Entomological Science
Volume 10, Issue 4 pp. 373-393
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Taxonomic notes on the paper wasps of the genus Ropalidia in the Indian subcontinent (Hymenoptera: Vespidae)

Jun-ichi KOJIMA, 

Jun-ichi KOJIMA

Natural History Laboratory, Faculty of Science, Ibaraki University, Mito, Japan;

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Kishore LAMBERT, 

Kishore LAMBERT

Department of Zoology, Malabar Christian College, Calicut, Kerala, India; and

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Lien T. P. NGUYEN, 

Lien T. P. NGUYEN

Institute of Ecology and Biological Resources, Vietnamese Academy of Science and Technology, Hanoi, Vietnam

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Fuki SAITO, 

Fuki SAITO

Natural History Laboratory, Faculty of Science, Ibaraki University, Mito, Japan;

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First published: 21 December 2007
https://doi.org/10.1111/j.1479-8298.2007.00237.x
Citations: 7
Jun-ichi Kojima, Natural History Laboratory, Faculty of Science, Ibaraki University, Bunkyo 2-1-1, Mito 310-8512, Japan. Email: [email protected]

Abstract

The taxonomy of Ropalidia wasps in the Indian subcontinent is revised, recognizing 26 species in the subcontinent. Their diagnostic characteristics are summarized in a key to species. New synonymies proposed in the present study are: R. bicolorata shiva Das and Gupta, 1989 under R. bicolorata van der Vecht, 1962; R. colorata sordida van der Vecht, 1941 under R. colorata van der Vecht, 1941; R. rodialipa Lambert and Narendran, 2005 and R. anupama Lambert and Narendran, 2005, both under R. cyathiformis Fabricius, 1804; R. jacobsoni flavoscutellata Das and Gupta, 1989, and R. bangalorica Lambert and Narendran, 2005, both under R. jacobsoni du Buysson, 1908; R. travancorica Lambert and Narendran, 2005, under R. marginata Lepeletier, 1836; R. sridharani Lambert and Narendran, 2005, under R. rufocollaris Cameron, 1900; Ropalidia rufoplagiata nursei van der Vecht, 1941 under R. rufoplagiata Cameron, 1905; Icaria lugubris Smith, 1858, under R. sumatrae Weber, 1801; and a revised synonymy is Icaria pendula Smith, 1857, under R. variegata Smith, 1852. The new replacement name Ropalidia kasaragodensis Lambert and Narendran is proposed for R. indica Lambert and Narendran, 2005, non van der Vecht, 1941.

INTRODUCTION

Of somewhat more than 180 Ropalidia species currently recognized, 27 species have been recorded from the Indian subcontinent (Pakistan, India, Bangladesh, Nepal, Bhutan, Sri Lanka, and the eastern part of Myanmar), including 16 species so far known to be endemic to the subcontinent (Kojima & Carpenter 1997; Kojima 2006). Because of its well-known historical geology in terms of plate tectonics, the social wasp fauna in the Indian subcontinent is of special interest from the view point of the biogeography of social wasps, especially for the groups occurring in Madagascar, the Indian subcontinent and Australia – that is, eastern elements of the Gondwana. The paper wasp genus Ropalidia Guérin-Méneville, 1831, is one of the two such vespid genera; the other is the cosmopolitan genus Polistes Latreille, 1802.

Das and Gupta (1989) made an extensive taxonomic study on the social wasps in the Indian subcontinent and recognized 22 species in Ropalidia. Gusenleitner (2001) added R. sculpturata from Nepal and R. tamila from Tamil Nadu in India (Gusenleitner 2004) to the Indian subcontinent fauna of Ropalidia. Recently, Lambert et al. (2005a,b) described six species from India, mainly referring to keys in Das and Gupta (1989). Unfortunately, the key to Ropalidia species in Das and Gupta (1989) is at least in part barely workable, because those authors adopted the subgeneric concepts applied by van der Vecht (1941, 1962) and Richards (1978), which often prevents correct identification at the species level (Kojima 1997). In the present paper, we give taxonomic notes on the Ropalidia species occurring in the Indian subcontinent, together with a key to species.

MATERIALS AND METHODS

Specimens examined are deposited in the Biologiezentrum des Oberösterreichischen Landesmuseums, Linz, Austria (BOLM), the Department of Zoology, University of Calicut, Kerala, India (DZUC), Malabar Christian College (MCC), Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (MCZ), Nationaal Naturhistorisch Museum, Leiden, the Netherlads (RMNH), Natural History Collection, Ibaraki University, Mito, Japan (IUNH), US National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA (USNM), and Western Ghats Regional Station, Zoological Survey of India, Kozhikode, India (ZSIK).

Abbreviations of collector names are as follows: L. K., Lambert Kishore; P. S. N., P. S. Nathan; K. V. K., Karl Von Krombein; J. C. B., J. C. Bridwell.

Morphological and color characters were observed on pinned specimens under a stereoscopic dissecting microscope. The following morphometric characters and views of observation are often ill-defined in the literature. In the present study they are defined as follows: widths of eye and gena are measured as the maximum for each in strictly lateral view of the head; length of first metasomal tergum is measured in lateral view as distance from the posterior end of the basal slit for the reception of the metasomal suspensory ligament to the posterodorsal end of the tergum; “dorsal view of first metasomal tergum” refers to the view perpendicular to the dorsal face of the posterior widened part of the tergum.

KEY TO ROPALIDIA SPECIES OF THE INDIAN SUBCONTINENT

The present key is applicable to both females and males unless the sex is specified. Color characteristics are also used for some species, but they may not be applicable to some local forms occurring outside the Indian subcontinent.

  • 1

    Propodeum with paired, longitudinal, basal carinae (Figs 23–25); propodeal orifice very narrow (Fig. 28)    2. R. marginata species group

  • –

    Propodeum without such carinae (Figs 26, 27); propodeal orifice wider, more or less rounded dorsally (Figs 26,27)    6

  • 2

    Propodeum with strong punctures between basal carinae; striations lateral to carinae strong (Fig. 25). Second metasomal segment entirely black or dark reddish-brown, but often with yellow or brownish-yellow apical band; the yellow band, if present, narrow and not swollen anteriorly in lateral sides of the tergum, or faint and interrupted     3

  • –

    Propodeum with week transverse striation and scattered shallow punctures between basal carinae; striation lateral to carinae weak (Figs 23,24). Second metasomal segment reddish-brown to brown, with distinct yellow apical band usually swollen anteriorly on lateral sides of the tergum    4

  • 3

    Body larger; body length (head + mesosoma + first two metasomal segments) more than 15 mm; forewing length more than 12 mm. Posterolateral corner or propodeum with thin lamella absent (Fig. 29). First metasomal tergum in lateral view strongly swollen dorsally behind level of spiracle and highest before level of posterior margin of the sternum (Fig. 29). Male antennal flagellum with distinct tyloids, serrated beneath; terminal flagellomere curved, with apex not sharply pointed (see Gusenleitner 2004: Fig. 1)        R. magnaminavan der Vecht, 1941

  • –

    Body smaller; body length less than 14 mm; forewing length less than 11 mm. Popodeum with each posterolateral corner raised into thin lamella (23–28, 29–36). First tergum in lateral view more or less gradually swollen dorsally toward level of posterior margin of the sternum (Fig. 30). Male antennal flagellum with tyloids modified into flat plates, not serrated beneath; terminal flagellomere curved, sharply pointed apically (Figs 1,2; also see Gusenleitner 2004: fig. 2)        R. tamilaGusenleitner, 2004

  • 4

    Yellow apical band of second tergum narrow, with its width approximately or less than one-fifth length of the tergum (Figs 31,32; possibly applicable only to Indian subcontinent populations)        R. marginata (Lepeletier, 1836)

  • –

    Yellow apical band of second tergum wide, with its width near the middle more than one-fourth length of the tergum (Figs 33,34)    5

  • 5

    First metasomal tergum proportionally slender (Figs 33,34), with its maximum width in dorsal view approximately 0.4-fold as wide as that of second tergum (Fig. 34). Male terminal antennal flagellomere strongly curved, blunt apically, approximately threefold as long as its basal width (Fig. 5). Aedeagus of male genitalia broadly spatulate apically (see van der Vecht 1962: fig. I.a)     R. spatulatavan der Vecht, 1962

  • –

    First metasomal tergum proportionally wider (Figs 35,36), with its maximum width in dorsal view nearly half as wide as that of second tergum. Male terminal antennal flagellomere less strongly curved and bluntly pointed at apex, approximately twofold as long as its basal width (Fig. 4). Aedeagus not spatulate apically        R. brevitaDas and Gupta, 1989

  • 6

    Basal angle of second submarginal cell distinctly less than 90° (Figs 18,19)    7 R. stigma species group

  • –

    Basal angle of second submarginal cell larger than 90° (Figs 20–22)    13

  • 7

    Propodeum with median depression distinct and reticulately punctured; lateral sides of propodeum also reticulately punctured     8

  • –

    Propodeum barely punctured, only finely, transversely striate; posterior face barely depressed medially or if depressed, only as fine median furrow    9

  • 8

    Female gena narrow, in lateral view about half as wide as eye (Fig. 11); female antenna with first flagellomere more than threefold as long as its apical width (Fig. 6)    R. kasaragodensis Lambert and Narendran, nom. nov.

