identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
F64087B8FFD0FFC9AB9E6C8FDC07FB15.text	F64087B8FFD0FFC9AB9E6C8FDC07FB15.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chrysorthenches Dugdale 1996	<div><p>GENUS  CHRYSORTHENCHES DUGALE, 1996</p><p>Chrysorthenches Dugdale, 1996: 34 . Type species:  Orthenches porphyritis Meyrick, 1886, by original designation.</p><p>Dugdale (1996) defined  Chrysorthenches in relation to  Orthenches and suggested the following apomorphies for the former genus: (1) in the male genitalia, the gnathos and socii absent; (2) the anellus as a spinulose sheath and often with strong apical thorns; (3) the vinculum plus saccus T-shaped; (4) the posterior margin of the saccus convex; (5) the male phallus uniformly cylindrical; (6) the male sternum VIII with a V-shaped lobe (Fig. 2G); (7) the male tergopleural lobe arising obliquely and fused dorsally (Fig. 2G, arrow); and (8) the larval abdominal segment VII with spiracle and seta SD1 on a common pinaculum or scobinate field.</p><p>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.</p></div>	https://treatment.plazi.org/id/F64087B8FFD0FFC9AB9E6C8FDC07FB15	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sohn, Jae-Cheon;Kobayashi, Shigeki;Yoshiyasu, Yutaka	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.text	F64087B8FFD0FFCEABB569E5DFBDFA79.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chrysorthenches callibrya (Turner 1923) Sohn & Kobayashi & Yoshiyasu 2020	<div><p>CHRYSORTHENCHES CALLIBRYA SPECIES GROUP</p><p>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.</p><p>Thorax – Forewing narrow, greenish; venation (Fig. 1F) 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 1A+2A near one-third of the length. Hindwing venation (Fig. 1F) 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; 1A+2A sinuous, close to CuP at distal one-third, with basal fork one-eighth of length; 3A straight.</p><p>Male genitalia (Fig. 2A–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. 2D–F).</p><p>Female genitalia (Fig. 3A, 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.</p>KEY TO THE SPECIES-GROUPS OF  CHRYSORTHENCHES 1. Ductus bursae in female genitalia narrow and entirely sclerotized; larval mesothoracic L 1 and L 2 setae on separate pinacula ......................................................................................................  C. callibrya species-group - Ductus bursae in female genitalia partly sclerotized or entirely membranous; larval mesothoracic L 1 and L 2 setae on same pinaculum ............................................................................................................................. 2 2. Ductus seminalis entirely sclerotized; ductus bursae shorter than corpus bursae ........................................ ..................................................................................................................................  C. argentea species-group  - Ductus seminalis partly sclerotized; ductus bursae longer than or as long as corpus bursae ............................... ..........................................................................................................................................  C. porphyritis species-group<p>Distribution: Eastern Australia, Thailand and Japan.</p></div>	https://treatment.plazi.org/id/F64087B8FFD0FFCEABB569E5DFBDFA79	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sohn, Jae-Cheon;Kobayashi, Shigeki;Yoshiyasu, Yutaka	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.text	F64087B8FFDCFFD9A83569D8D966FC33.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chrysorthenches smaragdina Sohn 2020	<div><p>CHRYSORTHENCHES SMARAGDINA SOHN,  SP. NOV.</p><p>(FIGS 1D, 2C, F, H)</p><p>lsid: urn:lsid:zoobank.org:act: F6F3149C-CEC9-4378- 8A65-D43429E2E7F8</p><p>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).</p><p>Description: Adult (Fig. 1D). 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.</p><p>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.</p><p>Male genitalia (Fig. 2C, 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. 2H). Phallus (Fig. 2F) 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.</p><p>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.</p><p>Distribution: Thailand.</p><p>Etymology: The epithet is derived from the Greek σμαράγδι, ‘ smarágdi ’, emerald, referring to the broad green patch on the forewing of this new species.</p><p>PHYLOGENETICS</p><p>Our cladistic analyses of the morphological characteristics of 13 yponomeutoids resulted in a single most parsimonious tree (Fig. 15A: 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. 15A). The other clade is divided into two subclades, but the supports are weak.</p><p>BIOGEOGRAPHY</p><p>Optimal reconstruction of our DIVA analysis requires two dispersals (Fig. 15B). 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.</p></div>	https://treatment.plazi.org/id/F64087B8FFDCFFD9A83569D8D966FC33	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sohn, Jae-Cheon;Kobayashi, Shigeki;Yoshiyasu, Yutaka	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.text	F64087B8FFC0FFDFA88A6914DF1FFA60.