taxonID	type	description	language	source
F64087B8FFD0FFC9AB9E6C8FDC07FB15.taxon	description	Two additional characters: the presence of the antennal scape with pecten, but no awning, and the valva of the male genitalia with an outer tuft of specialized scales or setae distally, were suggested as apomorphies for Chrysorthenches by Dugdale (1996). These were excluded from this study as proposed from our cladistics analysis. With our additions, Chrysorthenches comprises 13 species in three species-groups.	en	Sohn, Jae-Cheon, Kobayashi, Shigeki, Yoshiyasu, Yutaka (2020): Beyond Wallace: a new lineage of Chrysorthenches (Lepidoptera: Yponomeutoidea: Glyphipterigidae) reveals a journey tracking its host-plants, Podocarpus (Pinopsida: Podocarpaceae). Zoological Journal of the Linnean Society 190: 709-736
F64087B8FFD0FFCEABB569E5DFBDFA79.taxon	description	Description: Head – Vestiture of vertex appressed but rough around temporal area. Antenna filiform in both sexes, two-thirds as long as forewing costa. Labial palpus slightly ascending, 4 × longer than antennal scape; second labial palpomere with ventrodistally denser scale tuft; third palpomere longest. Maxillary palpi three-segmented, as long as antennal scape. Temporal and occipital areas with piliform scales. Thorax – Forewing narrow, greenish; venation (Fig. 1 F) with Sc reaching margin slightly before middle of costa; R arising from near basal two-fifths of radius; Rs 1 - 3 reaching margin above apex; Rs 1 arising from anterior margin of accessory cell near basal two-fifths, slightly divergent from R; Rs 2 slightly convergent to Rs 1; Rs 2 and Rs 3 basally separate, divergent; Rs 3 and Rs 4 slightly divergent; Rs 4 reaching margin right below apex; M 2 parallel to M 1 and M 3; CuA 1 and CuA 2 sinuous, close to each other at middle; CuP vestigial as fold in basal five-sevenths; basal fork of 1 A + 2 A near one-third of the length. Hindwing venation (Fig. 1 F) with Sc + R 1 reaching margin at middle of costa; Rs reaching margin above apex; M stem vestigial in basal five-sixths; M 1 slightly close to Rs at middle; M 1 and M 2 slightly divergent in distal half; M 2 nearly parallel to M 3; M 3 and CuA 1 divergent; CuA 1 parallel to CuA 2; CuP present; 1 A + 2 A sinuous, close to CuP at distal one-third, with basal fork one-eighth of length; 3 A straight. Male genitalia (Fig. 2 A – F, H) – Uncus elongate, bifid apically; tuba analis broad, sparsely setose apically; subscaphium densely setose; tegumen subrectangular, setose laterally. Valva obovate or subrectangular, densely hairy, with large, sparselysetose; membranous disc basally, sparsely setose. Anellus densely spinulate. Saccus elongate, digitate, enlarged subapically. Phallus slightly sinuous, broadened basally, opening obliquely at apex, with needle-like and spiniform cornuti; carina strongly sclerotized, spiniform; vesica with a needle-like cornutus, a spinulate, digitate cornutus and five spiniform cornuti (Fig. 2 D – F). Female genitalia (Fig. 3 A, B) – Ovipositor telescopic; papillae anales with setose, digitate protrusions dorsoapically. Abdominal segment IX with a pair of setose lobes ventrally. Ductus bursae slender, entirely sclerotized. Ductus seminalis arising on accessory sac of corpus bursae. Corpus bursae with accessory sac at middle; signum absent. Distribution: Eastern Australia, Thailand and Japan.	en	Sohn, Jae-Cheon, Kobayashi, Shigeki, Yoshiyasu, Yutaka (2020): Beyond Wallace: a new lineage of Chrysorthenches (Lepidoptera: Yponomeutoidea: Glyphipterigidae) reveals a journey tracking its host-plants, Podocarpus (Pinopsida: Podocarpaceae). Zoological Journal of the Linnean Society 190: 709-736
