Corresponding author: Vadim V. Zolotuhin ( v.zolot@mail.ru ) Academic editor: Roman Yakovlev
© 2020 Vadim V. Zolotuhin, Sergey N. Pugaev, Tran Thieu Du.
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.
Citation:
Zolotuhin VV, Pugaev SN, Du TT (2020) A review of Apha floralis species group (Lepidoptera: Eupterotidae). Acta Biologica Sibirica 6: 611-635. https://doi.org/10.3897/abs.6.e59529
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5 species of the genus Apha Walker, 1855 are included here under the floralis group, three of which are described as new: A. zephyrus sp. nov. (type locality: China, Yunnan, Dianceng Shan, 2.200 m), A. witti sp. nov. (type locality: China, Sichuan, Qionglai Shan, 1.400 m) and A. chloralis sp. nov. (type locality: Vietnam, Cha Val, Nam Giang, Quang Nam, 546 m). Types of all species are illustrated, biology and morphology of preimaginal instars are discussed for A. kantonensis Mell, [1930] and A. chloralis sp. nov. A new synonymy for Apha Walker, 1855 and Preptothauma Draudt, 1931 (including a single mislabeled species P. oxydiata Draudt, 1931), syn. nov. is established.
Lepidoptera, Eupterotidae, China, Vietnam, Apha floralis group, taxonomy, morphology, new species
This article forms part of a planned revision of the genus Apha Walker, 1855, a group of tropical monkey moths which are endemic to southeastern Asia but not known to extend to Sundaland.
During work with the Vietnamese members of the family, identification proved to be problematic; reference data were fragmentary and often controversial, there were no data on genitalic characters of some species, and great taxonomic confusion in species interpretation was experienced. This lack of data prompted us to undertake this investigation.
According to the literature, the genus Apha Walker, 1855 consists of 6 species and 5 subspecies known mainly from China; their status is not always clear-cut and the status of the subspecies not always correct. Preliminary investigation of material from various museums, particularly those housing the types, and also of our own material, has shown that the genus includes a total of 16 species, 7 of which are not described. Of the latter, 3 are described in this article and the status of the remaining 4 taxa should be changed.
The genus Apha is divided into 3 groups based on similarities in genitalic morphology discussed hereunder. These groups are: floralis Butler, 1881 – valvae without a saccular lobe (except for A. witti sp. nov., which has only a weakly developed one), subdives Mell, [1930] – valvae with a saccular lobe, vesica bears an appendix, and tychoona Butler, 1871 – valvae with a saccular lobe, vesica without an appendix (
Material for this article comprises eggs and caterpillars of all instars reared by the junior author in Hanoi ex ovo from the original material taken by him in Central Vietnam. Additional material was obtained from the collections of various museums. Their abbreviations, as used hereunder, are given below:
CSLL – collection of Swen Löffler – Lichtenstein, Germany; IEBR – Institute of Ecology and Bio-Resources – Hanoi, Vietnam;
Further abbreviations used are: TL = type locality; | = a new line in the citations of the labels of holotypes and lectotypes.
The genitalic preparations illustrated were made using standard dissecting techniques and mounted in Euparal on glass slides. The preparations were photographed under magnification using Canon PowerShot A570 and Olympus Camedia C-750 camera with Soligor Adapter Tube for Olympus and Slide Duplicator for Digital 10 dptrs modified for object glasses. All figures in the article were produced by Sergey Pugaev using Adobe Photoshop 7.0 from photographs taken partly by himself, and also by Dieter Stüning, Tran Thieu Du, Vadim V. Zolotuhin and Alexey Prozorov. The final version of illustrations is also produced by Sergey N. Pugaev.
