Ropalidia wasps (Insecta: Hymenoptera; Vespidae

Natural History Bulletin of Ibaraki University 5: 31–60, 9 February 2001

Ropalidia wasps (Insecta: Hymenoptera; Vespidae, Polistinae) in New Guinea and its adjacent islands

(first part)

Jun-ichi Kojima Natural History Laboratory, Faculty of Science, Ibaraki University, Mito, 310-8512 Japan

Abstract The first part of a taxonomic revision of the Old World polistine genus Ropalidia Guérin-Minéville, 1831, in New Guinea and its adjacent islands is given. Lectotypes of the following species are designated: Icaria wollastoni Meade-Waldo, 1912, Icaria bispinosa Meade-Waldo, 1912, and Icaria torrida Smith, 1863. A new species is described in the Ropalidia loriana-group. Nomenclatural changes include: synonymies of Icaria sexmaculata Cameron, 1911, Ropalidia pratti Cheesman, 1952, R. maculiventris humboldti Cheesman, 1952, and R. mondoensis Cheesman, 1952, under R. maculiventris Guérin-Méneville, 1831, NEW SYNONYMIES; synonymies of Icaria bispinosa Meade-Waldo, 1912, and Ropalidia fasciata erratica Cheesman, 1952, under R. australis (de Saussure, 1853), NEW SYNONYMIES; synonymy of R. fasciola Richards, 1978, under R. dispila (Cameron, 1913), NEW SYNONYMY; synonymy of R. ivorina Cheesman, 1952, under R. jaculator (Smith, 1871), NEW SYNONYMY; synonymy of R. novaeguineae insolens Cheesman, 1952, under R. semihyalinata (Meade-Waldo, 1912), NEW SYNONYMY; synonymies of R. wudai Cheesman, 1952, and R. wudai trullissima Cheesman, 1952, under R. brunnea (Smith, 1859), NEW SYNONYMIES; synonymies of R. domestica Chessman, 1952, and R. sepicana Richards, 1978, under R. nigra (Smith, 1859), NEW SYNONYMIES; synonymy of R. integra Cheesman, 1952, under R. deminutiva Cheesman, 1952, NEW SYNONYMY; synonymy of Polybia papuana Cameron, 1913, under R. albobalteata (Cameron, 1906), REVISED SYNONYMY; synonymy of R. canaria Cheesman, 1952, under R. marginata jucunda (Cameron, 1898), NEW SYNONYMY. Key words Hymenoptera, Vespidae, Polistinae, Ropalidia, New Guinea, lectotype designation, new species, synonymy

Introduction Ropalidia Guérin-Méneville, 1831, comprising

about 180 species, is one of the largest polistine genera, and is spread over the Ethiopian, Oriental and Australian regions (Kojima and Carpenter, 1997). The genus, together with Parapolybia de Saussure, 1854, Polybioides du Buysson, 1913, and Belonogaster de Saussure, 1854, forms a monophyletic tribe Ropalidiini (Carpenter, 1991, 1993), which is confined to the Old World. Despite the fact that nearly one-third (55 species) of the Ropalidia species that are currently treated as valid species are distributed in New Guinea and its adjacent islands, taxonomy of Ropalidia in the area has yet been very poorly studied compared with congeners in the Oriental region (van der Vecht, 1941a, 1962; Das and Gupta, 1984, 1989) and in Australia (Richards, 1978; Kojima, 1999a, b), as well as with the other three genera of the Ropalidiini (Parapolybia: van der Vecht, 1966; Polybioides: Bequaert, 1918; van der Vecht, 1966; Belonogaster: Richards, 1982; Hensen and Blommers, 1987; Kojima, 2001).

The first record of Ropalidia from New Guinea is of Guérin-Méneville (1831, in Duperrey et al., “1826”[-

1838]), who described Ropalidia maculiventris from “Dory” [=Manokwari]. Ten years later, Le Guillou (1841a) described the second Ropalidia species from Toriton Bay in northwestern New Guinea as Polistes lefebvrei. Around the mid-nineteenth century, de Saussure (1853-58) described two species in his genus Icaria de Saussure, 1853, from the area, and Smith (1859, 1860, 1863, 1865, 1871) described 14 species under the genera Icaria, Polybia Lepeletier, 1836, and Odynerus Latreille, 1802; all but two of Smith’s species are currently treated as valid species in Ropalidia (Kojima and Carpenter, 1997). About the end of nineteenth and early twentieth century, du Buysson (1909a, b, 1913) and von Schulthess (1913) respective-ly described Polybia loriana, Icaria cohni, Polybia novaeguineae and Parapolybia novaeguineae from New Guinea and the adjacent islands. Around the same period, Cameron (1898, 1906a, b, 1911, 1913) describ-ed 20 Ropalidia species from this region under the genera Icaria, Polistratus Cameron, 1906, Polistes Latreille, 1802, Polybia, Odynerus and Ancistrocerus Wesmael, 1836, of which six are currently considered as valid species. Meade-Waldo (1911, 1912; also Meade-Waldo and Morley, 1914) described five species

32 J. Kojima

in Icaria, all of which are currently considered valid, and studied distribution and synonymic relationships of ten species having been described by other authors. However, neither Cameron nor Meade-Waldo made any revisional studies on the Ropalidia fauna in New Guinea nor provided even a list of species recorded in New Guinea by their times.

Bequaert (1932) described Ropalidia leopoldi from Angi-Kita in northwestern New Guinea, and listed 37 species, nearly all the species that had been described from New Guinea. However, he did not make any detailed taxonomic comparisons. Van der Vecht (1934) studied the types of Cameron’s species and concluded that 11 “species” were not valid, and he (1941b) made a brief note on the New Guinean Ropalidia. He also treated six species or subspecies from Papuan region in his monographs on Malayan species of Ropalidia (van der Vecht, 1941a, 1962). Although Cheesman (1952) attempted to study thoroughly the Ropalidia fauna in New Guinea, with descriptions of 18 new taxa, her work was based on only specimens that she herself collected and those in the collections in the University Museum, Oxford, and The Natural History Museum, London, and referred to only 14 of 37 species that Bequaert (1932) listed. Richards (1978) described five additional Ropalidia species from New Guinea in the monograph of the Australian social wasps, and provided brief taxonomic notes on some New Guinean species so far described. We have described further three species from New Guinea (Kojima, 1989; van der Vecht and Kojima, 1996). Thus, as summarized above, the Ropalidia fauna in New Guinea and its adjacent islands, one of the centers of divergence of this genus, has never been studied thoroughly.

It had been the intension of the late Prof. Dr. J. van der Vecht “to discuss the species of this area in a future paper” (van der Vecht, 1962: 4). I started, jointly with him, the taxonomic study of the New Guinean Ropalidia nearly 20 years ago. However, the study was interrupted half way through due to his illness and death (in 1992) (see van Achterberg, 1992). Recently I had several opportunities to visit the Nationaal Natuur-historisch Museum, Leiden, including one-year stay at the museum in 1996, and reexamined a large part of the materials that we studied before.

At the beginning of the preparation of this taxonomic revision, I intended to publish it as a joint paper with the late van der Vecht. However, I decided later to take responsibility for all the nomenclatural changes, because many conclusions presented here are very different from those on which we agreed before and/or the late van der Vecht reached. Most of these discrepancies between my conclusions given in this paper and those we previously reached are due to the following three reasons. First, when we started this

study, either van der Vecht or I followed the subgeneric division defined by Richards (1978), which was shown to be supported by no robust grounds (Kojima, 1997a, b) and even led us some taxonomic confusions at the species level (for examples, see the relationships among R. semihyalinata, R. novaeguineae and R. novaeguineae insolens). Second, some of the late van der Vecht’s conclusions were based on the examination of the types housed in the museums and institutions in Europe that he made in 1933 and 1934 (see van der Vecht, 1934, 1941a); several new taxa have been described since then. Third and the most important, the recent development and sophistication of the phylogenetic species concept led me to concur that subspecies have no place in a phylogenetic system and that diagnosable taxa are to be treated as species (Nelson and Platnick, 1981; Nixon and Wheeler, 1990; Carpenter and Kojima, 1997).

The present paper, as the first part of a compre-hensive taxonomic revision of the New Guinean Ropalidia, treats the results of the research on the type materials of the previously described taxa, and gives a taxonomic revision of the R. loriana-group. The remaining species will be treated in the second part of this series, which will be published in the next issue of the present bulletin.

Materials The present study is based mainly on the specimens

housed in the Nationaal Natuurhistorisch Museum, Leiden (abbreviated as NNM), but also on those in the following museums and institutions:

American Museum of Natural History, New York (AMNH);

Australian National Insect Collection, CSIRO, Canberra (ANIC);

Bernice P. Bishop Museum, Honolulu (BISH); California Academy of Science, San Francisco

(CAS); Hope Entomological Collection, University Mu-

seum, Oxford (OUM); Institut Royal des Sciences Naturelles de Belgique,

Brussels (IRSNB); Museum of Comparative Zoology, Harvard Uni-

versity, Cambridge, Massachusetts (MCZ); Muséum d’Histoire Naturelle, Genève (MHN); Museum National d’Histoire Naturelle, Paris

(MNHN); Museum Zoologicum Bogoriense, Bogor (MZB); Natural History Laboratory, Ibaraki University,

Mito (IUNH); The Natural History Museum, London (BMNH); South Australian Museum, Adelaide (SAM); Zoölogisch Museum, Amsterdam (ZMA).

Ropalidia of New Guinea 33

The New Guinea mainland is divided into the following four sectors, mainly for the convenience in reference (Fig. 1): “North-Western (abbreviated as NW) New Guinea” is the part of Papua (formerly Irian Jaya) north of the central mountains, including the Vogel Kop Peninsula; “South-Western (SW) New Guinea”, the part of Papua south of the central mountains; “North-Eastern (NE) New Guinea”, the part of Papua New Guinea north of the central mountains; and “South-Eastern (SE) New Guinea”, the part of Papua New Guinea south of the central mountains, including Owen Stanley Range. The specimens examined are listed according to this subdivision of the mainland and the names of adjacent islands. When exact localities are not indicated on the labels or are not recognized, the specimens are listed under “New Guinea”.

The name-bearing types (holotypes and lectotypes) of the previously described species are listed with the complete information on the attached labels; for some species, such information is available elsewhere (Kojima and Carpenter, 1997; Kojima and van Achterberg, 1997), but is repeated here for convenience. When label data are quoted, a slash (/) denotes the beginning of a new line on the label; however, I failed to record the line breaks for labels of the specimens in the OUM and BMNH, and their data are given without indicating the line breaks. My notes on the label data and citations are given in brackets.

Methods The adult morphology except for the terminal sterna

and male genitalia was observed on pinned specimens

under a stereoscopic dissecting microscope with an ocular micrometer. When the terminal sterna and male genitalia were examined they were dissected out, briefly cleared in KOH and mounted on glass slide in glycerin to observe under a stereoscopic dissecting microscope and/or a compound microscope. Termi-nology of the male genitalia follows Kojima (1999c).

Fig. 1. Map of New Guinea and it adjacent islands, showing the four sectors of the main island defined in

the present paper.

The following morphometric characters and views of observation are often ill defined in the literature. In the present study they are defined as follows (Figs. 2–7): width of clypeus (CW in Fig. 2) is measured as a distance between the uppermost points where clypeus touches eyes; height of clypeus (CH in Fig. 2) is measured as a distance from the dorsal emargination to the apex; distance between inner eye margins at vertex (OV in Fig. 2) refers to the distance between them at the level of anterior ocellus in frontal view of head, and that at clypeus (in most case equivalent to CW in Fig. 2) refers to the distance at the level where inner eye margins approach each other most closely; inter-antennal (AA in Fig. 2) and antennocular (AO in Fig. 2) distances are measured as horizontal distances at the level of middle of antennal socket; widths of eye (EW in Fig. 3) and gena (GW in Fig. 3) are measured as the maximum for each in strictly lateral view of the head; distance between posterior ocelli (POD in Fig. 5) and distance between posterior ocellus and inner eye margin (OOD in Fig. 5) are measured in dorsal view of head; length of scape (SL in Fig. 4) is measured excluding the radicle; length of first metasomal tergum (T1L in Fig. 6) was measured in profile as a distance between the posterior end of the basal slit for the reception of the mesosomal suspensory ligament and the posterodorsal

34 J. Kojima

Figs. 2–7. Female of Ropalidia maculiventris, showing definitions of morphometric characters and views of

observation. 2, 3, head in frontal (2) and lateral (3) views; 4, right antenna; 5, vertex in dorsal view; 6, apical part of propodeum and first two metasomal segments in lateral view; 7, first metasomal segment in ventral view. OV, distance between inner eye margins at vertex; AA, interantennal distance; AO, antennocular distance; CW, width of clypeus; CH, height of clypeus; EW, width of eye; GW, width of gena; SL, length of antennal scape; POD, distance between posterior ocelli; OOD, distance between posterior ocellus and inner eye margin; T1L, length of first metasomal tergum; T2L, length of second metasomal tergum (or segment); MT2H, height of second metasomal segment; ADV, anterodorsal view of first metasomal tergum; DV, dorsal view of first metasomal tergum; PL, length of posterior widened part of first metasomal sternum; PW, width of posterior widened part of first metasomal sternum.

end of the tergum; length of second metasomal tergum (or segment) (T2L in Fig. 6) was measured in profile as a distance from the bottom of the basal depression or “neck” to the posterodorsal margin of the tergum;

widths of first and second metasomal segments refer to their maximum widths in dorsal view; height of second metasomal segment (MT2H in Fig. 6) is measured as the maximum height in lateral view; length of posterior


widened part of first metasomal sternum (PL in Fig. 7) is measured as a distance between the hypothetical line on the level where ventral margins of the first tergum approach each other most closely and the hypothetical line connecting the posterolateral ends of the sternum, and its width (PW in Fig. 7) refers to the maximum posterior width. The “dorsal view of first metasomal tergum” (DV in Fig. 6) refers to the view perpendicular to the dorsal face of the posterior widened part of the tergum, and “anterodorsal view of first metasomal tergum” (ADV in Fig. 6) is perpendicular to the first metasomal sternum.

Only major references are listed in this paper, and other references can be found in a catalog of ropalidiine species (Kojima and Carpenter, 1997; or on a web-site http://www.sci.ibaraki.ac.jp/~jkrte/wasp/ropa/top.html). For each original citation, the name of the taxon, author name, and year of publication are followed by the sex(es), if stated, and the quotation of the type locality. In cases in which I did not examine the types, the type depositories are also given.

Species are arranged according to the species-groups based on an on-going cladistic analysis of the genus Ropalidia (Kojima and Carpenter, unpublished).

Designation of the lectotypes In any studies of New Guinean Ropalidia before

Bequaert (1932), the holotypes were not designated and the numbers of specimens examined were rarely mentioned. Kojima and Carpenter (1997) designated the lectotypes of 16 New Guinean taxa described by Smith (1859, 1860, 1863), Cameron (1906a, b, 1911, 1913) and Cheesman (1952). The lectotypes of the following three taxa are herewith designated and labeled so.

Icaria wollastoni Meade-Waldo, 1912: Meade-Waldo (1912: 449) described this species based on four females from Mimika River collected in July 1910. Cheesman (1952: 7) inadvertently referred to one of them as “holotype”, stating “1♀ (holotype), 3♀♀, S.W. New Guinea, Mimika R., 1911 (A. F. R. Wollaston)”. Without specifying an individual specimen by its label data and/or characters, her statement cannot be regarded as the designation of the lectotype (Article 74.3 in the International Code of Zoological Nomenclature). The following specimen is herewith designated as the lectotype of I. wollastoni: female (BMNH), labeled “Holotype” (BM type label), “New Guinea, Mimika R., A. F. R. Wollaston, 1911-229 July 1910”, “Icaria (Icariastrum) wollastoni sp. n., Type, ♀, G. Meade-Waldo det.”, and “B.M. TYPE HYM 18.838”. I examined two other specimens in the BMNH with the same collection data as the lectotype and they

are paralectotypes, but I failed to locate the last specimen.

Icaria bispinosa Meade-Waldo, 1912: Meade-Waldo (1912: 449) described this species based on four females from Mimika River collected in July and August 1910. Cheesman (1952: 9) treated this taxon as a subspecies of Ropalidia fasciata (Smith, 1859) and listed the specimens she examined as follows: “1♀ (holotype), 1♀ (paratype), Mimika River, 1911 (A. F. R. Wollaston)”. Again her statement cannot be regarded as the designation of the lectotype. I could find only one specimen of Meade-Waldo’s syntypes, and here designate it as the lectotype of I. bispinosa: female (BMNH), labeled “Holotype” (BM type label), “New Guinea Mimika R. A. F. R. Wollaston, 1911-229 Aug. 1910”, “Icaria (Icariastrum) bispinosa sp. n., Type, ♀ , G. Meade-Waldo det.”, “fasciata Sm. v. bispinosa M. W., L. E. Cheesman det. 1950”, and “B.M. TYPE HYM 18.851”.

Icaria torrida Smith, 1863: Smith (1863: 42) described I. torrida from “Ceram” without giving any indications for the number of specimens he examined. The OUM houses two females labeled “Cer” [= Ceram] (circular) and “Icaria torrida Smith”, that are obviously from the type series. One of them is here designated as the lectotype of I. torrida, and the other specimen (metasoma detached and glued on the circular label) is a paralectotype.

Summary of synonymies Synonymies that are newly proposed in the present

paper and those that were previously published and are confirmed in the present study are summarized below. Ropalidia maculiventris Guérin-Méneville, 1831: =

Icaria sexmaculata Cameron, 1911, NEW SYNONYMY; = Ropalidia pratti Cheesman, 1952, NEW SYNONYMY; = Ropalidia maculiventris humboldti Cheesman, 1952, NEW SYNONYMY; = Ropalidia mondoensis Cheesman, 1952, NEW SYNONYMY.