  • –

    Female gena wider, in lateral view distinctly more than half as wide as eye; female antenna with first flagellomere distinctly less than threefold as long as its apical width (Fig. 7)        R. birmanicavan der Vecht, 1941, R. santoshaeDas and Gupta, 1989, R. sculpturata Gusenleitner 2001. (For taxonomic status of these three taxa, see the comments for each species.)

  • 9

    Suture between second tergum and sternum indistinct except in posterior margin (Fig. 41)    R. hongkongensis de Saussure, 1854

  • –

    Suture between second tergum and sternum distinct throughout the segment (Figs 42,43)     10

  • 10

    Female gena as wide as eye. Second metasomal tergum in dorsal view gradually swollen posteriorly in anterior one-third (Fig. 45). First tergum brown, except for black basal half; second tergum with paired, anterolateral, yellow spots         R. nigritaDas and Gupta, 1989

  • –

    Female gena narrower than eye. Second tergum in dorsal view gradually swollen posteriorly in anterior two-thirds (Fig. 44)     11

  • 11

    Female antenna slender; first flagellomere more than threefold as long as its apical width (Fig. 8); second flagellomere longer than wide; terminal flagellomere distinctly longer than its basal width. First tergum more than 2.6-fold as long as wide        R. rufocollaris (Cameron, 1900)

  • –

    Female antenna thicker; first flagellomere nearly threefold as long as its apical width; second as long as wide; terminal slightly longer than its basal width. First metasomal segment less than 2.4-fold as long as wide     12

  • 12

    Ventral metapleuron smooth, without punctures.    R. stigma (Smith, 1858)

  • –

    Ventral metapleuron sparsely, shallowly punctured    R. mathematica (Smith, 1860)

  • 13

    Ventral corner of pronotum obliquely cut off (Fig. 14). First tergum abruptly swollen dorsally near posterior margin of basal slit to receive metasomal suspensory ligament (Fig. 46).    R. rufoplagiata (Cameron, 1905)

  • –

    Ventral corner of pronotum gradually narrowed ventrally (Figs 15–17). First tergum in lateral view gradually swollen posteriorly (Figs 47–50,52,54–56,58,60)    14

  • 14

    Forewing with two submarginal cells (Fig. 22). Mesosoma black, with pronotum, scutellum and metanotum mostly red        R. bicoloratavan der Vecht, 1962

  • –

    Forewing with three submarginal cells (Figs 20,21)    15

  • 15

    Propodeal valvula large, semicircular or broadly rounded-triangle in outline, in lateral view covering most part of propodeal teeth forming propodeal orifice (Figs 47,48,50)    16

  • –

    Propodeal valvula small, in lateral view barely covering propodeal teeth forming propodeal orifice (Figs 55,56).    19

  • 16

    Posterior margin of head in dorsal view broadly and shallowly emarginate medially (Fig. 12). Pronotal carina barely sinuate backward at humeral angle (Fig. 15). Propodeal valvula large, nearly circular (Fig. 50). First metasomal segment long, with posterior widened part of the tergum in dorsal view more or less gradually swollen posteriorly, then narrowed again near apical margin (Figs 51,53)        17, R. fasciata species group

  • –

    Posterior margin of head in dorsal view barely emarginate medially (Fig. 13). Pronotal carina strongly sinuate backward at humeral angle (Fig. 16). Propodeal valvula rounded-triangle (Figs 47,48). First metasomal segment short, with posterior widened part of the tergum in dorsal view gradually swollen toward apical margin, or parallel-sided or only slightly narrowed near posterior margin        18, R. variegata species group

  • 17

    Second metasomal segment obliquely cut off at posterior end, with tergum longer than sternum (Fig. 50)    R. fasciata (Fabricius, 1804)

  • –

    Second metasomal segment vertically cut off, with tergum and sternum nearly equal in length (Fig. 52)    R. coloratavan der Vecht, 1941

  • 18

    Median furrow of propodeum usually indistinct (Fig. 26). First metasomal tergum in lateral view with dorsal margin more or less evenly convex, highest at level of posterior margin of the sternum (Figs 47,48)        R. variegata (Smith, 1852)

  • –

    Median furrow of propodeum usually distinct (Fig. 27). First metasomal tergum in lateral view with dorsal margin more strongly convex in posterior half, highest slightly posterior to level of posterior margin of the sternum (Fig. 49)         R. jacobsoni (du Buysson, 1908)

  • 19

    Border between punctured posterodorsal area and unpunctured anteroventral area of mesepisternum distinct, often marked by carina. Posterior margin of first metasomal sternum not deeply emarginate    20

  • –

    Border between punctured posterodorsal area and unpunctured anteroventral area of mesepisternum indistinct; punctures often extending into anteroventral part, though very sparsely. Posterior margin of first sternum deeply emarginate medially (Figs 57,59)    21

  • 20

    Small species, body length (head + mesosoma + first two metasomal segments) less than 7.5 mm. First metasomal segment short, in lateral view with dorsal margin arising from posterior margin of basal slit for reception of metasomal suspensory ligament (Fig. 54); second segment in lateral view vertically cut off posteriorly (Fig. 54). Male flagellum with developed tyloids, more or less serrated ventrally; terminal flagellomere weakly curved (Fig. 9). Body largely yellow    R. cyathiformis (Fabricius, 1804)

  • –

    Larger species, body length more than 9 mm. First metasomal tergum in lateral view with dorsal margin arising after some distance from posterior end of basal slit for reception of metasomal suspensory ligament (Fig. 55); second segment in lateral view obliquely cut off posteriorly, with tergum shorter than sternum (Fig. 55). Male flagellum with tyloids of slightly raised longitudinal ridge; terminal flagellomere bullet-shaped (Fig. 55). Body black; first tergum red or orange or black        R. sumatrae (Weber, 1801)

  • 21

    Head and mesosoma shiny, with very scattered, indistinct punctures. First metasomal segment relatively short (Fig. 57); dorsal margin of tergum in lateral view arising from posterior end of basal slit for reception of metasomal suspensory ligament (Fig. 56)    R. montanaCarl, 1934

  • –

    Head and mesosoma dull, with microscopic fine punctures in addition to rather large, shallow, flat-bottomed punctures. First metasomal segment longer (Fig. 59); dorsal margin of tergum in lateral view arising after short distance from posterior end of basal slit for reception of metasomal suspensory ligament (Fig. 58)     22

  • 22

    Occipital carina strongly widened ventrally, occupying about one-sixth width of gena at widest part (see Kojima 1996)         R. sericea (Cameron, 1905)

  • –

    Occipital carina not widened, more or less fine throughout its length     23

  • 23

    Punctures on metasoma very shallow, minute and sparse. Clypeus entirely yellow (only sometimes with median black spot); scutum with paired longitudinal yellow bands; second metasomal tergum with large yellow spot on each lateral side        R. ornaticeps (Cameron, 1900)

  • –

    Punctures on metasoma distinct. Clypeus mostly black; scutum entirely black, without yellow spots; second tergum black    R. scitula (Bingham, 1897)

Details are in the caption following the image
Figure 23–28
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Propodeum of Ropalidia of the Indian subcontinent. (23) R. marginata; (24, 28) R. brevita; (25) R. tamila; (26) R. variegata; (27) R. jacobsoni. (23, 25) Posterolateral view; (24, 26, 27) posterodorsal view; (28) propodeal orifice. Scale lines: 0.5 mm.

Details are in the caption following the image
Figure 29–36
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Characters of the Ropalidia marginata group. (29) R. magnanima; (30) R. tamila; (31, 32) R. marginata; (33, 34) R. spatulata; (35, 36) R. brevita. (29, 30, 35) Posterior part of propodeum and first metasomal segment, lateral view; (31, 33) posterior end of propodeum and first two metasomal segments, lateral view; (32, 34) first two terga, dorsal view; (36) first tergum, dorsal view. Scale lines: 1 mm.

Details are in the caption following the image
Figure 1–13
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Characters of Ropalidia of the Indian subcontinent. (1, 2) R. tamila; (3) R. marginata; (4) R. brevita; (5) R. spatulata; (6, 11) R. kasaragodensis Lambert and Narendran, nom. nov. (7) R. birmanica; (8) R. rufocollaris; (9) R. cyathiformis; (10) R. sumatrae; (12) R. fasciata; (13) R. variegata. (1–10) Antenna (1, 9, 10) flagellum, male; (2–5) apical part of flagellum, male; (6, 8) basal part of flagellum, female; (7) flagellum, female); (11) female head in lateral view; (12, 13) female head in dorsal view. Scale lines: 0.5 mm.

Details are in the caption following the image
Figure 18–22
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Forewing characters of Ropalidia. (18) R. rufocollaris; (19) R. sculpturata; (20) R. colorata; (21) R. scitula; (22) R. bicolorata. (18–21) Submarginal cells, with an arrow indicating basal angle of the second submarginal cell; (22) apical part of forewing. Scale lines: 0.5 mm.

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Figure 37–45
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Metasomata of the Ropalidia stigma group. (37, 38) R. birmanica; (39, 40) R. sculpturata; (41) R. hongkongensis; (42) R. rufocollaris; (43, 44) R. stigma; (45) R. nigrita. (37–43) First two segments; (37, 39, 41–43) lateral view; (38, 40) dorsal view; (44, 45) second tergum, dorsal view. Scale lines: 1 mm.

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Figure 14–17
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Pronotum in lateral view. (14) Ropalidia rufoplagiata; (15) Ropalidia fasciata; (16) Ropalidia variegata; (17) Ropalidia montana. Scale lines: 0.5 mm.

