Biodiversity Data Journal : Taxonomic Paper
Taxonomic Paper
Harmonia manillana (Mulsant), a new addition to Indian Coccinellidae, with changes in synonymy
expand article infoJ. Poorani, Roger G Booth§
‡ ICAR-National Research Centre for Banana, Thogamalai Road, Thayanur Post, Trichy, India
§ The Natural History Museum, Cromwell Road, SW7 5BD, London, United Kingdom
Open Access



Harmonia dunlopi (Crotch), a rare lady beetle species, was originally described from 'India' by Crotch (1874). But information on subsequent sightings of this species is absent and it has not been reported by anyone from India and its neighbouring countries ever since its original description. Because of this, Indian records of H. dunlopi were suspected to be probably misidentifications of H. dimidiata (F.), a species common in northern and northeastern India and also widely distributed in the Oriental region.

New information

A single male specimen of a species collected in recent surveys from Arunachal Pradesh, India, was suspected to be H. dunlopi. Comparison of this specimen with the collections at the Natural History Museum, London, confirmed that it belonged to Harmonia manillana (Mulsant), hitherto known from Malaysia, Indonesia and the Philippines. Harmonia manillana is a highly polymorphic species with many synonyms and based on examination of the type material, the following nomenclatural changes are proposed. Harmonia dunlopi was found to be only a color variant of H. manillana and hence it is reduced to a junior synonym of H. manillana (syn. nov.). Harmonia decussata (Crotch 1874) is removed from synonymy with H. manillana and reinstated as a valid species (stat. rev.) and H. flavomarginata Bielawski 1968 is a new junior synonym of H. decussata (syn. nov.). This is the first record of H. manillana for India and South Asia. The male genitalia of H. manillana are illustrated and compared with those of H. dimidiata, the more common Indian species, to facilitate its recognition.


Harmonia manillana, Harmonia dunlopi, new synonymy, Coleoptera, Coccinellidae


Harmonia dunlopi (Crotch 1874) was originally described as Leis dunlopi Crotch with "India" as the type locality. It has not been seen in any major Indian collections at the Zoological Survey of India, Kolkata, and the National Pusa Collection at the Indian Agricultural Research Institute, New Delhi. Its name has not been mentioned by any workers from India ever since the original publication. Poorani (2002) included it in her checklist of the Indian fauna of Coccinellidae and indicated that Indian records of H. dunlopi could be probably misidentifications of Harmonia dimidiata (Fabricius 1781) as suggested by the second author.

During recent surveys in Arunachal Pradesh, northeastern India, a single male specimen of a species thought to be H. dunlopi was collected and examined along with several specimens of H. dimidiata. This specimen was found to match Malaysian specimens of H. manillana (Mulsant) in the collections of the Natural History Museum, London. The male genitalia of the Indian specimen were also found to be identical to those of H. manillana illustrated by Bielawski (1962). This is the first record of H. manillana for South Asia.

Iablokoff-Khnzorian (1982) listed many synonyms of H. manillana, which were also followed by Gordon (1987) and Coutanceau (2008). Harmonia manillana is externally quite variable and examination of the type material of these species in the Crotch Collections at Cambridge and the Natural History Museum, London, by RGB indicates there is a need to revisit and change some of these synonymies. The following changes and new synonymies are proposed here: Harmonia decussata Crotch (1874) is a valid species and is removed from synonymy with H. manillana (stat. rev.) and Harmonia flavomarginata Bielawski (1968) is a new junior synonym of H. decussata (syn. nov.). Harmonia dunlopi is considered by us as a mere color variant of H. manillana and synonymized with H. manillana here (syn. nov.). A brief diagnostic description is given for H. manillana based on the sole Indian specimen examined and the male genitalia of both H. manillana and H. dimidiata are illustrated to facilitate their identification in the event of any subsequent collection.

Materials and methods

The specimen examined was part of the collections made in Pasighat and nearby places in the north eastern Indian state of Arunachal Pradesh, in November 2014 and it is deposited in the reference collections of the National Bureau of Agricultural Insect Resources, Bangalore. Photographs of the habitus and the male genitalia were taken using a Leica M205A Stereo microscope. Composite images were generated from image stacks using the software Combine ZP and touched up in Photoshop Elements 11.

The following acronyms are used for the repositories mentioned in this paper:

BMNH - Natural History Museum, London:

MNHUB - Museum für Naturkunde der Humboldt Universität, Berlin

UCCC - University of Cambridge Crotch Collections, Cambridge

Taxon treatment

Harmonia manillana (Mulsant, 1866)


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.

Material   Download as CSV 
  1. scientificName:
    Harmonia manillana (Mulsant)
    ; taxonomicStatus:
    ; kingdom:
    ; phylum:
    ; class:
    ; order:
    ; family:
    ; continent:
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    Arunachal Pradesh
    ; municipality:
    ; locality:
    Pasighat College of Horticulture & Forestry
    ; verbatimLocality:
    College of Horticulture & Forestry
    ; samplingProtocol:
    Yellow pan trap
    ; eventDate:
    ; year:
    ; month:
    ; individualCount:
    ; sex:
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    ; preparations:
    Male genitalia
    ; identifiedBy:
    J Poorani
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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.)

Figure 1.

Harmonia manillana (Mulsant)

aAdult, dorsal habitus  
bAbdominal postcoxal line  
cMale genitalia: Tegmen, lateral view  
dMale genitalia: Tegmen, ventral view  
eMale genitalia: Penis  
fMale 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.

Figure 2.

Harmonia dunlopi (specimen examined at ZSI, Kolkata)

aAdult, lateral view  
bAdult, 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.

Figure 3.

Harmonia dimidiata (F.)

aAdult, dorsal view  
Figure 4.

Harmonia dimidiata: variants

aWithout pronotal spots  
bWith macula on pronotum  
cWith enlarged spots  
dWithout 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.

Taxon discussion

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.


This work was carried out under the Network Project on Insect Biosystematics funded by the Indian Council of Agricultural Research, New Delhi.

Author contributions

J. Poorani - Collection, identification, imaging, manuscript preparation

R.G. Booth - Confirmation of identity, examination of types, manuscript preparation


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