Ropalidia wollastoni (Meade-Waldo, 1912): = Icaria sericea Cameron, 1911, non 1905, synonymized by Kojima and Carpenter (1997: 60).

Ropalidia australis (de Saussure, 1853): = Icaria bispinosa Meade-Waldo, 1912, NEW SYNONY-MY; = Ropalidia fasciata erratica Cheesman, 1952, NEW SYNONYMY.

Ropalidia conservator (Smith, 1860): = Polybia limatula Smith, 1863, synonymized by van der Vecht (1966: 6); = Icaria parvimaculata Cameron, 1911, non 1907, synonymized by van der Vecht (1934: viii); = Icaria insularis Cameron, 1911, synonymized by van der Vecht (1934: viii); =

36 J. Kojima

Ropalidia conservator goodfellowi Cheesman, 1952, synonymized by Richards (1978: 59); = Icaria waigeuensis Cameron, 1913, synonymized by Meade-Waldo and Morley (1914: 405).

Ropalidia dispila (Cameron, 1913): = Ropalidia fasciola Richards, 1978 (replacement name for Icaria fasciata Smith, 1859, non Eumenes fasciata Fabricius, 1804), NEW SYNONYMY.

Ropalidia jaculator (Smith, 1863): = Ropalidia ivorina Cheesman, 1952, NEW SYNONYMY.

Ropalidia novaeguineae (von Schulthess, 1913): = Polybia novaeguineae du Buysson, 1913, synony-mized by van der Vecht (1966: 6).

Ropalidia semihyalinata (Meade-Waldo, 1912): = Ropalidia novaeguineae insolens Cheesman, 1952, NEW SYNONYMY.

Ropalidia brunnea (Smith, 1859): = Polistratus cariniscutis Cameron, 1906, synonymized by van der Vecht (1934: viii); = Ropalidia wudai Cheesman, 1952, NEW SYNONYMY; = Ropalidia wudai trullissima Cheesman, 1952, NEW SYNONYMY.

Ropalidia conspicua (Smith, 1863): = Odynerus mysolicus Dalla Torre, 1889 (unnecessary replacement name for Odynerus conspicuus Smith, 1863); = Ropalidia conspicua alleni Cheesman, 1952, synonymized by Kojima and Carpenter (1997: 16) without indicating that their treatment was a new synonymy [Richards (1978: 57) treated alleni as a junior synonym of R. brunnea (Smith, 1859), possibly erroneously taking “conspicua alleni” as “cariniscutis alleni”; see also Kojima and Carpenter, 1997: 16].

Ropalidia nigra (Smith, 1859): = Icaria morosa Smith, 1863, synonymized by Cheesman (1952: 10); = Icaria spilostoma Cameron, 1906, synonymized by van der Vecht (1934: viii); = Odynerus sariensis Cameron, 1906, synonymized by van der Vecht (1934: viii); = Odynerus confraternus Cameron, 1911, synonymized by van der Vecht (1934: viii); = Ancistrocerus catharinae Cameron, 1913, synon-ymized by van der Vecht (1934: viii); = Ropalidia domestica Chessman, 1952, NEW SYNONYMY; = Ropalidia sepicana Richards, 1978, NEW SYN-ONYMY.

Ropalidia festina (Smith, 1865): = Icaria zonata Cameron, 1906, synonymized by Meade-Waldo (1911: 100).

Ropalidia albobalteata (Cameron, 1906): = Polybia papuana Cameron, 1913, REVISED SYNONYMY; = ? Icaria (Icariola) fluviatilis Meade-Waldo, 1912; = ? Ropalidia fluviatilis petulans Cheesman, 1952.

Ropalidia catharinae (Cameron, 1913): = ? Ropalidia fluviatilis petulans Cheesman, 1952.

Ropalidia lefebvrei (Le Guillou, 1841): = ? Icaria reactionalis Saussure, 1853.

Ropalidia deminutiva Cheesman, 1952: = Ropalidia integra Cheesman, 1952, NEW SYNONYMY.

Ropalidia impetuosa (Smith, 1860): = Icaria spilocephala Cameron, 1906, synonymized under R. gregaria by van der Vecht (1941a: 104); = Icaria cohni du Buysson, 1909, synonymized under R. gregaria by van der Vecht (1941a: 104); = Ropalidia gregaria inquieta Cheesman, 1952, synonymized under R. gregaria spilocephala by Richards (1978: 86); = Ropalidia gregaria var. tolerans Cheesman, 1952, synonymized under R. gregaria spilocephala by Richards (1978: 86). Designating the lectotype of Icaria gregaria de Saussure, 1854, I (Kojima, 2001: 5) showed that “Ropalidia gregaria” of authors, including van der Vecht (1941a) and Richards (1978), is not the true R. gregaria and adopted R. impetuosa for “R. gregaria” of authors; van der Vecht (1941a: 104) synonymized Icaria impetuosa under R. gregaria.

Ropalidia gracilis (Smith, 1859): = Icaria deceptor Smith, 1863, synonymized by Cheesman (1952: 18).

Ropalidia marginata jucunda (Cameron, 1898): = Icaria pruinosa Cameron, 1906, synonymized by van der Vecht (1941a: 104); = Ropalidia canaria Cheesman, 1952, NEW SYNONYMY.

Misidentifications of New Guinean Ropalidia in the literature

The following misidentifications of New Guinean Ropalidia published in the literature were proved during the course of the present study. Cameron (1906a: 63) recorded a female from

Manokwari as probably “Icaria irritata Smith”. The specimen is in the ZMA and belongs to R. conservator (Smith).

Cameron (1913: 76) recorded “Icaria irritata Smith” from Waigeo, stating that “A male taken in January is probably this species . . .” This specimen is in the NNM and actually the female of R. australis (de Saussure).

Von Schulthess (1932: 39) recorded “Polistes tasmaniensis de Saussure” from Sakoemi. The specimen is in the IRSNB and is a female of R. conservator (Smith).

Cheesman (1952: 11) recorded two females, each from Japen and Waigeo, under “Ropalidia irritata”. I located the female from Japen in the BMNH and this specimen is certainly not R. irritata, but probably an undescribed species; details will be given in the second part of the present series.

Kojima and Spradbery (1987): “Ropalidia leopoldi” (p. 606) is not R. leopoldi and an undescribed species,


which will be treated in the second part of this series; “Ropalidia loriana” (p. 607) is R. longipetiolata (Cameron) as I have already pointed out elsewhere (Kojima, 1998: 218); “Ropalidia sp. possibly R. irritata (Cameron, 1863)” [error in author name; this should be “Smith”] (p. 603) is R. irritata.

Spradbery and Kojima (1989): “Ropalidia kurandae” (p. 639) is an undescribed species, which will be treated in the second part of this series; “Ropalidia leopoldi” (p. 649) and “Ropalidia loriana” (p. 633) are the materials from the same colonies as those in Kojima and Spradbery (1987); and “sp. close to R. bispinosa” (p. 644) is R. australis (de Saussure).

Kojima and Kojima (1994): “Ropalidia leopoldi” (p. 52) is the material from the same colony as in Kojima and Spradbery (1987) and Spradbery and Kojima (1989).

Taxonomy of the Ropalidia loriana-group The R. loriana-group includes three species, R.

loriana (du Buysson, 1909), R. longipetiolata (Cameron, 1911) and a species described in this paper. They are, so far known, endemic to the New Guinea mainland. Richards (1978) placed R. loriana and R. longipetiolata in the subgenus “Icarielia” only based on weak sculpture, but these species are very different from the other species that he placed in “Icarielia” (see Kojima, 1996, 1997b). The diagnostic characters (not all of them is apomorphic) of the R. loriana-group are: body surface barely sculptured; mid-femur slender throughout its length (Fig. 33); propodeal valvula not carinate basally; first metasomal segment elongate and slender (Figs. 12, 13, 26, 27, 36–38); tergum and sternum of second metasomal segment not fused, but overlapping (Fig. 36).

Ropalidia longipetiolata (Cameron) (Figs. 8–21)

Icaria longipetiolata Cameron, 1911: 189, female – “Lorentz”. — van der Vecht, 1941a: 105 (subfamily Polybiinae). — Kojima and Carpenter, 1997: 4 (designation of lectotype).

Ropalidia longipetiolata; Bequaert, 1932: 50 (“probablement un Parapolybia”). — Kojima, 1996: 328.

? Ropalidia loriana (du Buysson); Richards, 1978: 58 (in subgenus Icarielia), 128 (key).

Ropalidia loriana; Kojima and Spradbery, 1987: 607, figs. 37–41 (larva). — Spradbery and Kojima, 1989: 633 (nest; colony population). — Spradbery, 1991: 351 (colony population).

Female — Body length (head + mesosoma + first two metasomal segments) 9–10 mm; forewing length 9.5–10 mm.

Head in frontal view about 1.2 times as wide as high; in dorsal view little more than twice as wide as long, strongly narrowed behind eyes, strongly emarginate posteriorly (Fig. 8), slightly wider than mesosoma including tegulae. Gena in lateral view widest at level of mid-height of eye (about 0.65 times as wide as eye), then gradually narrowing ventrally; occipital carina complete, smoothly and weakly curved. Ocelli proportionally larger than most of other Ropalidia species; posterior ocelli slightly closer to each other than to anterior ocellus; distance between posterior ocelli about 0.65× their diameter, and about 0.3× distance between posterior ocellus and inner eye margin; space between posterior ocelli flat (Fig. 9). Inner eye margins weakly converging ventrally, about 1.1 times further apart at vertex than at clypeus. Clypeus rather convex, pointed apically, slightly wider than high. Interantennal distance nearly the same as antenocular distance. Antennal scape slightly curved, a little more than four times as long as its apical width; third article slightly more than twice as long as its apical width, about as long as fourth and fifth articles combined; tenth and eleventh articles widest in flagellar articles, each distinctly wider than long, and about 1.3 times as wide as apical width of third article; terminal article bullet-shaped, about 1.5 times as long as its basal width (Fig. 10). Mandible rather stout, not twisted; teeth not elongate.

Mesosoma not thick; in dorsal view, nearly 1.5 times as long as wide. Pronotum in dorsal view with lateral margins diverging posteriorly, barely convex; pronotal carina raised into fine lamella, more strongly raised at humeral angles, not sinuate laterally. Scutum convex, nearly 1.1 times as long as its width between tegulae. Scutellum convex (Fig. 11); lateral margins of disc not carinate, barely diverging posteriorly. Disc of metanotum convex, with no lateral marginal carina, slightly produced posteromedially into rounded corner. Mesepisternum without carina; secondary entrance of first spiracle raised, followed posteriorly by depression, which is margined dorsally by fine carina. Posterior face of propodeum convex, without median furrow or depression; in posterodorsal view, lateral sides of propodeum converging posteriorly in nearly straight lines; propodeal orifice round above, about twice as long as wide; propodeal valvula rather large, with broadly rounded apex (Fig. 12), in profile covering ventral half of propodeal teeth.

First metasomal segment slender, about 3.5 times as long as wide, raised at spiracles; in dorsal view with basal parallel-sided part about three times as long as wide, weakly widening posteriorly from level slightly anterior to spiracles, slightly constricted near apical margin, and then weakly widening again at apex, apical width about twice as wide as width of basal parallel-

38 J. Kojima

sided part (Fig. 13); in lateral view dorsal surface very slightly arising posteriorly from posterior margin of basal slit for reception of mesosomal suspensory ligament, then weakly and smoothly convex in posterior one-third (Fig. 12); ventral margins of tergum weakly approaching each other before posterior divergence; posterior margin of sternum weakly emarginate medially. Second metasomal segment flattened dorsoventrally, about as long as wide, about 1.2 times as long as high; in dorsal view spiracle weakly protruding, smoothly expanding posteriorly in anterior half, then slightly narrowing towards apex; thyridium distinct, elongate oval on tergum and line on sternum. Terminal sternum with basal suboval membranous area armed with sparse, short hairs (Fig. 14) (in most Ropalidia species, this area is in peculiar bifurcate shape; see van der Vecht, 1968).

Body shining, covered with fine tomentum except in apical half of clypeus, and sparse short, erect, silvery hairs. Head and mesosoma with sparse, shallow, ill-defined punctures; metasoma unpunctured.

Color brown to dark brown; first metasomal segment (often with brown markings on posterior widened part of tergum), and basal part of second segment, yellow; specimens from Musgrave River with upper half of clypeus, base of antennal scape, ventral part of ocular sinus, gena (partly), yellowish brown, and most of legs (except tarsi), scutellum, metanotum, propodeum, yellow.

Male — Body length (head + mesosoma + first two metasomal segments) 9 mm; forewing length 9.5 mm.

Structure as in female except following characters. Head relatively thinner, about 1.2 times as wide as high, about 2.3 times as wide as long; posterior emargination in dorsal view shallower. Clypeus smaller, nearly 1.1 times as wide as high. Inner eye margins more strongly convergent below, about 1.25 times further apart at vertex than at clypeus. Eye more strongly swollen. Gena narrow, in lateral view about 0.3 times as wide as eye. Antennal scape about 3.7 times as long as its apical width; third article nearly 2.5 times as long as its apical width, slightly longer than fourth and fifth articles combined; terminal article about 1.8 times as long as wide; fourth to terminal articles with slightly raised tyloids (oval shaped rather than of narrow ridge).

Terminal sterna and genitalia as in Figs. 17-21. Parameral spine without hairs; Parameral Spine Index of Charnley (1973): (length of projection beyond lobe)/(total length of paramere and spine) × 100 = 14. Volsella as in most Ropalidia species (see Kojima, 1999c). Penis valves about 1.2 times as long as basal apodeme.

Color as in females from Musgrave River except brown scutellum.

Specimens examined — NW New Guinea: 1♂ (NNM), Vogelkop, Kebar Val., W of Manokwari, 550 m, 4-31.i.1962, S. Quate & L. Quate; 1♀ (NNM), Neth. Ind.-Amer. New Guinea Exp., Mountain slopes above Bernhard Camp, 750 m, 27.iii.1938, L. J. Toxopeus. SW New Guinea: ♀ (ZMA, lectotype designated by Kojima and Carpenter (1997: 4)), labeled “Z. Nieuw Guinea/Lorentz 1909-10/Noord rivier/IX. 09”, “Nest B?”, “longipetiolata [in Cameron’s handwriting]”, “Holotype”, “Icaria/longipetiolata/ ♀ Cam. (1911) /HoloTYPE/sec. J. v. d. Vecht 33 [in van der Vecht’s handwriting” and “Ropalidia ♀ /longipetiolata/(Cam.) /det. J. v. d. Vecht 1933”. NE New Guinea: 1♀ (NNM), Torricelli Mts., Sugoitei Vill., 900 m, 6-9.ii.1959, W. W. Brandt. SE New Guinea: 24♀ (12♀ in NNM, 9♀ in IUNH, 3♀ in AMNH), from a colony, Central District, Baruanumu, Musgrave R., 19.v.1973, J. P. Spradbery; 1♀ (NNM), Fly R., Olsobip, 400-600 m, viii.1969, J. & M. Sedlacek.

Ropalidia loriana (du Buysson) (Figs. 22–27)

Polybia loriana du Buysson, 1909b: 314, female – “Nouvelle Guinée S. E. Moroka 1300 m”

Parapolybia loriana; von Schulthess, 1913: 153 (key), 158, pl. 11 fig. 5 (female).

Ropalidia loriana; van der Vecht, 1966: 6.

Female — Body length (head + mesosoma + first two metasomal segments) about 10 mm; forewing length about 10 mm.

Structure and coloration as in female of R. longipetiolata, but different in the following characters. Head in dorsal view with lateral sides behind eyes hardly convex, more strongly convergent posteriorly (Fig. 22); gena in profile proportionally narrower, slightly more than half as wide as eye Space between posterior ocelli with slightly raised, short, longitudinal carina (Figs. 23, 24). Scutellum less convex (Fig. 25 vs. 11 for longipetiolata); posterior face of propodeum slightly more convex (Fig. 26 vs. 12 for longipetiolata). First metasomal tergum slightly more slender; in profile dorsal margin nearly parallel with its ventral margin in anterior half, smoothly and weakly convex in posterior half (Fig. 26); in dorsal view nearly parallel-sided in anterior half, and weakly swollen posteriorly from just in front of spiracle, then weakly constricted subapically (Fig. 27). Punctures shallower and sparser; scutellum and metanotum barely punctured. Body dark, blackish brown; antennal scape slightly paler; posterodorsal margin of pronotum sometimes with narrow band slightly paler; extreme apex of mesosoma, sometimes area around propodeal spiracle, first metasomal segment (sometimes posterior widened part of the tergum dark brown), base of second metasomal


segment, yellow to orange-yellow. Wings semihyaline, slightly infuscate along anterior margin.

Male — Not known.

40 J. Kojima

Specimens examined — NW New Guinea: 7♀ (NNM, 1♀ in IUNH), Neth. Ind.-American New Guinea Exped., Rattan Camp [3°30’S, 139°09’E], L. J. Toxopeus (1♀ : 1,150 m, 12.i.1939; 3♀ : 1,150 m, ii.1939; 2♀ : 1,200 m, 4.iii.1939; 1♀ : 1,100 m, iii.1939); 1♀(NNM), Neth. Ind.-American New Guinea Exped. Araucaria Camp [3°30’S, 139°11’E], 800 m, iii.1939, L. J. Toxopeus; 1♀ (NNM), Star Mts., Sibil Val., 1,245 m, 18.x-8.xi.1961, L. W. Quate; 1 ♀ (NNM), NE, Swart Val., Karubaka 1,300 m, 7.xi.1958, J. L. Gressitt. NE New Guinea: 1♀ (BISH), Torricelli Mts., Nengian Vill. [3°22’S, 142°05’E], 17-24.xi.1958,

W. W. Brandt. SE New Guinea: ♀ (MNHN, holotype; see also Carpenter, 1999: 36), labeled “N. GUINEA S. E./Moroka 1300 m/LORIA VII-XI 93”, “MUSEUM PARIS/G. Gestro 1909”, “TYPE”, “Polybia/Loriana Buyss./type/R. DU BUYSSON det 1909”.