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Figure 46–59
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Metasomata of the Ropalidia of the Indian subcontinent. (46) R. rufoplagiata; (47, 48) R. variegata; (49) R. jacobsoni; (50, 51) R. fasciata; (52, 53) R. colorata; (54) R. cyathiformis; (55) R. sumatrae; (56, 57) R. montana; (58, 59) R. scitula. (46, 48, 49, 51, 53, 57, 59) First segment (46, 48, 49) lateral view; (51, 53) dorsal view; (57, 59) ventral view; (47, 50, 52, 54–56, 58) first two segments, lateral view. Scale lines: 1 mm.

TAXONOMIC NOTES

Ropalidia marginata group
Ropalidia brevita Das and Gupta

Ropalidia spatulata van der Vecht (1962): 9 (partly).

Ropalidia brevita Das and Gupta (1984): 416. Nomen nudum.

Ropalidia brevita Das and Gupta (1989): 110, 121, 163.

Das and Gupta (1989) described this species as closely related to R. spatulata and R. marginata. In the morphological characters including the male antenna (see Figures 1–3) and genitalia, R. brevita is similar to R. marginata, while in coloration R. brevita is similar to R. spatulata. These three species certainly co-occur and can be distinguished from each other by the characters referred to in the key. Although the distinction is sometimes unclear, the first metasomal segment of R. brevitta tends to be more strongly swollen laterally in the posterior widened part (Fig. 36) than that of R. marginata (Fig. 32) and to have a shorter posterior widened part than that of R. spatulata (Fig. 34).

The late J. van der Vecht recognized the species as a subspecies of R. marginata and may have distributed specimens with his identification label of an unpublished subspecific name to various institutions/museums.

Specimens examined. India: Himachal Pradesh: 9♀ (MCZ), Kooloo (=Kullu), M. M. Carleton; Haryana: 6♀ (MCZ), Amballa, M. M. Carleton; Rajasthan: 2♂ (USNM), Abu; Goa: 6♀4♂ (USNM), Morumugao, J. C. B. (2♀, xii.1924; 2♀, viii.1925; 2♀4♂, ix.1925); Kerala: 2♀ (USNM), Walayar forest, S. Malabar, 1000 ft., ix.1951, P. S. N.

Distribution. India (Himachal Pradesh, Delhi, Haryana, Rajasthan, Uttar Pradesh, Sikkim, West Bengal, Assam, Orissa, Goa, Karnataka, Kerala).

Ropalidia marginata (Lepeletier)

Epipona marginata Lepeletier (1836): 541.

Icaria jucunda Cameron (1898): 46. Synonymized under Ropalidia marginata (Lepeletier, 1836) by Nguyen et al. (2006): 101.

Icaria pruinosa Cameron (1906): 228. Synonymized under R. marginata jucunda (Cameron, 1898) by van der Vecht (1941): 104.

Ropalidia marginata sundaica van der Vecht (1941): 122. Synonymized under Ropalidia marginata (Lepeletier, 1836) by Nguyen et al. (2006): 101.

Ropalidia canaria Cheesman (1952): 3, 19. Synonymized under R. marginata jucunda (Cameron, 1989) by Kojima (2001b): 36, 55.

Ropalidia travancorica Lambert and Narendran, in Lambert et al. (2005b): 273–275. Syn. nov.

Represented by several geographic color forms, R. marginata is distributed widely from Pakistan in the west, to New Guinea and Queensland in Australia, and some Pacific islands in the east. In the eastern part of the Lesser Sunda Islands and the southern part of Moluccas, R. marginata is replaced by the closely related species, R. laticintavan der Vecht, 1962, which co-occurs with R. marginata in the western part of the Lesser Sunda Islands (Kojima et al. 2007). In the southern part of Myanmar, a local color form currently treated as a formally named subspecies of R. marginata, rufitarsisvan der Vecht, 1941, is distributed, which, however, may eventually be revealed to be a local color variation of R. spatulata (see Nguyen et al. 2006).

Lambert and Narendran (in Lambert et al. 2005b) described R. travancorica based on a single female from Travancore in Kerala in the subgenus “Icarielia,” and referring to Das and Gupta (1989), compared the species with R. scitula but without seeing any specimens of R. scitula. Detailed comparison of the holotype of R. travancorica with specimens of nearly all the Oriental species of Ropalidia from various localities has led us to conclude that R. travancorica is a synonym of R. marginata.

Specimens examined. India: Rajasthan: 1♀1♂ (USNM), Abu; Karnataka: 5♀ (MCC), Indian Institute of Science Campus, Bangalore, 17.xi.1999, Agrahori; Kerala: 1♀ (holotype of Ropalidia travancorica, ZSIK), labeled “HOLO/TYPE” (circular label, margined with red), “INDIA: Kerala/Thiruvanathapuram/Palayam Kotta/Coll. Thovali/31.xii.1998 (collection date given in the original description is “3.xii.1998”),”“Ropalidia (Icarielia)/travancorica sp. n./Det. Narendran T.C. &/Lambert, 2004” and “1781;” 2♀ (MCC), Kasaragod, 5.x.2005, L. K.; 3♀ (USNM), Walayar forest, S. Malabar, 1000 ft., ix.1951, P. S. N.; Maharashtra: 1♀ (USNM), Bombay 1924, J. C. B.; Tamil Nadu: 1♂ (USNM), Nedundada, Tanjore, 1.vi, P. S. N.; 22♀1♂ (7♀1♂, USNM; 15♀, MCZ), Coimbatore, P. S. N. (15♀, v.1948; 3♀1♂, 2.ix.1951; 2♀, xi.1950; 2♀, ix.1951). Sri Lanka: Puttalam: 1♀ (USNM), 5.iii.1958, R. L. A. Perera; Matela: 2♀ (USNM), 3 miles south-east of Naula, S. L. Wood & J. L. Petty (14.vi.1975; 15.vi.1975); Kandy: 3♀ (USNM), Kandy 1800 ft., Peak View Motel, 15–24.i.1970, Davis & Rowe; 6♀ (USNM), Hantane Hill, 10.xii.1970, Piyadasa & Somapala; 4♀ (USNM), Udawattakele, K. V. K. et al. (510–580 m, 14–16.x.1977; 1–3.x.1973; 9–13.ii.1975; 510–580 m, 8–10.ix.1977, 1700 ft., 29–30.v.1976); 1♀ (USNM), Udawattakele, 2100 ft., 20–30.vii.1976, S. Karunaratne; 1♀2♂ (USNM), Peradeniya, 1♀, 1–15.iii.1971, Piyadasa & Somapala; 1♂, 3.iii.1974, Gans & Prasanna; 1♂, 1500 ft., 5.x.1976, G.F. Hevel et al.; 1♀ (USNM), Teldeniya, 10.v.1975, S. & PB. Karunaratne; Colombo: 2♀ (USNM), Colombo Museum garden (18.ii.1977, K. V. K.; 11–12.i.1977, K. V. K. et al.); 3♀ (USNM), Ratmalana, 27.x.1973, M & B Robinson; 19.i.1975, K. V. K. et al.; 15.ii.1975, K. V. K. et al.; 3♀ (USNM), Kalatuwawa, Y-M. Huang et al. 2♀, 12–15.viii.1975; 1♀, 6–11.viii.1975; 1♀ (USNM), Labugama Reservoire, 11.vii.1978, K. V. K. et al.; 1♀ (USNM), Gampaha Botanic Garden, 14.i.1977, K. V. K. et al.; Monaragala: 3♀ (USNM), Inginiyagala, collected by Malaise trap (2♀, 2–4-vi.1975, D.H. Messersmith et al.; 1♀, 7–8.ix.1975, D.M. Davies et al.); Ratnapura: 1♀ (USNM), Gilimale Jungle, 17.vi.1976, K. V. K. et al.; 2♀ (USNM), Udawalawe 1.viii.1973, Ginter Ekis; 5–6.vii.1975, D.H. Messersmith et al.; Galle: 2♀ (USNM), Kaneliya Jungle, 11–16.i.1975, K. V. K. et al.; 1♀ (USNM), Hiniduma, 20–28.ii.1958, R. L. A. Perera.

Distribution. Pakistan, India (Punjab, Uttar Pradesh, West Bengal, Rajasthan, Gujarat, Madhya Pradesh, Orissa, Maharashtra, Andhra Pradesh, Kernataka, Kerala, Tamil Nadu), Sri Lanka, Myanmar, Vietnam, Malay Peninsula, Borneo, Bangka, Sumatra, Java, Kariman Djawa, Bali, Lombok, Sumbawa, Sumba, Sulawesi, Talud Islands, Tukang Besi Islands, Philippine Islands, Mariana Islands, Palau Islands, Volcano Islands, New Guinea, New Britain, Australia (Thursday Islands, Queensland).

Ropalidia spatulata van der Vecht

Icaria ferruginea Fabricius; de Saussure (1853), in 1853–1858: 38; Horne (1870): 169; André (1889): 189; Bingham (1897): 386, 387; Rothney (1903): 107; Maxwell-Lefroy (1909): 215.

Ropalidia marginata indica van der Vecht (1941): 121 (new name for Vespa ferruginea Fabricius, 1793, non Vespa ferruginea Gmelin, 1790, non Icaria ferruginea of authors).

Ropalidia spatulata van der Vecht (1962): 9; Yamane and Yamane (1979): 4, 6, 32; Das and Gupta (1984): 422; Das and Gupta (1989): 110, 119, 163.

