Morphological methods | PaDIL Plant Biosecurity Toolbox

The forewing pattern of Lymantria is based on or derived from a series of more or less parallel, transverse dark bands or lines, usually in zig-zag shape with the segments between the veins inwardly curved and bow-shaped with their ends meeting at the veins in outwardly directed points. Shape and exact position of these transverse bands vary between species and are strongly sexually dimorphic, and their expression may differ greatly within a species. Generally, three bands are recognisable, a subasal, antemedial and postmedial one, with the antemedial band in Lymantria arising usually at or beyond the middle of the costa. There often is a submarginal row of dark marks of spots between the postmedial band and the margin, and a row of dark spots between the veins along the margin. At the end of the discal cell, at about the middle of an imaginary line from the wing base to the wing tip, there often is a dark, crescentic to V-shaped discal spot, with a dot or ring basal to it, distal to the subbasal band.

The male genitalia of Lymantria are very diverse throughout the genus, especially the valva structure, with those of the AGM and related species strongly sclerotised and relatively simple. The pointed uncus is usually continuous with the rest of the tegumen, but in L. nephrographa it is separated by a membranous region. The gnathos is absent, but some species including L. lunata have small, lateral, paired, finger-shaped tegumen processes. The saccus is tapering, often of diagnostic shape. The aedeagus is simple, straight or lightly sinuate, with the ductus entering subapically.

The female genitalia have an extensile ovipositor which, together with the 8^th segment, is as long or longer than ductus and bursa combined. The details of the sterigma are diagnostic, but dissection and slide preparation are difficult. AGM and the Australian Lymantria species lack a signum.

For images, see the Identification Key section.

Description of AGM

Male (Figs 1-3): Abdomen ochreous to brownish, never pinkish red. Forewings very variable, with ill-defined brownish to blackish markings on ground colour ranging from ochreous to dark grey-brown, with a broad darker band along distal margin unless entire wing dark; subbasal band indicated only near costa; antemedial band diffuse, strongly angled near middle of wing but often developed near costa only, postmedial band only weakly indicated at costa; submarginal spots variably developed V-shaped marks; marginal spots a variably developed series of dark spots; discal spot a well-defined, narrowly crescentic black mark, adjacent to antemedial band. Hindwings very variable, ranging from yellowish, usually with a darker band around margin, to grey-brown, often with an ochreous centre.

Female (Fig. 5): Abdomen white. Forewings with few and partly faint brownish markings on whitish to pale ochreous ground; usually a few faint darker spots near base; subbasal band present as a curved brown mark near costa; antemedial band from middle of costa and sharply angled, usually weakly developed; postmedial band a faint, thin, zig-zag line, angled and roughly parallel to antemedial band; often faint traces of a V-shaped submarginal marks, a row of small dark marks along outer margin; brown crescentic discal mark very prominent. Hindwing whitish to pale ochreous, with a series of dark small marginal spots, a faint, darker, suffused submarginal band and an indistinct darker mark at end of discal cell

Male genitalia (Fig. 7): Uncus continuous with tegumen; without lateral tegumen processes. Valva with costa continued as a long, straight, distally tapering, finger-like process, roughly in the same plane as the valva; valva proper widest at ventrodistal angle, with lateral margin inwardly oblique towards costal process; sacculus long and very narrow, pointed; juxta a simple transverse band; aedeagus long and slender.

Female genitalia (Figs 9-10): Ovipositor lobes and additional paired ventral lobes strongly sclerotised. Sterigma a weakly sclerotised broad ring, anteriorly smooth and posteriorly split into two rounded, spinulose lobes, with centre of ring leading to ductus bursae. Posteriormost part of ductus bursa with an elongate, barrel-shaped, ventrally split sclerotisation followed by wrinkled thickened membranous portion opening into corpus bursae.

Organisms that occur in Australia with which AGM might be confused

Four species of Lymantria occur in Australia, L. antennata Walker, L. pelospila (Turner), L. lunata Stoll and L. nephrographa Turner. The first three species are closely related to AGM, included in the same subgenus, and some of their males could be mistaken for AGM. Lymantria antennata and L. pelospila, both with flightless females with minute wing stumps, are either two distinct species with overlapping distribution and possibly hybridisation in north Queensland or a single cline in both wing pattern and genitalia morphology ranging from northern New South Wales around the Queensland coast to the Northern Territory and just into Western Australia. Molecular data will be needed to resolve this question. They are keyed out together, but descriptions are given for the typical form for both taxa. Lymantria nephrographa Turner is quite distinct and much less closely related to AGM. It is classified as a species incertae sedis within Lymantria by Schintlmeister (2004) rather than included in one of his 12 subgenera.

Given the wide variability of Lymantria wing pattern and the fact that crucial material needing identification will be extracted from pheromone traps which attract males only, the protocol is focused on male genitalia information. However, adults of both sexes and male and female genitalia are figured for AGM and the four Australian Lymantria species, and a brief diagnosis is given for all these species. The key is written for both sexes, with L. pelospila and L. antennata keyed out together.

Description of Lymantria lunata

Male: Abdomen at least partly pinkish red. Forewings with ill-defined pale to dark brownish to blackish markings on pale brownish grey ground; a few prominent dark spots near base of wing; three hardly angled transverse bands, each a series of very diffuse spots; a prominent submarginal row of dark diffuse spots preceded by a row of pale V-shaped blotches; a row of small dots along margin; discal mark conspicuous, well-defined, V-shaped, next to the antemedial band, with a small dark spot basal to it. Hindwings yellowish with a darker band around margin, with pink hue next to abdomen.

Female: Abdomen white with pale pink tinge. Forewings with well-defined ochrous to grey-brown markings on white ground; a few faint darker spots near base; subbasal band strongly angled twice, widest at costa; antemedial band straight, from beyond middle of costa to middle of dorsum, broad and gradually widening towards dorsum; postmedial band straight or lightly zig-zag, roughly parallel to but narrower than antemedial one; weak traces of a few submarginal spots, a row of elongate dark marks along outer margin; V-shaped mark prominent, connected to antemedial band. Hindwing whitish with black marginal dots and pink hue next to abdomen.