Figs. 8–27. Ropalidia longipetiolata (Cameron) (8–21) and R. loriana (du Buysson) (22–27). 8–14, 22–27, female;

15–21, male. 8, 22, head in dorsal view; 9, 23–24, ocellar area, 10, 15–16, right antenna; 11, 25, scutum, scutellum and metanotum in lateral view; 12, 26, propodeum and first metasomal segment in lateral view; 13, 27, first metasomal tergum in dorsa view; 14, terminal sternum; 17, terminal sterna, 18, inner aspect of paramere with volsella and digitus; 19, volsella and digitus; 20–21, aedeagus in ventral (20) and lateral (21) views. Scale lines: 1 mm, but 0.2 mm for Figs. 9, 23–24. Figs. 14, 17–21, without scales, based on unpublished sketches given by the late J. van der Vecht; the sketches were scanned and edited with the HP Paper Port program.

Ropalidia principalis new species (Figs. 28–41)

? Ropalidia longipetiolata; Richards, 1978: 58 (in subgenus Icarielia; list), 128 (key).


Female — Body length (head + mesosoma + first two metasomal segments) 11–12.5 mm; forewing length 10–11 mm.

Head in frontal view about 1.2 times as wide as high (Fig. 28); in dorsal view slightly more than twice as wide as long, distinctly wider than mesosoma including tegulae, deeply emarginate posteriorly, with lateral sides behind eyes convex, and strongly converging posteriorly (Fig. 30); in profile, gena widest at level of

mid-height of eye, little more than 0.7 times as wide as eye, gradually narrowing ventrally in ventral half; occipital carina complete, smoothly and weakly curved (Fig. 29). Ocelli large, arranged close to each other in nearly equilateral triangle; anterior ocellus slightly larger than posterior ones; distance between posterior ocelli about 0.6× their diameter and about 0.25× distance to inner eye margin; interocellar space flat. Inner eye margins weakly converging ventrally, about

Figs. 28–41. Ropalidia principalis new species. 28–38, female; 39–41, male. 28–30, 39–40, head in frontal (28,

39), lateral (29, 40) and dorsal (30) views; 31, 41, right antenna; 32, lateral view of scutum, scutellum and metanotum; 33, mid-leg (tarsus not drawn); 34, apical part of right fore wing; 35, propodeal orifice; 36, propodeum and first two metasomal segments in lateral view; 37, first two metasomal terga in dorsal view; 38, first metasomal segment in ventral view. Scale lines: 1 mm, but 0.5 mm for Fig. 35.

42 J. Kojima

1.2 times further apart at vertex than at clypeus (Fig. 28). Clypeus weakly convex, pointed apically, nearly 1.1 times as wide as high. Antenna weakly thickened apically; scape slightly curved, slightly more than 4 times as long as its apical width; third article rather short, slightly more than twice as long as its own apical width, about as long as fourth and fifth articles combined; fourth and fifth articles each about as long as wide; tenth article widest among flagellar articles, about 1.2 times as wide as long, and nearly 1.2 times as wide as fourth article; terminal article bullet-shaped, nearly 1.5 times as long as its basal width (Fig. 31). Mandible rather stout, not twisted; dorsal tooth truncate apically, remaining three teeth pointed apically and subequal in length.

Mesosoma not thick, in dorsal view about 1.7 times as long as wide. Pronotum in dorsal view broadly and weakly rounded anteriorly, with lateral sides diverging posteriorly in nearly straight lines; ventral corner gradually narrowed; pronotal carina fine, thinly raised throughout its length, hardly sinuate on lateral sides of pronotum. Scutum convex, with fine, very shallowly depressed line in anterior one-third to half. Disc of scutellum trapezoid, strongly convex (Fig. 32), with convex, margined laterally by neither carinae nor truncation. Posterior face of propodeum weakly convex (Fig. 36), without median depression or furrow, slightly depressed only just in front of orifice, smoothly curved down into lateral faces; propodeum in posterodorsal view with lateral sides converging posteriorly in nearly straight lines; valvula large, rounded, without basal carina, in profile covering most of propodeal teeth (Fig. 36); orifice rounded above, about twice as long as wide (Fig. 35). Mid femur with short fringe beneath, with sparse outstanding fine setae.

Metasoma slender. First segment elongate, 3.7-3.9 times as long as wide; in dorsal view, with basal parallel-sided part about three times as long as wide, weakly thickening posteriorly from level slightly anterior to spiracle, widest well before apex, then slightly narrowed, and very weakly widened again near apex, widest part about 2.2 times as wide as base (Fig. 37); in profile dorsal face very weakly arising in straight line from posterior end of basal slit, becoming weakly and smoothly curved near level of spiracle (Fig. 36); ventral margins of tergum weakly approaching each other before posterior divergence; posterior widened part of first sternum about as long as wide, with posterior margin emarginate medially (Fig. 38). Second segment shortly petiolate basally, about as wide as long (Fig. 37), about 1.1 times as wide as high; articulation of tergum hardly separated from “neck”.

Clypeus except apical one-third, front, pronotum, scutellum, and anterior half of metanotum with sparse, shallow punctures; vertex and gena barely punctured;

punctures on scutum more or less defined, with central postules; mesepisternum posteriorly with ill-defined sparse punctures, anteriorly unpunctured, border between them ill-defined; mesopleura and sides of propodeum with scattered, shallow punctures; metasoma barely punctured. Body with sparse silvery tomentum, which is dense on upper half of clypeus, sides of mesosoma and second to sixth metasomal segments.

Body pale ferruginous; antennal flagellum fuscous; mandible brown, except dark brown basal margin and teeth; vertex often dark brown; first metasomal tergum dark brown in apical one-third, with ill-defined pale-ferruginous apical band; second to sixth metasomal segments dark brow to blackish brown. All tarsi and hind tibia brown to dark brown. Wings slightly infuscate, more strongly infuscate along anterior margin.

Male — Body length (head + mesosoma + first and second metasomal segments): about 11 mm; forewing length about 10 mm.

Structure and color as in female except: head more transverse, nearly 1.3 times as wide as high (Fig. 39); inner eye margins more strongly converging below, nearly 1.25 times further apart at vertex than at clypeus; clypeus smaller, about as wide as high, covered with rather dense silvery tomentum; gena proportionally narrower, in profile little more than 0.3 times as wide as eye (Fig. 40); antennal scape shorter, about 3.5 times as long as its own apical width; flagellum more slender; third antennal article slightly shorter than fourth and fifth articles combined; terminal article bullet-shaped, about 1.5 times as long as its own basal width (Fig. 41); fourth to terminal articles with slightly raised, ill-defined tyloids.

Holotype — ♀ (NNM), from a colony, NW New Guinea “Neth. Ind.-American/New Guinea Expedit /Bernhard Camp [3°29’S, 139°13’E] B 100 m/9.iv.1939 L. J. Toxopeus”.

Paratypes — 1♂ (allotype) + 112♀1♂ (NNM, 4♀ in IUNH, 1♀in AMNH and 4♀in MZB; 1♀in the MZB has an additional label “Ropalidia/loriana/♀det. v. d. V.”), same data as the holotype.

Other specimens examined — NW New Guinea: 7♀ (MZB; 1♀in BMNH, identified as “longipetiolata” by the late van der Vecht), same data as the holotype; 4♀ (NNM), Maffin Bay [1°57’S, 138°51’E], 25.v.1944, E. S. Ross; 3♀ (NNM), Hollandia [= Jayapura], vi.1937, W. Stüber; 10♀ (NNM, 1♀ in BMNH), Mts. Bewani, iii.1937, W. Stüber; 1 ♀ (NNM), N. N. Guinean Exp., Mamberamo, Alb.-Bivak, 1926, W. Docters v. Leeuwen (with a label “Parapolybia loriana det. Schulthess 93”); 1 ♀ (AMNH), Maffin Bay,


25.xi.[19]44, E. S. Ross. NE New Guinea: 1♀ (NNM), Lae, v.1945, W. Harden.

Etymology — The loriana-group, to which the present species belongs, is the basal clade in Ropalidia (Kojima and Carpenter, unpublished), thus the group can be considered as the first (principalis) group in the genus.

Remarks — This species can be distinguished from the previous two species by the characters given the following key. The present species seems to have been identified as “longipetiolata” or “loriana”. For example, the late van der Vecht identified a female in the MZB as “loriana” and the specimens in the NNM as “longipetiolata”. Furthermore, he identified R. longipetiolata as “loriana” (see Spradbery and Kojima, 1989). He probably sent misidentified specimens to O. W. Richards, and Richards’ (1978: 128) key to species of “subgenus Icarielia” seems to be based on those specimens.

Key to the species in the R. loriana-group (females, but may be applicable to males)

1. Larger species; body length (head + mesosoma + first two metasomal segments) 11–12.5 mm, forewing length 10.5–11 mm. Dorsal surface of first metasomal tergum in profile weakly and smoothly arising from posterior margin of basal slit for reception of mesosomal suspensory ligament, and broadly and smoothly curved down to posterior margin in posterior half (Fig. 36). Interocellar space flat. Punctures on head and pronotum slightly deeper, those on scutum more or less defined and with central postules. Head yellowish brown or pale ferruginous ...................... R. principalis new species

– Smaller species; body length (head + mesosoma + first two metasomal segments) 9–10 mm, forewing length 9.5–10 mm. First metasomal tergum in profile more distinctly swollen dorsally near level of spiracle rather than evenly swollen posteriorly from posterior margin of basal slit for reception of mesosomal suspensory ligament. Punctures on head and mesosoma very sparse, shallow and ill defined. Head brown to dark brown...................................... 2

2. Head in dorsal view with lateral sides behind eyes barely convex (Fig. 22); gena in profile narrower, slightly more than half as wide as eye. Space between posterior ocelli slightly raised into short, longitudinal carina (Figs. 23, 24). First metasomal tergum in profile with dorsal and ventral margins more or less parallel in anterior half, then rather abruptly swollen dorsally just behind spiracle (Fig. 26) ...................................... R. loriana (du Buysson)

– Head in dorsal view with lateral sides behind eyes weakly convex (Fig. 8); gena in profile about 0.65

times as wide as eye. Space between posterior ocelli flat (Fig. 9). First metasomal tergum in profile weakly and gradually swollen posteriorly from its base, then more strongly swollen near spiracle (Fig. 12) .......... R. longipetiolata (Cameron)

Taxonomic notes mainly on types of the species so far described

Ropalidia maculiventris Guérin-Méneville and its closely related species

Ropalidia maculiventris Guérin-Méneville, 1831 and its very closely allied species appear to be widely distributed and rather common in New Guinea and some neighboring islands. It shows a remarkable variation not only in color pattern, but also in the degree of development of the posterolateral angles of the propodeum, which are called “propodeal apophyses”. Specimens from various localities have been described as different species and/or subspecies, and at this moment seven species and/or subspecies are recognized (see also Kojima and Carpenter, 1997): R. maculiventris, R. maculiventris pratti Cheesman, R. humboldti Cheesman, R. mondoensis Cheesman, R. sexmaculata (Cameron), R. fulvopruinosa (Cameron), and R. wollastoni (Meade-Waldo). The variability of the first five “species” and/or “subspecies”, as well as the occurrence of various transitional forms, indicate that they should be considered as a single diagnosable species. The remaining two forms are treated as species, as they are diagnosable with stable characters even only in color pattern.

Ropalidia maculiventris Guérin-Méneville Ropalidia maculiventris Guérin-Méneville, 1831: pl. 9 fig. 8 –

“Dory” [= Manokwari, New Guinea] (holotype in the Museo Civico di Storia Naturale, “Giacomo Doria”, Genova); 1838: 266 (Rhopalidia [!]). — Ducke, 1913: 331 (Rhopalidia [!]; correct name of Icaria maculipennis [!] de Saussure). — Bequaert, 1932: 50. — van der Vecht, 1941b: 28. — Guiglia, 1948: 177 (Rhopalidia [!]; type: a damaged specimen from “Austr. Voy. Coquille” ex coll. Guérin-Méneville in MCG). — Cheesman, 1952: 2, 4.

Icaria maculiventris; de Saussure, 1853-58: 23. Icaria sexmaculata Cameron, 1911: 188, female – “Bivak

Island”. NEW SYNONYMY. — Kojima and Carpenter, 1997: 5 (designation of lectotype).

Ropalidia sexmaculata; Bequaert, 1932: 51. — van der Vecht, 1941b: 28.

Ropalidia maculiventris humboldti Cheesman, 1952: 5, female – “Hollandia”. NEW SYNONYMY.

Ropalidia pratti Cheesman, 1952: 2, 6, female, male – “Papua, Kokoda”. NEW SYNONYMY. — Richards, 1978: 56 (in subgenus Ropalidia).

44 J. Kojima

Ropalidia mondoensis Cheesman, 1952: 2, 6, female – “Papua, Mafulu, 4000 ft”. NEW SYNONYMY. — Richards, 1978: 56 (in subgenus Ropalidia).

Ropalidia humboldti; Richards, 1978: 56 (in subgenus Ropalidia).

Ropalidia maculiventris pratti; Kojima and Spradbery, 1987: 601, figs. 1-7 (larva) — Spradbery, 1991: 351, 354 (colony population).

The late J. van der Vecht, having examined the holotype of Ropalidia maculiventris, suggested me to treat all of I. sexmaculata, R. pratti, R. maculiventris humboldti and R. mondoensis as subspecies of R. maculiventris. I examined the name-bearing types of all of these taxa except for R. maculiventris. All of them were so different in the marking pattern, body size and developmental degree of propodeal apophyses that they all could be treated as belonging to different species. However, examination of many additional specimens, as mentioned in the second part of this series, showed that these differences are intergraded and concluded to follow the late van der Vecht’s view; however, these taxa are all synonymized under R. maculiventris in stead of treating them as subspecies.

Types examined — NW New Guinea: ♀ (BMNH, holotype of Ropalidia maculiventris humboldti Cheesman), labeled “Holotype” (BM type label), “Dutch New Guinea, Humboldt Bay, Hollandia, iv.1936, L. E. Cheesman B.M.1936-271”, “L. E. Cheesman det. 1948, maculiventris ssp. humboldti Chees. Type”, and “B.M. TYPE HYM 18.832”. SW New Guinea: ♀ (ZMA, lectotype of Icaria sexmaculata Cameron, designated by Kojima and Carpenter (1997: 5)), labeled “Z. Nieuw Guinea /Lorentz 1909-10/Bivak Eiland [= Bivak I.; 5°00’S, 138°39’E]/II-’10 [last line vertically written]”, “6-maculata [in Cameron’s handwriting]”, “Icaria ♀/sexmaculata Cam/Holotype by design./J. v. d. Vecht 1933 [in van der Vecht’s handwriting]”, “Ropalidia /sexmaculata (Cam.)/det. J. v. d. Vecht ’34 [in van der Vecht’s handwriting]” and “Holotype”; 1♀ (ZAM, paralectotype of I. sexmaculata), Lorentz 1909-10, Noord Rivier, ix.[19]09. SE New Guinea: ♀ (BMNH, holotype of R. pratti Cheesman), labeled “Holotype” (BM type label), “Papua: Kokoda 1200 ft. vi.1933, L. E. Cheesman B.M. 1933-427”, “pratti Chees. Type, L. E. Cheesman det. 195”, and “B.M. TYPE HYM 18.839”; ♀ (BMNH, holotype of R. mondoensis Cheesman), labeled “Holotype” (BM type label), “Papua, Mafulu 4000ft, i.1934, L. E. Cheesman B.M. 1934-321”, “mondoensis Ch. Type L. E. Cheesman det. 195”, and “B.M. TYPE HYM 18.833”.

Ropalidia fulvopruinosa (Cameron) Odynerus (Leionotus) fulvopruinosa Cameron, 1906b: 225,

female – “Etna Bay”.

Ropalidia fulvopruinosa; Bequaert, 1932: 50. — Richards, 1978: 54 (nest), 56 (list), 63 (key), 67, figs. 12, 25, 33, 41 (in subgenus Ropalidia; redescription).

This species seems to be distributed in the southern coast of New Guinea, from southern coast of Vogel Kop Peninsula and Aru to the Fly River, and in Cape York Peninsula in Australia.

I examined, in addition to the holotype female (ZMA: second to sixth metasomal segments lacking, as mentioned in the original description), labeled “Nieuw Guinea-/Expeditie 1909 / [this slash is written on the label, not indicating line break] 5/Etna-baai [3°58’S, 134°40’E; NW New Guinea]” “Ropalidia [in van der Vecht’s handwriting] / Leionotus / fulvopruinosus / Cam. Type /New Guinea [in Cameron’s handwriting] /Holotype ♀ sec. J. v. d. Vecht ‘33 [in van der Vecht’s handwriting]”, specimens from several localities in New Guinea and Australia (they will be referred to in the second part of this series), and the most reliable character distinguishing R. fulvopruinosa from R. maculiventris is the combination of develop-mental degree of the propodeal apophyses and yellow spots on them. In R. fulvopruinosa, the apophyses are never well developed, and the yellow spots are always present on them. On the other hand, in R. maculiventris, the apophyses are usually well developed and yellow spots on them are found only when the apophyses are produced into sharp teeth.

Ropalidia wollastoni (Meade-Waldo) Icaria wollastoni Meade-Waldo, 1912: 448 (key), 449, female

(in subgenus Icariastrum) – “Mimika River”. Icaria sericea Cameron, 1911: 188, female – “Bivak Island”

(lectotype Amsterdam). Junior primary homonym of I. sericea Cameron, 1905. — Kojima and Carpenter, 1997: 5, 60 (designation of lectotype; syn. of Ropalidia wollastoni).

Ropalidia wollastoni; Bequaert, 1932: 51. — van der Vecht, 1941b: 28. — Cheesman, 1952: 2, 6 (redescription of the type).