Specimens examined. India: Kerala: 10♀ (MCC), Calicut, 30.x.1998, L. K.; 1♀ (MCC), Nilambur, 4.xi.1998, L. K.; 1♀ (MCC), Manjeri, 18.iv.1999, Jobiraj; 1♀ (MCC), Pulpally, 24.iv.1999, L. K.; 1♀, Trissur, 4.x.1999, K. Usha; 1♀, Trivandrum, 10.iii.2000, L. K.; 1♀, Sulthan Battery, 24.v.2000, L. K.; Tamil Nadu: 2♀ (MCZ), Coimbatore, xi.1949, P. S. N.; 1♀ (MCZ), Gudalur, Nilgiri Hills, 3500 ft., iv.1949, P. S. N.. Myanmar: 5♀ (USNM), Mandalay, H. M. Smith (8.vii.1951; 10.vii.1951; 29.vii.1951; 6.viii.1951; 15.ix.1951)

Distribution. Nepal, India (Kerala).

Ropalidia magnanima van der Vecht

Ropalidia magnanima magnanima van der Vecht (1941): 109, 125; van der Vecht (1962): 9; Das and Gupta (1984): 419; Das and Gupta (1989) 110, 116, 161.

Ropalidia magnanima albitarsis van der Vecht (1941): 125; van der Vecht (1962): 9; Das and Gupta (1984): 419; Das and Gupta (1989): 110, 116, 161.

Ropalidia magnanima anthracina Das and Gupta (1984): 419. Nomen nudum.

Ropalidia magnanima anthracina Das and Gupta (1989): 110, 115–116, 157, 159, 161.

Three subspecies have been recognized in R. magnanima, viz. the nominate subspecies described from Myanmar, albitarsisvan der Vecht, 1941 from Myanmar and Vietnam, and anthracinaDas and Gupta, 1989 from Myanmar. They might be color variations, but their subspecific status is tentatively maintained until more material become available to us.

Specimens examined. India: Kerala: 1♀ (MCC), Mukka, Kozhikode, 18.v.1982, K. N. Nair; 1♀ (MCC), Kannur, 3.ii.1995, Sureshan.

Distribution. India (Kerala), Myanmar, Vietnam.

Ropalidia tamila Gusenleitner

Ropalidia tamila Gusenleitner (2004): 1089.

Specimens examined. India: Kerala: 2♂ (RMNH), Poonmudi, 900 m, Trivandum, iv–v.1971, P. S. N.; 1♂ (RMNH), Tenmalai, Travancore, 12–15.v.1937.

Distribution. India. (Kerala, Tamil Nadu).

Ropalidia stigma group
Ropalidia birmanica van der Vecht

Ropalidia taiwana birmanica van der Vecht (1962): 23; Das and Gupta (1984): 425; Das and Gupta (1989): 135, 167.

Van der Vecht (1962) treated the specimens from Myanmar as forming a subspecies, birmanica, of R. taiwana Sonan 1935, based on the observation that they are, in the female “very similar to typical R. taiwana, but the second gastral (=metasomal) segment not obliquely cut off at end, as seen in profile,” and in the male “antennae slightly less modified than in typical taiwana . . . second gastral segment . . . only slightly obliquely cut off.”Nguyen et al. (2006) referred to four females from Vietnam (two from Sa Pa, Lao Cai Province; one from Pa Co, Mai Chau, Hoa Binh Province; and one from Lam Dong Province). They also recorded “typical”R. taiwana from Vietnam and remarked that the four females "5 belong to birmanica, which might be raised to species rank” and “this, however, should be ascertained with detailed comparison among specimens from various localities.” The species status of the taxon that we identified as “birmanica” seems to be in little doubt based on observations on nests and adult specimens collected in Vietnam (L. T. P. Nguyen et al. unpubl. data, 2006). However, its exact taxonomic status (birmanica or an undescribed species) should be decided only after a detailed comparison of specimens from various localities with the type specimen of R. taiwana birmanica.

Specimens examined. China: 2♀ (USNM, MCZ), Gang Keu, Fukien, South China, 25.viii.1936, L. Gressitt.

Ropalidia sculpturata Gusenleitner

Ropalidia sculpturata Gusenleitner (2001): 655.

Gusenleitner (2001), when he described R. sculpturata, compared it with a Sulawesian species, R. crassavan der Vecht, 1941, and an Indian species, R. santoshaeDas and Gupta, 1989, without examining specimens of these species. We examined the holotype of R. sculpturata and found that it is similar to the specimens that we have identified as R. birmanicavan der Vecht, 1962, including the shape of the metasoma (Figs 37–40), but it differs from the latter in having the disk of the metanotum slightly produced at each posterolateral corner and the posterior face of the propodeum more strongly punctured. As mentioned above, however, the exact taxonomic status of R. sculpturata should be discussed only after the concept of R. birmanica is established.

Specimens examined. Nepal: 1♀ (holotype of R. sculpturata, BOLM), labeled “NEPAL/ANNAPURNA HIMAL/LUMLE/17.-22.06.1999/A. KUDRNA JR. LGT.”, “Biologiezentrum Linz/Austria (LI)/ex Coll. J. Halada/Eigang 2001”, “Ropalidia/sculpturata nov. spec. ♀/J. Gusenleitner, det. 2001/Holotypus”, and “Holotypus” (red square).

Distribution. Nepal.

Ropalidia santoshae Das and Gupta

Ropalidia santoshae Das and Gupta (1984): 422. Nomen nudum.

Ropalidia santoshae Das and Gupta (1989): 111, 123, 156–159, 164.

No specimens of this taxon were available in the present study. Judging from the description by Das and Gupta (1989), R. santoshae is very similar to R. birmanica. However, again, a decision on their taxonomic status needs further detailed comparative study, including observations of the holotypes of both taxa.

Distribution. India (Sikkim, Arunachal Pradesh, West Bengal, Meghalaya).

Ropalidia kasaragodensis Lambert and Narendran, nom. nov.

Ropalidia indica Lambert and Narendran, in Lambert et al. (2005a): 1921. Junior primary homonym of Ropalidia marginata indicavan der Vecht (1941): 121.

Ropalidia indica is preoccupied by R. marginata indicavan der Vecht, 1941, which was a replacement name of Vespa ferruginea Fabricius, 1793, non Vespa ferruginea Gmelin, 1790 (=unrecognized species placed in Crabro (Sphecidae) by Dalla Torre (1897: 600)).

When they described this species, Lambert and Narendran (in Lambert et al. 2005a) compared the species with R. rufocollaris (Cameron, 1900; erroneously cited as “ruficollaris” as in Das & Gupta (1989: 111)). Ropalidia kasaragodensis undoubtedly belongs to the R. stigma species group, to which R. rufocollaris also belongs to. Within the species group, however, R. kasaragodensis is closer to R. birmanica, R. sculpturata or R. santoshae than to R. rufocollaris in having the propodeum distinctly punctured and having a distinct median depression. Ropalidia kasaragodensis is easily distinguished from these three taxa by the female gena very narrow, about half as wide as the eye in lateral view (at least 0.7-fold as wide as eye in R. birmanica, R. sculpturata and R. santoshae).

Specimens examined. 1♀ (holotype, OZNC), labeled “HOLO/TYPE” (circular label, margined with red), “INDIA: Kerala/Kasaragod/Coll. K. Gopi/26.i.1994,”“♀ Ropalidia (Anthreneida)/indica sp. nov./Det. Narendran T.C. &/L. Kishore 2004,” and “♀ Ropalidia (Anthreneida)/kasaragodensis nom. nov./Det Lambert &/DET NARENDRAN, T.C./5.vii.2005.”

Distribution. India (Kerala).

Ropalidia mathematica (Smith)

Polybia mathematica Smith (1860): 90.

Icaria nigroplagiata Cameron (1900): 498. Synonymized under Ropalidia mathematica (Smith) by Kojima et al. (2005: 175).

Ropalidia mathematica mathematica; van der Vecht (1941): 110, 130; Das and Gupta (1984): 420; Das and Gupta (1989): 111, 133.

Ropalidia mathematica binotata van der Vecht (1941): 131. Synonymized under Ropalidia mathematica (Smith) by Kojima et al. (2005: 175).

Ropalidia mathematica nigroplagiata; van der Vecht (1941): 104, 132; Das and Gupta (1984): 421; Das and Gupta (1989): 111, 133, 134, 166.

Ropalidia mathematica sumbaensis van der Vecht (1962): 20. Synonymized under Ropalidia mathematica (Smith) by Kojima et al. (2005): 175.

The taxonomy of R. mathematica is detailed in Saito and Kojima (2005) and Kojima et al. (2005).

No specimens from the Indian subcontinent were available to us.

Distribution. India (Utter Pradesh, Meghalaya, Tripura), Thailand, Vietnam, Bangka, Sumatra, Java, Karimon Djawa, Bali, Lombok, Sumbawa, Flores, Sumba, Timor, Sulawesi.

Ropalidia hongkongensis (de Saussure)

Icaria hongkongensis de Saussure, 1854, in de Saussure (1853–1858): 239.

Ropalidia hongkongensis hongkongensis; Das and Gupta (1984): 418; Das and Gupta (1989): 111, 132.

Ropalidia hongkongensis juncta van der Vecht (1941): 141; Das and Gupta (1984): 418; Das and Gupta (1989): 132, 166. Synonymized under R. hongkongensis (de Saussure) by Nguyen et al. (2006: 101).

No specimens from the Indian subcontinent were available for the present study.