Male genitalia: Uncus continuous with tegumen; with small, paired, finger-shaped processes laterally on tegumen. Valva with dorsal and ventral margins straight, roughly parallel, with a straight, finger-like process from middle of distal margin, roughly in the same plane as the valva; sacculus irregular, elongate and rather broad; juxta a simple transverse band; aedeagus long and slender.

Female genitalia: Ovipositor lobes and additional paired ventral lobes very weakly sclerotised. Sterigma a weakly sclerotised, subrectangular spinulose area with a pronounced ventral lip few a few bristles laterally and a central depression leading to ductus bursae. Posteriormost part of ductus bursae a longitudinally wrinkled funnel of thickened membrane.

Description of Lymantria pelospila

Male: Abdomen whitish or yellowish, often with a pink tinge but rarely pink. Forewings with brownish to blackish markings on white to pale ochreous ground, sometimes markings so pale as to be hardly visible; up to five dark spots in wing base; subbasal band strongest on costa, a zig-zag series of dark marks; antemedial band weakly developed or absent, sinuate and zig-zag from middle of costa; postmedial band prominent, deeply zig-zag and strongly sinuate from 3/5 costa, parallel with antemedial band in dorsal half; a few irregular submarginal spots and an often incomplete row of marginal dots. Hindwings yellowish white, rarely with pinkish hue next to abdomen.

Female: flightless, with only minute wingstumps.

Male genitalia: Uncus continuous with tegumen; without lateral tegumen processes. Valva rounded-rectangular, with a long, straight, finger-like process from close to valva edge protruding at right angle from valva, and with a sharp, toothed ridge from inner surface of valva basal to finger-like process; sacculus with truncate apex; juxta a simple transverse band; aedeagus long and slender.

Female genitalia: Ovipositor lobes and additional paired ventral lobes moderately sclerotised. Sterigma weakly sclerotised, entirely spinulose, medially split to form two low, posteriorly evenly rounded lobes with a dimple each at their base, with entrance to ductus bursae in between. Posteriormost part of ductus bursa of strongly wrinkled, thickened membrane gradually opening into corpus bursae.

Description of Lymantria antennata

Male: Abdomen nearly always pink. Forewings with brownish to blackish markings on ground colour ranging from whitish to grey; up to five dark spots in wing base; subbasal band strongest on costa, a zig-zag series of dark marks; antemedial band sinuate and zig-zag from middle of costa, often widened and ill-defined, extending towards postmedial band and sometimes fused with subbasal band on dorsum; postmedial band deeply zig-zag and strongly sinuate from 3/5 costa, parallel with antemedial band in dorsal half; very few submarginal spots and a usually complete row of marginal dots. Hindwings ranging from whitish to grey, usually with pinkish hue next to abdomen.

Female: flightless, with only minute wingstumps.

Male genitalia: Uncus continuous with tegumen; without lateral tegumen processes. Valva rounded, with a long, straight, finger-like process from inner surface of valva protruding at right angle from valva, at most with a low hump on inner surface of valva basal to finger-like process; sacculus with V-shaped apex; juxta a simple transverse band; aedeagus long and slender.

Female genitalia: Ovipositor lobes and additional paired ventral lobes moderately sclerotised. Sterigma weakly sclerotised, entirely spinulose, medially split to form two high, irregularly rounded lobes, with entrance to ductus bursae in between. Posteriormost part of ductus bursa of strongly wrinkled, thickened membrane gradually opening into corpus bursae.

Description of Lymantria nephrographa

Male: Abdomen black and white. Forewing with well-defined complex, fine, reddish grey to blackish pattern on whitish ground, with the many scattered U-shaped marks not obviously aligned into transverse bands but the discal spot prominent as an interrupted outline of a crescent with a minute ring basal to it; with a complete row of mostly U-shaped submarginal marks and regular black marginal spots followed by a white fringe. Hindwing with black scales except for a whitish band along costa and around margin with a series of black marginal spots.

Female: Similar to male, but pattern elements larger and more distinctly aligned into transverse lines, with the crescent-shaped discal spot partly fused with elements of the antemedial line. Hindwing blackish in basal half with a crescentic black mark across end of cell, rest whitish with a series of blackish marginal spots.

Male genitalia: Uncus separated from tegumen by membranous area; without lateral tegumen processes. Valva simple, pointed, elongate-rhomboid, without finger-like process; sacculus triangular, pointed; juxta an elliptic plate topped with a vertical transverse band; aedeagus short, medially much wider.

Female genitalia: Ovipositor lobes and additional paired ventral lobes moderately sclerotised. Sterigma two well-sclerotised, smooth elongate-triangular plates with a row of large bristles along posterior margins and edged darker along ventral midline, with an irregular, sclerotised domed plate in between at their base leading to corpus bursae.

Palaearctic Lymantria species

A table summarising the diagnostic information presented by Pogue and Schaefer, in their taxonomic review of the economically important Lymantria of the Palaearctic region, is shown in Table 1. below. The review has excellent illustrations of diagnostic characters, including wing maculation patterns and male and female genitalia, however these illustrations will not be available until the review is published. The review is in press and is expected to be published within months. When published, it will become the primary reference for the morphological identification of economically important Lymantria. I have prepared the following table summarising the pertinent information in that review on the morphological diagnosis of Lymantria species. In addition to training me in the diagnosis of economically important Lymantria species, Dr. Pogue expressed interest in collaborating with me to extend his recent review to produce a web-accessible illustrated online key to Lymantria species, and to include DNA barcode data for Lymantria species. Such collaborative research is needed to develop a universal identification system for Lymantria species, and we will pursue that project as soon as we can fund it. In addition I visited the Natural History Museum in London to discuss lymantriid diagnostics and systematics with Dr. Jeremy Holloway, who has done much work on the Australasian lymantriid fauna. I obtained a copy of his treatise of the lymantriids of Borneo (Holloway, 1999) and had two full days in the collection examining Bornean specimens of species that are not covered by Pogue and Schaefer's review, e.g. L. singapura which feeds on economically important Pinus spp., and L. marginalis which has been recorded from Eucalyptus. Both species are potential threats to Australia.

