Dtsch. Entomol. Z. 70 (2) 2023, 261-282 | DOI 10.3897/dez.70.102952 > PENSUFT. yee BERLIN An “American” silkmoth endemic to Himalayas, part I: life history and natural distribution of Antheraea compta Rothschild, 1899 (Lepidoptera, Saturniidae) Zhengyang Liu? 1 Zhangdian District, Zibo, Shandong Province 255000, China https://zoobank. org/DCDDAA 33-01C1-4E9B-9B5B-B579BA37D008 Corresponding author: Zhengyang Liu (saturniidae@qq.com) Academic editor: Wolfram Mey @ Received 3 March 2023 Accepted 8 June 2023 @ Published 4 July 2023 Abstract Females of the uncommon Antheraea compta Rothschild, 1899 were collected at the natural habitat in the Tibetan Sub-Himala- yas, descendants of which were reared on Quercus yunnanensis (Fagaceae) successfully, with all the preimaginal instars recorded morphologically. Correlated characters revealed a close relationship between this taxon and New World Antheraea spp., suggesting more attention and protection towards this key species is necessary in the future. Key Words chaetotaxy, ecoregion, evolution, Kachin, Meghalaya, morphology, protection, Quercus, SEM, silkworm, 7elea, Tibet Introduction Species of Antheraea Hubner, 1819 have long been known in the history of sericulture as excellent sources for non-mulberry silk (Peigler 1993). Research surrounding the genus has always been a focus 1n Lepidoptera, occupy- ing most of the scientific works on the family Saturniidae around the world because of their spectacular appearances and considerable economic benefits. After a systematic review, Nassig (1991) proposed to classify these silk- moths into three subgenera, 1.e., Antheraea, Antheraeop- sis Wood-Mason, 1886 and 7e/ea Htibner, 1819. Textiles called Chinese tussah (from Antheraea pernyi (Guérin- Méneville, 1855)), Japanese tensan (from Antheraea ya- mamai (Guérin-Meéneville, 1861)) and Indian tasar (from Antheraea paphia (Linnaeus, 1758)) make the subgenus Antheraea the center of attention, while the golden muga silk (from Antheraea assamensis (Helfer, 1837)) produced in Sub-Himalayas renders Antheraeopsis famous (Peigler 2020). The venerable polyphemus moth (Antheraea poly- phemus (Cramer, 1775)) within 7e/ea is a cultural icon in American entomology. From southern Siberia to the Indonesian islands, from central Europe to the Japanese archipelago, the subgenus Antheraea occupies almost all the humid broad-leaved forests from temperate to tropical Eurasia. Antheraeopsis flies in Asia only south of the level of Qinling Mountains, but in any case, populations within these two subgenera are all entirely limited to the Palearc- tic-Indo-Malayan regions. The main habitats of 7e/ea are in the Nearctic-Neotropical realms, but one putative mem- ber, namely Antheraea compta Rothschild, 1899 (Fig. 1), is endemic to the northeastern part of the Indian subcon- tinent (Peigler 1999), this disjunct distribution considered one of the core mysteries in saturniid evolution. The original type series of A. compta was stated to be from “Khasia Hills, Assam, 8 3', no @ [sic]’, regarded from the beginning to be an adelphotaxon allied to A. as- samensis within Antheraeopsis (Rothschild and Jordan 1899). Such judgment may have been based largely on the superficial features, especially the similar eyespots and the shared natural habitat. Subsequently, Rothschild and Jor- dan (1901) provided a color photo of a male 4. compta and supplemented “The @ is similar to the <, but has a short- er and broader forewing [sic]”. Watson (1912: pl. 2) first Copyright Zhengyang Liu. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 262 illustrated the female, captured at the type-locality. Pack- ard (1914: 197) cited this taxon without additional com- ments, while Conte (1919: 14-15) and Seitz (1926-1928: 511) continued to associate A. compta with A. assamensis. The first entomologist who recognized the problem was Bouvier (1936: 167) who definitively stated that there is a fairly high morphological similarity between the male genitalia of Himalayan A. compta and Central American Antheraea godmani (Druce, 1892), especially their three- lobed valvae, although he did not transfer the former into his checklist for 7elea. After a comparison with A. poly- phemus, Lemaire (1978: 207) further acknowledged the strong homology with A. compta about male genital struc- tures, a taxonomic opinion recognized in more recent works (Nassig 1991; Nassig et al. 1996; d’Abrera 2012: 126-127; Naumann and Loffler 2015), but Meister (2011: 144) still regarded A. compta as a member of the geronto- geous subgenus Antheraea, while Paukstadt et al. (2000, 2003) suggested that the issue requires further clarifica- tion. Arora and Gupta (1979: 30) drew the venations of Indian Antheraea spp. and noticed the forewing R1 [radial vein 1] of A. compta (“Maghalaya, Cherra Punyi, 1220— 1524 m., 1 4, 24.vii.1931 [sic]”) is relatively longer than that of A. assamensis, with a more medial arising point on the anterior margin of the discal cell, but those two authors did not discuss the New World taxa. Except for Khasi Hills, specimens of A. compta have also been collected from “Assam, Jaintia Hills... 12 Juill. 