The sequences of mitochondrial gene cytochrome c oxidase I (658 bp) are used here for taxonomic purposes. They are kept in the base of sequences of “The Barcode of Life Data System” international project (BOLD, URL: http://www.boldsystems.org/) in authors’ projects LBEOW and LBEOA (Table
№ | Species | Sample ID | BOLD Process ID | Locality | Comments |
---|---|---|---|---|---|
1 | Apha chloralis | EUPT-11-043 | LBEOA045-11 | Vietnam: Quang Nam, Nam Giang distr., Cha Val vill. | Paratype |
2 | Apha chloralis | EUPT-DR-8 | LBEOW100-10 | Vietnam: Quang Nam, Phuoc Son, Deo Lo Xo | Paratype |
3 | Apha chloralis | VZ-EUPTER-39 | LBEOW190-10 | Vietnam: Cao Bang, Nguyen Binh, Phia Den | Paratype |
4 | Apha floralis | EUPT-11-040 | LBEOA042-11 | Nepal: Western Prov., Annapurna Himal, Banthanti vill. | |
5 | Apha kantonensis | EUPT-11-041 | LBEOA043-11 | Vietnam: Ha Nam, Ninh Binh, Quan Dist., Bong-Cuc Phuong vill. | |
6 | Apha kantonensis | EUPT-11-042 | LBEOA044-11 | Cambodia: Kampot, Bokor Nature Park, Hill Station | |
7 | Apha witti | 2011.65 | LBEOW1015-11 | China: Sichuan, Siguliang Shan, Wolong Reserve | Paratype |
8 | Apha witti | 2011.66 | LBEOW1016-11 | China: Sichuan, Quonglai Shan | Holotype |
9 | Apha witti | 2011.67 | LBEOW1017-11 | China: Sichuan, Siguliang Shan, Wolong Reserve | Paratype |
10 | Apha witti | 2011.69 | LBEOW1019-11 | China: Sichuan, Quonglai Shan | Paratype |
11 | Apha zephyrus | VZ-EUPTER-6 | LBEOW096-10 | Vietnam: Lai Chau, Fan Si Pan Mts, 5 km N Sa Pa | Paratype |
12 | Apha zephyrus | 2011.68 | LBEOW1018-11 | Myanmar: Kachin, Chudu Raji | Paratype |
13 | Apha zephyrus | EUPT-11-044 | LBEOA046-11 | China: Yunnan, Diancang Shan | Holotype |
14 | Apha zephyrus | EUPT-11-045 | LBEOA047-11 | China: Yunnan, 18 km SW of Baoshan | Paratype |
15 | Apha zephyrus | SNB 2525 | SASNC441-11 | Myanmar: Kachin, Chudu Razi Hills, 30 miles E of Kawnglanghpu |
Moderate sized moths (fore wing length 24–29 mm in males, 30–35 mm in females), characterized by bright, contrasting pattern. The basal fascia of the fore wing is dark brown, dentate and sometimes indistinct. The discal spot is round, dark brown and is always present. The antemedial fascia is angled or curved, towards the costa approximately at a right angle. The apical patch of the fore wing is elongated, semicircular, and extends along the costa. It is pale in colour, usually with a dark centre. The postmedial fascia is yellow, usually of the same colour as the apical forewing patch, straight, and running towards the apex of the wing; it is fused with the apical patch in A. floralis, A. zephyrus, A. witti, A. kantonensis, and A. chloralis. The postmedial fascia is bordered basally by a well expressed, thin, dark shadow giving the impression that it is a double line; in some species the postmedial fascia is not clearly visible and weakly separable in colour from the distal part of the wing, but its basal shadow is always well defined. The submarginal fascia is zigzag-shaped, brown, sometimes indistinct. The hind wings are darker basally while the external field is bright yellow, usually without prominent pattern. The postmedial fasciae of the hind wings are of the same colour as those of the fore wings.
Uncus well developed, bilobed; the lobes usually strongly widened at the apex; gnathos very weakly sclerotized, distinct and consisting of two separate branches, not fused, but held together by a membrane; valvae are without saccular lobes, however these lobes are present in A. witti, but they are weakly developed with only a moderate sized prominence; the saccus is weakly developed, the aedeagus width is equal to the coecum length. The vesica is without an appendix and bears cornuti which are not extended longitudinally as in other species groups but form a compact basal circle.