F64087B8FFDCFFD9A83569D8D966FC33.taxon	description	(FIGS 1 D, 2 C, F, H) lsid: urn: lsid: zoobank. org: act: F 6 F 3149 C-CEC 9 - 4378 - 8 A 65 - D 43429 E 2 E 7 F 8 Diagnosis: This new species is similar to C. callibrya in the male genitalia, but differs from the latter in having the subrectangular valva (obovate in C. callibrya) and the subtriangular apex of the saccus (digitate in C. callibrya). Description: Adult (Fig. 1 D). Head – Vertex yellowish green, with greenish orange piliform scales on temporal and occipital areas; frons yellowish green. Antenna with scape yellowish green; flagellum pale greyish green, intermixed with dark brown scales basally and distally. Maxillary palpus with first palpomere dark brown; second and third palpomeres dark brown, tinged with pale greenish grey apically. Labial palpus with first palpomere yellowish green on outer surface, pale orange on inner surface, tinged with dark brown dorsally; second palpomere 2 × longer than first palpomere, dark brown, mottled with yellowish green on outer surface, pale orange on apex and inner surface; third palpomere 2.2 × longer than second palpomere, dark brown on outer surface, pale orange on inner surface. Thorax – Patagium dark yellowish green, tinged with dark brownish green medially; tegula dark purplish brown; mesonotum dark greenish grey, intermixed with dark brown scales anterolaterally; mesoscutellum dark brown. Foreleg with coxa dark greyish brown; femur dark brown; tibia and tarsomeres dark brown, with pale orange ring distally. Midleg with coxa and femur pale orange, intermixed with dark greyish brown scales; tibia and tarsomeres dark brown dorsally, pale orange ventrally, with pale orange ring distally. Hindleg with coxa and femur pale greyish orange, sparsely intermixed with brownish grey scales; tibia and tarsomeres dark brownish grey, with pale orange ring distally. Forewing length 5.2 mm (sample number = 1), dark brown, tinged with yellowish green narrowly along costal area and broadly along dorsal area; apical area dark greenish brown; costal strigulae dark brown, irregularly intermixed with small white bars; subbasal and median fascia bar-like, yellowish green, bordered with pale greyish green, and then with black; dorsal margin with dark brown strigulae; fringe pale greyish green. Hindwing greyish brown, paler to base; fringe brownish grey. Male genitalia (Fig. 2 C, F, H) – Uncus slightly concave apically, gradually broadened basally; setose area of tuba analis one-quarter of its length. Valva subrectangular; costa nearly straight in basal two-thirds, slightly curved in distal third; sacculus narrow, one-third as long as ventral margin of valva; membranous disc obliquely round, half as long as valva. Vinculum U-shaped; saccus as long as uncus, slightly narrowed at middle, narrowly round apically. Anellus densely spinose (Fig. 2 H). Phallus (Fig. 2 F) with broad, needle-like cornutus one-sixth the length of the phallus; spinulate, digitate cornutus twosevenths as long as the needle-like cornutus; elongate, spinose cornutal zone. Type: Holotype – ‘ HOLO- | TYPE’ (round label with red edges), ‘ HOLOTYPE | Chrysorthenches | smaragdina | Sohn’ (red label with black marginal lines), ‘ N. THAILAND: 1640 – 1685 m | Nan, Doi Phu Kha NP, | km 33.8 to 34.4, | 26 – 30. xii. 1991 ’, ‘ B. M. ♂ | Genitalia slide | No. 32892 ’, deposited in NHMUK. Distribution: Thailand. Etymology: The epithet is derived from the Greek σμαράγδι, ‘ smarágdi ’, emerald, referring to the broad green patch on the forewing of this new species. PHYLOGENETICS Our cladistic analyses of the morphological characteristics of 13 yponomeutoids resulted in a single most parsimonious tree (Fig. 15 A: tree length = 74, Ci = 67, Ri = 68). The resulting tree recovers strong support (Fig. 16: JK support = 100, Bremer support = 5) for the monophyly of Chrysorthenches against the outgroup, Orthenches chlorocoma. The backbone of the tree is divided into two clades. One clade corresponds to the C. callibrya species-group (including C. callibrya and C. muraseae) and is recovered as monophyletic (Fig. 16: JK support = 95, Bremer support = 4). The monophyly of the C. callibrya species-group is defined by one unambiguous character (Fig. 15 A). The other clade is divided into two subclades, but the supports are weak. BIOGEOGRAPHY Optimal reconstruction of our DIVA analysis requires two dispersals (Fig. 15 B). The result shows three possible scenarios for the ancestral distribution of Chrysorthenches: all areas covering (1) New Zealand – Tasmania – eastern Australia – East Asia, (2) New Zealand – eastern Australia – East Asia or (3) Tasmania – eastern Australia – East Asia. The Chrysorthenches callibrya species-group branched off from the ancestors and dispersed to East Asia. Subsequently, the C. porphyritis species-group diverged and occupied New Zealand and Tasmania. The Chrysorthenches argentea species-group radiated within New Zealand.	en	Sohn, Jae-Cheon, Kobayashi, Shigeki, Yoshiyasu, Yutaka (2020): Beyond Wallace: a new lineage of Chrysorthenches (Lepidoptera: Yponomeutoidea: Glyphipterigidae) reveals a journey tracking its host-plants, Podocarpus (Pinopsida: Podocarpaceae). Zoological Journal of the Linnean Society 190: 709-736