African Lymantria species

The insect collection at the US National Museum/USDA Systematic Entomology Laboratory has no material of African Lymantria species, thus Pogue and Schaefer (in press) did not discuss African species. Schintlmeister (2004) revised the entire genus, including African species, but did not provide much detail of the African species. Therefore it was necessary to visit African Lymantriidae specialist Dr. Ugo Dall'Asta at the Royal Museum for Central Africa in Tervuren, Belgium, for specialist advice. It turned out that the RMCA does not have much material from Madagascar, so it also proved necessary to visit the Muséum national d'Histoire naturelle in Paris to spend two days examining the Madagascaran Lymantria and related genera discussed by Griveaud (1977).

The African species treated by Schintlmeister (2004) in Lymantria subgenera Griveaudtria Schintlmeister and Pyramocera were treated in more detail by Griveaud (1977) as the genera Lymantica Collenette and Pyramocera Butler, respectively. Although noting that the type species of Lymantica quite clearly does not belong in Lymantria, Schintlmeister noted the similarity of the male and female genitalia of 22 African species and those of Lymantria and transferred them to Lymantria as a consequence. While the African species have not been recorded as pests to date, many African lymantriids feed on Acacia, and Australia, being home to the majority of Acacia species, could potentially be invaded by these species in the future. Thus it is necessary to take the African species into account when developing diagnostic methods or Lymantria.

Griveaud (1977) illustrated the male and female genitalia of most species of Lymantica and also provided black and white photographs of each species. However, Griveaud's key to Lymantica species is based almost entirely on descriptions of colour patterns, which can be difficult to interpret. Therefore it proved impossible to produce a diagnostic summary for the African species in the same manner as I did for the Palaearctic species. However I purchased the literature necessary for identification of African Lymantria and used the opportunity to familiarise myself with it with the help of Dr. Dall'Asta. In addition, I took advantage of the opportunity to make contact with potential research collaborators so that it will be possible to develop future collaborative research projects on lymantriid diagnostics.

Differentiation of Lymantria dispar egg batches from egg batches of Australian Lymantriidae and of Ochrogaster lunifer (Notodontidae)

Lymantria dispar females lay their eggs in large batches covered with scales from the abdomen. Several Australian lymantriid and one notodontid species are known to produce egg batches covered with abdominal scales, superficially potentially very similar to those of L. dispar. If an egg batch is found on a shipping container after it has arrived in Australia it could have been laid by a female of one of these Australian species. The structure of the abdominal scales viewed in a scanning electron microscope (SEM) seems to be diagnostic to some extent. Figs. 49-51 illustrate scales from egg batches of various populations of L. dispar, Fig. 52 those from L. mathura Moore and Fig. 53 from L. xylina Swihnoe, to give an indication of variation within the genus Lymantria. Fig. 58 illustrates abdominal scales of Orgyia thyellina Figs 54-59 illustrate scales removed from the abdomen of females of five Australian species with scale covered egg batches, four lymantriid species and the notodontid Ochrogaster lunifer.

Table 1. Diagnosis of the economically important Lymantria species of the Palaearctic region, summarised from Pogue & Schaefer (in press). Unambiguously diagnostic characters are shown in bold.