Beside the color pattern, such as wings very strongly infuscate, colored blue-black rather than dark brown, I could not find any characters that distinguish R. wollastoni from R. maculiventris or R. fulvopruinosa.

Types examined — SW New Guinea: 3 ♀ (BMNH, lectotype and paralectotypes of Icaria wollastoni Meade-Waldo), Mimika R., vii.1910, A. F. R. Wollaston [for the label data, see the section “Designation of the lectotypes” in this paper]; ♀ (ZMA, lectotype of Icaria sericea Cameron, 1911, designated by Kojima and Carpenter (1997: 5)), labeled “Z. Niuew Guinea/Lorentz 1909-10/Bivak Eiland I 10”, “sericea [in Cameron’s handwriting]”, “Icaria ♀ sericea Cam 1911/Holotype by design. J. v. d. Vecht 1933 [in van der Vecht’s handwriting]”, “= wollastoni M. Waldo


1912 [in van der Vecht’s handwriting]”, and “Ropalidia/wollastoni (M. Waldo)/= sericea (Cam.) 1911/nec. 1906/det. J. v. d. Vecht 1934”; 1♀ (ZMA; Kojima and Carpenter (1997: 5) stated that the lectotype of I. sericea was “probably unique”, but this specimen is evidently from the type series and is a paralectotype of I. sericea), labeled “Z. Nieuw Guinea/Lorentz 1909-10/Bivak Eiland IX 09”.

Other species in the R. maculiventris-group

Ropalidia bicolor (Smith) Icaria bicolor Smith, 1865: 90, female – “New Guinea”. Ropalidia bicolor; Bequaert, 1932: 50. — Cheesman, 1952: 2,

7 (redescription of the type). — Richards, 1978: 56 (in subgenus Ropalidia).

The OUM houses a single female labeled “Holotype” (circular), “N” [= New Guinea] (circular), and “Icaria bicolor Sm.” This is presumably the holotype of I. bicolor Smith, although Smith (1865: 90) did not give any indication for the number of specimens he examined. Smith (1871) did not mention this species in his catalog of the aculeata wasps in India and the Eastern Islands while he listed all the other species of his New Guinean Icaria.

Ropalidia irritata (Smith) Icaria irritata Smith, 1863: 42, female – “Mysol”. Ropalidia irritata; Bequaert, 1932: 51. — Cheesman, 1952: 2,

9 (redescription of type). — Richards, 1978: 56 (in subgenus Ropalidia).

The OUM houses two females: one is labeled “M”, “Icaria irritata Smith”, “Icaria irritata Sm.”, “Type locality = Misool, this true type is labeled “M” the other specimen is from “N” = New Guinea v. d. Vecht, 1958 [in van der Vecht’s handwriting]”; the other is labeled “Type Mysol” “N” (circular), “Icaria irritata Sm.” As suggested in van der Vecht’s label, the former is the true type (and the holotype) of Icaria irritata because “M” of Smith’s label indicates “Mysol” [= Misool].

Ropalidia incurva Cheesman Ropalidia incurva Cheesman, 1952: 2, 7, female, male -

“Papua, Mondo, 4000 ft”.

The holotype female in the BMNH is labeled “Holotype” (BM type label), “Papua: Mondo [8°35’S, 147°18’E; SE New Guinea] 5000 ft. ii.1934, L. E. Cheesman B.M.”, “L. E. Cheesman det. 1948, incurva Chees.” and “B.M. TYPE HYM 18.1033”. Cheesman (1952: 8) listed also a male from the same locality as the holotype, but I failed to locate this male specimen.

Ropalidia australis (de Saussure)

Icaria australis de Saussure, 1853, in de Saussure, 1853-58: 24, pl. 34 fig. 5, female – “Dorey. Nouvelle-Guinée”.

Icaria irritata Smith; Cameron, 1913: 76 [misidentification]. Icaria bispinosa Meade-Waldo, 1912: 448 (key), 449, female

(in subgenus Icariastrum) – “Mimika River”. NEW SYNONYMY.

Ropalidia australis; Bequaert, 1932: 50. Ropalidia bispinosa; Bequaert, 1932: 50. — Richards, 1978:

56 (in subgenus Ropalidia). Ropalidia fasciata bispinosa; Cheesman, 1952: 2, 9. Ropalidia fasciata erratica Cheesman, 1952: 2, 9, female –

“Cyclops Mt.” NEW SYNONYMY. Ropalidia erratica; Richards, 1978: 56 (in subgenus

Ropalidia).

In the MNHN, under “Icaria australia” there are three females and a male: a female labeled “MUS. PARIS/Dorey [= Manokwari]/Dumont d’Urv. 1841”, circle with “Dorey/d Urville”, “TYPE” and “Icaria/australis/type Sauss. [in du Buysson’s hand-writing]/H. DE SAUSSURE DET. [in print]”, that is evidently the holotype of Icaria australis (see also Carpenter, 1999: 34). A male labeled “MUS. PARIS /Nlle Guinie/allemande/J. Bequaert 1909”, “Finschhafen [NE New Guinea]/Duitsch Nieuw/Guinea” and “Icaria /australis /Sauss./R. DU BUYSSON det 1909”, and two females (but they are actually R. irritata (Smith)), labeled “MUS. PARIS/NOUV. GUINÉE/I. SALA-WATI [NW New Guinea]/RAFFRAY & MAINDRON /1878” are apparently of later origin.

Cameron (1906a: 62) and Meade-Waldo (1912: 448) recorded I. australis from “Moaif” and “Mimika River”, respectively, but the holotype of I. australis had not long been referred to until Carpenter (1999). As the identification labels for R. dispila (Cameron, 1913) specimens show, the late van der Vecht regarded that R. dispila was a synonym of R. australis (see the label data for the specimens listed under R. dispila). Actually, however, they are different species.

When he described I. bispinosa, Meade-Waldo (1912: 450) stated that “This species comes very near I. spilostoma, Cam. . . .”; however, I. spilostoma is a synonym of R. nigra (Smith, 1859), which is very different from R. australis in structure. Meade-Waldo’s remarks are based only on the coloration.

Cheesman (1952) treated Meade-Waldo’s (1912) “bispinosa” as a subspecies of R. fasciata (Smith, 1859) and described a new subspecies, R. fasciata erratica. She (p. 8) examined and redescribed the holotype of Icaria fasciata Smith, 1859, and referred to the differences only in marking pattern to treat bispinosa as a subspecies of R. fasciata, and in the structure of scutellum and metanotum as well as marking pattern to describe R. fasciata erratica. She, however, overlooked the facts that the gena and the first metasomal tergum of the types of I. bispinosa and R. fasciata erratica are distinctly proportionally wider than those of the type of

46 J. Kojima

I. fasciata. I could not find any differences among the holotypes of I. australis, I. bispinosa and R. fasciata erratica to treat them as different species.

Other specimens examined — Waigeo: 1 ♀ (NNM), “Waigeoe”, 5.i.[19]10, Mevr. de Beaufort (also labeled “Icaria irritata Sm. ♂ !, det. Cam. [in Cameron’s handwriting]” and “Ropalidia bispinosa (M.W.) var.? ♀ , det. J. v. d. Vecht, 34 [in van der Vecht’s handwriting]”). NW New Guinea: ♀ (BMNH, holotype of Ropalidia fasciata erratica Cheesman), labeled “Holotype” (BM type label), “Dutch New Guinea, Cyclops Mts. Camp. 1: 1200 ft. 22.v.1936 L. E. Cheesman B.M.1936-271”, “fasciata Sm. erratica Ch., Type, L. E. Cheesman det., 1951”, and “B.M. TYPE HYM 18.1038”. SW New Guinea: ♀ (BMNH, lectotype of Icaria (Icariastrum) bispinosa Meade-Waldo) from Mimika River, labeled as shown in the section “Designation of the lectotypes”.

Ropalidia dispila (Cameron) Icaria fasciata Smith, 1859: 167, female – “Aru”. Junior

secondary homonym of Ropalidia fasciata (Fabricius, 1804).

Icaria dispila Cameron, 1913: 77, female – “Waigeu”. Ropalidia dispila; Bequaert, 1932: 50. Ropalidia fasciata; Bequaert, 1932: 50. — Cheesman, 1952:

2, 8 (redescription of the type). Ropalidia fasciola Richards, 1978: 56. (in subgenus

Ropalidia). Replacement name for Icaria fasciata Smith, 1859, non Eumenes fasciata Fabricius, 1804 [Richards sated “Icaria fasciata Smith, 1858]. NEW SYNONYMY.

Although Smith (1859) does not give any indication for the number of specimens, the specimen in the OUM is presumably a unique specimen on which his description of Icaria fasciata was based, and thus the holotype. The specimen in the ZMA marked as type of Icaria dispila is evidently the holotype as Cameron (1913: 77) mentioned that “The 2nd and following segments are absent.”

Judging from the identification labels by the late van der Vecht, he regarded that both Icaria fasciata and I. dispila were the synonyms of Ropalidia australis; he, however, undoubtedly did not examine the holotype of Icaria australis in the MNHN, and thus recognized R. bispinosa Meade-Waldo as a good species (see the identification label for a specimen from Waigeo listed in the section for R. australis). I did not find any differences between the holotypes of I. fasciata and I. dispila to reject the late van der Vecht’s opinion that the two taxa belong to the same species. Here I propose to synonymize R. fasciola Richards under R. dispila (Cameron).

Specimens examined — Waigeo: 1♀ (ZMA, holotype of Icaria dispila Cameron; second to sixth metasomal segments lacking), labeled “Waigeoe 5. 1.

10”, “Mevr. de Beaufort leg”, “Holotype”, “Icaria [in Cameron’ handwriting]/dispila [in Cameron’s hand-writing] ♀ [in van der Vecht’s handwriting]/Cam. Type [in Cameron’s handwriting]/Holotype sec. J. v. d. Vecht ’33 [in van der Vecht’s handwriting]”. Aru: ♀ (OUM, holotype of Icaria fasciata Smith, 1859), labeled “Type” (BM type label), “Icaria fasciata Smith [in Smith’s handwriting]” (blue); 12♀ (NNM, IUNH), labeled “Ins. Aru” and “Museum Leiden, Ropalidia australis Sauss. ♀ , det. J. v. d. Vecht, ‘33”; 1♀ (NNM), labeled “Icaria fasciata Smith, Ins. Aru” and “Museum Leiden, Ropalidia australis Sauss. ♀, det. J. v. d. Vecht, ‘33”; 3♀ (NNM), Rosemberg leg. [1♀ with a label “Ropalidia australis Sauss., fasciata Sm., dispila Cam.”, and 1♀ with a label “Ropalidia australis (Sauss.), J. v. d. Vecht”]; 10♀ (NNM), labeled “Ins. Aru, Rosemberg” and “Museum Leiden, Ropalidia australis Sauss. ♀, det. J. v. d. Vecht, ‘33”.

Ropalidia mimikae (Meade Waldo) Icaria mimikae Meade-Waldo, 1912: 448 (key), 450, female

(in subgenus Icariastrum) – “Mimika River”. Ropalidia mimikae; Bequaert, 1932: 50. — Cheesman, 1952:

2, 9 (redescription of the type).

Although Icaria mimikae is the only species for which Meade-Waldo (1912) did not give the number of specimens he examined, a female in the BMNH labeled “Holotype” (BM type label), “New Guinea Mimika R. [SW New Guinea], A. F. R. Wollaston, 1911-229 aug. 1910”, “Icaria (Icariastrum) mimikae sp. n., Type, ♀, G. Meade-Waldo det.”, “Ropalidia mimikae Meade-Waldo, L. E. Cheesman det., 1951”, “B.M. TYPE HYM 18.852”, and “BM 79” (red) is evidently the holotype of I. mimikae.

This specimen is the only known specimen for R. mimikae; this species could be very rare.

Ropalidia conservator (Smith) Icaria conservator Smith, 1860: 130, female – “Dory”. Polybia limatula Smith, 1863: 43, female – “Mysol”. —

Meade-Waldo, 1911: 100 (syn. of Icaria conservator Smith)). — van der Vecht, 1966: 6 (syn. of Ropalidia conservator (Smith)). — Kojima and Carpenter, 1997: 5 (designation of lectotype).

Icaria irritata Smith; Cameron, 1906a: 63 [misidentification]. Icaria parvimaculata Cameron, 1911: 189, female – “Bivak

Island”. Junior primary homonym of I. parvimaculata Cameron, 1907. — van der Vecht, 1934: viii (syn. of Ropalidia conservator (Smith)). — Kojima and Carpenter, 1997: 5 (designation of lectotype).

Icaria insularis Cameron, 1911: 188, female – “Bivak Island”. — van der Vecht, 1934: viii (syn. of R. conservator (Smith)).

Parapolybia limatula; von Schulthess, 1913: 164 (syn. of Icaria conservator).


Icaria waigeuensis Cameron, 1913: 76, female – “Waigeu”. — Meade-Waldo and Morley, 1914: 405 (syn. of Icaria conservator Smith). — van der Vecht, 1934: viii (syn. of R. conservator (Smith)).

48 J. Kojima

Ropalidia conservator; Bequaert, 1932: 50. — van der Vecht, 1934: viii. — Cheesman, 1952: 2, 15 (redescription). — Richards, 1978: 54, 59, 130 (in subgenus Icarielia).

Ropalidia insularis; Bequaert, 1932: 50. Ropalidia parvimaculata; Bequaert, 1932: 50. Polistes tasmaniensis de Saussure; von Schulthess, 1932: 39

[misidentification]. Ropalidia conservator goodfellowi Cheesman, 1952: 2, 16,

female – “S. Dutch New Guinea, Mimika R.”. Ropalidia conservator waigeuensis; Cheesman, 1952: 2, 16. Ropalidia conservator form goodfellowi; Richards, 1978: 130. Ropalidia waigeuensis; Richards, 1978: 59 (syn. of R.

conservator (Smith)). Ropalidia conservator form waigeuensis; Richards, 1978:

130.

In addition to the specimens listed below, I examined many specimens from various localities (they will be referred to in the second part of this series); these specimens showed so large variations in the marking pattern and the shape of the first metasomal segment that the differences among the types of above listed taxa are intergraded.

Specimens examined — Waigeo: ♀ (ZMA, holotype of Icaria waigeuensis Cameron), with labels “Holotype”, “Waigeoe 31 XII 09”, “Mevr. de Beaufort leg”, and a label on which are Cameron’s handwriting “Icaria/waigionensis/Cam. Type [in Cameron’s hand-writing]” and van der Vecht’s handwriting “= Icaria /conservator/Smith”. Misool: ♀ (OUM, lectotype of Polybia limatula Smith, designated by Kojima and Carpenter (1997: 5)), labeled “M” [= Mysol], “Polybia limatula Smith” and “Lectotype desig. J. v. d. Vecht Aug. 1958”; 1♀ (OUM; Kojima and Carpenter (1997) did not refer to this specimen when they designated the lectotype of Polybia limatula, but this specimen is evidently from the type series of P. limatula and is now a paralectotype), labeled “M” and “Polybia limatula Smith”. NW New Guinea: ♀ (BMNH, holotype of Icaria conservator Smith), labeled “Type” (BM type label), “This must be regarded as the holotype of Ic. conservator Sm. no other specimens from Dory being preserved in Oxford! J. v. d. Vecht, 1934 [in van der Vecht’s handwriting]”, “conservator Smith”, “Dor” [= Dory] (circular), “Smith coll. press by Mrs. Farren White 99-303”, “B.M. TYPE HYM 18.849”, and “Holotype Icaria conservator Smith, 1860, Teste C. O’Toole, 1993”; 1♀ (IRSNB), Sakoemi [probably near Triton Bay], with an identification label as “Polistes tasmaniensis” by von Schulthess; 1♀ (ZMA), labeled “Manokwari 27 mei [19]03”, “Icaria irritata Sm. Sec. Cam. New Guinea [in Cameron’s handwriting]”. SW New Guinea: ♀ (ZMA, lectotype of Icaria parvimaculata Cameron, designated by Kojima and Carpenter (1997: 5)), with labels “Holotype [in van der Vecht’s handwriting]”, “Z. Nieuw Guinea/Lorentz 1909-10/Heurel Bivak/XI 09/750 m [last two lines are

vertically written]”, “parvimaculata [in Cameron’s handwriting]” and “Icaria ♀ /parvimaculata Cam /Holotype sec. J. v. d. Vecht ’33 [in van der Vecht’s handwriting]”; 1♀ (ZMA, paralectotype of Icaria parvimaculata; damaged specimen), labeled “Z. Nieuw Guinea/Lorentz 1909-10/Bivak Eiland/II-’10 [last line written vertically]”, “parvimaculata [in Cameron’s handwriting]” and “Icaria ♀ Cam. /parvimaculata /Paratype [in van der Vecht’s handwriting]”; ♀ (ZMA, holotype of Icaria insularis Cameron), labeled “Holotype”, “Z. Nieuw Guinea/Lorentz 1909-10/Heurel Bivak/XI-09/750 m [last two lines are written vertically]”, “insularis [in Cameron’s handwriting]”, “Icaria ♀/insularis Cam./HOLOTYPE/designated by/J. v. d. Vecht 1933 [in van der Vecht’s handwriting]”; ♀ (BMNH, holotype of Ropalidia conservator good-fellowi Cheesman), labeled “Type” (BM type label), “New Guinea Mimika R. A. F. R. Wollaston 1911-229”, “L. E. Cheesman det. 1948 conservator Sm. ssp. goodfellowi Chees.” and “B. M. TYPE HYM 18.850”; 2♀ (BMNH, paratypes of R. conservator goodfellowi), Mimika R., vii.1910, A. F. R. Wollaston.

Ropalidia novaeguineae (von Schulthess) Parapolybia novaeguineae “Saussure nom. mus.” von

Schulthess, 1913: 154 (key), 162 (description based on notes received from Meade-Waldo) – “Neu-Guinea”.