F64087B8FFC0FFDFA88A6914DF1FFA60.taxon	description	The systematic status of the Orthenches - group remains poorly understood. Dugdale (1996) suggested its association with Plutellidae, based on larval and pupal characteristics. However, the Orthenches - group does not share the synapomorphy for Plutellidae proposed by Kyrki (1984): i. e. the presence of a gnathal process surrounding the anal tube. Sohn et al. (2013) recovered an unidentified species of the Orthenches - group from South America (‘ CL 67 ’ in their phylogeny) that was nested in a glyphipterigid subfamily, Orthoteliinae. This suggests that the Orthenches - group, including Chrysorthenches, belong to the subfamily. Our COI - based maximum likelihood tree of ten yponomeutoids also supported their association with Glyphipterigidae (Supporting Information, Fig. S 1), although usefulness of one-locus phylogeny for family association is often limited. The Orthoteliinae have been redefined with the addition of putative yponomeutoids, formerly associated with Plutellidae, from the Southern Hemisphere (Heppner, 2005). 1: antennal scape: (0) with awning and pecten; (1) with pecten only. 2: maxillary palpi: (0) four-segmented; (1) one-segmented. 3: subtegular tuft: (0) narrow, sinuous; (1) broad, straight. 4: forewing chorda: (0) present, long; (1) present, short; (2) absent. 5: uncus: (0) long, linear; (1) short, broad or absent. 6: apex of uncus: (0) bifid, (1) unifid. 7: gnathos and socii: (0) present; (1) absent. 8: anellus: (0) trough-like with paired lateral sclerites; (1) a single spinulate sheath; (2) sheath divided transversely into proximal and distal parts. 9: spinulation on anellus: (0) absent; (1) uniformly spinulate; (2) with apical outstanding spinules. 10: vinculum and saccus: (0) V- or U-shaped; (1) T-shaped. 11: posterior margin of saccus: (0) straight or concave; (1) convex. 12: [modified] lobe on male sternum VIII: (0) paired; (1) single, V-shaped. 13: valva structure: (0) undivided; (1) divided transversely at right angle to costa; (2) divided obliquely. 14: outer scale-tuft of distal part of valva: (0) absent; (1) setose; (2) comprising broad persistent scales; (3) comprising scales and one seta; (4) as line of scales. 15: phallus shape: (0) basally swollen; (1) uniformly cylindrical. 16: apex of phallus: (0) simple; (1) with a ventral mesal process; (2) with lateroventral process. 17: apical carina of phallus: (0) absent; (1) acuminate; (2) acute; (3) hooked; (4) with a thorn patch. 18: number of cornuti: (0) four or more; (1) two or three; (2) one; (3) none. 19: tergopleural lobe: (0) arising perpendicularly, separate dorsally; (1) arising obliquely, largely fused dorsally. 20: sterigma: (0) sunken; (1) on a papilla. 21: antrum: (0) parallel-sided or barrel-shaped; (1) funnel-shaped. 22: ductus seminalis: (0) arising dorsally; (1) arising ventrally. 23: ductus bursae: (0) tubular; (1) sinuous, furrowed; (2) cumuloid, furrowed, wider than corpus bursae; (3) gradually widened to corpus bursae. 24: [modified] sclerotization of ductus bursae: (0) unsclerotized; (1) partly sclerotized; (2) entirely sclerotized. 25: length of ductus bursae: (0) longer than corpus bursae; (1) equal to corpus bursae; (2) shorter than corpus bursae. 26: [added] inception of ductus seminalis: (0) on ductus bursae; (1) on corpus bursae. 27: appendix bursae on corpus bursae: (0) absent; (1) present. 28: signum: (0) single; (1) double; (2) absent. 29: larval spiracle VIII: (0) separate from seta SD pinacula or scobinate zone; (1) included on SD pinacula or scobinate zone. 30: [added] larval thoracic L 1 and L 2 seta: (0) on same pinaculum; (1) on separate pinacula. GENERIC ASSOCIATION	en	Sohn, Jae-Cheon, Kobayashi, Shigeki, Yoshiyasu, Yutaka (2020): Beyond Wallace: a new lineage of Chrysorthenches (Lepidoptera: Yponomeutoidea: Glyphipterigidae) reveals a journey tracking its host-plants, Podocarpus (Pinopsida: Podocarpaceae). Zoological Journal of the Linnean Society 190: 709-736