Common Name	*Lymantria* Species	Distribution	Diagnosis	Notes
	Subgenus: Porthetria Hübner, 1819		* Males have the forewing ground colour brown or white, as in most species, with a full complement of fascia consisting of the basal, antemedial, median, postmedial, and subterminal lines. These lines usually are a series of connected chevrons, or curved spots that can vary with intensity and with shading between them. * All species are sexually dimorphic with the females larger and with a white forewing ground colour. * Female forewing shape is more of an elongate triangle than trianglulate in the male. The forewing fascia are solid lines and can be reduced in number or nearly absent. * Valve in male genitalia is characteristic having an elongate arm that is either an extension of the costa or can extend from the middle of the valve. * Tymbals on the third abdominal sternite are absent. * Female ovipositor is not telescopic.	Type species: Phalaena dispar Linnaeus, 1758:501; designated by Kirby, 1892:475
North American Gypsy Moth	dispar dispar	Eastern North America	· Males generally smaller in L .d. dispar than in L. d. asiatica and L. d. japonica. · Females also smaller, slightly reduced wing size, but not able to fly as in L. d. asiatica. · Larvae with area between D verrucae on abdomen a more solid colour with less of a white pattern on L. d. asiatica, whereas in L. d. dispar the white pattern is more prominent and the solid colour is less evident. · The small D1 verrucae have a black primary seta in L. d. dispar and this seta is white in L. d. asiatica. · Along the anterior margin of abdominal segments 1-7 are a pair of irregular shaped white spots with a gray center in L.dispar asiatica and in L. dispar dispar these spots are absent. · In Ukraine a 1.8% frequency of black-backed mutants was detected where the normal integument background colour is replaced with a solid black stripe down the entire length of the caterpillar.	* Females winged but flightless * No significant difference between L. d. dispar and L. d. asiatica in either male or female genitalia * Male flight diurnal * Highly polyphagous
European Gypsy Moth	dispar dispar	Europe, western Asia and N. Africa	See Above.	* Females winged but flightless * Nominate L. d. dispar is this Eurasian form
Asian Gypsy Moth	dispar asiatica	Asia mostly east of Urals, S & E Siberia, Russia, Korea, north and central China, and Tibet	* Hindwing in some male specimens can be a dark brown and lack the contrasting marginal band that is usually found in L. dispar. * Late stage larvae can be found which exibit the blackbacked form but it is very limited in its percentage of the population	* Flight capable females (fly to lights) * Differs from L. d. dispar in size, distribution, and the ability of the female to fly. L. d. asiatica is found from
Japanese Gypsy Moth	dispar japonica	Main islands of Japan, limited in Hokkaido	* Males are the largest of all [sub]species covered in the review of Pogue & Shaeferr (2006) and are the darkest brown of all close relatives. * The males of L.d. japonica are very similar to L d. asiatica, but L .d. japonica has a dark brown marginal band on the hindwing and in L. d. asiatica the wing lacks a definite band. * The female wings of L.d. japonica have a distinct brown cast and are not as white as L. d. asiatica. * In the male genitalia, the saccus of L.d. japonica tends to be wider than in L. d. asiatica and the apex of the sacculus is curved in L.d. japonica and not produced as in most specimens of L. d. asiatica. * The male genital capsule of L. d. japonica is approximately 1.25-1.5x larger than that of L.d. dispar. The dorsal margin of the juxta is concave to slightly convex in L.d. japonica and convex in L.d. dispar. * Larvae show considerable variation in colour and patterning, being intermediate in colouration and pattern between L. dispar asiatica and L. umbrosa. Some specimens have the narrow dorsal stripe with the paired white spots similar to L. dispar asiatica and others have broad white patches, but they are not as well developed as those in L. umbrosa. * Among late stage larvae, the black-backed mutant form is found to a limited extent and is very similar in appearance to that illustrated for L. d. asiatica	* Flight capable females * A biometrics study that included the Honshu form concluded that L. dispar japonica should be considered a distinct species from L. dispar (Pintureau, 1980a, b). * Keena and Moore (1998) demonstrated a partial incompatibility between crosses between L. dispar and L. japonica.
Hokkaido Gypsy Moth	umbrosa	Hokkaido, esp. eastern part	* There is no significant difference (from L. dispar) in either male or female genitalia * Forewing ground colour generally cream in umbrosa as compared to brown in both dispar. Markings similar to L. d. asiatica being less distinct than in L. d. dispar. * Female has more rufous cast to the longer scales of the wings and body. Also fewer forewing markings than either L. d. dispar or L. d. asiatica. * The hindwing has more white than either L. d. dispar or L. d. asiatica, and the dark margin contrasts with the lighter ground colour and shares this distinct band with L. d. dispar. *** The larvae are quite different in L.umbrosa than in either L. d. dispar or L. d. asiatica by possessing the wide dorsal stripe. The black-backed mutant form of larvae has never been found.** * In L. umbrosa the primary setae on the D2 verruca are dark brown and white, but are black in L. d. dispar and L. d. asiatica. * The underside is speckled black and white in umbrosa and in dispar and asiatica it is a solid colour with little or no pattern.	* Flight capable females * Smaller males * Crosses between L. d. dispar and L.umbrosa are male lethal. * Often feeding on Japanese larch, L. umbrosa would be limited in its chances of finding its preferred host, but its polyphagous nature would allow it to feed on a number of alternate hosts including a variety of oaks. Invasive potential is intermediate based on the ability of females to fly to lights at night. * In more severe winter weather, egg mass survival is greater below the snow line than above.
Indian Gypsy Moth	obfuscate	N. India, Pakistan & Afghanistan	* General colouration and pattern is characteristic of L. d.dispar, but smaller. * The forewing border is less distinct and resembles the overall colouration of the wing in L. obfuscata, whereas in L. d. dispar there is a definite dark brown border. * The hingwing border is more sharply defined in L. obfuscata than in L. d. dispar. *** Male genitalia with saccus a broader V-shape with a more pointed apex in L. obfuscata. In L. d. dispar the saccus is narrower and the apex tends to be broader.** * Sacculus with apex straight and not produced. In L. d. dispar sacculus generally has a produced apex. * In the female abdominal hairs that cover the normal egg masses there is a preponderance of hairs that are elbowed, or contain a rather abrupt bend, usually near the proximal end of the seta (Roonwal 1954).	* Female brachypterous, flightless. * This is the N. India component of Lymantria (Porthetria). Literature reports of L. obfuscata from S. India are based on a misidentification of L. ampla (Walker) and those findings should apply to that species (Chacko and Singh 1990). * L. obfuscata also is separated from L. dispar based on mtDNA data (Ball & Armstrong 2006) and pheromonal communication evidence (Gries, Schaefer et al. unpubl.). * Potential for invasion is minimal because of the inability of females to fly.