1922 [sic]” and recorded by Bouvier and Riel (1931: 53), subtrop- ical forests of the two areas were formerly in Assam, but be- long to Meghalaya today (Gupta 2000); furthermore, a male moth from the adjacent Hahim of Assam was reported by Peigler (1999). Bryk (1944) published a more distant record of specimens mentioned as A. compta discovered at “Kam- baiti, 2000 m, 9. VI-17.VI [sic]” [Kanpaikti Sub-Township, Kachin State] in northeastern Burma by the entomological expedition of René Malaise in 1934 (specimens viewed by dez.pensoft.net Zhengyang Liu: Life history and natural distribution of Antheraea compta D Figure 1. Living adults of Antheraea compta, dorsal views. A. 4, calm condition; B. <, frightened condition; C. 9, calm condition; D. &, frightened condition. Scale bar: 4 cm. Richard S. Peigler in Stockholm in 2005, personal commu- nication to Liu from Peigler on 31 Aug. 2022). The third core population of the species inhabits southeastern Tibet (Fig. 2), first noted as “Motuo [Metok] County... 2120 m, VII.2013 [sic]’ by Naumann and Loffler (2015) (a separate species? see below), and in 2019, living material from the same ecoregion was unsuccessfully reared on Quercus sp. (Fagaceae), died in L, without detailed description (Nau- mann and Nogueira 2021), but suggested the caterpillars “fit very well with typical habitus of 7elea larvae [sic]”. The above mentioned moths were collected at upland mid-altitudes (ca. 1000—2500 m) in the height of sum- mer, basically June-July (Bouvier and Riel 1931: 53: “11 Déc. 1903 [sic]” possibly invalid), so the species is considered to be a strictly univoltine Himalayan flyer. It must be admitted that specimens of A. compta are rare in collections worldwide, so obviously knowledge of its complete preimaginal stages is of great significance for our understanding of the evolutionary history of the ge- nus Antheraea or even the family Saturniidae. Material and method INSIZE 1108-150C 0-150 mm/0.01 mm (£0.02 mm) was used to measure general lengths. XINGYUN FA1204E 120 g / 0.0001 g (4 0.0002 g) for recording weights. BENETECH GM1365 (+ 2 RH%; + 0.3 °C) for recording air humidity and temperatures. BENETECH GM1020 (< 10000 Lux + 3%; > 10000 Lux + 4%) for recording illuminances. PHILIPS TL 6W (UV-A, peak: 365 nm), was used for fluorescence photographs. All color figures were photographed by a NIKON D5500 with SIGMA 10-20 mm //4—5.6 lens or LAOWA 60 mm //2.8—22 lens. ZEISS GeminiSEM 360 was used for the SEM observa- tions and photographs, based on sputter-coated samples of 2 ova,6L,, 1 L,, 1 L, and 1 cocoon of A. compta. The Dtsch. Entomol. Z. 70 (2) 2023, 261-282 263 Figure 2. The natural habitat of Antheraea compta in southeastern Himalayas. Médog County, Tibet Autonomous Region, China, 2122 m. 28 Jun. 2021. dez.pensoft.net 264 sources of botanical and other zoological materials will be given in the corresponding paragraphs below. Terms involving continuity in time or space, the cor- responding numbers use the subscript format, including: L_, = 1-6" larval instars; T, , = 1*-3™ thoracic seg- ments; A__,, = 1*-10" abdominal segments. Structures not having the above relationship between each other are cited: S = stemma [ocellus]; BaS = sensil- lum basiconicum; ChS = sensillum chaeticum; TrS = sen- sillum trichodeum; PIS = sensillum placodeum; CaS = sen- sillum campaniformium; DiS = sensillum digitiformium; StS = sensillum styloconicum. If necessary, ending with Arabic numerals (non-subscript) for coding (e.g., “S1”, “ChS1”). Classification of the sensilla mostly consults some lepidopterous works, i.ec., Dethier (1937, 1941), Schoonhoven and Dethier (1966), Grimes and Neunzig (1986a; 1986b) and Faucheux (1999: 247-280). Ryan (2002: 113-114) expressed that the term “chaetic” is ap- plicable to thicker-walled bristles or spines, the word “trichoid” means slender and hair-like, and both can be used for chemo- or mechanosensilla (Zacharuk and Shields 1991; Keil 1999; Shields 2008), so they are some- times hardly distinguishable externally. Schneider (1964) thought such definitions are strictly morphological, the former “distinguished by a specialized and flexible circu- lar membrane at the base [sic]’, and the latter “without any specialized basal cuticular ring serving as articulating membrane [sic]”. In addition, the word “sensillum (sen- silla)” is used in figures only for some specified structures which are borne on antennae, maxillary palpi, maxillary mesal lobes and labial palpi. Here are labelled other prima- ry setae and pores on labrum, mandibles and maxilla-hy- popharynx-labial complex into only “lateral (L)’, “medial (M)’, “dorsal (D)” and “ventral (V)’ in figures, but does not rule out that they may have special sensory functions. Larval chaetotaxy, described below as far as possi- ble, follows the universal terminology named in earlier Lepidoptera publications, primarily based on Heinrich SD-III T1 7T2-3 At A2 Zhengyang Liu: Life history and natural distribution of Antheraea compta (1916), Gerasimov (1935) and Hinton (1946), with also the reviews by Chu (1956), Stehr (1987), Piao and Lee (1998) and Hasenfuss and Kristensen (2003). Addition- al works treating the standard setal coding of the fam- ily Saturniidae were consulted, of which Pease (1960), Heppner and Wang (1987) and Rougerie and Estradel (2008) are examples. The abbreviations of general areas are as follows: O = ocellar [stemmatal |; SO = subocellar [substemmatal]; F = frontal; AF = adfrontal; C = clypeal; G = genal [microgenal/midgenal]; A = anterior; V = ven- tral; SV = subventral; L = lateral; M = medial; XD = tac- tile dorsal (on the anterior margin of prothoracic shield and near the dorsal midline); D = dorsal; SD = subdorsal; MD = microdorsal [vertical]; P = parietal [posterodorsal]. Combined with these directional terms, Arabic numerals (non-subscript) and lowercase letters respectively code related primary setae and pores of the head (e.g., seta “SOI”, pore “MDa”). The term “pore” for text below may appear to be depressed (pierce the cuticle, or not) or elevated. Further, the term “palpifer” is only used for the maxillae in this work, whereas “palpiger” is considered as a part of the labium, although some entomologists used the latter term within the maxillae (e.g., Peterson 1948: 119; Schoonhoven and Dethier 1966). Nassig (1989) and Deml and Dettner (2002) classified saturniid scoli focusing on their basal morphology and functionality (secretions), but this article only divides related primary setae into the following four conditions expressed by Roman numerals on the setal map of T —A,, (Fig. 3); each of them is considered as a “term” for modi- fying singular or plural nouns, hyphenated with a general area abbreviation (e.g., seta(e) “D-I’, chalaza(e) “D-I”, area(s) “D-I”’), distinguished in detail with final Arabic numerals (non-subscript) if necessary (e.g., seta(e) “D- 11”, seta(e) “D-I2”): I = For describing a single seta, or the structure/loca- tion in which it resides [uni-setal]. If used for the A3-6 A7 As As Ato Figure 3. L, of Antheraea compta, the setal map, lateral view, with only the primary chaetotaxy coded. The setae of cephalic regions and legs T, , are not shown, proleg A,, displays the mesal surface, the ventral midline constitutes the bottom margin of A, ,. dez.pensoft.net Dtsch. Entomol. Z. 70 (2) 2023, 261-282 plural, for example, several setae within the desig- nated area, then each seta is treated as an individual existing independently which could be objectively distinguished from others. II = For describing a pair of setae (n = 2), or the structure/location in which they reside collective- ly [bi-setal]. If used for the plural, for example, several setal pairs within the designated area, then each pair is treated as an individual existing inde- pendently which could be objectively distinguished from others. III = For describing a cluster of setae (n > 2), or the structure/location in which they reside collective- ly [multi-setal]. If used for the plural, for example, several setal clusters within the designated area, then each cluster is treated as an individual existing independently which could be objectively distin- guished from others. IV = For describing uncertain condition on the desig- nated structure/location [uni/bi/multi-setal]. Results Morphology of preimaginal instars Unless otherwise specified, numerical (metric) data was based on single normal individual in this chapter, and the fluorescence tests are only valid for naked eye vision. 265 Ova (Figs 4, 8A) Tri-axial ellipsoid, ca. 3.71 mm (length) ventral view; K. , ventral view; a—c: A, 9: d: sexual gland; L. 9, ventral view; a—c: A, ,,; d: sexual gland; M-T. Latero-medioapical views; M. °, scolus D-III of T,; N. 9, scolus D-III of A,; O. 9, 8-10? scoli D-III of A, (medially fused); P. 2, scolus SD-III of A,; Q. 2, scolus SD-III of A,; R. 2, scolus L-III of A; S. @, scolus SV-II of T,; T. 3, chalaza SV-I of A,. Scale bars: 2 cm (A, B); 1 cm (C, D); 5 mm (E, G, 1D; 2 mm (F, M-T); 1.5 mm (H); 2.5 mm (J-L). dez.pensoft.net Dtsch. Entomol. Z. 70 (2) 2023, 261-282 L, (Fig. 8G, H) General morphology close to L,, but the head capsule is paler and widened to 3.73 mm. The flash basal parts have enlarged the volume further on scoli SD-III of A, ,, es- pecially. Supplemented long setae with club-like apexes on scoli D-III of T,—A,, as well as on chalazae SV-I of A,_, and scoli SV-III. Dense coryneform bristles initiating to present on the posterior zones of the middorsal A, ,, colored as yellow vividly; they are minute and cross a transverse row combined by longer and pointed setae in the same color but borne on the anterior zones of A, .. Tricolored lateral plates of prolegs A,, observed in most of L, now shared in all individuals. More secondary hairs are exhibited principally on ventral integument. Some of them are based on dollar spots, such dots originating since L, are now widely distributed to level L, gradually colored from black to light brown. The marginal setae of A,, are pronouncedly elongated and orientated to the pos- terior side. The numbers of 50—55 crochets arranged as biordinal mesoseries on each planta. L, (Fig. 8I-L) The macroscopic pattern is essentially same as L,, but the 5.47 mm width head capsule begins to appear more yellow- ish in proportions, mainly enriched on its ventrolateral and frontal zones, but the dark sepia strip ornamented on ocellar areas are still visible. The color of cervacoria is more vivid, slanted, parallel dark olive-green streaks arising between the levels SD and Lof A, ,. For T,—A,,, the venter is clearly darker than dorsa, the dense dark spots are developing to the level SD with some secondary hairs, and the coxal apexes of legs T, , are colored lightly like T,. The clubbed setal apexes already occur on the longer bristles on prolegs, starting from this instar, each lateral plate of anal prolegs shows a very minute black dot on the yellow tint area with relatively fixed position, and each lateral margin of the anal shield formed into vitreum, enveloping an internal dark brown band. Plan- tar crochets are still biordinal mesoseries but increased to numbers of 54-60. Take L, as an example, for a fresh larva or during the pre-molting, the oblate setal tip could be seen white translucent, whereas the silver-reflective zones of sco- li are transparent just after ecdysis but fully colored after approximately about 1 8—22 hours (Fig. 8], L). L, (Fig. 8M-P) Head capsule (width = 7.33 mm) 1s paler distinctly, but the distribution patterns of colors are basically the same as L,. The cephalic setae are more strongly developed, the dotted integument reaching the level D and all the sec- ondary hairs turned into brownish to black with longer sizes. Parallel oblique stripes