Specimen examined. China: 1♀ (MCZ), Liamui, Hainan Island, 1.viii.1935, L. Gressitt.

Distribution. India (Assam, Meghayala), Myanmar, Vietnam, South China, Hong Kong, Bangka, Java.

Ropalidia nigrita Das and Gupta

Ropalidia nigrita Das and Gupta (1984): 421. Nomen nudum.

Ropalidia nigrita Das and Gupta (1989): 111, 130, 156–159, 166.

Specimens examined. Nepal: 1♂ (USNM), 9 km South of Kodari, 28.x.1977, G. F. Hevel.

Distribution. Nepal, India (Manipur), Vietnam.

Ropalidia rufocollaris (Cameron)

Icaria rufocollaris Cameron (1900): 497; Aiyar (1916): 714.

Icaria ruficollaris[!]Dalla Torre (1904): 74.

Ropalidia ruficollis[!]Dover (1925): 301.

Ropalidia rufocollaris atrata van der Vecht (1941): 139. Synonymized under R. rufocollaris (Cameron) by Nguyen et al. (2006): 102.

Ropalidia rufocollaris rufocollaris; Das and Gupta (1984): 421; (1989): 125, 164.

Ropalidia ruficollaris[!]Das and Gupta (1989): 111.

Ropalidia sridharani Lambert and Narendran, in Lambert et al. (2005b): 268–270. Syn. nov.

When they described R. sridharani, Lambert and Narendran (in Lambert et al. 2005b), referring to the key to Indian species of Ropalidia (Das & Gupta 1989), compared it with R. stigma (Smith, 1858). However, their figures of R. sridharani show the characters that the species may be keyed to R. rufocollaris in Das and Gupta (1989); that is, the first antennal flagellomere (=third antennal segment of Das and Gupta) is more than threefold as long as wide at apex (Fig. 8) and the lateral sides of the second metasomal tergum (=gastral tergite in Das and Gupta) are marked yellow basally. Reexamination of the holotype of R. sridharami has revealed it to be a synonym of R. rufocollaris (Cameron, 1900).

In India this species had previously been recorded only from the north-eastern and northern parts; the present study confirmed its occurrence in southern India.

Specimens examined. India: Mizoram: 1♀ (RMNH), Lushai Hills, Thingsat 2000–35000 ft., 9–10.ix.1960, F. Schmid; Kerala: 1♀ (holotype of R. sridharami, OZNC), labeled “HOLO/TYPE” (circular label, margined with red), “INDIA: Kerala/Malappuram Dt;/Nilambur/Coll. Sridharan/15.XI.2000,” and “♀ Ropalidia (Anthreneida)/sridharani sp. nov./Det. Narendran T. C. &/L. Kishore 2004/779;”Tamil Nadu: 1♀ (USNM), Nilgiri Hills, Gudalur, 3500 ft., iv.1949, P. S. N.

Distribution. India (Utter Pradesh, Sikkim, West Bengal, Meghalaya, Tripura, Assam, Kerala, Tamil Nadu), Tibet, Myanmar, Thailand, Laos, Vietnam.

Ropalidia stigma (Smith)

Polybia stigma Smith (1858): 114; Bingham (1897): 384.

Icaria artifex; Smith (1871): 379; Bingham (1897): 386, 389; Cameron (1905): 71; Cameron (1907): 26; Maxwell-Lefroy (1909): 215.

Parapolybia stigma; von Schulthess (1913): 164.

Icaria stigma; du Buysson (1913): 296.

Ropalidia artifex; Bequaert (1918): 247; Dover (1925): 301; (1931): 257.

Ropalidia stigma stigma; van der Vecht (1941): 110, 126; Das and Gupta (1984): 423; Das and Gupta (1989): 111, 127, 165.

Ropalidia stigma nigrolineata van der Vecht (1962): 18; Das and Gupta (1984): 422; Das and Gupta (1989): 127, 129, 165. Synonymized under Ropalidia stigma (Smith) by Nguyen et al. (2006): 103.

Ropalidia stigma rufa van der Vecht (1941): 130; Yamane and Yamane (1979): 4, 7, 32; Das and Gupta (1984): 422; Das and Gupta (1989): 127, 129, 165. Synonymized under Ropalidia stigma (Smith) by Nguyen et al. (2006: 103).

This species was often recorded in the literature under the name “artifex de Saussure,” which may have led Das and Gupta (1989: 125) to list India in the distribution range of Ropalidia artifex (de Saussure, 1853). Ropalidia artifex, however, has never been recorded from the Indian subcontinent.

Specimen examined. India: Goa: 1♂ (USNM), Mormugao, ix.1925, J. C. B.; Kerala: 1♀ (USNM), Walayar Forest, S. Malabar, 1000 ft., viii.1949, P. S. N.; 1♀ (MCC), Calicut University Campus, 10.iii.1999, L. K.; 1♀ (MCC), Calicut, 20.ii.2000, L. K.. Sri Lanka: Matale: 2♀ (USNM), Sigiriya, 17.vi.1975, D.H. Messersmith et al.; Kurunegala: 1♀ (USNM), Badagamuwa Jungle, 24–27.i.1975, K. V. K. et al.; Ampara: 1♀ (USNM), Ekgal Aru, 12.vi.1976, K. V. K. et al.; Kandy: 5♀ (USNM), Udawattakele, K. V. K. et al. 1–3.x.1973; 2100 ft., 9–13.ii.1975, 1700 ft., 29–30.v.1976; Sanctuary, 8–11.ii.1979; 1600 ft., 18–21.i.1977; 1♀ (USNM), Udawattakele, 14–20.iv.1975, S. & P. B. Karunaratne; 1♀ (USNM), Jambugastenne near Laksapana, 1000 ft., 27.ix.1970, O. S. Flint, Jr.; 1♀ (USNM), Gannorwa Timber Reserve, 4.vi.1976, K. V. K. et al.; 1♀ (USNM), Kandy Reservoire, 29.iii.1975, S. & P B. Karunaratne; Colombo: 1♀ (USNM), Arakawila Jungle, Padukka, 26.xi.1972, P. B. Karunaratne; 3♀ (USNM), Labugama Reservoire (400 ft., 2–3.x.1976, G. F. Hevel et al.; 1.xii.1976, G. F. Hevel et al.; 2–4.ii.1977, K. V. K. et al.); 1♀ (USNM), Gampaha Botanic Garden, 14.i.1977, K. V. K. et al.; Monaragala: 2♀ (USNM), Inginiyagala, D.H. Messersmith et al. 2–3.vi.1975; 1–5.vi.1975; 1♀ (USNM), Angunakolapelessa, 17–19.vi.1978, K. V. K. et al.; Ratnapura: 2♀ (USNM), Sinharaja Jungle, 3 miles south of Weddagala, K. V. K. et al. (8–12.ii.1977; 22–23.ix.1977); 2♀1♂ (USNM), Gilimale, K. V. K. et al. (♂, 17.vi.1976; ♀, Induruwa, 2.ii.1979 & 708.iii. 1979); Uggalakaltota, 23–26.vi.1978, K. V. K. et al.; Galle: 7♀ (USNM), Kaneliya (2♀, 13–13, Baumann & Cross; Udugama, 400 ft., 6–12.x.1973, K. V. K. et al.; 11–16.i.1975, K. V. K. et al.; Hiniduma, 500 ft., 11–12.ii.1972, K. V. K.; Matara: 1♂ (USNM, Enselwatte, 25.v.1975, S. L. Wood & J. L. Petty.

Distribution. Nepal, India (Utter Pradesh, Sikkim, Bihar, West Bengal, Assam, Meghalaya, Manipur, Tripura, Madhya Pradesh, Orissa, Maharashtra, Goa, Kerala), Sri Lanka, Myanmar, Thailand, Malay Peninsula, Vietnam, China (Hainan), Borneo, Sumatra, Java, Bali, Philippine Islands.

Ropalidia plebeja group
Ropalidia rufoplagiata (Cameron)

Icaria rufoplagiata Cameron (1905): 71.

Ropalidia rufoplagiata; van der Vecht (1941): 111, 165.

Ropalidia rufoplagiata rufoplagiata; van der Vecht (1962): 32; Das and Gupta (1984): 422; Das and Gupta (1989): 112, 137, 168.

Ropalidia gravelyi Dover and Rao (1922): 244. Synonymized under Ropalidia rufoplagiata (Cameron) by Kojima et al. (2002): 19.

Ropalidia rufoplagiata gravelyi; van der Vecht (1941): 168; Yamane and Yamane (1979): 32; Das and Gupta (1984): 422; (1989): 137, 138, 168.

Ropalidia rufoplagiata nursei van der Vecht (1941): 167; Yamane and Yamane (1979): 32; Das and Gupta (1984): 422; (1989): 137, 138. Synonymized under Ropalidia rufoplagiata Cameron by Kojima et al. (2002): 19.

Ropalidia nursei; Richards (1978): 57.

Das and Gupta (1989) described R. andamanensis based on a single female collected in Port Blair of the Andamans. The species could be a synonym of R. rufoplagiata, but a decision on its taxonomic status will be made only after the holotype of R. andamanensis is compared with specimens of R. rufoplagiata from various localities including the Andamans. For this reason, R. andamanensis is not treated in this paper.

Specimens examined. India: Karnataka: 1♀1♂ (IUNH), Bangalore, 27.ii.1991, R. Gadagkar.

Distribution. India (Uttar Pradesh, Karnataka, Kerala, Maharashtra), Andaman Islands, Myanmar, Thailand, Vietnam, Malay Peninsula, Sumatra, Bangka, Java, Lombok, Sumbawa, Timor.