Okinawan Gypsy Moth	albescens	Okinawa, S. Ryukyu Isls.	* Male forewing ground colour is variably suffused with brown gray, resulting in specimens that are white to mostly light brown. The lines of the forewing vary with intensity from being quite distinct to almost absent. * Forewing outer margin colour is dark gray as compared to brown in L. apicebrunnea. * The marginal band in the hindwing is also variable, being more distinct and well developed in darker individuals and only a diffuse spot near outer apex in lighter specimens (it is less evident in L. apicebrunnea). * L. albescens and L. apicebrunnea share the broad costal margin of the hindwing, but it is dark gray in L. albescens and brown in L. apicebrunnea. * The distal process in the male genitalia is straight in L. albescens and curved dorsally in L. apicebrunnea. The distal process is longer and the apex is slightly expanded in L. albescens compared to L. dispar. * Egg masses are indistinguishable from those of L. xylina except for a slight colour tone difference.	* Flight capable females * More S. distibution * Has been treated as a subspecis of L. dispar but differences in male genitalia sufficient to establish L. albescens as a distinct species. * Invasion potential into most of North America is minimal because L. albescens inhabitats limited areas of semi-tropical island habitats, but like L. xylina, might possibly successfully invade Florida, southern California or Hawaii, as well as other more tropical countries.
Tsushima Gypsy Moth	postalba [perhaps a subspecies of L. albescens]	N. Ryukyu Isls. & S. Kyushu	* L. postalba can be separated from L. albescens by its smaller forewing length, 21-27 mm, versus > 30 mm in L.albescens.[Molecular data needed] * Forewing ground colour in male is brown in L. postalba and white or white suffused with gray in L. albescens. * Male with white hindwing	* Flight capable female * L. postalba has a more northerly distribution than L. albescens. * Invasion potential appears minimal because of the limited native range, the remote island habitat, and minimal levels of commerce. On the other hand, behaviour of females is not well known but capable of flying to lights.
Non-dispar complex species:
	apicebrunnea	China (Guangdong, Sichuan and Yunnan Provinces)	* L. apicebrunea resembles L. xylina, but L. xylina lacks the brown forewing margin, has a pink neck, pink on the legs, and a pink underside which L. apicebrunnea lacks. * The labial palpus is larger and black in L. xylina and smaller and white in L. apicebrunnea. *** The saccus in the male genitalia is wider and stouter in L. xylina than the narrow saccus in L. apicebrunnea*. In the males there is some variation in the extent of the costal margin extending down the outer margin of the forewing. Specimens can possess faint brown spots along the veins near the outer margin.	* Larva Unknown. * Potential for invasion appears minimal but little is known about the biology and behaviour of this species.
Brown bordered gypsy moth	brunneoloma [new sp. Pogue & Schaefer, 2006]	China, Yunnan	* Males very similar to L. apicebrunnea, but lacking the small tuft of pink scales just behind the head on the pronotum, found in L. apicebrunnea. * Forewing length shorter in L. brunneoloma and the outer margin has a much wider brown border than in L.apicebrunnea. * The subterminal line is not as deeply scalloped in L. brunneoloma as it is in L. apicebrunnea. * The postmedial line is discernable only as a faint spot along the posterior margin of the forewing in L. brunneoloma, but it is a well defined scalloped line in L. apicebrunnea. *** The male genitalia has a straight dorsal process in the valve of L. brunneoloma and in L. apicebrunnea it is curved toward the apex.** * The saccus is short with a truncated apex in L. brunneoloma and in L. apicebrunnea it is elongate and narrow with a rounded apex.	* Female and larva unknown.
Casuarina tussock moth	xylina	Japan, Taiwan, and China (Fujian, Guangdong)	* L. xylina resembles L. apicebrunea, but L. xylina lacks the brown forewing margin, has a pink neck, pink on the legs, and a pink underside which L. apicebrunnea lacks. * The females of L. xylina and L. apicebrunnea are more similar than the males. The angulate postmedial line in L. apicebrunnea is somewhat more crenulate than the straighter line in L. xylina. * The labial palpus is larger and black in L. xylina and smaller and white in L. apicebrunnea. * The saccus in the male genitalia is wider and stouter in L. xylina than the narrow saccus in L. apicebrunnea.	* L. xylina, native to south China and Taiwan, represents one of the most potentially invasive species for semitropical locations such as Hawaii, Southern California, or Florida, or other countries on the Pacific Rim. * Females have the same behaviour of flying to lights at night as do all the "Asian gypsy moths" (i.e. L. d. asiatica, L.d. japonica, L. umbrosa and others) and in areas around ports may readily lay eggs on shipping containers or transoceanic vessels.
	detersa	Southern India (Ahmednagar, Bombay, Belgaum, Nagrishpur, Poona, S. Coorg)	* Similar to L. obfuscata. * In L. detersa the hindwing ground colour is dirty white and in L. obfuscata the hindwing ground colour is dark reddish brown. * The male genitalia is similar to L. obfuscata, but the juxta is different. Also the distal process is shorter and more robust in L. detersa and more elongate and slender in L. obfuscata.	* Females brachypterous, flightless. * The brown forewing ground colour and zig zag postmedial line are characters that are shared with some species related to L. dispar. * L. detersa is a tropical species.
	lunata	India, SE Asia, Indonesia, New Guinea, Philippines, and NE coast of Australia	* Male has a well marked light brown forewing and a cream coloured hindwing with or without a light brown marginal band. * The abdomen is pink with brown quadrate spots dorsally. * The female has white forewings with a broad brown postmedial line and a thin subterminal line. *** Male genitalia with a lateral process on the tegumen [perhaps close to L. mathura in different subgenus?]. Also the costal process on the valve is derived from the middle of the valve rather than distally in other species.**	* L. lunata has the capacity to erupt into massive outbreak populations as recorded by Corbet (1963) and in doing so represents a potentially invasive species in many other tropical areas of the world.