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Association	<div><p>FAMILY  ASSOCIATION</p><p>Affinities between  Chrysorthenches and  Orthenches were suggested by Dugdale (1996) who designated the  Orthenches -group for those two genera.The  Orthenches - group was defined by Dugdale (1996) based on one synapomorphy: the presence of a unisetose SV setal group on the larval abdominal segment IX. However, not all the members of the group have been examined for that character due to scant larval descriptions (Dugdale, 1996). In the present study, we examined the larvae of  C. muraseae and observed that they exhibit the synapomorphy of the  Orthenches -group. That finding strengthens the phylogenetic value of the character defining the group.</p><p>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 (‘CL67’ 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. S1), 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).</p><p>1: antennal scape: (0) with awning and pecten; (1) with pecten only.</p><p>2: maxillary palpi: (0) four-segmented; (1) one-segmented.</p><p>3: subtegular tuft: (0) narrow, sinuous; (1) broad, straight.</p><p>4: forewing chorda: (0) present, long; (1) present, short; (2) absent.</p><p>5: uncus: (0) long, linear; (1) short, broad or absent.</p><p>6: apex of uncus: (0) bifid, (1) unifid.</p><p>7: gnathos and socii: (0) present; (1) absent.</p><p>8: anellus: (0) trough-like with paired lateral sclerites; (1) a single spinulate sheath; (2) sheath divided transversely into proximal and distal parts.</p><p>9: spinulation on anellus: (0) absent; (1) uniformly spinulate; (2) with apical outstanding spinules.</p><p>10: vinculum and saccus: (0) V- or U-shaped; (1) T-shaped.</p><p>11: posterior margin of saccus: (0) straight or concave; (1) convex.</p><p>12: [modified] lobe on male sternum VIII: (0) paired; (1) single, V-shaped.</p><p>13: valva structure: (0) undivided; (1) divided transversely at right angle to costa; (2) divided obliquely.</p><p>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.</p><p>15: phallus shape: (0) basally swollen; (1) uniformly cylindrical.</p><p>16: apex of phallus: (0) simple; (1) with a ventral mesal process; (2) with lateroventral process.</p><p>17: apical carina of phallus: (0) absent; (1) acuminate; (2) acute; (3) hooked; (4) with a thorn patch.</p><p>18: number of cornuti: (0) four or more; (1) two or three; (2) one; (3) none.</p><p>19: tergopleural lobe: (0) arising perpendicularly, separate dorsally; (1) arising obliquely, largely fused dorsally.</p><p>20: sterigma: (0) sunken; (1) on a papilla.</p><p>21: antrum: (0) parallel-sided or barrel-shaped; (1) funnel-shaped.</p><p>22: ductus seminalis: (0) arising dorsally; (1) arising ventrally.</p><p>23: ductus bursae: (0) tubular; (1) sinuous, furrowed; (2) cumuloid, furrowed, wider than corpus bursae; (3) gradually widened to corpus bursae.</p><p>24: [modified] sclerotization of ductus bursae: (0) unsclerotized; (1) partly sclerotized; (2) entirely sclerotized.</p><p>25: length of ductus bursae: (0) longer than corpus bursae; (1) equal to corpus bursae; (2) shorter than corpus bursae.</p><p>26: [added] inception of ductus seminalis: (0) on ductus bursae; (1) on corpus bursae.</p><p>27: appendix bursae on corpus bursae: (0) absent; (1) present.</p><p>28: signum: (0) single; (1) double; (2) absent.</p><p>29: larval spiracle VIII: (0) separate from seta SD pinacula or scobinate zone; (1) included on SD pinacula or scobinate zone.</p><p>30: [added] larval thoracic L1 and L2 seta: (0) on same pinaculum; (1) on separate pinacula.</p><p>GENERIC  ASSOCIATION</p></div>	https://treatment.plazi.org/id/F64087B8FFC0FFDFA88A6914DF1FFA60	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sohn, Jae-Cheon;Kobayashi, Shigeki;Yoshiyasu, Yutaka	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.text	F64087B8FFC6FFD3ABDE68E9DF21FC15.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chrysorthenches callibrya	<div><p>Chrysorthenches callibrya was originally combined with  Diathryptica (type species:  D. proterva) by</p><p>Turner (1923). In the present study, we examined the type species of  Diathryptica and found three major differences: the chorda of the forewing, the division of the male valva and the sclerotization of the ductus bursae in the female genitalia. The  C. callibrya species-group includes  C. callibrya and two allied new species. These three species share the diagnostic characteristics of  Chrysorthenches as proposed by Dugdale (1996), including, in both sexes, the lack of a dense awning of scales on the antennal scape, the long apical segment of the labial palpus, the lack of socii and gnathos, the presence of a V-shaped mesal lobe on the male sternum VIII, and the absence of a sclerotized costa on the membranous distal part of the valva in the male genitalia. In addition, our examination of the larvae of  C. muraseae revealed four characteristics associating it with  Chrysorthenches: (1) seta V1 on the meso- and metathorax present at the ventral edge of each coxa, (2) a spiracle on abdominal segment VIII on the SD1 pinaculum, posterior to the SD1 seta, (3) abdominal segments VII and VIII with one SV seta and (4) abdominal segment IX with setae D1 and D2 on the same broad pinaculum. These shared characteristics justify a generic transfer of  Diathryptica callibrya to  Chrysorthenches and the assignment of the two new species mentioned in this study to the same genus. The former finding was also consistent with our COI phylogeny result (Supporting Information, Fig. S1).