Ropalidia bicolorata group
Ropalidia bicolorata van der Vecht

Paraicaria bicolor Gribodo (1892): 249. Secondary junior homonym of Ropalidia bicolor (Smith, 1865).

Ropalidia bicolorata bicolorata van der Vecht (1962): 38, 39; Das and Gupta (1984): 428; Das and Gupta (1989): 110, 153, 154, 173 (error: “bicolorata shiva” in map 26 should read “bicolorata bicolorata”).

Ropalidia bicolorata parvula van der Vecht (1962): 38, 39. Synonymized under R. bicolorata van der Vecht, 1962 by Nguyen et al. (2006): 100.

Ropalidia bicolorata shiva Das and Gupta, 1984: 428. Nomen nudum.

Ropalidia bicolorata shiva Das and Gupta (1989): 153, 154, 173 (error: “bicolorata bicolorata” in map 26 should read “bicolorata shiva”). Syn. nov.

Based on 46 female specimens from the easternmost part of India (Tripura, Assam and Manipur), Das and Gupta (1989) described R. bicolorata shiva. In their key to subspecies of R. bicolorata, the characters of R. b. shiva that distinguish it from the nominate subspecies are “Head and thorax with close and comparatively deeper (close superficial in the nominate subspecies) punctures; sides of propodeum closely but strongly (superficially in the nominate subspecies) rugoso-punctate; fore coxa black or with small yellow mark (black in the nominate subspecies).” At the same time, Das and Gupta (1989) mentioned that the “nominate subspecies was not available for study,” and therefore their comparison with the nominate subspecies seemed to be based only on the description by van der Vecht (1962). Although we have not yet seen any specimens of this species from India, we do not find, based on the description by Das and Gupta (1989), any reason for treating R. b. shiva as a diagnosable species. Hence, we propose to synonymize R. bicolorata shiva under the nominate species.

No specimens from the Indian subcontinent were available in the present study.

Specimens examined. Thailand: 3♀ (USNM), Chiang Mai, D. & E. Thurman (2♀, 28.ii.1952; 1♀, 14.x.1951).

Distribution. India (Assam, Manipur, Tripura), Myanmar, Thailand, China (Yunnan), Vietnam, Borneo.

Ropalidia fasciata group
Ropalidia fasciata (Fabricius)

Eumenes fasciata Fabricius (1804): 290.

Polistes bioculata Fabricius (1804): 278. Synonymized under Ropaidia fasciata (Fabricius) by Kojima (2001a: 8).

Icaria picta de Saussure, 1854, in de Saussure (1853–1858): 238. Synonymized under Ropaidia fasciata (Fabricius) by van der Vecht (1959: 245).

Icaria maculifrons Cameron (1903): 172. Synonymized under Ropalidia picta (de Saussure) by van der Vecht (1941: 104).

Icaria intermedia Cameron (1905): 70. Synonymized under Ropalidia picta (de Saussure) by van der Vecht (1941: 104).

Ropalidia fasciata Fabricius: van der Vecht (1959): 245; Yamane and Yamane (1979): 4, 10, 32; Das and Gupta (1984): 417; Das and Gupta (1989): 112, 140, 169.

Ropalidia faciata[!]Lambert et al. (2005a): 1921.

Ropalidia fasciata is widely distributed from India and Nepal in the east to the Ryukyus, Taiwan, Palawan, Borneo, and the Lesser Sunda Islands in the west. In the areas east of the eastern border of the distribution range of R. fasciata, namely, the Philippine Islands (except Palawan), Sulawesi, Maluku, New Guinea and northern part of Australia, the closely related species, R. impetuosa (Smith, 1860) is distributed. The two species may be distinguished with certainty only in the male.

Specimens examined. India: Tamil Nadu: 1♀ (USNM), Coimbatore, P. S. N. Sri Lanka: Colombo: Iswetakeiyawa, 0–50 ft., 8.v.1976, K. V. K. et al.; Ratnapura: 1♀ (USNM), Udawalawa, 5–6.vi.1975, D. H. Messersmith; Monaragala: 1♀ (USNM), Monaragala, 6.vi.1975, S. L. Wood & J. L. Petty.

Distribution. Nepal; India (Uttar Pradesh, Sikkim, Arunachal Pradesh, Assam, Tripura, Maharashtra, Tamil Nadu), Sri Lanka, Myanmar, Thailand, Vietnam, Malay Peninsula, Borneo, Sumatra, Nias, Bangka, Java, Karimon Djawa, Bali, Flores, Timor, South China, Palawan, Taiwan, Ryukyu Islands.

Ropalidia colorata van der Vecht

Ropalidia colorata colorata van der Vecht (1941): 111, 151; Yamane and Yamane (1979): 32; Das and Gupta (1984): 416; Das and Gupta (1989): 112, 141, 142, 170.

Ropalidia colorata var. sordidavan der Vecht (1941): 154. Syn. nov.

Ropalidia colorata sordida; Das and Gupta (1984): 417; (1989): 112, 141, 142, 170.

Van der Vecht (1941) described sordida as a variety of R. colorata based on a single female from Kooloo (=Kullu, Himachal Pradesh). Judging from the content of his work, van der Vecht certainly proposed the name, “sordida,” for an infrasubspecific entity (see the International Code of Zoological Nomenclature (ICZN), fourth edition, Article 45.6.4). Das and Gupta (1984), without seeing any specimen that has characters agreeing with those of sordida, adopted it as a valid name of a subspecies (see ICZN article 45.6.4.1).

Judging from the original description of sordida, it would be, as van der Vecht considered, no more than an individual color variation. Thus, we propose to synonymize R. colorata sordida under the nominate species.

Specimens examined. India: Himachal Pradesh: 2♀ (IUNH), Solan, c. 1500 m, 28–31.x.1978.

Distribution. Pakistan (north-western part), India (Himachal Pradesh, Utter Pradesh).

Ropalidia variegata group

Distinctions of species in the R. variegata group are often difficult and treatments of taxa in this species group largely depend on the species concept on which a given taxonomic system is based. As we have worked with the species in this species group distributed in other areas (Kojima et al. 2005; Saito & Kojima 2005), the taxa in the Indian subcontinent are discussed based on the phylogenetic species concept elaborated by Nixon and Wheeler (1990) and Wheeler and Platnick (2000).

In the Indian subcontinent, two species of this secies group are recognized; R. variegata (Smith, 1852) and R. jacobsoni (du Buysson, 1908). Das and Gupta (1989) distinguished the two species by the “median line of median groove (=median furrow) of the propodeum not distinct” in R. variegata while “distinct” in R. jacobsoni, “temple (=gena) as wide as eye in profile” in R. variegata while “narrower than eye” in R. jacobosoni, and by the the shape of the aedeagus. The condition of the median furrow of the propodeum may differ between the species, but the distinction is often difficult. The female specimens we examined had their gena always narrower than the eye, about 0.65–0.7-fold as wide as the eye. We were unable to recognize any distinct difference in the male genitalia between the species, even in the figures given in Das and Gupta (1989). Detailed comparison of the morphology among the specimens from various places (specimens from the localities other than in the Indian subcontinent are not listed) revealed that the two species can be distinguished by the shape of the first metasomal tergum as given in the key.

Ropalidia jacobsoni (du Buysson)

Icaria jacobsoni du Buysson (1908): 123.

Ropalidia jacobsoni jacobsoni: Das and Gupta (1984): 418; Das and Gupta (1989): 113, 146, 171.

Ropalidia jacobsoni flavoscutellata Das and Gupta (1989): 146, 147, 160, 171. Syn. nov.

Ropalidia bangalorica Lambert and Narendran, in Lambert et al. (2005a): 1920. Syn. nov.

Based on the examination of specimens from various localities, we reached the conclusion that R. jacobsoni flavoscutellata described by Das and Gupta (1989) based on three females and three males from Assam is no more than a local (or even within-population) color variation and propose to synonymize it under the nominate species.

When Lanbert and Narendran (in Lambert et al. 2005a) described R. bangalorica based on a single female from Bangalore, they compared it with R. fasciata, referring to the key in Das and Gupta (1989). Reexamination of the holotype of R. bangalorica did not reveal any significant differences that allow us to conclude R. bangalorica and R. jacobsoni belong to different species. Herewith we propose to synonymize R. bangalorica under R. jacobsoni.

Specimens examined. India: Maharashtra: 2♀ (USNM), Bombay (ix.1924, J. C. B.; 1.x.1924); Goa: 1♀ (USNM), Mormugao, ix.1926, J. C. B.; Karnataka: 1♀ (holotype of R. bangalorica, OZNC), labeled “HOLO/TYPE” (circular label, margined with red), “INDIA: Karnataka/Bangalore/Coll. O. K. Remadevi/12.viii.1996”, “Host: Inglisia/bivalvata (Coccidae)/Host plant: Sandal”, “♀ Ropalidia (Anthreneida)/bangalorica sp. n./Det. NARENDRAN T. C. &/Lambert 2004”, and “Ropalidia jacobsoni/(du Buysson, 1908)/det. J. Kojima 2005;” 1♀ (USNM), Mangalore, vi.1926, J. C. B.; Tamil Nadu: 7♀ (USNM), Coimbatore, P. S. N. (4♀, 5.ix.1947; 31.viii.1946; x.1951; i.1953); Kerala: 1♀ (USNM), Walayar Forest, 1000 ft., ix.1951, P. S. N.