	Subgenus: Lymantria Hübner, [1819]		* Most species have a white forewing with distinct black fascia and a distinct V-shaped spot at the distal end of the discal cell. Some species have a gray or grayish-brown forewing ground colour with less distinct fascia. * The hindwing fringe has a checked pattern. * The valve in the male genitalia has a produced costal process similar to that of the subgenus Porthetria, but usually shorter than the valve. * A basal process is present at the base of the valve and is variously shaped and appressed to the inner surface of the valve.	Type species: Phalaena monacha Linnaeus, 1758:501; designated by Moore, [1883]:99
Nun moth (in N. America) Black arches tussock moth (in Europe)	monacha	Eurasia, nearly continuous distribution across Asia.	* Similar to L. concolor but L. concolor with more strikingly marked forewings & more pink on abdomen. * Tegumen with lateral process present	* At the European end of its range there is a high percentage of melanistic adults while in Japanese and Korea only the non-melanistic form is found. * Studies of sex pheromone communication and periodicity of response support the designation of an "Asian form" and a "European form." It remains unclear if these two forms occupy the two discrete areas making up the apparent disjunct pattern of distribution across Eurasia. * This moth appears more of a serious pest in Europe than is L. d. dispar.
	pulverea [new sp. Pogue & Schaefer, ms]	Taiwan	*** Male genitalia with length of dorsal process of valve half as long in L. pulverea than in L. monacha*. Basal process of valve more or less fused with the valve and only represented by a small spine-like process in L. pulverea but is well developed and thumb-like in L. monacha. * Saccus a broad V-shape in L. pulverea but narrower and constricted in L. monacha.	* Most similar to L. monacha and difficult to distinguish without dissection
	concolor	Pakistan, N. India, Bhutan, Tibet, Nepal, Myanmar, Thailand, south China and Taiwan.	* L. concolor has a complete set of fascia that are rather robust. In L. monacha the fascia are not as complete and less distinct. *** Female abdomen is pale yellow in L. concolor and rosy pink in L. monacha*. Male genitalia are similar, but the dorsal process is shorter in L. concolor than in L. monacha, and the saccus is wider in L. concolor than in L. monacha.	* L. superans is actually the female of L. concolor. * Found up to 7,000 feet elevation
	minomonis	Northern India, China, Taiwan, Japan (Honshu, Shikoku, Ryokyu Islands including Okinawa)	* Most similar to L. pulverea in external characters. * Male genitalia most similar to L. monacha, but easily separated by size and shape of the dorsal process of the valve: In L. minomonis this process is shorter and thicker than in L. monacha. * The saccus is longer with more parallel sides in L. minomonis than in L. monacha.	* L. minomonis is more southerly distributed, seemingly replacing L. monacha, for which it has been confused in the past.
	similis	India (Assam, Calcutta and Sikkim), Bhutan, Myanmar, Thailand, China (Yunnan), Indonesia (Sumatra), Papua New Guinea, Philippines	* Male of L. similis shares overall forewing pattern of L. monacha, L. minomonis, and L. concolor, but is easily separated from these species. * L. similis is larger than L. monacha, the lines in the forewing are heavier in L. monacha than in L. similis, and the hindwing is white in L. similis and gray in L. monacha. * Not all of the forewing lines in L. minomonis are complete as they are in L. similis, the hindwing colour is gray in L. minomonis and white in L. similis, and the abdomen is pink in L. minomonis and white in L. similis. * The lines in the forewing are black and heavy in L. concolor, but thinner and browner in L. similis, the hindwing is gray in L. concolor and white in L. similis, and the abdomen is pink in L. concolor and white in L. similis. * Female of L. similis is most like that of L. minomonis, both having incomplete forewing lines, but the hindwing is gray in L. minomonis and white in L. similis.	* The female forewing markings are much reduced as compared to the male. The hindwing submarginal band is barely present in the female. The form niasica is a female of L. similis, but has a gray hindwing. * Subspecies L. monachoides has somewhat bolder and more defined forewing markings and a slightly more smoky gray hindwing than L. similis from India. * Subspecies L. loeffleri is identical to L. similis in forewing maculation and has a white instead of smoky gray hindwing. * The synonym, L. cara Butler, is the female of L. similis. * Potential for invasion appears quite minimal, but females are attracted to outdoor lighting.
	umbrifera	Taiwan and SW China (Jiangsu, Zhejiang, Hebei, Hunan and Jiangxi Provinces)	* Size of L. umbrifera is close to L. monacha, but easily distinguished by the gray forewing colour, diffuse dark marginal border in the hindwing, and pink abdomen. L. monacha has forewing ground colour white (as do most other species in subgenus Lymantria), hindwing gray with the border being barely visible and narrower, and a gray abdomen with only a slight hint of pink on some specimens. * Male genitalia similar but costal process of valve shorter in L. umbrifera than in L. monacha and saccus V-shaped in L. umbrifera and U-shaped in L. monacha.	* The forewing fascia are incomplete and are irrorated with dark gray scales giving the overall appearance of "fuzzy" or out of focus fascia. * Schintlmeister (2004) illustrates a female form that has all forewing fascia distinct and a bold checkered hindwing fringe.
	dissoluta	China (Anhui, Jiangsu, Jiangxi, Hubei, Hunan, Guangdong, incl. Hong Kong), Taiwan and Vietnam	* L. dissoluta is a small grayish-brown species with faint forewing markings, except for a distinct dark bar along the distal margin of the discal cell. A very slight hint of a marginal band is on the hindwing. * L. umbrifera and L. sinica have much more distinct forewing patterns.	* Males of L. dissoluta in Hong Kong are exceptionally small sized. * Since it may cause serious damage to pine species, it represents a potentially dangerous invasive to semi-tropical areas where pines are common.
	sinica	China (Shanghai, S. to Kwangtung and in Vietnam and Taiwan	* L. sinica resembles a pale gray L. monacha, but the spot at the end of the discal cell is pronounced in L. sinica. In L. monacha it is not pronounced but resembles the other forewing markings. * Male genitalia are distinct in L. sinica, with the bifurcate ventral process of the valve. No other Lymantria have this character. * Specimens from Vietnam and SW China (L.sinica albido, Schintlmeister 2004) have a white ground colour, instead of pale gray, in both the fore- and hindwings. The male genitalia are identical.	* This species has been recorded under many names, L. nebulosa, L. kosemponis, L. melanopogon, L. aprilis, L. formosana, L. roseola and perhaps others. It was mis-identified as L. umbrifera by Matsumura (1933), but should now be referred to as L. sinica Moore.