The papillae anales are ovoid with numerous setae. The posterior and anterior apophyses are almost equal in size, with similar length and width. The postvaginal plate is well developed, with a medial incision; the plate is often transverse (i.e. extended longitudinally across the segment). The antevaginal plate is also well developed, it is high and has a medial incision of different depth. The antrum is short, wide and weakly sclerotized. The ductus bursae is not long; it is membranous or very weakly sclerotized. The corpus bursae is globe-shaped with a single, claw-shaped signum at its equator.
Valvae are without saccular lobes, except for A. witti. The aedeagus is truncated. The antemedial fascia with angle of curvature about 90°.
The group occupies the southern area of the generic range and is known from northern India, Nepal, southern China, north-eastern Myanmar, Cambodia, and Vietnam (Figs
Species of this group develop at least two generations per year which can be seasonally dimorphic with regard to size and pigment saturation. They are confined basically to mountainous relief, known from altitudes of from 500 to 3000 m, but were also found at the lower altitudes of coastal biotopes. The caterpillars are covered with long hairs which are situated in tufts on verrucae but do not form specific brushes. They resemble large Arctiini caterpillars and feed on various deciduous trees and shrubs including members of Anacardiaceae, Annonaceae, Caprifoliaceae, Fabaceae, Malvaceae, Rosaceae, Rubiaceae, Salicaceae and Sapindaceae. The caterpillar stage lasts for 1 to 4-5 months. The pupa is enclosed in a thin, weak cocoon in the shelter of leaves, bark or on soil.
The group includes 5 species.
Apha floralis Butler, 1881, Illustrations of typical specimens of LepidopteraHeterocera in the collection of the British Museum 5: 64, pl. XCIV, figs 5, 6. TL: “India, Darjeeling”. Lectotype: male (BMNH), here designated [examined].
= Preptothauma oxydiata Draudt, 1931, Entomologische Rundschau 48(11): 121. TL: erroneously given as “West-Columbia, Altaquer, 500 m”. Holotype: female (ZMHU) [examined].
Lectotype, ♂, Darjiling, 79. 57 (BMNH). Paralectotype, ♀, Darjiling, 79. 57 (BMNH); f#, Darjeeling (
Male (Figs
Male genitalia (Figs
Female (Figs
Female genitalia (Figs
The males of the species differ from some of the other members of the group by having a well expressed “double” postmedial fascia on the hind wing. This “double” postmedial fascia can be also found in A. zephyrus and A. witti, but in both the latter species this fascia (not its basal shadow) is much more weakly expressed. The species A. floralis can be distinguished from A. zephyrus sp. nov. and A. witti sp. nov. by the smaller distance between the postmedial fascia and outer wing margin. Lobes of uncus are more slender and longer than in other species. Valvae with small apical spurs visible only under magnification. Aedeagus is straight in contrast to faintly curved in the other three species, cornuti on vesica are sparse, not dense as in other species. Saccus is small and not substantially distinguished.
Northern India and Nepal (Fig.
Mountain species producing at least two generations per year with flight periods in June-July and October; it can be found at altitudes of 1.700 – 2.500 m. Preimaginal stages and host plants are unknown.
Holotype, ♂, “China/Yunnan, | Dianceng Shan, | 25°41'N, 100°05'E, | 2.200 m, 15-30.VII.2004, | leg. Sinaev & his team” (MWM). Allotype, ♀, China, Prov. Nord-Yuennan, Li-kiang, 2.000 m, 3.VII.1935, H. Höne (
Male (Fig.
Male genitalia (Figs
Female (Fig.
Female genitalia (Fig.