Polybia novae-guineae du Buysson, 1913: 297, female – locality not stated, but evidently from New Guinea. Junior secondary homonym of Ropalidia novaeguineae (von Schulthess). — van der Vecht, 1966: 6 (syn. of Ropalidia novaeguineae (von Schulthess)).

Ropalidia novae guineae [!]; Cheesman, 1952: 3, 23 (redescription of a type of “Polybia novaeguineae du Buysson”).

Ropalidia novaeguineae; van der Vecht, 1966: 6. — Richards, 1978: 59 (in subgenus Icarielia; listed as R. novaeguineae (von Schulthess)), 129 (R. novaeguineae (R. du Buysson) in a key to species of the subgenus Icarielia).

Based on the notes received from Meade-Waldo, von Schulthess (1913, April) described Parapolybia novaeguineae, indicating “Neu-Guinea (Type Museum London)”. Meade-Waldo’s notes was evidently based on a female in the BMNH (second to terminal metasomal segments are separately attached to a small piece of card board), that is labeled “Type H. T” (circled with red), “New Guin.”, “Type. F. Sm. coll. 79.22”, “Polybia novae guineae Sauss. Type”, “Saussure’s name in ms. R. du Buysson det. Bull. ent. Soc. Fr. p. 297 (1913) G. Meade-Waldo det.” and “B.M. TYPE HYM 18.876”. Thus this specimen is the holotype of Parapolybia novaeguineae von Schulthess.

In July of the same year, du Buysson (1913) described Polybia novaeguineae, mentioning “. . . n. sp., in mus. SMITH 1879. . . . Cet insecte est unique et il porte de la main de SMITH l’étiquette: «Polybia Novae-


Guineae Sauss» . . .” A female in the OUM labeled “New Guinea” (circular), “Polybia Novae Guineae Sauss.”, “Polybia novae-guineae Saussure et Smith, type, det. 1912, R. du Buysson”, “Coll. Smith, 1879” is evidently the holotype of Polybia novaeguineae du Buysson. There are no distinct differences between the holotypes of Parapolybia novaeguineae von Schulthess and Polybia novaeguineae du Buysson that suggest they belong to different species; van der Vecht’s (1966: 6) synonymy is confirmed.

Cheesman (1952: 23) mentioned “novae guineae du Buyss., 1913 . . . Named by Saussure (Polybia novae guineae) but not described. Smith’s Collection. Collected by A. P. R. Wallace on Salawatti Is., W. New Guinea. Type, B.M. . . . a redescription follows. . . . 1♀ (holotype), Smith’s collection, New Guinea, 1♀ , labelled 62:91, which signifies 1862, Salawatti (Accession Register).” The specimen that she considered as the holotype of “novaeguineae du Buysson, 1913” is evidently the holotype of Parapolybia novaeguineae von Schulthess, 1913. Furthermore, neither von Schulthess (1913) nor du Buysson (1913) referred to the exact locality or the collector; Cheesman (1952) seemed to refer to the label datum for the specimen labeled “62:91” as if it were for the holotype of Parapolybia novaeguineae.

Richards (1978: 59, 129) placed this species in the subgenus “Icarielia”, but the species may be keyed to the “subgenus Ropalidia” with his key to the subgenera of Ropalidia (p. 51).

Ropalidia jaculator (Smith) Odynerus fallax Smith, 1863: 40, female – “Mysol”. Odynerus jaculator Smith, 1871: 377. Replacement name for

O. fallax Smith, 1863, non 1861. Odynerus tertius Dalla Torre, 1889: 125. Unnecessary

replacement name for Odynerus fallax Smith, 1863, non 1861 and de Saussure, 1852.

Ropalidia ivorina Cheesman, 1952: 3, 25, female – “Papua, Kokoda, 1200 ft.” NEW SYNONYMY. — Richards, 1978: 58, (in subgenus “Icarielia”), 129 (key to species in the subgenus “Icarielia”).

Ropalidia jaculator; Kojima and Carpenter, 1997: 6.

Since Smith (1871) replaced his Odynerus fallax with Odynerus jaculator this species had not long been referred to until Kojima and Carpenter (1997: 6); exceptions are Dalla Torre (1889, 1894, 1904), but he certainly did not see any specimens of this species. The holotype of Odynerus fallax that Kojima and Carpenter (1997: 6) examined is in the OUM and labeled “M” [= Mysol] (circular), “Odynerus fallax Smith”, “Icaria tertia DT”, and “Type 091 Hope Dept. Oxford”. I compared this specimen with the holotype of Ropalidia ivorina (BMNH), labeled “Type” (BM type label), “Papua: Kokoda [SE New Guinea] 1200 ft VI 1933 L.

E. Cheesman B. M. 1933-427”, “ivorina Chees. L. E. Cheesman det. 1951” and “BM TYPE HYM 18.1036”, and a paratype (BMNH) of R. ivorina from the same locality as the holotype, and found no specific differences among them. I herewith propose to synonymize R. ivorina under R. jaculator.

Ropalidia semihyalinata (Meade-Waldo) Icaria semihyalinata Meade-Waldo, 1912: 449 (key), 451,

female (in subgenus Icariola) – “Mimika River”. Ropalidia semihyalinata; Bequaert, 1932: 51. — Cheesman,

1952: 3, 24 (semihyalineata [!]; redescription). — Richards, 1978: 56 (in subgenus Ropalidia).

Ropalidia novaeguineae insolens Cheesman, 1952: 3, 24, female – “Japen, Mt. Baduri, 1000 ft.”. NEW SYNONYMY. — Kojima and Carpenter, 1997: 6 (designation of lectotype).

Ropalidia insolens; Richards, 1978: 58 (in subgenus Icarielia), 129 (key).

Meade-Waldo (1912: 451) clearly indicated that he examined a single female for his Icaria semihyalinata and the holotype in the BMNH is labeled “Holotype” (BM type label), “New Guinea, Mimika R. [SW New Guinea], A. F. R. Wollaston, 1911-229 July 1910”, “Icaria (Icariola) semihyalinata sp. n. Type, G. Meade-Waldo det.”, and “B.M. TYPE HYM 18.874”.

I failed to locate the specimens from the type series of Ropalidia novaeguineae insolens other than the lectotype designated by Kojima and Carpenter (1997: 6; erroneously mentioning “Ropalidia insolens Cheesman, 1952”), which is labeled “Type” (BM type label), “Dutch New Guinea, Japen Is., Mt. Baduri, 1000 ft., viii.1938, L. E. Cheesman, B.M. 1938-593”, “L. E. Cheesman det. 1948, insolens Chees.”, and “B.M. TYPE HYM 18.875”. A headless female (BMNH) identified as insolens by Cheesman is from “Humboldt Bay”, the locality listed by Cheesman (1952: 24). However, the collection date of this specimen is “vi 1937”, while Cheesman mentioned “Humboldt Bay, ii 1936”, thus this specimen is not a paralectotype.

I could not find any specific differences among the holotype of I. semihyalinata, the lectotype of R. novaeguineae insolens and a female identified as “insolens” by Cheesman; R. novaeguineae insolens is a synonymy of R. semihyalinata. On the other hand, they are different from the types of R. novaeguineae in the shape of the first metasomal segment (details will be given in the second part of this series).

Cheesman (1952) described R. novaeguineae insolens in the “Group VIII”, which included also R. novaeguineae, R. semihyalinata and R. ivorina. Richards (1978) raised insolens to the species rank and placed it in “subgenus Icarielia”, in which he placed also R. novaeguineae and R. ivorina. On the other hand,

50 J. Kojima

Richards (1978) placed R. semihyalinata in “subgenus Ropalidia”.

Species in the R. bidens-group Ropalidia bidens van der Vecht

Ropalidia bidens van der Vecht, 1996, in van der Vecht and Kojima, 1996: 99, female, male – “N. W. New Guinea, Klamono Oilfds [1°08’S, 131°30’E].”

The holotype female (NNM) is kept in the type collection separately from the ordinary specimens, and labeled “NW New Guinea/Klamono Oilfds/18-24.viii. 1948/M. A. Lieftinck”. The paratype male and female in the BISH are labeled as follows: male: “NEW GUINEA: NETH./Bodem, 100 m, 11 km/SE of Oerberfaren/July 7-17, 1959” and “T. C. Maa/Collector /BISHOP”; and female paratype: “NEW GUINEA: NETH/Boden, 100 m, 11 km/SE of Oerberfaren/July 7-17, 1959/T. C. Maa”

Ropalidia clypeata Kojima Ropalidia clypeata Kojima, 1996, in van der Vecht and

Kojima, 1996: 103, male, female – “New Guinea, Sivipi near Sasambata, Popondetta subdist.”

The holotype male (NNM) is kept in the type collection separately from the ordinary specimens, and labeled “Sivipi near Sasambota/Popondetta Subdist. /Papua 1 Nov. 1963/D. K. McAlpine”, “Austr. Mus. Collection” and “Museum Leiden/ex. collectie/J. v. d. Vecht”. I failed to locate the lost female paratypes.

Species in the R. nigra-group

Ropalidia bambusae Richards Ropalidia bambusae Richards, 1978: 54 (nest), 57 (list), 65

(larva in key), 122 (key), 123, female, male, nest (in subgenus Polistratus) – “New Guinea: Morobe District, Wau”.

This species is certainly a valid species. Types examined — NE New Guinea: ♀ (BMNH,

holotype), labeled “Holotype” (BM type label), “NG42 nest, New Guinea: Morobe district, Wau, 5.i.1973, O.W. Richards”, and “Ropalidia bambusae Richards ♀, Holotype”; 1♂ (NNM, paratype), same data as the holotype; 1♀ (NNM, paratype), nest NG6, Wau, 20.xii.1973, O. W. Richards.

Ropalidia brunnea (Smith) Icaria brunnea Smith, 1859: 167, female – “Aru”. Polistratus cariniscutis Cameron, 1906a: 59, [female] –

“Manokwari”. — van der Vecht, 1934: viii (syn. of R. brunnea (Smith)).

Ropalidia brunnea; Bequaert, 1932: 51. — van der Vecht, 1934: viii. — Cheesman, 1952: 2, 14 (redescription of the type) [error: year of publication, 1864]. — Richards, 1978: 57 (in subgenus Polistratus; list; syn.: cariniscutis (Cameron), alleni Cheesman) [alleni as syn., apparently by error; see also Kojima and Carpenter, 1997: 16], 121 (key).

Ropalidia brunnea var. cariniscutis; van der Vecht, 1934: viii. Ropalidia cariniscutis; Bequaert, 1932: 51. — Richards,

1978: 121 (syn. of R. brunnea (Smith)). Ropalidia wudai Cheesman, 1952: 2, 14, female – “Papua,

Kokoda, 1200 ft.” NEW SYNONYMY. — Richards, 1978: 57 (in subgenus Polistratus).

Ropalidia wudai trullissima Cheesman, 1952: 2, 15, female – “Dutch New Guinea, Cyclops Mts., 930 ft.” NEW SYNONYMY. — Richards, 1978: 57 (in subgenus Polistratus).

The OUM houses a female that was referred to by Cheesman (1952: 14) and is labeled “Paratype O. U. M.”, “M”, “N. G. Ins.” and “Icaria brunnea Sm”. “M” is written in Smith’s handwriting and indicates that the specimen was from “Mysol” [= Misool]; thus the specimen is not from the type series of I. brunnea. A female in the BMNH labeled “Aru” is evidently the holotype of I. brunnea.

Cameron (1906a: 59) did not mention the sex and the number of specimens when he described Polistratus cariniscutis. A female in the ZMA is certainly the unique specimen as Kojima and Carpenter (1997: 12) mentioned.

In addition to the holotypes of I. brunnea and P. cariniscutis, I examined the types of Ropalidia wudai and R. wudai trullissima. I did not find any structural differences to treat them as different species, or differences in the marking pattern that may diagnose any of them. I synonymize R. wudai and R. wudai trullissima under R. brunnea.

Types examined —NW New Guinea: ♀ (ZMA, holotype of Polistratus cariniscutis Cameron), labeled “Manokuwari 2 mei 03”, “Ropalidia brunnea (Sm.) var. cariniscutis Cam. ♀/det J. v. d. Vecht ’34 [in van der Vecht’s handwriting]” and “Polistratus cariniscutis Cam. Type/New Guinea [in Cameron’s handwriting] /Holotype by designation J. v. d. Vecht 1933 [in van der Vecht’s handwriting]”; ♀ (BMNH, holotype of Ropalidia wudai trullissima Cheesman), labeled “Holotype” (BM type label), “Dutch New Guinea, Cyclops Mts. Sabron 930 ft. iv.1936, L. E. Cheesman B.M. 1936-271”, “L. E. Cheesman det. 1949, wudai Chess. ssp. trullissima Chess.”, and “B.M. TYPE HYM 18.861”; 1♀ (BMNH, paratype of R. w. trullissima), same data as the holotype of R. w. trullissima. Aru: ♀ (BMNH, holotype of Icaria brunnea Smith), labeled “Holotype” (BM type label), “Aru” (blue, circular), “Icaria brunnea Smith” (blue), “Smith coll. press by Mrs. Farren White 99-303”, and “B.M.TYPE HYM


18.863”. SE New Guinea: ♀ (BMNH, holotype of Ropalidia wudai Cheesman), labeled “Holotype” (BM type label), “Papua: Kokoda 1200 ft. viii.1933, L. E. Cheesman B.M. 1933-427”, “L. E. Cheesman det. 1948, brunnea Sm. wudai Chess.”, and “B.M. TYPE HYM 18.862”; 1♀ (BMNH, paratype of R. wudai), same collection data as the holotype of R. wudai.

Ropalidia conspicua (Smith) Odynerus conspicuus Smith, 1863: 40, female – “Mysol” Odynerus mysolicus Dalla Torre, 1889: 124. Replacement

name for Odynerus conspicuus Smith, 1863, non de Saussure. [Unnecessary replacement, incorrectly considering de Saussure, 1870, as de Saussure, 1857].

Ropalidia conspicua; Cheesman, 1952: 2, 13 (redescription of the type).

Ropalidia conspicua alleni Cheesman, 1952: 2, 13, female – “Mysol”. — Kojima and Carpenter, 1997: 16 (syn. of R. conspicua, without indicating a new synonymy).

Ropalidia alleni; Richards, 1978: 57 (syn. of R. brunnea (Smith) [this treatment was apparently based on erroneously taking “conspicua alleni” as “cariniscutis alleni”]).

Ropalidia mysolica; Richards, 1978: 57 (in subgenus Polistratus; Odynerus conspicuus as a synonym).

Carpenter (1999: 34) referred to the holotype of Odynerus conspicuus Smith, stating “In the OUM are two females marked as types, labeled with type numbers 097 and 098. The first has no locality label, and the determination label states, “By mistake described/as an Odynerus.” Number 098 is from “New Guinea”, and has a labed [sic] . . . stating, “Not a type!” As the locality is incorrect, number 098 is a pseudotype. Number 097 is therefore apparently the holotype.” I examined both specimens and number 097 is labeled “Holotype O.U.M” (circled with red), “M” [= Mysol] (circular label), “Holotype”, “Icaria conspicuus by mistake described as an Odynerus” (blue label), and “Type 097 Hope Dept. Oxford”, and number 098 is with labels “New Guinea” (circular), “Odynerus conspicuus Smith” (blue), “Icaria”, “Not a type!” and “Type 098 Hope Dept. Oxford”. Carpenter overlooked the locality label for number 097 specimen, which definitely supports his interpretation.

Although Cheesman (1952: 13) stated that the holotype of R. conspicua alleni was “labeled by Smith “Odynerus conspicuus type, Mysol Island”, this speci-men in the BMNH is actually labeled “Type” (BM type label), “S” (circular), “Smith coll. press by Mrs. Farren White 99-303”, “Odynerus conspicuus Smith, New Guinea, Mysool”, “conspicua Sm. ssp. alleni Chees., L. E. Cheesman det. 1950”, and “B.M.Type HYM 18.860”. Cheesman (1952) overlooked the locality label “S” [= Salwatty = Salwati]; NW New Guinea]. Furthermore, the fourth label does not include “type”.

Ropalidia melania Richards Ropalidia melania Richards, 1978: 55 (nest), 57 (list), 65

(larva in key), 122 (key), 126, female, nest (in subgenus Polistratus) – “New Guinea: Morobe District, Wau”.

This species certainly a valid species, distinguished from other species in the R. nigra-group by a wide median impression at the occipital margin as noted in the original description by Richards (1978).

Types examined — NE New Guinea: ♀ (BMNH, holotype), labeled “Holotype” (BM type label), “Nest NG13, New Guinea, Morobe district, Wau, 24.xii.1972, O.W. Richards”, “Ropalidia melania Richards, ♀ , holotype”; 1♀ (paratype, NNM), same collection data as the holotype.

Ropalidia nigra (Smith) Icaria nigra Smith, 1859: 167, female – “Aru”. — Meade-

Waldo and Morley, 1914: 406 (syn.: Ancistrocerus catharinae (Cam.) as var.). — Kojima and Carpenter, 1997: 4 (designation of lectotype).

Icaria morosa Smith, 1863: 42, female – “Waigiou”. — Cheesman, 1952: 10 (syn. of Ropalidia nigra (Smith)).

Icaria spilostoma Cameron, 1906a: 62, female, male – “Manokwari”. — van der Vecht, 1934: viii (syn. of Ropalidia nigra (Smith)). — Cheesman, 1952: 10 (syn. of Ropalidia nigra (Smith)). — Kojima and Carpenter, 1997: 5 (designation of lectotype).

Odynerus sariensis Cameron, 1906a: 63, female – “Siari”. — van der Vecht, 1934: viii (syn. of Ropalidia nigra (Smith)). — Kojima and Carpenter, 1997: 5 (designation of lectotype).

Odynerus confraternus Cameron, 1911: 192, female – “Lorentz River”. — van der Vecht, 1934: viii (syn. of Ropalidia nigra (Smith)). — Kojima and Carpenter, 1997: 5 (designation of lectotype).

