FEF9.taxon	description	Figure 20. Linearized neighbour-joining (NJ) tree based on Kimura’s two-parameter model using a combined sequence of mitochondrial cytochrome c oxidase subunit I (COI) and NADH dehydrogenase subunit 5 (ND 5) genes. Identical sequences were treated as a single operational taxonomic unit (OTU). Numbers indicate bootstrap values from NJ (top left) and maximum parsimony (top right) analyses, and posterior probabilities from Bayesian analysis (bottom). Only bootstrap values> 50 % and Bayesian posterior probabilities> 0.50 are shown. The abbreviations of OTUs correspond to those in Table 1. Molecular clocks of 1.1 – 1.2 ¥ 10 - 8 substitutions per site per year for mitochondrial genes were applied for the divergence time estimations (Brower, 1994). The classifications of Tsukada (1991) and from the present study are shown on the right. 218 – 219; Swinhoe, 1893: 286; Moore, 1898: 123, pl. 238, figs 1 (♂), 1 a (♂), 1 b (♀), 1 c (♀); Bingham, 1905: 280, fig. 53 (♂); Tytler, 1911: 59; Matsumura, 1919: 573, pl. 44, figs 4 (♀), 5 (♂); Evans, 1924: 904, pl. 19, F. 18.17 (1 ♂ 1 ♀); Matsumura, 1929 *: 23; Corbet, 1945: 182; Corbet, 1956: 233, pl. 7, fig. 92 (♂ genitalia); Doggett, 1974: 37 (1 ♂); Eliot, 1978: 200, 202, 437, 541, fig. 99 (♂ genitalia); G. & B. Johnston, 1980: 164 (♂); D’Abrera, 1985: 357 – 359; Mani, 1986: 134, 136; Wu, 1988: 135, pl. 12, figs 4 (♂), 5 (♀); Koiwaya, 1989: 89, 147, fig. 548; Eliot, 1992: 187, 188, 408, 557, fig. 99 (♂ genitalia), pl. 27, figs 23 (♂), 24 (♀); Igarashi & Fukuda, 1997: 437 – 438, pl. 198, figs ♂ UP, ♂ UN, ♀ UP, ♀ UN (+ larva, pupae, and habitat); Aoyama, 1998: 132, fig. 3 (♂), 133, fig. 5 (♂), 282; Watanabe, 1998: 83, fig. ♀, 148; Chou, 1998: 136, pl. 56, figs 1 (♂), 2 (♂ UN); Nishiyama, 1999: 118, pl. 64 (2 ♂ 1 ♀); Bascombe et al., 1999: 15, 54, 55, 344, 345, 382, 391, 392, figs 9.141 (wing venation), 9.142 B (♂ genitalia).	en	Yago, Masaya, Yokochi, Takashi, Kondo, Mariko, Braby, Michael F., Yahya, Bakhtiar, Peggie, Djunijanti, Wang, Min, Williams, Mark, Morita, Sadayuki, Ueshima, Rei (2012): Revision of the Euthalia phemius complex (Lepidoptera: Nymphalidae) based on morphology and molecular analyses. Zoological Journal of the Linnean Society 164 (2): 304-327, DOI: 10.1111/j.1096-3642.2011.00772.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2011.00772.x
194CAF38FF92B04370FAFE48FADCFC3B.taxon	distribution	Distribution: Borneo.	en	Yago, Masaya, Yokochi, Takashi, Kondo, Mariko, Braby, Michael F., Yahya, Bakhtiar, Peggie, Djunijanti, Wang, Min, Williams, Mark, Morita, Sadayuki, Ueshima, Rei (2012): Revision of the Euthalia phemius complex (Lepidoptera: Nymphalidae) based on morphology and molecular analyses. Zoological Journal of the Linnean Society 164 (2): 304-327, DOI: 10.1111/j.1096-3642.2011.00772.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2011.00772.x