This taxon is a sister species of A. floralis and is its vicariant, being found allopatrically in southern China, northern Vietnam and north-eastern Myanmar. The costal margin of the fore wing is straight as in A. floralis, it distinguishes them from A. kantonensis and A. chloralis. The yellow postmedial fascia of the hind wing is not distinctly visible (but its basal shadow is well expressed) and does not transect the middle of the wing as in A. floralis but is closer to a wing base. The apical fore wing patch is yellow, and encloses a distinct black spot. The submarginal fascia terminates apically as two dots on the vein. It also differs from related species in genitalic characters, especially by the wide valvae base, the narrowed apex which terminates with one, two or three small spurs, and the small, medial depression on a straight costa; the lobes of the uncus are narrowly separated from each other in contrast to A. floralis and A. kantonensis. From A. chloralis it also differs in the shape of the uncus lobes. Vesica with cornuti different from A. floralis but similar to A. kantonensis and A. chloralis; the field with cornuti is divided into two parts by a ring lacking any cornuti. Position of the postmedial fascia in the hind wing distinguishes the females of the species from those of A. floralis: in A. floralis it divides the wing into two, rather equal halves and therefore its end point on the hind wing costa does not correspond to the fore wing’s postmedial fascia at the tornus. In contrast, in A. zephyrus, both fore and hind wing fasciae come together at almost the same point. The signum is a bit shorter than in A. floralis. The species A. zephyrus differs from A. witti by the presence of a dark small spot near the forewing apex situated close to the costa and by the blurred dark submarginal marking near the costa of the hind wing (A. witti has no such markings); the length of the rami in the A. zephyrus male antennae are diagnostically half the length of those of A. witti. Transversal hind wing fascia diagnostically angled in M1 zone in both sexes in A. zephyrus.
China: Yunnan, Jiangxi, Sichuan, Xizang; northern Vietnam; north-eastern Myanmar (Fig.
This high altitude, montane species develops one generation per year; the moths are on the wing from May to August depending on the elevation at which they occur. They are known from 1200–3200 m. Preimaginal stages and host plants are unknown.
In Greek mythology, Zephyrus or Zephyr (Greek: Ζέφυρος), is the god of the West Wind. He was reported to have had several wives in different stories, one of which was the nymph Chloris, upon whom he bestowed the domain of the flowers. The Romans later raised her status to goddess and gave her the name of Flora – the Latin name for flowers.
Holotype, ♂, China/ Sichuan, | Qionglai Shan, | 31°13'N, 102°23'E, | May 2006, 1.400 m, | leg. Siniaev & his team (MWM). Allotype, ♀, China/Sichuan, Qionglai Shan, 31°13'N, 102°23'E, May 2006, 1400 m, leg. Siniaev & his team (MWM). Paratypes, 6 ♂♂, China, Sichuan, Qionglai Shan, 31°13'N, 102°23'E, May 2006, 1.400 m, leg. Siniaev & his team (MWM); 4 ♂♂, China, Sichuan, Wolong Reserve, Siguliang Shan, 31°09'N, 103°06'E, May 2006, 1.500-1.800 m, leg. Sinaev & his team (MWM); ♂, China, Sichuan, Dayu, Xiling, 1.900 m, 22.IV.2001, Nativ. coll. (NSMT).
Male (Fig.
Male genitalia (Figs
Female (Fig.
1 – male lectotype Apha floralis Butler, 1881, Darjiling, 79. 57 (BMNH); 2 – male A. floralis Butler, 1881, East-Nepal, Surke Danda, 3 km NE Sukeyer, Lali Kharka 18.V.1997, 2.600 m, lg. Hreblay & Szecsenyi (MWM); 3 – female A. floralis Butler, 1881, Nepal Annapurna Himal, 2.450 m, 83°43'E, 28°22'N, 23-24.VI 1996, leg. Gy. M. Lszl & G. Ronkay (MWM); 4 – female holotype Preptothauma oxydiata Draudt, 1931, West-Columbia, Altaquer, 500 m., März-Juni 1927 (
Female genitalia (Fig.
A sister species to A. zephyrus and A. floralis; all of them very similar externally. The species A. witti can be separated from A. floralis by the position of the postmedial fascia of the hind wing which is located more basally in A. witti; the submarginal fascia of the hind wing is excavate in A. witti, but is dentate in A. floralis. In A. witti the rami of the male antennae are twice as long as those of A. zephyrus (Figs
China: Sichuan (Fig.
This high altitude, montane species develops one generation per year; moths are on the wing from April to May depending on the elevation at which they occur, they are known from 1.200 – 1.900 m. The preimaginal stages and host plants are still unknown.