Icaria confraternus; Meade-Waldo and Morley, 1914: 506. Ancistrocerus catharinae Cameron, 1913: 78, female –

“Waigeu”. — Meade-Waldo and Morley, 1914: 406 (var. of Icaria nigra). — van der Vecht, 1934: viii (syn. of Ropalidia nigra (Smith)). — Kojima and Carpenter, 1997: 4 (designation of lectotype).

Icaria nigra var. catharinae; Meade-Waldo and Morley, 1914: 406.

Ropalidia confraterna; Bequaert, 1932: 50. Ropalidia nigra; Bequaert, 1932: 50. — van der Vecht, 1934:

viii (syns.: Icaria spilostoma Cameron, Odynerus sariensis Cameron, Odynerus confraternus Cameron, Ancistrocerus catharinae Cameron). — Cheesman, 1952: 2, 10 (redescription of the type female, description of male; syns.: Icaria morosa Smith, Icaria spilostoma Cameron; distribution [erroneously recorded from Makassar in Celebes]). — Richards, 1978: 57 (in subgenus Polistratus; syn.: spilostoma (Cameron)), 121 (key).

Ropalidia spilostoma; Bequaert, 1932: 51. Ropalidia morosa; Bequaert, 1932: 51. Ropalidia domestica Cheesman, 1952: 2, 12, female, male –

“Papua, Kokoda”. NEW SYNONYMY. — Richards,

52 J. Kojima

1978: 54 (nest), 57 (in subgenus Polistratus), 122 (holotype stated to be female; redescription).

Ropalidia sepicana Richards, 1978: 57 (list), 122 (key), 127, figs. 22, 27, female, male (in subgenus Polistratus) – “New Guinea: Sepik district, Sepik River, Kondanggei”. NEW SYNONYMY.

I examined many specimens from various localities in addition to the types of all the six species here treated as synonyms of R. nigra (Smith) and that of Icaria nigra. Despite the fact that some of these taxa can be diagnosed by minor characters when only the type specimens are concerned, additional specimens showed that they are intergraded as given in the second part of this series. Van der Vecht (1934: viii) synonymized Cameron’s four species under R. nigra (smith), and Cheesman (1952: 10) synonymized Smith’s species I. morosa under R. nigra. They have been, since then, treated as the synonyms of R. nigra (Smith). I agree with their treatments. Cheesman (1952: 12), however, described Ropalidia domestica in comparison with R. nigra; besides minor differences in the marking pattern, a distinct difference recognized in her descriptions is “width of 3rd [antennal flagellar article] 1/3 of 9th” (p. 10) for the type of I. nigra and “3rd segment width 2/3 that of 9th” (p. 12) for R. domestica. The types (lectotype and paralectotype) of I. nigra, however, do not have antennae so strongly dilated apically.

Richards (1978: 57) listed only I. spilostoma as a synonym of R. nigra and regarded R. domestica and his species R. sepicana as good species; he did not refer to the other species of Cameron and Smith that had been synonymized by van der Vecht (1934) and Cheesman (1952), respectively. The grounds on which Richards recognized R. domestica and R. sepicana as good species are given in his key to species of the “subgenus Polistratus” (Richards, 1978: 121-122). In the couplet 3, he mentioned that R. nigra has the eyes “bare”, the widened part of the first metasomal tergum “4 times as wide as long” and “half as wide as tergite 2”, and the scutum “entirely dull”. On the other hand, he stated that R. domestica and R. sepicana have their eyes “usually with a few hairs”, and the widened part of their first metasomal tergum is “5 times as wide as long”. However, the types (lectotype and paralectotype) of Icaria nigra Smith have eyes possessing sparse hairs. Furthermore, their first metasomal tergum is wider than half the width of the second tergum. Richards’ measurements of the parts of metasomal segments are so poorly defined that nobody can repeat his measurements.

Types examined — Waigeo: ♀ (OUM, holotype of Icaria morosa Smith), labeled “Wag” [= Waigiou] (circular), “Icaria morosa Smith” and “= nigra Smith, L. E. Cheesman, 1949”; ♀ (ZMA, lectotype Ancistrocerus catharinae Cameron, designated by

Kojima and Carpenter (1997: 4)), labeled “Waigeoe/4. 1. 10”, “Mevr de/Beaufort/Leg” and “Icaria [in van der Vecht’s handwriting]/Ancistrocerus [in Cameron’s handwriting]/Beauforti/Cam. Type [in Cameron’s hand-writing]/Holotype sec. J. v. d. Vecht ’33 [in van der Vecht’s handwriting]”. NW New Guinea: ♀ (BMNH, lectotype of Icaria spilostoma Cameron, designated by Kojima and Carpenter (1997: 5)), labeled “Manokwari, 23 mei 03”, “Icaria spilostoma Cam. Type [in Cameron’s handwriting]”, “B.M. TYPE HYM 18.873a” and “Ropalidia spilostoma (Cam.) Lectotype [in Richards’ hand-writing]”; 2♂ (ZMA, paralectotype of Icaria spilostoma), Manokwari, 2.v.[19]03; 1♂ (ZMA, paralectotype of Icaria spilostoma), Manokwari, 15.v.[19]03; 1 ♀ (ZMA, lectotype of Odynerus sariensis Cameron, designated by Kojima and Carpenter (1997: 5)), labeled “Holotype”, “Sari [= Siari; 1°58’S, 131°58’E]/12. II 03” and “Odynerus /Sariensis/Cam. Type/New Guinea [in Cameron’s handwriting]/Holotype sec. J. v. d. Vecht [in van der Vecht’s handwriting]”. Aru: ♀ (OUM, lectotype of Icaria nigra Smith, designated by Kojima and Carpenter (1997: 4)), labeled “Aru” (circular), “Type” (BM type label), “Icaria nigra Smith”, “Icaria nigra Sm.” and “Lectotype v. d. Vecht desig. 1958”; 1♀ (OUM; Kojima and Carpenter (1997: 4), when they designated the lectotype of Icaria nigra, stated “probably unique”, but this specimen is presumably from the type series and is a paralectotype of I. nigra), labeled “I. Aru” (circular), “Icaria nigra Isl. Aru”, “Smith coll. press by Mrs. Farren White 99-303” and “Icaria nigra Sm”. SW New Guinea: ♀ (ZMA, lectotype of Odynerus confraternus Cameron, designated by Kojima and Carpenter (1997: 5)), labeled “Z. Nieuw Guinea/Lorentz 1909-10/Noord Rivier IX. 09”, “HOLOTYPE”, “Odynerus confraternus Cam. Type [in Cameron’s handwriting; “Odynerus” is lined on it with pencil and “Icaria” in van der Vecht’s handwriting is above it]/HOLOTYPE ♀ SEC J. v. d. Vecht 1933 [in van der Vecht’s handwriting]”. NE New Guinea: ♀ (ANIC, holotype of Ropalidia sepicana Richards), labeled “NEW GUINEA /Kandanggei,/Sepik River/29 Feb. 1964/D. H. Colless” and “Ropalidia / sepicana /♀ Rich. / holotype [in Richards’ handwriting]”; 1♀ (NNM, paratype of R. sepicana), same collection data as the holotype of R. sepicana. SE New Guinea: ♂ (BMNH, holotype of Ropalidia domestica Chessman), labeled “Holotype” (BM type label), “Papua: Kokoda 1200 ft. iv.1933, L. E. Cheesman B.M. 1933-427”, “L. E. Cheesman det. 1949, domestica Chess. ♂”, and “B.M. TYPE HYM 18.1037”; 2♀ (BMNH, paratypes of R. domestica), same collection data as the holotype of R. domestica; 1♀ (BMNH, paratype of R. domestica), same collection data as the holotype of R. domestica but vi-vii.1933.


Ropalidia leopoldi and its related species (or species in the R. flavopicta-group)

Ropalidia leopoldi Bequaert Ropalidia leopoldi Bequaert, 1932: 47, female - “Nouvelle-

Guinée hollandaise. Lac d’Angi-Gita” [= Anggi Gita; 1°23’S, 133°58’E]. — Richards, 1978: 58 (in subgenus Icarielia; list), 129 (key).

Icaria spec.; von Schulthess, 1932: 40 (Nouvelle-Guineé, Lac de Angi-Gita). This species is included in a key to species of

“subgenus Icarielia” by Richards (1978: 129). As discussed in detail in the second part of this series, in New Guinea there could be several species of this species-group. I failed to locate any specimens of R. leopoldi that Richards (1978) used to produce the key.

Types examined — NW New Guinea: ♀ (IRSNB, holotype), labeled “Lac d’Angi-Gita/(Nouvelle Guinée) /10-III-1929/Prince Léopold”, “Ropalidia/leopoldi J. Beq./Holotype [in red]/Dr. J.C.Bequaert det., 1932 [in Bequaert’s (?) handwriting except the printed last line]”, and “cf. Mem. Mu. Hist. Nat. Belg. (hors serie) IV 5 (1932) p.47, fig. 1, A-D”; 40♀ (paratypes, IRSNB; 2♀ in MCZ), same collection data as the holotype.

Ropalidia deminutiva Cheesman Ropalidia deminutiva Cheesman, 1952: 3, 23, female – “S. of

Humboldt Bay, inland 16 miles, Mt. Nomo”. — Richards, 1978: 54 (diminutiva [!]; nest), 58 (in subgenus Icarielia), 129 (key).

Ropalidia integra Cheesman, 1952: 3, 22, female, “male” – “Papua, Kokoda, 1200 ft”. NEW SYNONYMY. — Richards, 1978: 58 (in subgenus Icarielia), 129.

Richards (1978: 129) noted that “All the specimens [of deminutiva] . . . labelled Bougainville, Mt. Nomo. I cannot trace Mt Nomo, but I suspect that Bougainville is an error, possibly for Mt. Bougainville . . . in West Irian.” I examined the holotype and a paratype of R. deminutiva in the BMNH: they have the labels “W. New Guinea: Mt. Nomo [2°42’S, 139°59’E; NW New Guinea], S. of Mt. Bougainville, 700 ft. ii.1936” and “L. E. Cheesman B.M. 1936-271”; the holotype is also with “Type” (BM type label), “Ropalidia diminutiva TYPE [in Cheesman’s (?) handwriting] Det. L. E. Cheesman 195”, “B.M. TYPE HYM 18.1034”; and the paratype is with additional labels “cotype”, “Ropalidia deminutiva Chees. Det. L. E. Cheesman”. Richards (1978) misread the locality label.

Richards (1978: 129) referred to the types of R. integra Cheesman and stated that “The specimen in the BMNH labeled type is a female from Kokoda [SE New Guinea] but is, I think, a specimen of R. deminutiva Cheesman. The paratype, though labelled male, is actually a female without a gaster, also coming from

Kokoda. It is a very different species and agrees better with the description. I have transferred the type label to this specimen”. However, the type label was not transferred, but Richards merely added his own labels to the specimens: one of the two females is labeled “Type” (BM type label), “Papua, Kokoda, 1200 ft. IX. 1933, L. E. Cheesman, B.M. 1934-321”, “L. E. Cheesman det. 1948, integra Chees.”, “B.M. TYPE HYM 18.1035” “BM ‘79” (red), “This is not type of integra OWR, 1974 [in Richards’ handwriting]”; and the other is with “Paratype” (circled with yellow), “Papua: Kokoda, 1200 ft. viii.1933., L. E. Cheesman, B.M. 1933-427”, “L. E. Cheesman det. 1948, integra Chees.”, “♂”, “This is the real type of integra OWR ‘74, ♀”. There is no grounds to accept Richards’ interpretation even if the specimen labeled “Paratype” were to agree better with the original description than the specimen labeled “Type” does. Only one fact that may support Richards’ interpretation is that Cheesman (1952: 23) listed the material as “1♀ (holotype), 1♂ (paratype) . . . viii., ix. 1933”. If this means that “♀ and ♂ was collected in August and September, respectively”, the holotype must be labeled “viii. 1933”. The specimen labeled “paratype” was collected in “viii. 1933” and that labeled “type” was collected in “ix.1933”. However, judging from the collection data given for other species, it is apparent that Cheesman (1952) listed the dates of collection chronologically.

The holotype of R. integra should be the specimen collected in “ix.1933” and a specimen labeled “♂” is the paratype. The holotype agrees well in structure with the holotype of R. deminutiva Cheesman, and R. integra is synonymized under R. deminutiva. The paratype of R. integra is not, as Richards pointed out, conspecific with the holotype of R. integra, but its definitive affiliation could not be determined because its metasoma is missing. Richards’ (1978: 128) “R. integra” in his key to species of “subgenus Icarielia” was based on the paratype of R. integra, which lacks the metasoma.

Species in the R. lefebvrei-group

Ropalidia lefebvrei (Le Guillou) Polistes lefebvrei Le Guillou, 1841a: 325, female – “Triton-

Bay”; 1841b: 322. ?Icaria lefebvrei; de Saussure, 1853-58: 241 (unidentified

species). Icaria lefebvrei; Smith, 1857: 99 [Kojima and Carpenter

(1997: 33) listed the pagination as “95” by error]. Ropalidia lefebvrei; Bequaert, 1932: 50. ? Icaria reactionalis de Saussure, 1853, in de Saussure, 1853-

58: 28, female – “La Nouvelle-Guinée” (coll. Guérin-Méneville, current depository unknown).

? Ropalidia reactionalis; Bequaert, 1932: 51. — Cheesman, 1952: 3, 22 (group of festina; female).

54 J. Kojima

Le Guillou published the name Polistes lefebvrei twice in 1841. Although the exact dates of these two publications are not known, I take Le Guillou (1841a) as the original description of P. lefebvrei; in Le Guillou (1841b: 322) the author name of P. lefebvrei is clearly indicated as “Le Guillou” (see also Kojima and Carpenter, 1997: 33; Carpenter, 1999: 36). In the MNHN, there is a female labeled “MUSEUM PARIS/Triton bay [3°45’S, 134°10’E; NW New Guinea]/LE GUILLOU”, “114”, “Type”, “314 - Polistes ne 18/Lefebvrei - Leg./Triton bay:” and “Ann. Soc. Ent. 1841/n. 322/no. 26/par le 8 Le Guillou”, that is evidently the holotype of P. lefebvrei (see also Carpenter, 1999: 36).

De Saussure (1853-58: 241) referred to Icaria lefebvrei (Le Guillou) as an unidentified species, and this is the only citation of I. lefebvrei by de Saussure. Thus, de Saussure should not have seen the type of Polistes lefebvrei. Du Buysson (1911: 229) referred to the holotype of Polistes lefebvrei, but he never cited Icaria reactionalis de Saussure, 1853. Although Bequaert (1932: 50 and 51) cited both species simply listing them as species so far recorded from New Guinea, it seems highly probable that he did not see the types of either taxa. Cheesman (1952: 21) cited I. reactionalis when she redescribed Icaria festina Smith, 1864, as a subspecies of Ropalidia reactionalis (de Saussure), but she did not examine the type(s) of I. reactionalis. A female in the MHN labeled with a blue circle with “voyage/d’Cap/d’Urville [probably in de Saussure’s handwriting]”, “Ropalidia/spec. [in van der Vecht’s handwriting]/det. J. v. d. Vecht 1967” and “Ropalidia lefebvrei /comp. w / HOLOTYPE / det. J. Kojima 1999” is R. lefebvrei and agrees, in general, with the original description of Icaria reactionalis. This specimen, however, has the scutum colored ferruginous with a pair of yellow longitudinal bands and the propodeum is entirely yellow, while de Saussure (1853-58: 28) stated for I. reactionalis that “Corselet jaune; métathorax [= propodeum] ferrugineux, avec deux lignes jaunes longitudinales”. In the original description of I. reactionalis, the body parts are referred to in an unusual order; that is, de Saussure (1853-58: 28) mentioned “Corselet . . . métathorax [= propo-deum] . . . écusson [= scutellum]”. This fact may suggest that “métathorax” in the description of I. reactionalis could be an erroneous writing in the manuscript or erroneous typesetting for “mésothorax”. Even if this interpretation is correct, this specimen in the MHN could not be regarded as the type of I. reactionalis. De Saussure (1853-58: 29) mentioned “La Nouvelle-Guinée” as the locality and “Collection de M. Guérin-Méneville” as the depository: the specimen does not have a locality label and hymenopteran specimens in the collection of Guérin-

Méneville were not deposited in the MHN (see Horn et al., 1990). I failed to locate the type(s) of I. reactionalis.

Ropalidia gemmea Cheesman Ropalidia gemmea Cheesman, 1952: 3, 22, female, male –

“Papua, Owen Stanley Mts., Kokoda, 1200 ft.” — Richards, 1978: 58 (in subgenus Icarielia), 129 (key).

I examined: the holotype (in the BMNH) labeled “Papua: Kokoda [SE New Guinea], 1200 ft., ix.1933, L. E. Cheesman, BM 1934-321” “L. E. Cheesman det. 1950, TYPE, gemmea Chees.” and “B.M. TYPE HYM 18.1032”; a paratype female (BMNH) from the same locality as the holotype; and a headless paratype (NNM) labeled “Para-/type” (BM type label), “PAPUA: Kokoda. /1,200ft. vi.1933/L. E. Cheesman. /B.M.1933-427”, “Museum Leiden/ex. coll. J. v. d. Vecht”, “L. E. Cheesman det. 1950/gemmea Chees”. They are presumably conspecific, and I concluded that R. gemmea is a valid species.

Ropalidia catharinae (Cameron) Icaria catharinae Cameron, 1913: 77, female – “Waigeu”. —

Meade-Waldo and Morley, 1914: 406. — van der Vecht, 1934: viii.

? Ropalidia catharinae; Bequaert, 1932: 50. ?Ropalidia fluviatilis petulans Cheesman, 1952: 18, female –

“Japen Is.”