F64087B8FFC6FFD3ABDE68E9DF21FC15.taxon	description	Our cladistic analysis supported the monophyly of the C. callibrya based on one synapomorphy, the entirely sclerotized ductus bursae (24: 2 in Table 1), and three homoplastic characters: the bifid uncus (6: 0 in Table 1), the ventrally arising ductus seminalis (22: 1 in Table 1) and an enception of the ductus seminalis on the corpus bursae (26: 1 in Table 1). The fast and slow character optimizations in the cladistics study recognized two additional synapomorphies: the presence of a short chorda on the forewing (4: 1 in Table 1) and larval thoracic L 1 and L 2 setae arising on separate pinacula (30: 1 in Table 1) (Supporting Information, Fig. S 2). The larval features of C. muraseae differed from those of the C. argentea and the C. porphyritis species-groups in the presence of thin SD 1 setae on the mesothorax and the abdominal segment VIII, and an SV setal group on abdominal segments I and II bisetose. These characters can also serve as synapomorphies of the C. callibrya species-group, but more information on larval characters is needed to confirm their phylogenetic value. Our cladogram (Fig. 15 A) for 12 species of Chrysorthenches differed from that presented by Dugdale (1996). The most critical dissimilarity was in the position of C. polita (Philpott, 1918), which was placed in the C. argentea species-group in our study but in the C. porphyritis species-group by Dugdale (1996). The positions of C. glypharcha (Meyrick, 1919) and C. phyllocladi Dugale, 1996 were also discordant between the two studies. All these differences may be the result of our modifications and additions of characteristics to the data matrix presented by Dugdale (1996). Thus, we analysed another data matrix (J. - C. Sohn, unpublished) that included the same characterset and coding as that used by Dugdale (1996). The analysis still resulted in a different cladogram from that described by Dugdale (1996), possibly due to the additions of C. callibrya and C. muraseae. In fact, the relationships among the species-groups in Chrysorthenches are ambiguous, because those depend on the characteristics of the ductus bursae and the ductus seminalis, which are membranous and thus versatile. In accordance with this ambiguity, the backbone relationships of Chrysorthenches were poorly supported by the results of our study (1 – 2 range in Bremer supports: Fig. 16). Thus, the phylogenetic relationships within Chrysorthenches need further attention. PODOCARPACEAE ASSOCIATION	en	Sohn, Jae-Cheon, Kobayashi, Shigeki, Yoshiyasu, Yutaka (2020): Beyond Wallace: a new lineage of Chrysorthenches (Lepidoptera: Yponomeutoidea: Glyphipterigidae) reveals a journey tracking its host-plants, Podocarpus (Pinopsida: Podocarpaceae). Zoological Journal of the Linnean Society 190: 709-736
F64087B8FFC6FFD3ABDE68E9DF21FC15.taxon	description	The trophic associations between Chrysorthenches and Podocarpaceae are noteworthy, given the limited numbers of insects that utilize these plants. Other than Chrysorthenches, few lepidopterans feed on Podocarpaceae and they include macroheterocerans such as Erebidae (Lymantriinae), Geometridae and Lasiocampidae and some microlepidopterans (Tortricidae, Gracillariidae, Lecithoceridae and Pyralidae) worldwide (Okelo, 1972; Singh et al., 1978; Oku, 1979; Murase, 2005; Costa & Boscardin, 2014; Liu et al., 2018). Most of these moths are generalist larval feeders, but Makivora hagiyai Oku, 1979 (Tortricidae) is a specialist on Podocarpus. Chrysorthenches are comparable to Milionia Walker, 1854 (Geometridae) in that all or nearly all members are associated with Podocarpaceae. Yasui (2001) found that Milionia were able to sequester the phytochemicals of Podocarpus for protection against predatory stink bugs. Like Milionia, the adults of Chrysorthenches are colourful, but it is unknown if they can also take advantage of a chemical defence system. BIOGEOGRAPHY AND HOST – PLANT TRACKING The high trophic fidelity of Chrysorthenches with Podocarpaceae hints that the radiation of Chrysorthenches may have been affected by the host-plants. Recent studies have suggested that Podocarpaceae originated in Gondwana during the Triassic – Jurassic periods (Biffin et al., 2011; Rothwell et al., 2012; Escapa et al., 2013). Furthermore, Lu et al. (2014) estimated the origination of the extant podocarp genera to be in the Early Cretaceous. The largest genus of Podocarpaceae, Podocarpus, is one of the representative groups in the Antarctic flora that originated in the cold and wet climate of southern Gondwana (Page, 1990; Mill, 2003). Quiroga et al. (2016) dated the divergence of two subgenera of Podocarpus as within the Late Cretaceous – Early Palaeogene. The surviving lineages of Podocarpaceae radiated into the tropical regions, not earlier than 30 million years ago or the Late Eocene (Cernusak et al., 2011). Extant species of Chrysorthenches occur only in New Zealand, eastern Australia, Tasmania, South-East Asia and Japan (Fig. 17). The highest diversity among the Chrysorthenches species (eight of 13 total species) is observed in New Zealand. This, from the viewpoint of traditional dispersal biogeography, would suggest that New Zealand is the centre of origin for Chrysorthenches. However, the result of our DIVA analysis (Fig. 15 B) favoured a broad distribution of ancestral Chrysorthenches that subsequently split according to palaeogeographical changes. Regarding their presence in Tasmania and Australia, the Chrysorthenches – conifer association may pre-date the opening of the Tasman Sea, which began about 80 million years ago (Molnar et al., 1975). The distributional range of Chrysorthenches occupies only a small proportion of the distribution of Podocarpaceae. This difference may indicate that Chrysorthenches evolved long after the Podocarpaceae radiation that pre-dated the splitting of the Gondwanan subcontinents. Another, less plausible, explanation would be the extensive extinction of Chrysorthenches, except in the Australasian region. Direct evidence for this hypothesis does not exist to our knowledge, but a leaf-mine trace left by a larva that was presumed to belong to Chrysorthenches in Wilf et al. (2005) may indicate their existence on other Gondwana subcontinents until at least 52 million years ago. The Chrysorthenches callibrya species-group differs from the other two congeneric species-groups as the distribution of the former is not restricted to the Australasian region (Fig. 17). Moreover, three species of the species-group have disjunctive distributions: eastern Australia for C. callibrya, Thailand for C. smaragdina and Japan for C. muraseae. Our cladogram recovered this species-group as the earliest diverging with respect to other Chrysorthenches (Fig. 17). This poses questions, such as: why have no members of the species-group been reported from west and north Australia, Papua New Guinea and other islands spanning the Wallacea zone? Further inventory of Chrysorthenches in the Australasian region may help to fill these gaps. The collective distributional range of the C. callibrya species-group corresponds to that of the island arc system connecting Australia and East Asia. This island system has facilitated trans-Wallacean radiation in many organisms through faunal exchanges between Australia and Asia during 15 – 20 million years ago (Sklenarova et al., 2013). The C. callibrya species-group may have followed this route, but their direction was distinctively northward, as reconstructed from our DIVA analysis (Fig. 15 B). Most biogeographic studies in Australia and Asia have suggested southward dispersals (De Jong, 2001), although there are a few examples indicating northward radiations; for example, the plant family Proteaceae (Truswell et al., 1987) and skipper butterflies of the Taractroceragroup (De Jong, 2001).	en	Sohn, Jae-Cheon, Kobayashi, Shigeki, Yoshiyasu, Yutaka (2020): Beyond Wallace: a new lineage of Chrysorthenches (Lepidoptera: Yponomeutoidea: Glyphipterigidae) reveals a journey tracking its host-plants, Podocarpus (Pinopsida: Podocarpaceae). Zoological Journal of the Linnean Society 190: 709-736