The species is named in honor of the eminent entomologist late Dr. Thomas J. Witt (Munich, Germany) He has made many outstanding contributions to the study of Lepidoptera worldwide and is well known for his support of young scientists; he is the founder of the largest Lepidoptera collection in Europe and it is thanks to this collection that the new species was revealed.
Apha kantonensis Mell, [1930], Deutsche entomologische Zeitschrift 1929 5: 428, fig. 43, figs 54; pl. 8: 9, 10; pl. 12: 2. TL: China, “Südkwangtung [= South Guangdong prov.], Lo fao shan”. Holotype (by original designation): male (ZMHU) [examined].
Holotype, ♂, China, Kwangtung, [Lo fao shan] (
Male (Figs
Male genitalia (Figs
Female (Figs
Female genitalia (Fig.
The brown shadow of the postmedial fascia reaches the wing margin at the apex in contrast to A. floralis, A. zephyrus and A. witti which terminates approximately 5 mm from the wing apex. It is very similar to A. chloralis in colour and pattern, but in the latter species the submarginal fasciae are vague or absent in the hind wings; in the male genitalia the lobes of the uncus are spatulate.
Localized but rather common in Hong Kong (Roger Kendrick, pers. comm., see also http://www.hkwildlife.net/viewthread.php?tid=12308). Moths are on the wing as two to four generations per year: two are obligate – the first with emergence from October to January; the second from April to May. Intermediate generations are known to occasionally occur. The species is typically coastal and is not known from the interior of the continent. In contrast to A. chloralis sp. nov., it does not occur in the higher mountains and is mostly known from lowlands (100 m) up to 1.025 m. The biology of the species was studied by
Caterpillars grow fast; the spring generation develops in only 31 – 33 days. The pupa is sheltered between leaves in a dense but soft cocoon of 25-35 mm length, densely covered with larval setae. Male pupae reach 22 mm, the females 28 – 29 mm. Antennal cases of both sexes are more than twice as thick at the level of mid legs bases. Cremaster is straight. Pupal (Fig.
South-eastern China: Guangdong; Vietnam; Cambodia (Fig.
Holotype, ♂, Vietnam, | Cha Val, Nam Giang, Quang | Nam, 15°35'21.9"N, | 107°29'01.9"E, RTS, 546 m, VD, | 28.V.2009, Tran Thieu Du leg. (coll. S. Pugaev) | [RTS means "secondary forest" and VD means "come to light trap"] (MWM, GU 35.635). Paratypes, ♂, Vietnam, Cha Val vill., Nam Giang, Quang Nam, 15°35'21.9"N, 107°29'01.9"E, RTS, 546 m, VD, 28.V.2009, Tran Thieu Du leg (IEBR); ♂, Vietnam, S. Vietnam, Quang Nam Prov., Nam Giang Distr., Cha Val vill., secondary forest, 400 m [15°35'21.9"N, 107°29'01.9"E], 24.VI.2009, Tran Thieu Du leg (MWM, GU 17.744); ♂, Central Vietnam, Quang Nam Prov., Phuoc Son Distr., Phuos My Comm., Deo Lo Xo, 17.VII 2009, Du Thieu Tran leg. (coll. S. Pugaev); ♂, N. Vietnam, Cao Bang Prov., Phi Oak Mts., Nguyen Binh Distr., Thanh Cang comm., Phia Den vill., 22°34'N, 105°52'E, 1030 m, pupa 10.XI.2009, ex pupa 22.XII.2009, leg. S. Pugaev (MWM, GU 35.634).
Eggs: C. Vietnam, Prov. Da Nang, comm. Hoa Vang, Ba Na Mts, Da Nang, 15°59.5'N, 107°59'E, 1.500 m, 08.XII 2009, leg. Tran Thieu.