Distribution. India (Delhi, Utter Pradesh, Rajasthan, Maharashtra, Karnataka, Kerala, Tamil Nadu, Assam), Myanmar, Sumatra, Bangka, Java, Lombok, Sulawesi.

Ropalidia variegata (Smith)

Epipona variegata Smith (1852): 48.

Icaria variegata; de Saussure (1854, in de Saussure 1853–1858): 237, pl. 4, figure 3a; Horne (1870): 169, pl. 20 figures 8 and 9; Bingham (1897): 386, 388.

Icaria pendula Smith (1857): 98. Synonymized under Ropalidia variegata (Smith) by van der Vecht (1941). Revised synonymy.

Ropalidia variegata; Bequaert (1918): 247; Dover and Rao (1922): 244; Dover (1925): 302; (1929): 47; (1931): 257; van der Vecht (1941): 104.

Ropalidia variegata variegata; van der Vecht (1941): 112, 154; Yamane and Yamane (1979): 4, 11, 32; Das and Gupta (1984): 425; Das and Gupta (1989): 113, 144, 170.

Ropalidia pendula; Richards (1978): 58.

Smith (1857) described Icaria pendula from Bareily (=Bareilly in Utter Pradesh) and van der Vecht, on examination of the holotype, synonymized it under R. variegata. Without giving any grounds for treating it as a good species, Richards (1978) listed I. pendula as a species. Following van der Vecht (1941), we propose to treat I. pendula as a synonym of R. variegata.

Specimens examined. India: Kashimir: 1♀ (USNM), Kalabar, ix.1946, P. S. N.; 1♀ (USNM), Tamil Nadu: 1♀ (USNM), Kurumbagaram, Karikal, 22.v.1946, P. S. N.; 18♀2♂ (MCZ, USNM), Coimbatore, P. S. N. (14♀2♂, MCZ, no date; 4♀, USNM: 31.viii.1946; 5.ix.1947; viii.1948; i.1953; Kerala: 3♀ (USNM), Walayar Forest, 1000 ft., 8.ix.1947, P. S. N.

Distribution. Pakistan, Nepal, India (Kashimir, Punjab, Delhi, Uttar Pradesh, Bihar, West Bengal, Gujarat, Madhya Pradesh, Maharashtra, Karnataka, Tamil Nadu, Kerala), Myanmar, Malay Peninsula, China, Sulawesi.

Ropalidia cyathiformis group
Ropalidia cyathiformis (Fabricius)

Eumenes cyathiformis Fabricius (1804): 289.

Icaria ceylonica Cameron (1898): 48; Aiyar (1916): 713. Synonymized under R. cyathiformis (Fabricius, 1804) by van der Vecht (1941: 104).

Icaria cagayanensis Ashmead (1905): 3. Synonymized under R. cyathiformis (Fabricius, 1804) by van der Vecht (1941).

Icaria bilineata Cameron (1905): 72. Synonymized under R. cyathiformis (Fabricius, 1804) by van der Vecht (1941).

Ropalidia cyathiformis; van der Vecht (1941): 104, 112, 158; Yamane and Yamane (1979): 4, 14, 32; Das and Gupta (1984): 416; Das and Gupta (1989): 112, 143, 170.

Ropalidia rodialipa Lambert and Narendran, inLambert et al. (2005a): 1922–23. Syn. nov.

Ropalidia anupama Lambert and Narendram, inLambert et al. (2005b): 270–273. Syn. nov.

We examined the holotype and paratype of R. rodialipa and found that they are females but not males as mentioned by Lambert et al. (2005a); in their figures, the antenna was drawn to have 13 articles, with the terminal flagellomere very short (see fig. 12), but in contrast the metasoma was of the female, with six segments. They compared R. rodialipa with R. jacobsoni (du Buysson, 1908); the holotype differs from R. jacobsoni in the characters mentioned by Lambert et al. (2005a) as well as the pronotal carina barely sinuate laterally (in R. jacobsoni, it is strongly sinuate backward on lateral sides). We have not recognized any characteristics allowing us to consider R. rodialipa as distinct from R. cyathiformis.

Lambert and Narendram (in Lambert et al. 2005b) described R. anupama based on three females from Kerala and compared it with R. malaiseivan der Vecht, 1962 and R. scitula (Bingham, 1897), both belonging to the R. flavopicta group (Kojima 1996). We have reexamined the holotype of R. anupama and found no significant characteristics that allow us to distinguish R. anupama from R. cyathiformis.

Specimens examined. India: Maharashtra: 1♂ (USNM), Bombay, 23.ix.1924; Goa: 1♀1♂ (USNM), Mormugao, ix.1925, J. C. B.; Kerala: 2♀ (holotype and paratype of R. rodialipa, DZUC), holotype labeled “HOLO/TYPE” (circular, with red margin), “INDIA: Kerala/Elathur/Steedharan/19.viii.2000”, “♂ Ropalidia (Anthreneida)/radialipa sp. nov./Det. Narendran T. C. &/L. Kishore, 2004”, and “Ropalidia cyathiformis/♀ (Fabricius, 1804)/det. J. Kojima 2005”; 1♀ (holotype of R. anupama, DZUC), labeled “HOLO/TYPE” (circular, with red margin), “INDIA: Kerala/Cali. Uni. Campus/T. C. Narendran/6.ix.2000”, “♀ Ropalidia (Icarielia)/anupama sp. nov./Det. Narendran T. C. &/L. Kishore, 2004”. Sri Lanka: Kandy: 2♀ (USNM), Peradeniya Botanical Garden, i.1971, Piyadasa & Somapala.

Distribution. Nepal, India (Utter Pradesh, Arunachal Pradesh, Bihar, Assam, Madhya Pradesh, Maharashtra, Goa, Karnataka, Kerala), Sri Lanka, Myanmar, Thailand, Vietnam, Malay Peninsula, Java, Bali, Lombok, Sumba, Sulawesi, Philippine Islands.

Ropalidia sumatrae group
Ropalidia sumatrae Weber

Vespa sumatrae Weber (1801): 103.

Vespa mutillata Illiger (1802): 189. Synonymized under Icaria sumatrae (Weber, 1801) by de Saussure (1854, in de Saussure 1853–1858: 241).

Polistes pubescens Fabricius (1804): 279. Synonymized under Icaria sumatrae (Weber, 1801) by de Saussure (1854, in de Saussure 1853–1858: 241).

Eumenes formicaria Fabricius (1804): 288. Synonymized under Icaria speciosade Saussure, 1855 by Schulz (1912: 88); under Ropalidia sumatrae (Weber, 1801) by van der Vecht (1941: 104).

Anthreneida coronata White (1841): 321. Synonymized under Ropalidia sumatrae (Weber, 1801) by van der Vecht (1941: 104).

Icaria speciosa de Saussure (1855): 374. Synonymized under Ropalidia sumatrae (Weber, 1801) by van der Vecht (1941: 104)

Icaria lugubris (Smith, 1858): 115. Syn. nov.

Icharia[!]marangensis Gribodo (1892): 243. Synonymized under Ropalidia sumatrae (Weber, 1801) by van der Vecht (1941: 104).

Icaria rufinoda Cameron (1904): 121. Synonymized under Icaria speciosa by Meade-Waldo and Morley (1914: 406); under Ropalidia sumatrae (Weber, 1801) by van der Vecht (1941: 104).

Ropalidia sumatrae (van der Vecht (1941): 104), Treubia 18: 104.

Ropalidia sumatrae sumatrae (van der Vecht (1962)): 35, pls. 1 and 3; Das and Gupta (1984): 424; (1989): 112, 136, map 20.

Ropalidia krishna Dover and Rao (1922): 246. Synonymized under Ropalidia lugubris (Smith, 1858) by Dover (1925: 302).

Ropalidia lugubris; Dover (1925): 302; Richards (1978): 57.

Ropalidia sumatrae lugubris; van der Vecht (1941): 104, 185; Yamane and Yamane (1979): 32; Das and Gupta (1984): 425; Das and Gupta (1989): 136, 167.

Van der Vecht (1941) treated R. lugubris as a subspecies of R. sumatrae, mentioning that R. lugubris“is the Bornean representative of R. sumatrae; it differs from the typical form in having the first abdominal (=metasomal) segment black instead of red . . . in some of the specimens I have seen the first segment is partly red . . . occurrence of this form in Bengal needs confirmation.”Richards (1978), without mentioning any reason, listed R. lugubris as a good species. Das and Gupta (1989), following the view of van der Vecht, treated it as a subspecies of R. sumatrae and listed the holotype of R. krishna Dover and Rao, 1922, under R. sumatrae lugubris, which is the only the specimen that they examined. They also examined four female specimens from Myanmar and a female specimen from unknown origin that they identified as R. sumatrae sumatrae. They mentioned for R. sumatrae lugubris that its first metasomal segment is “black but sometimes reddish” (although they examined a single specimen) and that it differed from the nominate subspecies in the condition of punctation on the propodeum and second tergum. Although no specimens from the Indian continent were available for us, examination of specimens from various localities including Borneo (partly listed below) have led us to synonymize R. lugbris under R. sumatrae.

Specimens examined. Malaysia: 2♀ (USNM), Selangor (Kepong, viii.1949; Ulu Langget, 17.vii.1957, Usamru). Singapore: 4♀3♂ (USNM) (1♀, v.1949, N. L. H. Krauss; 1♂, ii.1949, N. L. H. Krauss; 3♀2♂, Baker). Borneo: 2♀ (USNM), Muara Kaman, Samarinda, xi.1950, A. M. R. Wagner. Sumatra: 5♀ (MCZ), Pematong Siantar, vi.1937, C. T. & B. B. Brues.