Cameron (1913) described two “catharinae” of New Guinean Ropalidia, respectively in Icaria and Ancistrocerus. Meade-Waldo and Morley (1914: 406) treated Ancistrocerus catharinae as a variety of Icaria nigra Smith, and stated “All the species published with the specific name “catharinae” in Cameron’s paper . . . bear labels with the specific name “beauforti” on the type-specimens”, without referring to the taxonomic status of Icaria catharinae. Bequaert (1932) listed “R. catharinae (Cameron, 1913)” as a species distributed in New Guinea, but he did not indicate which “catharinae” he referred to. Van der Vecht (1934: viii) stated “Icaria catharinae . . . is niet synoniem met . . . Ancistrocerus catharinae . . . ; of deze sort reeds vroeger beschreven was, hoopt Spr. nog nader te onderzoeken” [Icaria catharinae is not a synonym of Ancistrocerus catharinae; the speaker [= the late van der Vecht] wish to make further investigation if this species was already described previously].

In the ZMA is a female marked as “Holotype” [red label, detached metasoma glued], with labels “Waigeoe [= Waigeo] 29 XII 09”, “Mevr. de Beaufort leg” and “Icaria/Catharinae/Beauforti/Cam. type/HoloTYPE sec. J. vdV. ’33 [in Cameron’s handwriting, but “Catharinae” and the last line are in van der Vecht’s handwriting]”, that is evidently the holotype of Icaria


catharinae. I examined the types of almost all taxa described from New Guinea and its adjacent islands, and I concluded that R. catharinae (= Icaria catharinae) is a valid species.

Cheesman (1952: 18) described R. fluviatilis petulans based on three females from Japen Island. In the drawer in the BMNH that contains the two paratypes of R. fluviatilis petulans is a label “Type missing”; I searched the holotype in vain. The both paratypes are labeled “Dutch New Guinea, Japan Is. Mt. Baduri 1000 ft. viii 1938 L. E. Cheesman BM. 1938-593” and “L. E. Cheesman det. 1949 fluviatilis M. W. ssp. petulans Chees”. These two specimens apparently belong to different species; one agrees well with the holotype of Icaria catharinae, and the other with the holotype of Polistes albobalteatus Cameron, 1906. Unless the holotype of R. fluviatilis petulans is eventually found, the exact taxonomic status of this taxon cannot be determined.

Ropalidia festina (Smith) Icaria festina Smith, 1865: 90, female – “New Guinea”. Icaria zonata Cameron, 1906a: 61, female – “Moaif”. —

Meade-Waldo, 1911: 100 (syn. of I. festina). — van der Vecht, 1934: viii (syn. of R. festina (Smith)). — Kojima and Carpenter, 1997: 5 (designation of lectotype).

Ropalidia festina; Bequaert, 1932: 51. — van der Vecht, 1934: viii.

Ropalidia reactionalis festina; Cheesman, 1952: 3, 21 (redescription of the type).

Ropalidia zonata; Cheesman, 1952: 3, 22. — Richards, 1978: 58 (in subgenus Icarielia), 129 (key).

? Ropalidia festina; Richards, 1978: 58 (in subgenus Icarielia; list), 129 (key).

Although Smith (1865) did not give any indication for the number of specimens, a female in the OUM labeled “Holotype” (BM type label), “N” [= New Guinea] (circular) and “Icaria festina Smith” is presumably the holotype of Icaria festina Smith, 1865. I compared this specimen with the lectotype and paralectotypes of Icaria zonata Cameron, 1906 (the lectotype designated by Kojima and Carpenter (1997: 5) is in the ZMA and labeled “Holotype”, “Moaif [prob. NW New Guinea]/26 VI - 4 VII”, “Icaria /zonata/Cam. Type/New Guinea [in Cameron’s handwriting]/ ♀ HOLOTYPE/Sec. J. v. d. Vecht 33 [in van der Vecht’s handwriting]” and “Ropalidia zonata ♀ (Cam.) det J. v. d. Vecht 1974”), and I confirmed the synonymy of I. zonata under R. festina proposed by Meade-Waldo (1911: 100) and van der Vecht (1934: viii).

Richards (1978) treated both species as valid species and included them in his key to species of the “subgenus Icarielia” (p. 129), stating that the first metasomal tergum is “about as long as mid femur” in R. festina and “clearly shorter than mid femur, wider” in R. zonata. However, I wonder if Richards examined

the types of Icaria festina and I. zonata. The late van der Vecht distributed at least to the BMNH the specimens of an undescribed species (this will be treated in the second part of this series) with his identification label as “R. festina”; Richards could use those specimens when he produced the key.

Ropalidia albobalteata (Cameron) Polistes albobalteatus Cameron, 1906a: 61, female –

“Manokwari”. — van der Vecht, 1934: viii (syn. of R. festina (Smith)).

? Icaria fluviatilis Meade-Waldo, 1912: 449 (key), 451, female (in subgenus Icariola) – “Mimika River”.

Polybia papuana Cameron, 1913: 77, female – “Waigeu”. REVISED SYNONYMY. — van der Vecht, 1934: viii (syn. of R. festina (Smith)); 1966: 6. — Kojima and Carpenter, 1997: 6 (designation of lectotype).

? Ropalidia fluviatilis; Bequaert, 1932: 51. — Cheesman, 1952: 2, 17 (redescription; distr.), 26 (fluvialis [!]). — Richards, 1978: 58 (in subgenus Icariola; list).

? Ropalidia fluviatilis petulans Cheesman, 1952: 2 (petulaus [!]), 18, female – “Japen Is., Mt. Baduri, 1000 ft.”

Ropalidia papuana; van der Vecht, 1966: 6. ? Ropalidia petulans; Richards, 1978: 58 (in subgenus

Icariola). Ropalidia albobalteata: Kojima and van Achterberg, 1997:

166 (“they [albobalteata and festina] differ at species level”).

Cameron (1906a: 61) stated “3. Polistes albobalteatus sp. nov. Manokwari, February. One example”, thus a female in the ZMA labeled “Mano-kwari [NW New Guinea]/14-28 II 03”, “Polistes /albobalteatus/Cam. Type/New Guinea [in Cameron’s handwriting] /♀Holotype sec. J. v. d. Vecht ‘33 [in van der Vecht’s handwriting]”, “Ropalidia/festina (Sm.)/var. albobalteata (Cam.)/det. J. v. d. Vecht 34 [in van der Vecht’s handwriting]” and “Holotype” is certainly the holotype of P. albobalteatus. While van der Vecht (1934: viii) synonymized P. albobalteatus under R. festina (Smith), Kojima and van Achterberg (1997: 166) noted that they are different species. As detailed in the second part of this series, comparison of the holotypes of I. festina and P. albobalteatus together with the examination of additional specimens led me to conclude that they are different species.

Van der Vecht (1934: viii) synonymized Polybia papuana Cameron, 1913, under R. festina, but later he (1966: 6) transferred it to the genus Ropalidia and treated it as a valid species. I did not find any specific differences between the holotype of P. albobalteatus and the lectotype of P. papuana (ZMA, designated by Kojima and Carpenter (1997: 6), and labeled “Holotype”, “Waigeoe [= Waigeo]/25 XII 09”, “Mevr. de Beaufort leg” and “Polybia/papuana/Cam. Type [in Cameron’s handwriting]/Holotype ♀ sec J. v. d. Vecht

56 J. Kojima

33 [in van der Vecht’s handwriting]”) and propose to synonymize P. papuana under R. albobalteata.

Meade-Waldo (1912: 452) described Icaria (Icariola) fluviatilis based on a single female from “Mimika River”. The drawer in the BMNH, in which the holotype should be, contains a note “headless type loaned to v.d. Vecht”. When I visited the NNM in 1999, I located the holotype of I. fluviatilis among the specimens brought from the late van der Vecht’s house after his death [the specimen is tentatively kept at the IUNH and I will return it to the BMNH on my responsibility]; the specimen is headless and labeled “Type” [BM type label]; “New Guinea./Mimika R./A. F. R. Wollaston. /1911-229./aug 1910” [in print but last line in handwriting]; “B. M. TYPE/HYM./18.859”; “BM ’79”, and “Icaria/(Icariola) /fluviatilis sp. n./Type /G. Meade-Waldo det”. Although I could not find any differences between the holotypes of Polistes albobalte-atus and Icaria fluviatilis to consider them as different species, headlessness of the I. fluviatilis holotype prevents me from making a definitive conclusion.

The BMNH houses two females identified as “fluviatilis” by Cheesman; they could be the specimens referred to in Cheesman (1952: 17), and one is from Kokoda and the other is from Humboldt Bay. While the specimen from Kokoda and one of the paratypes of R. fluviatilis petulans, as noted before, are conspecific with the holotype of P. albobalteatus, the specimen from Humboldt Bay seems to me to be R. gemmea (because of the orientation of the head, I could not determine its definite specific affiliation).

Ropalidia bensoni Richards Ropalidia bensoni Richards, 1978: 55 (nest), 59 (list), 65

(larva in key), 129 (key), 130, female, larva, nest (in subgenus Icarielia) – “New Guinea: Morobe district, Wau”.

This species is undoubtedly a valid species. Types examined — NE New Guinea: ♀ (BMNH,

holotype), labeled “Holotype” (BM type label), “Nest NG 3, New Guinea, Morobe district, Wau, 12-16. Dec. 1972, O. W. Richards”, “Ropalidia bensoni Richards, ♀, Holotype [in Richards’ handwriting]”; 1♀ (NNM, paratype), same collection data as the holotype.

Species in the R. impetuosa-group Ropalidia impetuosa (Smith)

Icaria impetuosa Smith, 1860: 131, female – “Batchian; Amboyna”. — van der Vecht, 1941a: 104 (syn. of R. gregaria (de Saussure)). — Kojima and Carpenter, 1997: 4 (designation of lectotype).

Icaria spilocephala Cameron, 1906b: 230, female – “Etna Bay”. — van der Vecht, 1941a: 104 (syn. of R. gregaria

(de Saussure)). — Kojima and Carpenter, 1997: 5 (designation of lectotype).

Icaria cohni du Buysson, 1909b: 306, male – “Bougainville”. — van der Vecht, 1941a: 104 (syn. of R. gregaria (de Saussure)). — Richards, 1978: 86 (kohni [!]; syn. of R. gregaria spilocephala (Cameron)).

Ropalidia impetuosa; Bequaert, 1932: 51. — Richards, 1978: 58 (in subgenus Icariola), 86 (possible syn.: Ropalidia gregaria spilocephala (Cameron)).

Ropalidia gregaria; van der Vecht, 1941a: 104 (syn.: Icaria cohni du Buysson, I. impetuosa Smith, I. spilocephala Cameron), 111 (key), 149 (distribution [including subspecies spilocephala]). — Cheesman, 1952: 3, 20 (male). — Spradbery, 1991: 349 (colony population).

Ropalidia cohni; Bequaert, 1932: 51. Ropalidia spilocephala; Bequaert, 1932: 51. Ropalidia gregaria inquieta Cheesman, 1952: 3, 20, female,

male – “Papua, . . . Mafulu, 4000 ft.” — Richards, 1978: 86 (designation of lectotype [error in locality: Mafalu]; syn. of R. gregaria spilocephala).

Ropalidia gregaria var. tolerans Cheesman, 1952: 3 (as subspecies), 21 (as var.), female – “Cyclops Mts. Sabron”. — Richards, 1978: 86 (designation of lectotype; syn. of R. gregaria spilocephala).

Ropalidia gregaria gregaria; Richards, 1978: 53 (nest), 62 (key), 86 (in subgenus Icariola). — Spradbery, 1991: 351 (colony population).

Ropalidia gregaria spilocephala; Richards, 1978: 53, 62 (key), 86 (syns.: Icaria kohni [!] du Buysson, Ropalidia gregaria inquieta Cheesman, Ropalidia gregaria var. tolerans Cheesman; possibly syn. of Icaria impetuosa Smith; “I examined the holotype male [!] in the Zoological Museum, Amsterdam.”). — Spradbery and Kojima, 1989: 636 (nest; colony population). — Spradbery, 1991: 351 (colony population).

During my recent research in the MHN, I located the types of Icaria gregaria de Saussure, 1854. They are different species of “Ropalidia gregaria” of authors, and I proposed to apply R. impetuosa (Smith) to “R. gregaria” of authors (see Kojima, 2001: 5).

Specimens examined — Halmahera: 3♀ (NNM, 1♀ in IUNH), between Gita & Payahe, Tayawi, 4.iii.1995, R. de Vries; 4♀ (NNM), nr. Payahe, Tayawi, R. de Vries (2♀ : 10.iii.1995; 2♀ : 14.iii.1995); 1♀ (NNM), road Payahe-Weda, E. of Payahe, 15.iii.1995, R. de Vries. Bacan: ♀ (OUM, lectotype of Icaria impetuosa Smith, designated by Kojima and Carpenter (1997: 4)), labeled “Bac” and “Lectotype v. d. Vecht des. 1958”. Buru: 1♀ (ZMA), Station 5, iv.[19]21, L. J. Toxopeus; 1♀ (ZMA), Station 9, 18.v.1921, L. J. Txopeus. Ambon: 1♀ (OUM, paralectotype of I. impetuosa), labeled “Amb”; 1♀ (NNM), 10 km NE Pago, 200 m alt. 3°38’S, 128°16’E, 21.iii.1988, R. Hensen; 2♀1♂ (NNM), Ambon City, 200 m alt., 3°41’S, 128°10’E, 20.iii.1988, R. Hensen. Seram: 1♀ (NNM), 15 km NNE of Tehoru, Hatumete, 50 m alt., 3°17’S, 129°36’E, 24.iii.1988, R. Hensen. NW New Guinea: ♀ (ZMA, lectotype of Icaria spilocephala


Cameron, designated by Kojima and Carpenter (1997: 5)), labeled “Nieuw Guinea/Expeditie 1904/[slash]5 /Etna baai”, “Icaria / spilocephala /Cam. Type / New Guinea [in Cameron’s handwriting] /♀ Holotype sec. J. v. d. Vecht 1933 [in van der Vecht’s handwriting]” and “Holotype”; 1♀ (ZMA), Baliemvalley, 5 km W. Wamena, 1,500 m, 23.x.1993, A. J. de Boer, A. L. M. Rutten & R. de Vos; ♀ (BMNH, lectotype of Ropalidia gregaria inquieta Chessman, designated by Richards (1978: 86)), labeled “Lectotype” (BM type label), “Dutch New Guinea, Cyclops Mts., Sabron, 930 ft., v.1936, L. E. Cheesman, B.M. 1936-271”, “Ropalidia gregaria var. tolerans det. L. E. Cheesman”, “Ropalidia gregaria (Sauss.) var. tolerans Cheesm. Lectotype”. NE New Guinea: 3♀3♂ (BMNH), Madang Dist., Finisterre Mts., Damanti, 3,550 ft., 2-11.x.1964, M. E. Bacchus; 2♀ (BMNH), Madang Dist., Finisterre Mts., Moro C., 5,550 ft., 30.x.-15.xi.1964, M. E. Bacchus; 5♀ (CAS), Madang Province, Bundi, 5°45’S, 145°15’E, 10-12.iii.1987, W. J. Pulawski; 8♀ (CAS), Madang Province, Simbai, 5°17’S, 144°26’E, W. J. Pulawski; (6♀: 25-26.iii.1987; 1♀: 27.v.1988; 1♀: 31.v.1988); 10♀ 1♂ (CAS), Madang Province, Tep Tep, 5°55’S, 146°30’E, 1,900 m, M. Wasbauer (6♀: 27.xi.1987; 4♀1♂ : 29.xi.1987); 11♀ (CAS), Madang Province, Karisokora, 7.5 air km W. Bundi, 5°44’S, 145°10’E, ca. 1,800 m, 17.v.1988, E. Koima & P. Yomane; 2♀ (CAS), Madang Province, Brahman Catholic Mission, 6°45’S, 145°23’E, 21.x.1987, M. Wasbauer; 1 ♀ (SAM), Wareo, L. Wagner; 10 ♀ (SAM), Finsch Haven, L. Wagner; 1♀ (SAM), Komba, L. Wagner; 2♀2♂ (IUNH), foot of Mt. Missim, Wau, Morobe, 11.vii.1985, J. & K. Kojima; 1♀2♂ (IUNH), Wau, 23.xii.1973, T. Okazawa; 5♀ 4♂ (IUNH), Bulolo Watut, 21.xii.1973, T. Okazawa; 1 ♀ (IUNH), Kundiawa, 27.xii.1973, T. Okazawa. SE New Guinea: ♂ , (BMNH, lectotype of Ropalidia gregaria var. tolerans Cheesman, designated by Richards (1978: 86)), labeled “Lectotype” (BM type label), “Papua, Mafulu, 4000 ft. xii.1933, L. E. Cheesman, B.M. 1934-321”, “ ♂ ’, “gregaria Sauss. inquieta Ch. L. E. Cheesman det. 195”, “Ropalidia gregaria (Sauss.) spp. inquieta Cheesm. ♂ lectotype”; 2♀ (IUNH), Waitape, 20.i.1974, T. Okazawa; 7♀1♂ (SAM), Mt. Lamington, 1,300-1,500 feet, C. T. McNamara; 1♂ (BMNH), Laloki, 6.xii.1980, J. W. Ismay; 6♀ (CAS), Port Moresby, W. J. Pulawski (2♀: 1-2.iv.1987; 1♀: 29-30.iii.1987; 3♀: 22-23.vi.1988); 1♀1♂ (CAS), Central Province, Lake Iaraguma, 20 km NW Port Moresby, 21.vi.1988, W. J. Pulawski. New Ireland: 1 ♀ (BMNH), Nama Kainai, xi.1934, J. L. Froggatt. Bougainville: ♂ (MNHN, lectotype of Icaria cohni du Buysson, designated by Kojima (2001: 5)), labeled “MUSEUM PARIS/I. Bougainville/(leg. Cohn)/Alfken 1909”, “TYPE”, “Icaria/Cohni Buyss. /types/R. DU

BUYSSON det 1909”; 1♂ (MNHN, paralectotype of I. cohni); 2♀ (SAM), A. H. Voyce. Solomon Islands: 1♀ (CAS), Tenaru River, Guadalcanal, i.[19]45, G. E. Bohart; 1♀ (NNM), Guadalcanal, Lunga Estate, on nest, 9.vii.1933, H. T. Pagden; 9♀1♂ (NNM), Tulgi, H. T. Pagden (3♀, nest on Hibiscus, 1.vii.1933; 1♂, 2.ix.1933; 3♀, 23.vii.1934; 3♀, on nest, 16.ix.1934); 1♀ (ZMA), Guadalcanal, Honiara, xi.1967.