	lucescens	Japan (Hokkaido, Honshu, Kyushu) and Korea	* In Japan L. luscescens is most similar to L. umbrifera in overall forewing colouration and maculation. Differences include: forewing length is larger in L. lucescens; abdomen in L. lucescens is various shades of gray with only a hint of light pink scales laterally, whereas in L. umbrifera the abdomen is pink; the discal spot in the hindwing is more distinct in L. lucescens than in L. umbrifera. * Male genitalia are distinct between these two species placing them in different species groups. The most obvious difference is the extremely elongate and narrow saccus in L. lucescens; in L. umbrifera the saccus is a broad triangle.	* Inoue (1957) synonymized L. takamukui and L. aomoriensis with L. lucescens. The holotype of L. aomoriensis was examined and it is slightly smaller than the other specimens, but fits the description of L. lucescens. * Invasion potential is moderate because of the ability of females to fly considerable distances and their tendancy to respond to outdoor lighting.
	*Subgenus: Beatria* Schintlmeister, 2004**		* Shape of the male forewing is characteristic of this subgenus: triangulate, with straight margins and a sharp obtuse angle where the outer and posterior margins meet. * Tymbals are absent in the male abdomen. * The hindwing in the female is white with a wide, black marginal band that can be either solid or with a few white spots. * Male genitalia have a triangular valve with a curved, elongate dorsal projection.	* Type species: Phalaena Bombyx beatrix Stöll, 1790:173; original designation.
Dark mango tussock moth	marginata	Indonesia, India, Myanmar, N. to Tibet and China	* Female forewing ground colour can be white or light brown, with black or dark brown fascia respectively. * Male of L. atemeles has two forms, one with white forewing ground colour, the other with brownish-gray forewing. *** Hindwing in L. marginata is brown with small white patches along margin. In L. atemeles** the hindwing is yellow with a broad black marginal band. * Male genitalia in L. marginata have an elongate, curved dorsal process on the valve which lacks a subapical process. In L. atemeles, the dorsal process on the valve has a small subapical process.	* Considerable sexual dimorphism in L. marginata with the small dark male and the large white female. * Referred to in earlier literature as "L. beatrix" e.g. Beeson, 1941, "L. pusilla form nigra" or simply "L. nigra" in India, and "L. ganaha". * Hollaway (1999) considers L. beatrix Stoll and L. marginata Walker as similar but distinct species with the same food habits.
Orangewinged tussock moth	Atemeles	Malaysia, Thailand, Cambodia, and Vietnam	* Hindwing in L. atemeles is yellow with a broad black marginal band, in L. marginata the hindwing is brown. * Female is more heavily marked and somewhat smaller in L. atemeles than in L. marginata. *** Male genitalia - see above (L. marginata**)	* Forewing fascia in males can become browner in older specimens while the orange hindwings are unique. * This tropical species may have invasive potential among tropical regions providing they share mango cultivation (larva eats mango leaves).
	*Subgenus Nyctria* Schintlmeister, 2004**		* Male forewing ground colour is variable including white, yellow, greenish-gray, and brown, with a dark brownishgray pattern. * Female forewing has a white ground colour with a dark brown to brownish-gray pattern. * Hindwing flushed with pink and with a broken submarginal band that can vary from bold to faint. * Male genitalia have a lateral digitate process arising from the ventral margin of the tegumen and the valves are deeply divided.	* Type species: Lymantria mathura Moore, 1865:501; original designation
Pink gypsy moth, Rosy Russian moth	mathura	Widespread in E. Asia, N. to Hokkaido, Japan, Nepal, India (Ussuri and Amur), and S. to Vietnam and Sri Lanka	* L. mathura and L. flavida are very similar but have subtle differences. * Male forewing shape is more pointed in L. mathura and more rounded in L. flavida; veins are white in L. mathura and yellow in L. flavida; and the hindwing the fringe is white in L. mathura and yellow in L. flavida. * Female has narrow V-shaped reniform spot at the end of the discal cell in the forewing; this spot is much wider in L. flavida. * In L. mathura the pink on the dorsal surface of the abdomen extends approximately 2/3 the length of the abdomen, in L. flavida it extends to about half the abdominal length.	* Males of L. mathura often have melanic forms. The white of the forewing is pale gray and the wing markings are black. The hindwing is dark brown with black markings. * Synthetic lures have been successfully used in Japan, but have thus far failed to be effective in S. Korea. We had thought this was due to problems in the pheromone chemistry but we must now wonder if the two moth populations are even conspecific. * Polyphagous, hosts include mango.
	flavida [new sp. Pogue & Schaefer, ms]	Japan (Okinawa)	* There are very subtle differences in the male genitalia of L.mathura and L. flavida similar to those in Lymantria (Porthetria) species. The lateral process of the tegumen is shorter and the apex is round in L. flavida, but in L. mathura the lateral process is longer and its apex is slightly narrowed. The ventral process of the valve has parallel sides in L. flavida, but in L. mathura the sides slightly converge toward the apex resulting in the process being wider at the base than at the apex. * The female ovipositor is shorter in L. flavida than in L. mathura. The ostium bursae in the female genitalia is a large ovate shape in L. flavida and is a smaller ventrally produced flap in L. mathura. * Larvae of L. flavida have white setae on XD1 verruca and these setae are black in L. mathura. On A5 there are a pair of curved bars on L. flavida and in L. mathura there is a yellowish-brown stripe angled from the D verruca on A4 to the L verruca on A5.	* The type series were all reared. No melanic forms were reared, but it is possible that some populations may contain melanics. * The separation of the Okinawan L. flavida from the more widespread L.mathura based on mtDNA analysis by Ball & Armstrong (2006) raises questions as to the subtle differences that might exist concerning the pheromone chemistry of these moths.
	*Subgenus: Collentria* Schintlmeister, 2004**		* Schintlmeister (2004) defines the subgenus Collentria as having a violet-brown forewing ground colour, a brown pattern, and a rounded apex. There is a prominent discal spot and a series of three dashes in the tornal area of the forewing that are part of the subterminal line. These markings are also present on the female. * Male genitalia are quite variable and those of L. grisea Moore and L. fumida Butler are easily separated.	* Type species: Lymantria grisea Moore, 1879:55; original designation