Distribution. India (West Bengal), Myanmar, Thailand, Vietnam, China (Yunnan), Borneo, Bangka, Sumatra.

Ropalidia flavopicta group
Ropalidia montana Carl

Icaria montana Carl (1930): 22. Nomen nudum.

Ropalidia montana Carl (1934): 675, figures 1–6 and 8–13.

Specimens examined. India: Tamil Nadu: 39♀ (MCZ, USNM), Nilgiri Hills, P. S. N. (10♀ (MCZ) 1♀ (USNM), Singara, 3400 ft., v.1948; 1♀ (MCZ), Mungo Range, 3800 ft., v.1949; 9♀ (MCZ), Gudalur, 3500 ft., iv.1949; 18♀ (MCZ), Cherangoole (1♀, 3400 ft., vi.1948; 8♀, 3500 ft., iv.1949; 5♀, 3500 ft., x.1950; 4♀, 3500 ft., xi.1950)); 2♀ (MCZ), Kurumbagaram, Karikal, v.1950, P. S. N.; 33♀ (USNM), Anamalai Hills, P. S. N. (1♀, 4–5.ix.1946; 2♀, ix.1946; 3♀, 21.vi.1946; 1♀, 28.vi.1946; 24♀, 3100 ft., v.1951; 1♀, 4–5000 ft., 28.vi.1946; 1♀, Cinchana, 1050 m) iv.1956.

Distribution. India (Kerala, Karnataka, Tamil Nadu).

Ropalidia ornaticeps (Cameron)

Icaria ornaticeps Cameron (1900): 496.

Ropalidia flavopicta ornaticeps; van der Vecht (1962): 49; Yamane and Yamane (1979): 32; Das and Gupta (1984): 427; Das and Gupta (1989): 150, 151, 172.

Ropalidia ornaticeps; Yoshikawa et al. (1969): 167, pl. 51; Kojima (1996): 325, 326, 336.

The taxonomy of Ropalidia flavopicata (Smith, 1857) and its closely related species, including R. ornaticeps, was revised by Kojima (1996). No specimens from the Indian subcontinent were available to us.

Distribution. India (Assam, Tripura), Myanmar, Thailand, Cambodia, Vietnam, Malay Peninsula.

Ropalidia scitula (Bingham)

Icaria scitula Bingham (1897): 387, 392.

Ropalidia scitula; van der Vecht (1941): 110, 142; Yamane and Yamane (1979): 32; Das and Gupta (1984): 428; Das and Gupta (1989): 113, 152, 172.

This species has so far been recorded from north-eastern India and northern Myanmar. Northern Thailand is newly added for its distribution range.

Specimens examined. India: Arunachal Pradesh: 1♂ (RMNH), Amatulla 1300–2000 ft., Kameng, 23.v.1961, F. Schmid. Thailand: 1♀ (USNM), Chengmai, 24.vi.1936, C. Tongyai.

Distribution. India (Arunachal Pradesh, Sikkim, West Bengal, Meghalaya), Myanmar, Thailand (new record).

Ropalidia sericea (Cameron)

Icaria sericea Cameron (1905): 73; Aiyar (1916): 714.

Ropalidia sericea; Richards (1978): 57.

Since the original description of Icaria sericea, based on the single specimen, no additional specimens have been collected for this species (see Kojima 1996).

Distribution. India (Sikkim).

DISTRIBUTION PATTERN OF ROPALIDIA SPECIES IN THE INDIAN SUBCONTINENT

In total 26 species of Ropalidia are currently recognized to occur in the Indian subcontinent. They can be divided into the following five groups in terms of their distribution patterns:

  • 1)

    Species endemic to the Indian subcontinent.The following ten species may be categorized in this group with regards to their distribution patterns, and they could be further divided into three subgroups with reference both to the mode of colony foundation (independent founding, in which a colony is founded by inseminated female(s) without aid of workers; swarm founding, in which a colony is founded by workers and associated queens) and phylogentic relationships with the species categorized into other distribution-pattern-based groups.

  • 1–1)

    Independent founding, phylogenetically related to the species with wide distribution ranges.

This subgroup includes three species: R. brevita, R. spatulata and R. colorata. The first two species are closely related to each other and to R. marginata, the related species with a wide distribution range; and R. colorata is closely related to R. fasciata, which has a wide distribution range.

1–2) Independent-founding species, phylogenetically related to the species that occur from the Indian subcontinent to the northen part of Indochina along the eastern slope of Himalaya.

This subgroup consists of four species: one is R. tamila, which is undoubtedly closely related to R. magnanima; and the remaining three species are R. santoshae, R. sculpturata and R. kasaragodensis, all of which are in the R. stigma group and are closely related to each other and to R. birmanica, which is related to R. taiwana distributed from North Myanmar through Vietnam and South China to Taiwan.

1–3) Swarm-founding species.

Ropalidia montana, R. scitula and R. sericea may belong to this subgroup. The former species is distributed in the southern part of India, where the species make large colonies consisting of 5000–60 000 adult female wasps (Yamane et al. 1983; Jeanne & Hunt 1992). The species may possibly be closely related to an Australian swarm-founding species, R. romandi Le Guillou, 1841. Swarm founding, or even any biology, of R. scitula and R. sericea is not known, but these species may be swarm-founders judging from their morphological similarities to the species that are known to be swarm-founders; these two species are known only from the north-eastern part of the Indican subcontinent (R. scitula is known also from northern Thailand).

  • 2)

    pecies occurring from the Indian subcontinent in the west to the northen part of Indochina and/or South China in the east along the eastern slope of Himalaya.

Ropalidia magnanima of the R. marginata groups and three species of the R. stigma group (R. birmanica, R. nigrita, and R. rufocollaris) belong to this group.

  • 3)

    Species distributed from the north-eastern part of the India subcontinent to Indochina and the Malay Peninsula.

Only R. ornaticeps could be categorized in this group. The species belongs to the R. flavopicta group and may closely related to R. flavopicta (Smith, 1857), which occurs in South China, Vietnam, Malay Pensular, Borneo and Sumatra.

  • 4)

    4) Species distributed widely from the Indian subcontinent to the Sunda Islands.

All of the ten species categorized in this distribution-pattern-based group except R. bicolorata and R. sumatra are evidently independent founders. Ropalidia bicolorata and R. sumatra may be swarm founders, but details of their colony founding modes are yet unknown. Of the ten species, three species (R. mathematica, R. hongkongensis, R. stigma) belong to a phylogenetically defined group, the R. stigma group, and two species (R. jacobsoni, R. variegata) belong to the R. variegata group. The other species in these two species groups are known to occur in Australia but not in New Guinea. Ropalidia rufoplagiata belongs to the R. plebeja group and R. fasciata to the R. fasciata group; other species of both species group are known to occur in Sulawesi, New Guinea and nothern part of Australia. The remaining three species, R. bicolorata, R. cyathiformis and R. sumatrae, may not have any closely related species that are known from New Guinea or Australia.

  • 5)

    Species occurring widely from the Indian subcontinent to Australia and Pacific islands.

Only R. marginata is included in this group.

As far as we know, the species in the R. flavopicta group, including R. montana and R. ornaticeps, are all swarm founders, which may use a chemical trail pheromone when they migrate a new nesting site in a swarm of workers and queens (Jeanne 1991; Kojima 1994) and thus may barely disperse across large water bodies such as channels and wide rivers. The swarm-founding Ropalidia wasps of the R. flavopicta group are distributed from the Indian subcontinent to New Guinea and northern part of Australia, but they are absent in Sulawesi, Moluccas and the eastern part of the Lesser Sunda Islands. Such a disjunct distribution pattern would strongly suggest their Gondwana origin and eastward dispersal into the Sunda Islands from the Asian continent (see also Kojima et al. 2005, 2007).

Similar disjunct distribution patterns are found in the R. marginata group, R. stigma group, R. variegata group and R. plebeja group. These four species groups are distributed from the Indian subcontinent to northern Australia (and further eastward to Pacific Islands in the R. marginata group); no species of the R. marginata group has yet recoded from Halmahera (Kojima et al. 2005) and no species of the R. stigma group and R. variegata group from Hamlahera and New Guinea (Saito & Kojima 2005), and no species of the R. plebeja group has yet been recorded from Moluccas (Kojima et al. 2005). An assumption of the Gondwana origin and subsequent eastward dispersal to the Sunda Islands from the continental Asia seems to be widely applicable to Indo-Australian species of Ropalidia.

Phylogenetic analyses in the genus and within species groups are, however, required to test this kind of biogeographic assumption. Furthermore the Gondwana origin hypothesis may explain little about the distribution pattern of species groups which includ species occurring only in the Indian subcontinent and South-East Asia, such as R. sumatrae group and R. bicolorata group. Again, a detailed phylogeny at species level is required as a reference for further discussion on the biogeography of Indo-Australian Ropalidia.

ACKNOWLEDGMENTS

The present study was partly supported by the Japan Society for the Promotion of Science to J. K. (no.s 11833001 and 11691161) and to F. S. (no. 18·5900). J. K. thanks A. S. Menke, J. M. Carpenter and F. Gusenleitner for arranging loans of specimens of the USNM, MCZ and BOLM, respectively. L. K. is grateful to T. C. Narendran for various advice and suggestions.

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