lying around spiracles A, , are more evident, each spiracle is outlined by pale aqua- marine margin. Scoli XD-III and SD-III on each side of prothoracic shield are highly fuse to be a unified base out- wardly. The band inlaid inside the vitric margin of anal 273 shield is now darker. The shining silvery bases of scoli D-II of T,—-A,, and SD-III of T,—A, are already spectac- ular (largest on scoli SD-III of A, , still), their epicuticles are extraordinary smooth in stark contrast to the green integument, the latter is densely covered noncellular pro- cesses (Fig. 8O, P). Proleg plantae are surfaced with lat- eral dark gray and medial white with 61-64 crochets in biordinal mesoseries. Each of the lateral plates of prolegs A,, 1s constituted by the wrinkled bright yellow-green area, the smooth dark ochre posterior margin, and a nar- row black stripe between the former two. L, (Figs 8Q, R, 9, 10) The final larval instar with giant body, head capsule is 10.32 mm width and the overall color pattern is near L, but more pallid (Fig. 9C, G). Combined with the high-sat- uration goldenrod cervacoria and the pale lemon coxae, the front margin of T, forms a bright yellow annular zone, which showed intensely fluorescent enrichment under the excitation by UV 365 nm (Figs 9G, 10B). Fluorescence was not detected in the flash bases with the clubbed se- tae (Fig. 10C), but weakly appeared on the yellow-green regions of the central anal shield and the lateral plates of prolegs A,, (Fig. 10D), the wrinkles of the latter are more obvious than those during the prior stage (Fig. 9E). Both scoli XD-HI and SD-III have different numbers of prima- ry setae during L, . which appeared as short spines, but all of these are elongated into hair-like now (Fig. 9C, G). Similar to L, ., for all scoli, only partial setae on T—A, have clubbed tips (Fig. 9M—T), statistic on the single larva displayed that most setal quantities of the corresponding sites are same as L,, but that on 12 scoli are decreased and 2 scoli are increased (Table 2). The rod-like part of the clubbed seta is smooth helical, the flaky end is paved by shallow pits densely but strongly reflection (Fig. 8Q, R). There is a significant difference in general integumenta- ry color, ie., 2 2Y are more yellowish than 3 4, and the shiny scoli of females are tending to be purple-red but that of males look more purple-blue (Figs 9M-—T, 10A). Observed the male sexual gland [“Herold’s gland” to be a slight mark localizing on ventral midline near to A,/A, boundary (Fig. 9K), the female sexual gland [“Ishtwata’s gland”] arising as two pairs of dots on ventral A, and A, (Fig. 9L). Distinguished from L,, the dark spots are added to the middorsal area except the prothoracic shield, and the secondary hairs are increased on the whole body at pres- ent, but except the sizes, the glassy margins of anal shield as well as the spiracles are morphologically almost un- changed (Fig. 9F, H). Vesicular processes were observed on the distomedial areas of each tibia and tarsus, and all the medial setae (both primary and secondary) of legs T, , form white thick spines (Fig. 91), unfortunately, it lacked a leg observation for L, .. Plantae of A, , and A,, bear 59-65 crochets within biordinal mesoseries (e.g., Fig. 9J). The liquid feces after feeding ended (all larval instars fed from Q. yunnanensis) is bronzed mucus (Fig. 10E), after that, 1 4 was recorded the weight as 14.77 g. dez.pensoft.net 274 Zhengyang Liu: Life history and natural distribution of Antheraea compta Figure 10. L, of Antheraea compta. A. Lateral view; a: 3; b: 9; BED. 3, under 365 nm UV; B. Head and T —A,, ventrolateral view; C. A., lateral view; D. A cle of cocoon; G. <, spinning the external part [floss] of cocoon. Scale bars: 2 cm (A); 1 cm (B, F, G); 5 mm (C-E). lateral view; E. ~, the last defecation [liquid defecation] from A.,, lateral view; F. @, spinning the pedun- 10? 10° Table 2. L, of Antheraea compta, a statistic based on primary chalazae/scoli which have clubbed setae, with the format “short 6699 spines + elongated hairs (how many clubbed ones)’. The “<” is lower, the is same, and the “>” is higher than the normal value of the corresponding locations during L, (see Table 1). Merged cells means the scoli are medially fused. Segments\Areas SV-I/III L-Il SD-II XD/D-III XD/D-II SD-II L-Il SV-I/II i 0+4(1)- 1+7(6)- 0+4(1)- 0 +4(2)- 0 +3(2)- 0+4(2)- 2+7(4)- 0 +4(0) T, 0+1(1)< 4 +2(2)- 3 +3(3)- 5 +3(3)- 5 +3(3) 3 +2(2)< 4 +2(2)- 2+10)- ie 2+1()- 4 +2(2)- 3 +3(3)- 5 +3(3)- 2+4(4)< 3 +2(2)< 4 +2(2)- 2+1()- A, 0+1(1)- 4 +2(2)- 3+1(0)- 3 +2(2)- 1+1(1)< 4+1(1)? 4+1(1)< 0+1(1)- A, 0+1(1)- 3+1(1)< 3+10)- 2 +2(2)* 3+10)- 2 +2(2)- 4 +2(2)- 0+1(1)- A, 3 +3(3) > 3+10)- 3+1()- 3+10)- 3+1()- 4 +2(2)- A 3 +2(2)< 3+1(1)- 3+1()- 3 + 2(2)> 3 +10) 4 +2(2)- iy 3+2(2)< 3+10)- 3+1(1)- 3+1(1)- 3+1()- 4 +2(2)- Ay 4 +2(2)- 3+10)- 3+1(1)- 3 +0(0) « 3+10)- 4 +2(2)- A, 4 +2(2)- 3+1(1)- 2 +2(2)- 3+10)- 3+1()- 4 +2(2)- A, 3 +2(2)< 3+10)- 2 + 2(2)- 2 +2(2)- 3+10)- 3 +2(2)< x 3 +3(3)- 2 +2(2)- 2 +2(2)- 4 +2(3)- Pupae (Figs 11, 12C) The epicuticle is dark maroon to black except the trans- lucent epicranial plate [frontoclypeus]. Females are larg- er (lengths: 3 ¢¢ = 40.58 mm, 39.64 mm, 38.68 mm; 2 92 = 42.63 mm, 44.48 mm) and heavier (weights of during overwintering diapause, measured on 06 Feb. 2023: 3 dd = 8.16 g, 7.46 g, 7.54 g, 2 99 = 9.29 g, 8.51 g) than males. The antennal margins in females are more flattened than males, the pair of antennae of the latter are not medially touching along the ventral mid- line, the maxillae and legs T, , are visible in both sexes (Fig. 11C, E). Spiracles T, is gapped on the