	grisea	Nepal, NE India, Myanmar, N. Thailand, SW (Yunnan) and E. China, and Taiwan	* L. grisea is a much paler species overall than L. fumida. The male forewing ground colour is white in L. grisea and brown in L. fumida and the pattern is virtually identical. * The male hindwing in L. grisea is white with barely a hint of a marginal band and brown with a marginal band in L.fumida. *** The male genitalia of L. grisea is unique in Lymantria with the shape of the valve and the elongate pointed projections on the dorso-medial margin of the tegumen.**	* Schintlmeister (2004) synonymizes L. roseola on Taiwan with L. grisea kosmeponis Strand on Taiwan. He lowers the rank of L. servula Collenette from China and L. kosemponis Strand to subspecies. We are taking a conservative approach and treating all of these entities as L. grisea because there is variation in the male genitalia, and the images in Schintlmeister (2004) are of too poor a quality to make reasonable comparisons.
Red-bellied tussock moth; Yellowstriped caterpillar	fumida	Japan (Honshu, Shikoku, Kyushu), Korea, China (as far SW as Yunnan Province)	* There is some variation in the forewing ground colour in both sexes. The males are generally darker than the females, but light coloured males are similar to dark females. In males the forewing ground colour can be a rich, deep brown to brown; the females can have a similar brown ground colour or it can be white, irrorated with brown scales. * L. fumida is darker than L. grisea as discussed above. * The male genitalia are quite different between L. fumida and L. grisea. The valve has multiple projections in L.fumida and only two very small projections in L. grisea. The saccus is very narrow in L. fumida and very wide in L. grisea. The vesica in L. fumida is covered with minute cornuti and these are absent in L. grisea.	* Schintlmeister (2004) treats L. fumida caliginosa Collenette as a good species. It is hard to compare the male genitalia of L. fumida and L. caliginosa because they are not presented as comparable views. Also, the male genitalia of L. fumida is distorted due to flattening of the preparation in Schintlmeister (2004, Fig. 652). * L. fumida is rather stenophagous on Pineaceae and Cupressaceae and in forests with suitable host trees, for example on Larix trees in Honshu, Japan, L. fumida may be abundant and cause complete defoliation. In such habitats, L.fumida is in direct competition with both L. dispar japonica and L. monacha for food, but they maintain a clear pheromone channel.
	*Subgenus: Spinotria* Schintlmeister, 2004**		* This subgenus forms a compact group characterized by a ventro-medially fused valve that is divided into dorsal and ventral arms. The ventral arms are longer than the dorsal. The saccus is evenly curved ventrally.	* Type species: Bombyx serva Fabricius, 1793:474; original designation
Ficus tussock moth	serva	Taiwan, Hong Kong, south China (Yunnan Province), Nepal and India	* L. serva is similar to L. hreblayi in overall forewing colour and pattern. The reniform spot is more distinct in L. serva than in L. hreblayi. The fine brown line that separates the margin of the hindwing from the fringe in L. serva is absent in L. hreblayi and the spots on the fringe in L. hreblayi are more distinct and darker than in L. serva. * Male genitalia different from L. hreblayi (see below).	* More needs to be done to fully clarify the sex pheromone communication systems of L. iris and L. serva that coexist and compete with one another on Ficus spp. on Taiwan.
	laszloronkayi	northern areas of Vietnam, Laos, Thailand and China (Yunnan and Sichuan Provinces)	* L. laszloronkayi has the same general indistinct pattern as L. serva, but has a bold black bar subbasally below M vein, which is absent in L. serva. * Male genitalia in L. laszloronkayi differ from L. serva in the shape of both the dorsal and ventral processes. The dorsal process is short and triangular in shape in L. laszloronkayi and in L. serva it is longer, with a spine-like apical projection.	* This species was in a mixed series of L. albolunulata Moore. When dissections were made, there were major differences in the male genitalia. The ventral process of the valve in L. albolunulata is of only slightly wider at base than at apex, but in L.laszloronkayi this process is bulbous basally and is significantly narrower toward apex.
	hreblayi	northern Vietnam, Yunnan, and Sichuan Provinces in China	* L. hreblayi has the same general indistinct pattern as L. serva, but is larger. The reniform spot, which outlines the veins at the end of the discal cell is distinct in L. serva, in L. hreblayi the reniform is obscured by the suffusion of black scales that make the other forewing pattern elements indistinct. * This species resembles L. serva, but the male genitalia are quite different. The ventral process is roughly the same width from base to slightly narrower apex in L. hreblayi, this process in wider and the apex has a spine-like apical process, in L. serva.
	iris	China and Taiwan (perhaps Vietnam and NE India - see notes)	* L. iris has a more brownish forewing ground colour and is smaller than L. bantaizana, which has a grayish ground colour. The black spots on the fringe of the wings are more distinct in L. bantaizana being much less obvious in L. iris. * L. obsoleta, L. iris, and L. serva have distinct male genitalia. The laterally angled ventral process is unique for L. iris.	* Originally described as a subspecies of L. obsoleta, both L. obsoleta and L. iris were then synonymized under L. serva (Swinhoe 1923). Schintlmeister (2004) was the first to examine the genitalia of Lymantria (Spinotria) and determined that L. obsoleta, L. iris, and L. serva were all good species. * It appears that L. iris is distributed in Taiwan, southern China, Vietnam, and northeastern India (Schintlmeister 2004), but confusing the matter is evidence that a real L. serva exists from India eastward to Myanmar, and perhaps further.
	bantaizana	Japan (restricted to Honshu but recently confirmed Hokkaido (Schaefer et al. 2006)	* L. bantaizana is the only representative of this subgenus in Japan. It is a gray species versus white or brown in other species from Japan. Both sexes are similar in L. bantaizana. * Male genitalia differ from L. iris by the ventral process of the valve being straight, with a slight medially curved apex. *** The male genitalia of L. bantaizana and L. laszloronkayi are very similar, but the cornuti are larger and more conspicuous in L. bantaizana than in L. laszloronkayi**.	* Schintlmeister (2004) treated L. bantaizana as a subspecies of L. grisescens Staudinger and described a new subspecies, L. grisescens goergneri Schintlmeister. The genitalia are all very different, much more so than other closely related species in Lymantria.
	albolunulata	SW China to the Himalayan regions of N.India, including Bangladesh and Thailand	* L. albolunulata is similar to L. serva, but the forewing pattern is darker and more contrasting in L. albolunulata than the more washed out appearance of L. serva. The female abdomens of L. serva are pink as compared to brown with some pink in L. albolunulata. * The male genitalia of L. albolunulata are most similar to L. elassa. The ventral process is perhaps more bulbous basally and the apex is less produced in L. albolunulata than in L. elassa.