Ypthima, IN SATYRINA

SYSTEMATICS OF YPTHIMA IN SATYRINA View in CoL

We propose that Y. multistriata (MRCA) speciated vicariantly to generate three subspecies, Y. m. multistriata , Y. m. niphonica and Y. m. ganus, in addition to Y. akragas and Y. masakii . These are members of the motschulskyi group of Shirôzu & Shima (1979). Ypthima riukiuana is classified into the sordina group, but these Ryukyu three-ocellus species, including Yunnan sordina group species, also appear to be derived by vicariant speciation from the MRCA. Shima (1988) treated these groups as the single motschulskyi group.

Ypthima esakii View in CoL occurs sympatrically with Y. m. multistriata View in CoL in Taiwan, and Y. esakii View in CoL and Y. m. multistriata View in CoL appear to be synonymous based on the present research ( Figs 2 View Figure 2 and 3 View Figure 3 ; another Y. esakii View in CoL specimen in Table S1 has the same sequence as Y. m. multistriata View in CoL specimens), which refines the morphological classification of Shirôzu & Shima, 1979 and Takáhashi & Kiuchi (2015).

Ypthima motschulskyi View in CoL ( obscura View in CoL group), collected in northern China, represents a basal divergence in subgenus Ypthima View in CoL clade with three ocelli ( Fig. 3 View Figure 3 ). Two specimens were collected in Dandong, a city bordering North Korea. Ypthima motschulskyi View in CoL (synonym of Y. obscura View in CoL ) is sympatric with Y. multistriata ganus View in CoL ( motschulskyi View in CoL group) in Korea, and these species (and groups) names have been confused (c.f. Hiura, 1970, Shirôzu & Shima, 1979, Dubatolov & Lvovsky, 1997, Uémura & Koiwaya, 2000), but it is now shown to be a genetically differentiated species. Ypthima iris View in CoL (megalomma group) was collected in Yunnan, China, and represents basal divergence of the subgenus Ypthima View in CoL clade ( Fig. 3 View Figure 3 ).

Ypthima huebneri View in CoL ( huebneri View in CoL group; type species of genus Ypthima View in CoL ), collected in Vietnam, is characterized by four ocelli, forming a sister to sampled members of the major three-ocellus clade, collectively subgenus Ypthima View in CoL ( Fig. 3 View Figure 3 ). The Ypthima View in CoL grouping of Shirôzu & Shima (1979) and Shima (1988) was based on the number of ocelli ( Fig. 2 View Figure 2 bottom) as well as male and female genitalia, androconia, labial palpi, male foreleg and other criteria. In their cladogram that is based on Hennig (1966), the huebneri View in CoL group is in the subgenus Ypthima View in CoL .

The larger Y. praenubila View in CoL with four ocelli is included in the praenubila View in CoL group ( Shirôzu & Shima, 1979; renamed as the chenu group by Shima, 1988). Although it is placed in the subgenus Ypthima ( Shima, 1988) View in CoL , it appears to constitute a basal clade of the praenubila View in CoL group, and it forms a sister with the second major clade of subgenus Thymipa View in CoL ( Figs 2 View Figure 2 and 3 View Figure 3 ). In the cladgram made by Shima (1988), the arctous View in CoL group containing Y. arctous View in CoL , distributed in Australia, and the asterope View in CoL group containing Y. asterope View in CoL , distributed over a wide area including Africa and Pakistan, form basal clades within the subgenus Ypthima View in CoL . We confirmed this by estimating a Bayesian inference tree using available COI -5P data (DDBJ/GenBank). In this tree, the praenubila View in CoL group clade was distinct from the major clade of subgenus Ypthima View in CoL including Y. arctous View in CoL and Y. asterope View in CoL .

The other four-ocellus species are smaller, included in the tappana group by Shirôzu & Shima (1979) and Shima (1988). The tappana group is a sister of the sakra group with five ocelli with larger form, and both are included in the subgenus Thymipa major clade ( Figs 2 View Figure 2 and 3 View Figure 3 ). The smaller philomela group ( Shirôzu & Shima, 1979; Shima, 1988) with five ocelli is also a component of the major clade of the subgenus Thymipa , and a sister of the tappana + sakra group ( Figs 2 View Figure 2 and 3 View Figure 3 ). We have analysed all the three groups belonging to the subgenus Thymipa , and the Thymipa clade is a sister of praenubila group ( Figs 2 View Figure 2 and 3 View Figure 3 ). These sister relationships are not strictly concordant to the cladogram of Shima (1988).

Interestingly, the South African Ypthimomorpha itonia ( Sourakov & Emmel, 1997) with seven ocelli (additionally appeared in M2 and Sc + R1; Fig. 2 View Figure 2 , bottom) falls in this subgenus Thymipa clade, and it is a sister of the philomela group ( Figs 2 View Figure 2 and 3 View Figure 3 ). This result may suggest that Ypthimomorpha should be synonymized with Ypthima s.l. (as suggested by Sourakov & Emmel, 1997, but at the subgeneric level). However, it may be also possible to devise a generic or subgeneric level grouping using one of the many synonymized genera of Ypthima (note that Ypthimomorpha van Son, 1955 is a name more junior than these candidates including Thymipa ).

The type species of Ypthima (by subsequent designation) is four-ocellus Y. huebneri Kirby, 1871 , and at a minimum the motschulskyi + sordina group and two other species with three ocelli in that clade would belong to this group of ‘typical’ Ypthima , as named subgenus Ypthima by Shima (1988). The morphological differences of the South African Ypthimomorpha , belonging to the other subgenus Thymipa clade containing Y. baldus ( Figs 2 View Figure 2 and 3 View Figure 3 ), seem to be based at least partly on the absence of characters (R1 arises from the radial stalk beyond the upper angle of the cell, and an accessory plate is absent in the male valve and the female posterior plate is missing from the ostium: van Son, 1955). However, its final instar larva appears identical to those of Y. argus ( Sourakov & Emmel, 1997) . The close genetic relationship of Ypthimomorpha with the group containing Y. baldus and Y. argus casts doubt on Sourakov & Emmel’s suggestion of a closer sister relationship of Ypthimomorpha with other African species. Further sequencing of other ypthimine genera with available names is necessary to resolve the question of genera and subgenera within the satyrine subtribe Ypthimina .

We chose Pseudonympha magus (subtribe Ypthimina ) (DDBJ/GenBank) as outgroup species ( Figs 2 View Figure 2 and 3 View Figure 3 ). As a whole, Ypthima are not effectively monophyletic. By comparison with this outgroup species, two major clades – of the subgenus Ypthima and the subgenus Thymipa adding the praenubila clade – are highly distinct, and the genus Ypthima could be revised to comprise two or three different genera as suggested and defined by Shima (1988).