boundary of T/T, (Fig. 11F), spiracle A, , are functional and large, but that of A, is closed as a remnant. A pair of forewing tubercles [adult spurs] is rendered on the dorsolateral ar- eas of T, specifically (Fig. 11F). The biserial larval sco- li D-III had been reduced to smooth vestiges on pupal dez.pensoft.net stage, in only one case that such bulges on T,—A, are obvious (Fig. 11H). The unsclerotized joints are junction zones between A,/A,, A/A,, and A./A_, the tip of A,, is rounded without cremaster (Fig. 11G). Genital pores are navel-shaped scars apertured midventrally for males (A,) and females (A, [ostium bursae] and A, [ostium oviduc- tus]) (Fig. 11B, D). Cocoons (Figs 10F, G, 12) Large (max-lengths without peduncles: 3 gd = 63.49 mm, 52.14 mm, 56.44 mm; 2 99° = 50.37 mm, 60.41 mm), single layered (thicknesses = ca. 0.37— 1.16 mm) and fully closed (no pre-formed exit), the external part was woven by floss for attaching to leaves (Fig. 10G), the general shape is irregular polyhedron but the silky pupal cell is prolate spheroid. White col- ored in dry environment (weak fluorescence under UV Dtsch. Entomol. Z. 70 (2) 2023, 261-282 Figure 11. Pupae of Antheraea compta. A. a: <, lateral view; b: @, ventral view; c: @, dorsal view. d: °, lateral view; e: 2, ventral view; f: 2, dorsal view; B. 3, ventral view; a—d: A___; e: genital pore; C. 4, head and T 7-10? e, f: genital pores; E. 9°, head and T 1-2? ventrolateral view; F. 4, T _» ventrolateral view; D. °, ventral view; a—d: A, ||; dorsolateral view; a: spiracle T,; b: forewing tubercle; G. 3, pos- terolateral view; a, b: A, ,,; H. G, dorsolateral view; a-e: T,—A,. Scale bars: 1 cm (A); 2 mm (B, D); 5 mm (C, E); 3 mm (F-H). 9-10? 365 nm, barely visible) but metamorphosing into brown tones in wet (fluorescence disappeared). 2 Jd and 1 9 spun on erect vegetations with wide and loose pedun- cles of inferior structure (e.g., Figs 10F, 12A-C), 1 and 1 2 spun cocoons within fallen leaves and branches on the ground without peduncle (e.g., Fig. 12D). Each filament is a double-strand flat belt, wrinkles formed by longitudinal stretching could be observed on epi-seri- cin (Fig. 12F), the part of fibroin is combined by dense elongated fibers longitudinally to be almost non-porous within (less voids) (Fig. 12E). Microscopically sur- veyed three pieces (ca. 6.5—8.4 mm_*) of silk-layer sliced from random sites of a cocoon, showing the external surface is rougher, the filaments are ca. 83.8—94.4 um wide and ca. 4.7—9.3 um thick (Fig. 12H). The inter- nal surface [pupal cell] is smooth and flat relatively, widths of the filaments are ca. 56.4—92.8 um with the thicknesses of ca. 2.3—3.9 um (Fig. 121). Many crystals (mainly calcium oxalate) granulated from the prepupal discharge liquor are scattered or concentrated in the silk-layer (Fig. 12G), due to gravity, the precipitation is more obvious in the outer surface of the cocoon that lies closer to the ground. Rearing report During my first expedition on Tibetan Nyingchi, 1 9 of A. compta was collected on 28 Jun. 2021 from Médog 2122 m (Fig. 2), her 60 ova were hatched into 37 larvae in the ex- perimental site in Kunming City of Yunnan Province, 1940 m, during mid Jul. to early Aug., these fresh L, rejected any Lauraceae, 1.e., Cinnamomum camphora, Cinnamomum burmannii, Machilus spp., Lindera sp., Neolitsea sp., Lit- sea pungens and Litsea populifolia, moreover, Magnolia delavayi (Magnoliaceae), Prunus cerasoides (Rosaceae), Salix babylonica (Salicaceae) and Liquidambar formosana (Altingiaceae) were likewise not accepted. Finally, three individuals nibbled the leaves of Quercus variabilis and Q. yunnanensis, but only one grew into the 3“ pre-molting with the latter, and because it was reared outdoors without a cage, this larva was preyed upon by a Myophonus caeru- leus (Aves: Passeriformes, Muscicapidae). After the first failed attempt, I recaptured 3 99 adults with some males (Fig. 1) from the same natural habitat; during the rainy nights of 25—26 Jun. 2022, many large Quercus lamellosa were noticed at the site. Successively, 3 (coded as Gi, laid on 26 Jun.), 102 (coded as Gui, laid dez.pensoft.net 2/76 Zhengyang Liu: Life history and natural distribution of Antheraea compta A 4 = ; Figure 12. Cocoons of Antheraea compta. A-C. @, spun on the erect vegetation, lateral view; A. Original condition; B. Removed some leaves and branches; C. Opened; D. &, spun on the bottom of the net cage under a fallen leaf, lateral view [the underside]; E. so Me? filament of the external part, vertical view; G. <, the crystals precipitated from the prepupal discharge liquor, vertical view; H. 2, filaments of the external part, cross sectional view; a: transverse [horizontal] plane; b: diagonal plane; c: sagittal plane; F. ¢ V9 filaments of the external surface, vertical view; I. @, filaments of the internal surface, vertical view. Scale bars: 3 cm (A—D); 10 um (E); 20 um (F); 2 um (G); 100 um (H, I). dez.pensoft.net Dtsch. Entomol. Z. 70 (2) 2023, 261-282 on 25 Jun.