Caria manillana Mulsant 1866: 170 (Type locality: 'Manilla', Philippines; Lectotype, UCCC).-Bielawski 1962: 197.-Gordon 1987: 14 (lectotype designation).

Leis atrocincta Mulsant 1866: 175 (Type locality: 'Manilla', MNHUB).-Crotch 1874: 120 (as var. atrocincta).-Coutanceau 2008: 7.

Neda paulinae Mulsant 1866: 203 (Type material: ?MNHUB).-Crotch 1874: 120 (as Caria paulinae).-Bielawski 1962: 197.-Iablokoff-Khnzorian 1982: 486.-Gordon 1987: 14.-Coutanceau 2008: 7.

Leis dunlopi Crotch 1874: 121 (Type locality: “India”; Lectotype, UCCC).-Iablokoff-Khnzorian 1982: 486.-Gordon 1987: 14 (lectotype designation).-Coutanceau 2008: 7. New Synonym.

Leis cerasicolor Crotch 1874: 121 (Holotype, UCCC).-Iablokoff-Khnzorian 1982: 486.-Gordon 1987: 14.-Coutanceau 2008: 7.

Leis aterrima Crotch 1874: 121 (Holotype, UCCC).-Iablokoff-Khnzorian 1982: 486.-Gordon 1987: 14.-Coutanceau 2008: 7.

Leis papuensis Crotch 1874: 121 (Lectotype, UCCC).-Iablokoff-Khnzorian 1982: 486.-Gordon 1987: 14 (lectotype designation).-Coutanceau 2008: 7.

Leis papuensis var. suffusa Crotch 1874: 121 (Lectotype, UCCC).-Korschefsky 1932: 275.-Gordon 1987: 14 (lectotype designation). Syn. nov.

Length: 6.5 mm. Form (Fig. 1a) hemipherical, strongly convex, dorsum glabrous except head with silvery white hairs around clypeal margin. Dorsal side bright reddish-testaceous, pronotum with a median black macula on posterior margin above scutellum, elytra with 11 black spots, spots on each elytron arranged in a 1-2-2-1/2 pattern, one below anterior margin, two transverse spots arranged just above midline (one lateral and one discal), the second pair positioned around apical third and smaller than the first pair of spots, the last spot sutural and reaching apex; ventral side reddish testaceous except metaventrite medially blackish. Abdominal postcoxal line (Fig. 1b) incomplete with a semi-circular associate line, ventrite 5 apically shallowly emarginate, ventrite 6 slightly more deeply emarginate. Male genitalia (Fig. 1c, d, e, f) with penis guide of tegmen in ventral view (Fig. 1d) basally broadest, progressively narrowed towards a tubularly produced apex, shorter than parameres; parameres with lateral and inner margins covered with dense pubescence. Penis (Fig. 1e, f) with a prominent, stout capsule, penis apex (Fig. 1f) as illustrated. (This description is based on the single specimen examined from India.)

a

b

c

d

e

f

Figure 1.

Harmonia manillana (Mulsant)

a: Adult, dorsal habitus  
b: Abdominal postcoxal line  
c: Male genitalia: Tegmen, lateral view  
d: Male genitalia: Tegmen, ventral view  
e: Male genitalia: Penis  
f: Male genitalia: Penis apex  

Harmonia manillana is externally highly variable and Bielawski (1962) described the nominate form in detail and illustrated the male and female genitalia. In the sole Indian male examined here and the specimens of H. manillana from Malaysia at BMNH, on each elytron the spots are characteristically arranged in a 1-2-2-1/2 pattern and the pronotum has a larger median spot. The male genitalia in H. manillana are diagnostic. The male genitalia of the Indian specimen fully match the illustrations given by Bielawski (1962) and Iablokoff-Khnzorian (1982).

The nominate form of Harmonia dunlopi is very similar to H. dimidiata and likely to be confused with it as observed by Bielawski (1962) and Iablokoff-Khnzorian (1982). Crotch (1874) described H. dunlopi as having a 1-2-1-1 elytral pattern and observed that an additional sutural spot was present in some specimens. The lone Indian specimen appears to have this extra spot observed by Crotch. The illustration given by Iablokoff-Khnzorian (1982) for H. dunlopi also shows a tiny extra spot next to suture in the third row, which corresponds to the Indian specimen. The only specimen of H. dunlopi examined by JP in the collections of the Zoological Survey of India, Kolkata, was collected in "Haruhasa Mt. Sambawa" (Indonesia) and identified by A.P. Kapur as Leis dunlopi var. nov. (compared with type) (Fig. 2). This specimen was much larger with smaller elytral spots compared to the present example. Further specimens of H. manillana from “Haruhasa, Mt. Sambawa” in BMNH were also determined by A.P. Kapur as dunlopi var. nov. and are from the same series as the specimen shown in Fig. 2.

a

b

Figure 2.