Species in the R. marginata-group Ropalidia marginata jucunda (Cameron)

Icaria jucunda Cameron, 1898: 46, female – “New Guinea”. Icaria pruinosa Cameron, 1906b: 228, female – “Digoel”. —

van der Vecht, 1941a: 104 (syn. of R. marginata jucunda). —Kojima and Carpenter, 1997: 5 (designation of lectotype).

Ropalidia jucunda; Bequaert, 1932: 50. Ropalidia pruinosa; Bequaert, 1932: 51. Ropalidia marginata jucunda; van der Vecht, 1941a: 104, 124. Ropalidia canaria Cheesman, 1952: 3, 19, male - “Papua,

Kokoda”. NEW SYNONYMY. — Richards, 1978: 58 (in subgenus Icariola).

In the latest citations, four subspecies are recognized in R. marginata (Lepeletier, 1836), based primarily on color pattern: nominate subspecies, jucunda (Cameron, 1898), rufitarsis van der Vecht, 1941, and sundaica van der Vecht, 1941 (see Kojima and Carpenter, 1997). Although such differences in color pattern could be intergraded among marginata, jucunda and sundaica, I leave at this moment them as subspecies of R. marginata. Their taxonomic status (color variation or diagnosable species) can be discussed only after a detailed comparative study of the R. marginata-group, which includes also R. spatulata van der Vecht, 1962, R. brevita Das and Gupta, 1989, R. laticincta laticincta van der Vecht, 1962, R. laticincta floresiana van der Vecht, 1962, and R. clavata (de Saussure, 1853) [= Icaria duchaaussoyi Gribodo, 1896; see Kojima, 2001: 5].

I examined the types of Icaria jucunda Cameron, I. pruinosa Cameron, and R. carinata Cheesman, and concluded that they all belong to the same taxa.

Specimens examined — New Guinea: ♀ (OUM, holotype of Icaria jucunda Cameron), labeled “Icaria jucunda Cam. Type, New Guinea [Cameron’s handwriting]”, “Ropalidia ferruginea (F.) ♀ var. jucunda (Cam.), det. J. v. d. Vecht”, “12”. SW New Guinea: ♀ (ZMA, lectotype of Icaria pruinosa Cameron, designated by Kojima and Carpenter (1997: 5)), labeled “Nieuw Guinea/Expeditie 1904/Digoel”, “Icaria/pruinosa/Cam. Type/New Guinea [in Cameron’s handwriting] /Holotype sec. J. v. d. Vecht 1933 [in van der Vecht’s handwriting], and “Ropalidia ferruginea (F.)/var. jucunda (Cam.)/det. J. v. d. Vecht”; 2♀

58 J. Kojima

(NNM), Neth. N. G. Exp. 1959, Tanah Merah, 17 m alt., 5.ix.1959. NE New Guinea: 33♀ (IUNH, 1♀ in NNM), Kundiawa, 27.xii.1973, T. Okazawa. SE New Guinea: ♂ (BMNH, holotype of Ropalidia canaria Cheesman), labeled “Type” (BM type label), “Papua: Kokoda 1200 ft. vii.1933, L. E. Cheesman B.M. 1933-427”, “♂ ”, “L. E. Cheesman det. 1948, canaria Chess.”, “B.M. TYPE HYM 18.840”, and “R. marginata jucunda (Cam.) det. J. v. d. Vecht ‘79”; 1♀1♂ (NNM), Port Moresby. 1889, L. Loria; 2♀ 2♂ (CAS), Port Moresby, W. J. Pulawski (1♀1♂ , 29-30.iii.1987; 1♀1♂ , 1-2.iv.1987); 2♀ (NNM), Port Moresby, Boroko, 5.ix.1959, J. I. H. Sient Ivany; 9♀2♂ (NNM), Kapakapa, 1891, L. Loria; 2♀ (NNM), Popondetta, 8.ii.1962, A. Catley; 1♀ (NNM), Central District, Rouna Falls, 24.vi.1973, J. P. Spradbery.

Species in the R. stigma-group Ropalidia mathematica torrida (Smith)

Icaria torrida Smith, 1863: 42, female – “Ceram”. Ropalidia torrida; Bequaert, 1932: 51. Ropalidia mathematica torrida; van der Vecht, 1941a: 104,

133 (distr.); 1962: 20 (in subgenus Anthreneida; distr.; “Perhaps this form will eventually prove to be closer to R. socialis or unicolor than to R. mathematica.”). The R. stigma-group is distributed from India to

Australia, but no species in this species-group has been known from New Guinea mainland. This species-group includes torrida and unicolor, and still awaits an intensive taxonomic study. In the present paper, the taxonomic status of torrida and unicolor follows those in the latest citations. Other taxa included in the R. stigma-group are (their taxonomic status is according to the latest citations; see Kojima and Carpenter, 1997): R. artifex artifex (Smith, 1854), R. a. fuscata van der Vecht, 1941, R. darwini Richards, 1978, R. elegantula Richards, 1978, R. hongkongensis hongkongensis (de Saussure, 1954), R. h. juncta van der Vecht, 1941, R. mathematica mathematica (Smith, 1860), R. m. binotata van der Vecht, 1941, R. m. nigroplagiata (Cameron, 1900), R. m. sumbaensis van der Vecht, 1962, R. nigrita Das and Gupta, 1989, R. rufocollaris rufocollaris (Cameron, 1900), R. r. atrata, van der Vecht, 1941, R. santoshae Das and Gupta, 1989, R. socialis socialis (de Saussure, 1862), R. s. trimaculata van der Vecht, 1962, R. stigma stigma (Smith, 1858), R. s. nigrolineata van der Vecht, 1962, R. s. rufa van der Vecht, 1941, and R. thailandia Gusenleitner, 1994.

Types examined — Seram: 2♀ (OUM, lectotype and paralectotype), labeled as given in the section “Designation of the lectotypes”.

Ropalidia unicolor (Smith)

Icaria unicolor Smith, 1859: 168, female – “Key Island” [= Kai].

Ropalidia unicolor; Bequaert, 1932: 51 (“probablement un Parapolybia”). — van der Vecht, 1962 (in subgenus Anthreneida) — Richards, 1978: 58 (in subgenus Icariola).

Ropalidia mathematica unicolor; van der Vecht, 1941a: 133.

In the OUM is a female labeled “Kei” (circular), “Icaria unicolor Smith” (blue label), “Icaria unicolor Sm.”, that is presumably the holotype of Icaria unicolor Smith, as noted by van der Vecht (1941a: 133).

Species in the R. gracilis-group Ropalidia gracilis (Smith)

Icaria gracilis Smith, 1859: 167, female – “Aru”. — Dalla Torre, 1904: 73 [error: year of publication, 1868].

Icaria deceptor Smith, 1863: 42, female – “Mysol”. Ropalidia gracilis; Bequaert, 1932: 50 (gracillis [!];

“probablement un Parapolybia”). — Cheesman, 1952: 3, 18 (syn.: Icaria deceptor Smith, 1863; redescription of type). — Richards, 1978: 58, 99 (in subgenus Icariola).

Ropalidia deceptor; Bequaert, 1932: 50. — Cheesman, 1952: 18 (syn. of Ropalidia gracilis (Smith, 1859)) — Richards, 1978: 53 (nest), 58 (list), 61 (key), 99, figs. 20, 34 (in subgenus Icariola).

Smith (1859: 167) described Icaria gracilis from Aru, but he did not refer to it when he later (1863: 42) described Icaria deceptor from “Mysol” (= Misool). Cheesman (1952: 18) synonymized Icaria deceptor under Ropalidia gracilis; she listed the holotype of only I. gracilis, and thus it is not clear whether her synonymy was based on the comparison of the types of I. gracilis and I. deceptor. Richards (1978: 99) examined the types of both I. gracilis and I. deceptor and treated them as valid species. I examined the types of I. gracilis and I. deceptor; both are presumably the unique original specimens (the holotypes). The holotype of I. gracilis differs from the holotype of I. deceptor in the following characters as Richards (1978: 99) noted: punctuation more closely and stronger; posterior widened part of first metasomal tergum in dorsal view more parallel-sided (Fig. 43 vs. 45); second metasomal segment proportionately longer (Figs. 42 and 43 vs. 44). The shape of the basal parallel-sided part of the first metasomal tergum is, however, not so different as Richards noted. Additional specimens from several localities in New Guinea show that the shape of the first metasomal tergum is so variable (Figs. 46–52) that the differences between the holotypes of I. gracilis and I. deceptor are intergraded. The punctuation is also so variable that this character is barely used to distinguish the two taxa. Furthermore, the marking pattern does not diagnose them. I follow Chessman’s (1952) synonymy.


In Australia, R. gracilis and its closely related species, R. gracilenta Richards, 1978, are distributed; the former is distributed in northern part of Australia’s Queensland and the latter from northern part of Queensland to northern part of New South Wales (Kojima, 1999b). The characters shared by these two species are the gena proportionately wider (in terms of

the gena width/eye width ratio) in the male than in the female (Figs. 53, 54; see also Kojima, 1987) and the pronotum with ventral corner obliquely cut off (Figs. 55, 56). Similar states in both characters are also found in the R. rufoplagiata-group (Fig. 57; see also Kojima, 1999a). Richards (1978: 61) stated that R. gracilenta could be distinguished from R. gracilis [as R. deceptor] by the presence of a strong angular projection on the

dorsal surface of the hind coax in R. gracilenta. I (Kojima, 1999a) found that the states of this character are variable in R. gracilenta, while the condition of the propodeum is more reliable to distinguish the two species: posterior face of propodeum distinctly and widely excavated and in lateral view slightly convex in R. gracilenta, while only shallowly depressed medially

and in profile slightly concave or nearly straight in R. gracilis (Fig. 55).

Figs. 42–64. Ropalidia gracilis (42–56), R. cristata (57–60), R. turneri (61–63) and R. maculiventris (64). 42–52, 54–64,

female; 53, male. 42–43, holotype of Icaria gracilis; 44–45, holotype of I. deceptor. 42–44, 48, first two metasomal segments in lateral (42, 44, 48) and dorsal (43) views; 45–47, 49–52, 60, 63, first metasomal segment in lateral view; 53–54, 59, 62, head in lateral view; 55, mesosoma in lateral view; 56–57, 64, ventral part of pronotum in lateral view; 58, 61, ocellar area. Scale lines: 2 mm, but 1 mm for Figs. 56–57, 60, 63–64, and 0.5 mm for Figs. 58 and 61.

Specimens of R. gracilis examined — New Guinea: 1♀ (OUM), labeled “N” [= New Guinea] (circular), “Icaria gracilis Smith” (blue), and “not a type”; 1♀ (NNM), labeled “N”, “compared with type” and “R. deceptor”; 1♀ (BMNH), labeled “N” and “Icaria gracilis Smith”; 1♀ (BMNH), New Guinea.

60 J. Kojima

Waigeo: 1 ♀ (BMNH), Mt. Nok, vi.1938, L. E. Cheesman. Misool: ♀ (OUM, holotype of Icaria deceptor Smith), labeled “M” [= Mysol] (circular), “Icaria deceptor Smith” and “= gracilis Smith, L. E. Cheesman, 1949”; 3♀ (NNM), Fokal, 0-75 m, 8.ix.-20.x.1948, M. A. Lieftinck. NW New Guinea: 1♀ (NNM), Torricelli Mts., 200-1000’, i.[19]39, E. L. Cheesman; 1♀ (SAM), Torricelli Mts., 1000’, i.[19]39, E. L. Cheesman; 1♀ (NNM), Humboldt Bay; 1♀ (NNM), Neth. Ind.-Amer. New Guinea Exp., Bernhard Camp, 50 m, 16.ix.1939, L. J. Toxopeus. Aru: ♀ (OUM, holotype of Icaria gracilis Smith), labeled “Aru” (circular), “Icaria gracilis Smith” (blue label), “Type [in Richards’ (?) handwriting]”; 1♀ (NNM), Mus. Leiden Snellius Exp., Aroe Eilanden, Manoem-baai en omgeving [= Manoem Bay and environs], 11-44.x.1929. SW New Guinea: 2♀1♂ (NNM, IUNH), 11 km S. Bupul, 20 m, 7°39’S, 140°53’E, 9.iv.1988, R. Hensen. SE New Guinea: 1♀ (SAM), Mt. Lamington, 1,300-1,500 feet, C. T. McNamara; 1♀ (BMNH), Kokoda, 1200 ft., ix.1933, L. E. Cheesman. Australia: 2♂ (BMNH), North Queensland, R. C. L. Perkins [identified as “deceptor”].

Species in the R. rufoplagiata-group Ropalidia cristata Kojima

Ropalidia cristata Spradbery, 1985: 17. Nomen nudum. Ropalidia cristata Kojima, 1989: 143, female, male, larva –

“PAPUA NEW GUINEA . . . Ower’s Corner, Sogeri, Central Province”. — Spradbery and Kojima, 1989: 633 (colony population; nest; ethology). — Spradbery, 1991: 350 (colony population).

This species is closely related to R. turneri Richards, 1978, distributed in north-eastern Australia. Among the characters I (Kojima, 1989) remarked to distinguish R. cristata from R. turneri, the state of the propodeal concavity, the shape of the clypeus and the condition of the pronotal carina were revealed to be less reliable after examination of many additional specimens of R. turneri (mainly from the Atherton Tableland, Queens-land, Australia; deposited in the IUNH). The following key gives the diagnostic characters, including those newly recognized, to distinguish between R. cristata and R. turneri.

1. Ocelli relatively close to each other, distance between posterior ocelli less than 1.5× their diameter; space between posterior ocelli hardly punctured (Fig. 58); gena in profile widened ventrally to level of about one-third from eye top, then posterior margin of gena (or occipital carina) run down in nearly parallel to posterior margin of eye, recurved anteriorly near the base of mandible (Fig. 59); posterior lamella of the first metasomal

tergum wide and not strongly depressed, in profile length of lamella distinctly longer than depth of depression (Fig. 60). Papua New Guinea………...… …. ................................................. R. cristata Kojima

– Ocelli less close to each other, distance between posterior ocelli 1.7× or more their diameter; space between posterior ocelli with a few punctures (Fig. 61); gena in profile smoothly widened ventrally to the level of mid of eye, then smoothly narrowed toward mandibular base, occipital carina broadly and regularly curved (Fig. 62); posterior lamella of the first metasomal tergum narrow, strongly depressed, in profile length of lamella nearly the same as depth of depression (Fig. 63). Northeastern Australia...................................... R. turneri Richards

Specimens of R. cristata examined — NE New Guinea: 1♀ (NNM, paratype), Wau, Morobe Province, 1250 m, 11.i.1950, J. Sedlacek; 9♀ (IUNH, paratypes), Wau, 23.xii.1973, nest 731223-01, T. Okazawa; 3♀ (IUNH), Wau, 1.vii.1985, nest RPNG8501, J. Kojima; 7♀ (IUNH), Wau, 12.vii.1985, nest RPNG8504, J. Kojima; 1 ♀ (IUNH), Wau, 14.vii.1985, nest RPNG8509, J. Kojima. SE New Guinea: ♀ (ANIC, holotype), labeled “PAPUA NEW GUINEA/Central District/Ower’s Corner/28 August 1973/J. P. Sprad-bery”; 3♀ (NNM, paratypes), same collection data as the holotype; 2♀ (NNM, paratype; BMNH), Ower’s Corner, 3.ix.1973, J. P. Spradbery; 2 ♂ (NNM, paratype; BMNH), Uberi, Kokoda Trail, 26.iii.1974, J. P. Spradbery; 1♀ (BMNH), Kokoda, 1,200 ft., v.1933, L. E. Cheesman.

Acknowledgments My special thanks are due to the late J. van der Vecht,

who had encouraged me to make an intensive study of Ropalidia, one of the largest and most interesting polistine genera; he always offered me valuable suggestions on the biology not only of the Ropalidia but also of the Vespidae in general. I am very pleased to have a chance to publish some more of his work. I thank C. van Achterberg for his help in various ways during the present study, without which the present study would not have been accomplished. My one-year stay in the Nationaal Natuurhistorisch Museum in 1996 was supported by the Nederlandse Organisatie voor Wetenschappelijk Onderzoek (NWO) and Japan Society for the Promotion of Science (JSPS). My thanks are also due to my family for their patience allowing me to spend most time of the one-year stay in the Netherlands in taxonomic work. I also wish to express my thanks to J. M. Carpenter for critical reading of an earlier draft, and to the following persons who assisted me in examining the specimens and/or provided me with the specimens for this study: A. D. Austin, B. Bregge, J. Cardale, J. M. Carpenter, J. Casewetts-Weulersse, M. C. Day, P. Dessart, S. Hartini, S. Heydon, W. Hogeness, T. Huddleston, S. Kahono, M. Lachaise, S. Lewise, I. Löbl, E. G. Matthews, A. Menke, B. Merz, I. D. Naumann, G. M.


Nishida, W. A. Noerdjito, D. Notton, L. Tarel, C. O’Toole, W. Pulowski, J. P. Spradbery, C. Taylor, C. Villemont, R. de Vries and Sô. Yamane. Beside the financial support from the NWO and JSPS, the present study was partly supported by Grants-in-Aid from the JSPS (nos. 11833001 and 11691161).

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Received 1 January 2001; revised 16 January 2001; accepted 20 January 2001.

Ropalidia wasps (Insecta: Hymenoptera; Vespidae

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