-01 Jul.) and 16 (coded as Giil, laid on 26—30 Jun.) eggs severally into paper envelopes. Each of Gi and Gili was oviposited naturally into a single tier without longitudinal accumulation, Gii was the same condition in the early period but later heaped up to lump because of larger quantity and lack of available space for adult abdo- men. The material was brought back to the same site in Kunming indoor and then grouped on three mesh pieces, a cotton pad was placed below with interval ca. 1 cm, wa- ter sprayed the ova every day, the circumambient air was ca. 17-21 °C with ca. 80-95% RH. This method helped prevent water from accumulating around the eggs; excess water drained through the mesh openings to the cotton pad below, while the space between helped ventilate the bottom of the eggs to prevent mold. At Beijing Time 06:00 to 16:00 every day, 3 larvae of Gi hatched on 13 Jul. (ovum period 17 days), 81 larvae of Gu hatched during 12—19 Jul. (ordered as: 10, 27, 12, 5, 15, 6, 5, 1; ovum period 17—18 days) and 14 larvae of Giii hatched during 13—18 and 20 Jul. (ordered as: 3, 4, 3, 1, 1, 1, 1; ovum period 17-18 or 20 days, but the last in- dividual was unhealthy and died on the day of hatching). Newly hatched L, usually fed on eggshells preferentially. Checking unhatched material on 31 Jul., 17 ova of Gil and 1 ovum of Giil were devoid of embryonic develop- ment and shriveled; mature embryos had died in 4 ova of Gu and 1 ovum of Gill. Fresh L, were moved to a semi-open balcony for rear- ing, without cage protection. The plant stems were inserted into the bottle filled with water, and the bottleneck blocked with paper towels to prevent larval drowning. Some new- ly hatched individuals did not bite the leaves of Quercus schottkyana and Magnolia denudata within two days, so all larvae were given Q. yunnanensis, which was readily accepted. These caterpillars had weak positive phototaxis, usually did not climb to the apex of host plant and did not exhibit gregarious behavior, but there were often 2—5 scat- tered L, ona single leaf. They rested on the undersides of the leaves and moved to the edges only when feeding, if a branch had fresh terminal buds and thick mature leaves, newly hatched larvae did not distinguish between them, as they are perfectly capable of gnawing on harder leaves, which is apparently related to their larger head with pow- erful mandibular adductors. Although they did not have strict requirements for air humidity during the larval stage (kept in ca. 64-98% RH), the temperature should not ex- ceed 28 °C, otherwise the larvae will die in large num- bers. When the environment reached 26 °C, they usually unclasped the legs T, , and prolegs A, , from host plant and suspended in air to enhance ventilation for heat dissi- pation. This may cause many larvae to fall to the ground. The whole larval stage accompanied a strong need for drinking water, and the leaves must be sprayed with water several times each day to ensure normal development of caterpillars, except on rainy days. A. compta is a rather difficult species to rear in cap- tivity with a very high mortality in L,, in this study, only four individuals of Gi and five of Gili survived to the 1* pre-molting. However, two larvae within the latter group 2dF died during pharate and two died in early L,. In the five surviving L,, the individual “Gilil” was caged in outdoor and other four still fed semi-openly, until they cocooned respectively. Although the weather and nutritional con- ditions are identical, ontogenic time-consumptions are quite different in the same gender (Table 3). Before entering each pre-ecdysis, larvae always moved away from their last feeding site. In late L, they cut the petiole to make the leaf fall to the ground or break it off completely to eliminate signs of feeding, which apparent- ly helps to avoid some natural enemies which could find the caterpillar location by defoliation. Some individuals would sometimes entangle silk in selected locations to an- chor themselves before shifting into apolysis state; such behavior was not consistent, but each of them always pref- erentially fed on their exuviae except the head capsule. During L, ,, larvae usually rested on the leaf midrib af- ter a period of feeding of the lamina and secondary veins. For L, to medium L, they rested intermittently near the petiole and gradually nibbled along the midrib. Since late L, the weight only allowed the activities on branches, be- fore feeding a new leaf, the larva often first bit a large pit on the petiole and then ate along the basal area, if nec- essary, gnawed more gaps on midrib for further folding the leaf apex closer to itself. For mature larvae, feeding behavior mostly occurred at night, probably to avoid de- tection by birds. A continuous observation on single L, was recorded for reference (Suppl. material 1). All larval instars were sensitive to light, 1.e., in resting states, they always faced toward the side where the illu- minance is higher (Suppl. material 2), and this behavior would draw predators’ attention to the bright yellow ring on the front of T,, whose fluorescent was sufficient to in- dicate it plays a role in defensive warning. These cater- pillars were extremely alert, and every slight shake would interrupt the feeding behavior. Under this state, larvae usually detached their legs T, , and prolegs A, (some- times A, ,) from the host plant, which may be an effec- tive mimicry to make the lateral views more like leaves, sometimes clicking the mouthparts as a popping noise. The most alarmed individuals would swing from side to side, even triggering biting, whereas the special clubbed setae may help to enhance the spatial scope of tactile cue Table 3. Developmental data of different individuals of Anther- aea compta, fed from Quercus yunnanensis. Stages\Codes Gil, 6 Giii2, ss Giii3, ss Gitid, 2s Gili, 2 Oviposited 30 Jun. 27 Jun. 28 Jun. 28 Jun. 25 Jun. Hatched into L, 17 Jul. 14 Jul. 15 Jul. 15 Jul. 12 Jul. 1*' pre-ecdysis 27 Jul. 25 Jul. 26 Jul. 26 Jul. 23 Jul. Molted into L, 29 Jul. 27 Jul. 28 Jul. 28 Jul. 25 Jul. 2™ pre-ecdysis 05Aug. O7Aug. O8Aug. O8Aug. 06Aug. Molted into L, 07 Aug. 09 Aug. 10 Aug. 10 Aug. 08 Aug. 3"¢ pre-ecdysis 14 Aug. 16 Aug. 18 Aug. 18 Aug. 17 Aug. Molted into L, 16 Aug. 18 