Male (Figs
9 – male holotype A. kantonensis [1930], China, Kwangtung, [Lo fao shan] (
Male genitalia: 15 – holotype Apha floralis Butler, 1881, GU-93, Darjiling, 79. 57 (BMNH); 16 – A. floralis Butler, 1881, MWM-11.922, Nepal, Annapurna Himal, 1.700 m, 1 km N of Tal, 84°23'E, 28°28'N, 08.VI.1996, leg. Hreblay & Szaboky (MWM); 17 – holotype A. zephyrus sp. n., MWM-16.858, China/Yunnan, Dianceng Shan, 25°41'N, 100°05'E, 2.200 m, 15-30.VII.2004, leg. Sinaev & his team (MWM); 18 – paratype A. zephyrus sp. n., SMFL-2135.10, East-Tibet Tongme-Pelung, 2.000 m, 24.06.96 (SMFL); 19 – holotype A. witti sp. n., MWM-17.770, China, Sichuan, Qionglai Shan, 31°13'N, 102°23'E, May 2006, 1.400 m, leg. Siniaev & his team (MWM); 20 – paratype A. witti sp. n., MWM-17.745, China, Sichuan, Wolong Reserv, Siguliang Shan, 31°09'N, 103°06'E, May 2006, 1.500-1.800 m, leg. Siniaev & his team (MWM).
Female genitalia: 21 – paratype A. floralis Butler, 1881,
Male genitalia (Figs
Male genitalia: 24 – holotype A. kantonensis Mell, [1930],
The paler yellow ground colour distinguishes the species clearly from all other congeners except the externally similar A. kantonensis. Both species are found flying allopatrically and can only be reliably separated genitalically. The apex of the uncus lobes is tapered whereas it is spatulate in A. kantonensis, the distance between two first spurs on the valvae apex is much greater than in A. kantonensis has. The apical lobes of the uncus of A. kantonensis are more slender with apex expanded and round while in A. chloralis they are stouter, with medial widening and pointed apex; the valve of A. kantonensis has 3 spines of the same size while in A. chloralis the ventral spine is much larger than the dorsal and medial spines.
Female genitalia: 28 – female allotype A. zephyrus sp. n.,
In Greek mythology, Chloris (or Flora in Roman mythology) was one of the wives of the god of the West Wind Zephyrus. “Chloris” also means a greenish yellow colour typical of the ground colour of the moth.
Eggs (Figs
Preimaginal stages of A. chloralis: 35 – eggs; 36 – head of the caterpillars L6; 37, 38– L1; 39, 40 – L2; 41, 42 – L3; 43 – L4; 44, 45 – L5; 46, 47 – L6; 48 – L7; 49 – pupa.
L1 (Figs
L2 (Figs
L3 (Figs
L4 (Fig.
L5-L7 (Figs
Pupa (Figs
Rearing from eggs was carried out in this study (Figs
Molecular data undertaken (Fig.
The tree for Apha species based on COI data showing highly supported clusters for all species recognized..
The figured dendrogram shows that the complex studied which is called here A. floralis group is perhaps not monophyletic but paraphyletic, and therefore can’t be separated in a rank of a subgenus. At the same time, it is easily recognizable morphologically because of very diagnostic coloration (mostly of a hind wing) and thus this term can be used for provisory designation of this complex.
Work in the different museums was made possible by the support of their curators: Geoff Martin and Martin Honey (BMNH, London, Great Britain), Dr. Axel Hausmann (
We are grateful to Roger Kendrick (Hong Kong) for information on the biology of Apha kantonensis in Hong Kong, and Swen Löffler (Lichtenstein, Germany) for valuable data on specimens of his collection.
The article was kindly corrected linguistically by the late John Joannou (Pretoria, South Africa).
Images of Apha kantonensis Mell from Hong Kong are taken from the Internet through the link: http://www.hkwildlife.net/viewthread.php?tid=12308.
Financial support for the senior author was granted by Thomas-Witt-Stiftung in 2010 and 2011. The expedition to Central Vietnam was supported for all authors by the Russian Fund of Basic Investigations RFFI № 08-04-90300-Viet_a and RFFI № 11-04-93001-Viet_a.
The images of the type specimens housed in BMNH are figured here courtesy of the Trustees of the Museum.
The work is part of a program of the Department of Biology & Chemistry (State Pedagogical University of Uljanovsk) on the investigation into the biodiversity of moths.