Harmonia dunlopi (specimen examined at ZSI, Kolkata)

a: Adult, lateral view  
b: Adult, dorsal view  

Harmonia dimidiata (Fig. 3a) is orange-yellow to bright red with a pair of black spots on pronotum, often fused into a single marking with a median emargination and 13 black spots on elytra arranged in a 1-3-2-1/2 pattern. The pronotal spots and apical elytral spot are sometimes absent in some examples (Figs 3a, 4a, d). The elytral color pattern is variable with the spots enlarged (Fig. 4c) or posterior two-thirds of elytra black and anterior portion yellowish, with the humeral black spots present (Fig. 4d) or absent. The abdominal postcoxal lines are incomplete with an associate line as in H. manillana. The metaventrite in the Indian specimen of H. manillana is almost black, whereas in H. dimidiata, the ventral side is reddish testaceous. The male genitalia (Fig. 3b) in H. dimidiata are superficially similar to those of H. manillana, but the penis guide is distinctly more elongate and narrower than that in H. manillana with a rounded apical projection and the penis capsule is elongate with a much longer and narrower outer arm. The parameres in H. manillana are stouter and shorter and rather abruptly narrowed in the apical third whereas in H. dimidiata, parameres are more or less uniformly wide throughout and apically truncate. The spermatheca and the infundibulum in H. dimidiata are illustrated in Fig. 3b. The spermatheca of H. manillana illustrated by Bielawski (1962) and Iablokoff-Khnzorian (1982) appears to be different from that of H. dimidiata and clearly shows a basal constricted projection of the cornu though the infundibulum appears to be similar in both species. Harmonia dimidiata is widely distributed in north, northwestern and northeastern India and several other South and southeast Asian countries in the Oriental region.

a

b

Figure 3.

Harmonia dimidiata (F.)

a: Adult, dorsal view  
b: Genitalia  

a

b

c

d

Figure 4.

Harmonia dimidiata: variants

a: Without pronotal spots  
b: With macula on pronotum  
c: With enlarged spots  
d: Without pronotal and elytral spots  

India: Arunachal Pradesh (new record); Philippines; Malaysia; Indonesia (Iablokoff-Khnzorian 1982; Coutanceau 2008).

Harmonia manillana appears to be extremely rare in India. Discovery of H. manillana from India is important as it could be taken as a confirmation of the type locality of H. dunlopi (synonymized here with the former). This is the first record of H. manillana from mainland India and its absence in Indian and international collections is an indication that it is probably a very rare species and it probably needs to be listed as such in Indian faunal lists. It is worth noting that this was the only specimen collected along with several specimens of H. dimidiata from the same locality.

Two species, Harmonia decussata (Crotch 1874: 161) and H. flavomarginata Bielawski (1968), are wrongly synonymized with H. manillana by Iablokoff-Khnzorian (1982), and are valid names. Coutanceau (2008) appears to have followed Iablokoff-Khnzorian in his checklist of world species of Harmonia, where both are listed as synonyms of H. manillana. Crotch (1874) described Callineda decussata from various localities and the syntype series in UCCC and BMNH is very mixed, but none is H. manillana. The lectotype of C. decussata, designated by Gordon (1987), is here confirmed as a valid species, Harmonia decussata (Crotch, 1874) (stat. rev.). This lectotype was examined by RGB in 1989 and found to represent the same species that Bielawski (1968) described as Harmonia flavomarginata (syn. nov.). This also appears to be the same species that Chazeau (1989) described as Harmonia incognita, but a formal synonymy of the latter must await further study.

Crotch (1874) listed Leis suffusa as a variety of L. papuensis (now a synonym of H. manillana) and Korschefsky (1932) listed it as an aberration of papuensis in his catalogue. Gordon (1987) designated a lectotype for L. suffusa and mentioned that it appeared to be a synonym of H. manillana though it was not formally designated so by Iablokoff-Khnzorian (1982). We formally synonymize L. suffusa with H. manillana here (syn. nov.).

Iablokoff-Khnzorian (1982) described both H. manillana and H. dunlopi with rather inadequate / poor illustrations. He illustrated the adult and the female genitalia of H. dunlopi and indicated that it could be probably synonymous with H. dimidiata. He also mentioned that the specimen he examined from the Crotch Collection at the University of Cambridge (UCCC) was a holotype. However, Crotch (1874) clearly mentioned about additional specimens. Gordon (1987) also noted this when he designated a lectotype from Crotch's material at UCCC.

Crotch’s original description of Leis dunlopi listed material from India (Dublin, B.M.). Gordon (1987) noted the locality datum on the Lectotype as India. However, the specimen in BMNH, a paralectotype, bearing Crotch’s name label in his own handwriting “Dunlopi ns” is from the Philippines and not India, and this specimen has the color pattern as shown in Fig. 1a. Crotch’s Lectotype was examined by RGB in 1988/89 and the paralectotype was examined again during the preparation of this paper, enabling the synonymy of H. dunlopi with H. manillana to be confirmed.