Aug. 20 Aug. 20 Aug. 19 Aug. 4" pre-ecdysis 23 Aug. 26Aug. 28Aug. 30Aug. 28 Aug. Molted into L, 26 Aug. 29 Aug. 31 Aug. O1 Sep. 31 Aug. 5" pre-ecdysis 05 Sep. 10 Sep. 14 Sep. 15 Sep. 17 Sep. Molted into L, 09 Sep. 15 Sep. 18 Sep. 19 Sep. 21 Sep. Feeding ended 03 Oct. 09 Oct. 12 Oct. 16 Oct. 23 Oct. Spun cocoon 04 Oct. 11 Oct. 13 Oct. 17 Oct. 24 Oct. dez.pensoft.net 278 (Suppl. material 3). Even so, no secretions exude from the scoli under stimulated conditions, and no urticating reaction was detected on human skin after touch. Larvae never left the host plant voluntarily if the leaves were plentiful. However, they quickly climbed or fell to the ground after feeding ended [liquid defecation] (hap- pened at night usually), whereupon they were caged with upright living vegetation and dead leaves at the bottom for observation. Almost all individuals moved restlessly for about 24 hours before choosing a spinning location. The species undoubtedly constructs cocoons away from the host plant in the wild, either in low shrubs under the oak canopies or in the leaf litter on the ground. The flight time in the natural habitat and captivity and reproductive behavior of adults will be recorded and pub- lished after the fresh adults emerge from existing pupae. Discussion and conclusion This article provides the first report of the complete life history of the enigmatic A. compta, whose lateral stripes of mature larvae are a feature close to that of North American Antheraea oculea (Neumoegen, 1883) (Tuskes et al. 1996: pl. 6; Lampe 2010: 121) and A. polyphemus (Bouseman and Sternburg 2002: 51; Hall 2021), the scale and distribution of the shiny silver spots closely resemble those in the Mexican Antheraea montezuma (Sallé, 1856) and Antheraea meridiana Naumann & Nogueira, 2021 (Paukstadt and Paukstadt 2020; Naumann and Nogueira 2021), while the tricolored lateral plate of anal proleg of Neotropical A. godmani (Nassig et al. 1996; Goossens et al. 2015) is similarly shared in A. compta. These obser- vations preliminarily suggest that Himalayan A. compta should be assigned to the subgenus Te/ea. Due to L, rejecting Q. schottkyana in captivity, it is inferred that Q. /amellosa is observed in the Tibetan hab- itat, and more cycle-cup oaks may not be the correct host plants to A. compta, namely the section Cyclobalanopsis. Additionally, in the eastern Himalayas, native specimens Quercus lodicosa, Quercus lanata, Quercus engleriana, Quercus semecarpifolia, Quercus aquifolioides, Quer- cus gilliana, Quercus rehderiana and Quercus senescens within the section //ex, as well as Quercus acutissima of the section Cerris were also identified in recent works (Zhou and Sun 1996; Denk and Grimm 2009; Yang and Zhou 2015: 130; Lahiri et al. 2017), the three sections have been confirmed to form a monophyletic subgenus Cerris (Denk et al. 2017; Hipp et al. 2019). Because of the observed rejection of Q. variabilis (a single larva re- luctantly ate a small gap, but did not continue), its sis- ter taxon Q. acutissima would probably also be rejected. There is currently no experimental result on infrasubge- neric //ex so it is uncertain whether A. compta can par- tially accept some species of the subgenus, but probably, Q. lodicosa and Q. lanata would be accepted as foods by such caterpillars; the former tree is common in south- eastern Tibet and the latter is dominant from Médog to Tengchong County, Yunnan (bordered with Myanmar dez.pensoft.net Zhengyang Liu: Life history and natural distribution of Antheraea compta Kanpaikti), all having humid mid-altitudinal distribu- tions (Zhou et al. 1995). In any case, the above species are Clearly distinguished from Q. yunnanensis, a decidu- ous oak within the subgenus Quercus. Another roburoid white oak Quercus griffithii is widely known in north- eastern India (Negi and Naithani 1995: 50-53; Singh et al. 2009; Singh et al. 2015), likewise naturally occurring in mountains from northern Yungui Plateau to sections of Indochina (Huang et al. 1999; Menitsky 2005: 47-49), with a high probability to be a primary host to 4. compta at the corresponding biotopes. A. assamensis 1s a well-known Lauraceae feeder in the wild (Seidel and Peigler 2018; Devi et al. 2021) but ac- cepted beech (Lampe 2010: 358) and oak (Crotch 1956: 55) in captivity. Interestingly, the current known host- plants of A. godmani are limited to oaks (Meister 2011: 148), while A. compta is also oligophagous and feeding on certain Quercus only. This means that the empty cocoons collected from Assam soalu trees Litsea monopetala and then sent to Peigler (1999) were actually spun by muga silkworms. Therefore, Jolly’s (1981) conclusion that “A. compta is almost identical with A. assamensis [sic] is completely incorrect, while native Indian material for study of chromosomes (Gupta and Narang 1981) and silk (Luikham et al. 2017) using the former name but based on Lauraceae host plants are surely misidentifications to A. as- samensis or its sister species Antheraea castanea Jordan, 1910. In fact, no Indian author has ever provided an adult image to prove the true identity of so-called “4. compta”, except Arora and Gupta (1979) who described a pinned male from the British India period. In recent years, there have been a few indigenous records from the country but limited to only checklists without reliable morphological description and often confusingly called “wild muga” or “oak tasar” (e.g., Bhatia et al. 2010; Devi et al. 2011; Ku- mar et al. 2016; Gogoi and Goswami 2016; CSB 2018; Marepally 2018: 4; Boro and Borah 2020; Kumar et al. 2020; Keisa et al. 2022). This confusion suggests that the current