</p><p>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 L1 and L2 setae arising on separate pinacula (30: 1 in Table 1) (Supporting Information, Fig. S2). The larval features of  C. muraseae differed from those of the  C. argentea and the  C. porphyritis species-groups in the presence of thin SD1 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.</p><p>Our cladogram (Fig. 15A) 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.</p><p>PODOCARPACEAE ASSOCIATION</p><p>Chrysorthenches is distinguished from other lineages of the  Orthenches -group by a trophic association with conifers. Larval host-plants are known for only nine of the 12 species in  Chrysorthenches and for two congeners whose larval hosts were inferred from vegetation in which the adult moths were observed (Dugdale, 1996). These records indicate that all species of  Chrysorthenches, except for  C. virgata (Philpott, 1920), which feeds on  Cupressaceae, and  C. smaragdina, whose larval hosts are unknown, are associated with  Podocarpaceae . Among the  Podocarpaceae, the majority of  Chrysorthenches species utilize the largest genus of that family,  Podocarpus . The members of the  C. callibrya species-group seem also to be associated with  Podocarpus . The host-plants of  C. muraseae are reported from the present study, while an association of  C. callibrya with  Podocarpus could be inferred from Dugdale’s (1996) field observation at Charlotte Pass, New South Wales, Australia. It is likely that  C. smaragdina also feeds on  Podocarpus, given the host associations of two other species in the same species-group and the occurrence of  Podocarpus in Thailand.</p><p>Chrysorthenches utilize seven genera of  Podocarpaceae and those genera are not necessarily closely related (Fig. 16). This may suggest that most, if not all, of their host associations have resulted from sequential colonization, not co-evolution, as Dugdale (1996) has already pointed out. Podocarpaceae-feeding species of  Chrysorthenches were associated with only one or two plant genera, while  Podocarpus was the genus on which most species of  Chrysorthenches feed (Fig. 16). The  C. callibrya species-group, earliest diverging in  Chrysorthenches, also uses  Podocarpus as a larval host. Taken together, these observations may suggest that ancestral  Chrysorthenches colonized  Podocarpus and later shifted to other podocarp genera. Among the  Podocarpus -feeding  Chrysorthenches, the New Zealand species are associated exclusively with the  Australis subclade in the subgenus  Podocarpus . On the other hand, the host-plants of the  C. callibrya species-group belong to two  Podocarpus subgenera (Fig. 17).</p><p>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 &amp; 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.</p><p>BIOGEOGRAPHY AND HOST–PLANT TRACKING</p><p>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).</p><p>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. 15B) 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.</p><p>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.</p><p>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. 15B). 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).</p><p>The  Chrysorthenches callibrya species-group may have evolved as a result of their colonization of  Podocarpus in the Cenozoic Era. Given the distributions of their sister groups, it would seem plausible that ancestors of the  C. callibrya species-group evolved as one of the lineages resulting from the radiation of  Chrysorthenches before the separation of New Zealand and Australia in the Middle–Late Cretaceous. In such a scenario, this lineage would have dispersed toward South-East Asia, as represented by the occurrence of  C. smaragdina in Thailand. Such an event could have happened only after the  Podocarpus species had radiated into tropical Asia in the Late Eocene, about 30 million years ago (Cernusak et al., 2011) and after the first opportunity for faunal exchange between Australia and Asia approximately 25 million years ago (De Jong, 2001). Consistent with these requirements, the emergence of host-plant clades for the  C. callibrya species-group were estimated to occur in the Eocene–Oligocene periods (Quiroga et al., 2016). Pre-existence of host-plants was a prerequisite for the  C. callibrya species-group crossing the Wallacea zone, like other lepidopteran examples (Beck et al., 2006).  Chrysorthenches muraseae may represent the currently understood terminus of the sequential radiation for the  C. callibrya species-group. It is known that the island arc system crossing the Wallacea zone reached Japan in the Late Miocene or in the Pliocene (De Jong, 2001).</p><p>Recent advances in biogeography allow the differentiation of dispersal from vicariance and the determination of possible divergence dates using molecular data (Trewick, 2000; Trewick &amp; Wallis, 2001; Waters &amp; Roy, 2004; de Queiroz, 2005). This type of approach would be necessary to better explain the curious distribution of  Chrysorthenches .</p></div>	https://treatment.plazi.org/id/F64087B8FFC6FFD3ABDE68E9DF21FC15	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sohn, Jae-Cheon;Kobayashi, Shigeki;Yoshiyasu, Yutaka	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