population of the species may be nearly extirpat- ed because virtually no local researchers have actually collected A. compta. Coincidentally, in works in which illustrated moth photographs corresponding to the scien- tific names, no specimen of A. compta was recaptured in recent surveys of northeastern India (Gogoi et al. 2014; Kakati and Chutia 2009; Kalita and Dutta 2014; Lalhmin- gliani 2015; Sondhi et al. 2021a) or even its type-locali- ty Meghalaya (Shangpliang and Hajong 2015a, 2015b), whereas also does not turn up in the long-term monitoring of the Himalayas and Indian subcontinent (Shubhalaxmi 2018; Chandra et al. 2019; Sondhi et al. 2021b). To date, no Chinese publications included this species, and no re- cords from Bhutan, Nepal and Uttarakhand are available, whereas Schiissler (1933: 175) added Sikkim to the distri- bution without supporting data. Actually, the moths flying in southeastern Tibet were separated taxonomically from Meghalaya A. compta, the former was named Antheraea discata Naumann & LOf- fler, 2015, whose type locality is the same as the origin of my material. Unfortunately, the authors of A. discata Dtsch. Entomol. Z. 70 (2) 2023, 261-282 overlooked the study of specimens from mountainous Burma, and although they discovered some morphologi- cal and COI differences between Médog and Khasi Hills, the Myanmar samples are still crucial in resolving the phylogenetic relationships in this case. Unlike the natural barrier of the Assam Valley, the mountains of Sagaing- Kachin are connected to the Shillong Plateau and Himala- yas, making it a logical corridor for the ancestral colonial activity of these moths. This suggests that the “species boundary” of A. discata is possibly fuzzy (transitional or gradual) or even not able to be located. Therefore, this paper rejects to use the name subjectively and treats it as “data-deficient” temporarily, however, it does not im- ply any effective taxonomic (synonymous) treatment for this taxon here. There is still a possibility in the future that the three ecoregional populations will be accepted as different species or subspecies to form a complex; the Sub-Himalayan genus Sinobirma Bryk, 1944 is a known case (Rougerie et al. 2012). At present, the sources of published specimens of A. compta are localized within the following three re- stricted ecoregions of Indo-Malayan realm, the zoning based on Dinerstein et al. (2017): ¢ Meghalaya subtropical forests (Rothschild and Jor- dan 1899; Rothschild and Jordan 1901; Bouvier and Riel 1931: 53; Arora and Gupta 1979; Peigler 1999; Naumann and Loffler 2015). ¢ Northern triangle subtropical forests (Bryk 1944). ¢ Eastern Himalayan broadleaf forests (Naumann and Loffler 2015; Naumann and Nogueira 2021; this article). Diverse evidence stresses the importance and necessity for conservation of wild A. compta in the current situa- tion. As one of the central clues regarding the evolution- ary history of the economically important genus Anther- aea, its ecological and taxonomic significance 1s critical. Larval feeding preference and morphological charac- ters shared by the montane A. godmani, A. montezuma and A. meridiana with the Himalayan A. compta seem- ingly suggest that those Neotropical species are closer to the New World ancestor than are the Nearctic species (A. polyphemus, A. oculea). Three centuries of intense study of the genus Antheraea make more rigorous com- parisons possible, and detailed phylogenetic studies on preimaginal morphologies and complete mitochondrial genomes of A. compta with related species are in prepa- ration, to further confirm its evolutionary status. Acknowledgments I extend my sincerest thanks to the saturniidologist Dr. Richard S. Peigler (San Antonio), for his invaluable assis- tance with literature, manuscript review, and editing my English. His expertise in sericulture has greatly inspired and guided this article. Dr. Stefan Naumann (Berlin) re- viewed the manuscript and offered related advice, along 279 with fine editing by Dr. Wolfram Mey (Berlin). And I would like to thank Museum fiir Naturkunde (Berlin) for waiving the publication costs of this article. References Arora GS, Gupta IJ (1979) Taxonomic studies on some of the Indian non-mulberry silkmoths (Lepidoptera: Saturniidae: Saturniinae). Memoirs of the Zoological Survey of India 16(1): 1-63. [pls. 1-11] Bhatia NK, Bhat MM, Khan MA (2010) Tropical tasar, utilization and conservation of natural resource for tribal development. The Ecos- can 1(Special Issue): 187-198. Boro P, Borah SD (2020) Biodiversity of sericigenous insects in north-eastern region of India — a review. Journal of Entomology and Zoology Studies 8(4): 269-275. 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The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for oth- ers, provided that the original source and author(s) are credited. Link: https://do1.org/10.3897/dez.70.102952.suppl1 Supplementary material 2 Light sensitivity (the darker side = ca. 544- 1185 Lux, the brighter side = ca. 9300-13600 Lux) of Antheraea compta L, (speed x 12) Author: Zhengyang Liu Data type: video (.mov file) Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons. org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow us- ers to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/dez.70.102952.suppl2 Supplementary material 3 Defensive behavior of Antheraea compta L, Author: Zhengyang Liu Data type: video (.mov file) Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons. org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow us- ers to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/dez.70.102952.suppl3