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 1838]), who described Ropalidia maculiventris from “Dory” [=Manokwari]. Ten years later, Le Guillou Ropalidia Guérin-Méneville, 1831, comprising (1841a) described the second Ropalidia species from about 180 species, is one of the largest polistine genera, Toriton Bay in northwestern New Guinea as Polistes and is spread over the Ethiopian, Oriental and lefebvrei. Around the mid-nineteenth century, de Australian regions (Kojima and Carpenter, 1997). The Saussure (1853-58) described two species in his genus genus, together with Parapolybia de Saussure, 1854, Icaria de Saussure, 1853, from the area, and Smith Polybioides du Buysson, 1913, and Belonogaster de (1859, 1860, 1863, 1865, 1871) described 14 species Saussure, 1854, forms a monophyletic tribe Ropalidiini under the genera Icaria, Polybia Lepeletier, 1836, and (Carpenter, 1991, 1993), which is confined to the Old Odynerus Latreille, 1802; all but two of Smith’s species World. Despite the fact that nearly one-third (55 are currently treated as valid species in Ropalidia species) of the Ropalidia species that are currently (Kojima and Carpenter, 1997). About the end of treated as valid species are distributed in New Guinea nineteenth and early twentieth century, du Buysson and its adjacent islands, taxonomy of Ropalidia in the (1909a, b, 1913) and von Schulthess (1913) respective- area has yet been very poorly studied compared with ly described Polybia loriana, Icaria cohni, Polybia congeners in the Oriental region (van der Vecht, 1941a, novaeguineae and Parapolybia novaeguineae from 1962; Das and Gupta, 1984, 1989) and in Australia New Guinea and the adjacent islands. Around the same (Richards, 1978; Kojima, 1999a, b), as well as with the period, Cameron (1898, 1906a, b, 1911, 1913) describ- other three genera of the Ropalidiini (Parapolybia: van ed 20 Ropalidia species from this region under the der Vecht, 1966; Polybioides: Bequaert, 1918; van der genera Icaria, Polistratus Cameron, 1906, Polistes Vecht, 1966; Belonogaster: Richards, 1982; Hensen Latreille, 1802, Polybia, Odynerus and Ancistrocerus and Blommers, 1987; Kojima, 2001). Wesmael, 1836, of which six are currently considered The first record of Ropalidia from New Guinea is of as valid species. Meade-Waldo (1911, 1912; also Guérin-Méneville (1831, in Duperrey et al., “1826”[- Meade-Waldo and Morley, 1914) described five species 32 J. Kojima in Icaria, all of which are currently considered valid, study, either van der Vecht or I followed the subgeneric and studied distribution and synonymic relationships of division defined by Richards (1978), which was shown ten species having been described by other authors. to be supported by no robust grounds (Kojima, 1997a, However, neither Cameron nor Meade-Waldo made any b) and even led us some taxonomic confusions at the revisional studies on the Ropalidia fauna in New species level (for examples, see the relationships among Guinea nor provided even a list of species recorded in R. semihyalinata, R. novaeguineae and R. novaeguineae New Guinea by their times. insolens). Second, some of the late van der Vecht’s Bequaert (1932) described Ropalidia leopoldi from conclusions were based on the examination of the types Angi-Kita in northwestern New Guinea, and listed 37 housed in the museums and institutions in Europe that species, nearly all the species that had been described he made in 1933 and 1934 (see van der Vecht, 1934, from New Guinea. However, he did not make any 1941a); several new taxa have been described since detailed taxonomic comparisons. Van der Vecht (1934) then. Third and the most important, the recent studied the types of Cameron’s species and concluded development and sophistication of the phylogenetic that 11 “species” were not valid, and he (1941b) made a species concept led me to concur that subspecies have brief note on the New Guinean Ropalidia. He also no place in a phylogenetic system and that diagnosable treated six species or subspecies from Papuan region in taxa are to be treated as species (Nelson and Platnick, his monographs on Malayan species of Ropalidia (van 1981; Nixon and Wheeler, 1990; Carpenter and Kojima, der Vecht, 1941a, 1962). Although Cheesman (1952) 1997). attempted to study thoroughly the Ropalidia fauna in The present paper, as the first part of a compre- New Guinea, with descriptions of 18 new taxa, her hensive taxonomic revision of the New Guinean work was based on only specimens that she herself Ropalidia, treats the results of the research on the type collected and those in the collections in the University materials of the previously described taxa, and gives a Museum, Oxford, and The Natural History Museum, taxonomic revision of the R. loriana-group. The London, and referred to only 14 of 37 species that remaining species will be treated in the second part of Bequaert (1932) listed. Richards (1978) described five this series, which will be published in the next issue of additional Ropalidia species from New Guinea in the the present bulletin. monograph of the Australian social wasps, and provided brief taxonomic notes on some New Guinean Materials species so far described. We have described further three species from New Guinea (Kojima, 1989; van der The present study is based mainly on the specimens Vecht and Kojima, 1996). Thus, as summarized above, housed in the Nationaal Natuurhistorisch Museum, the Ropalidia fauna in New Guinea and its adjacent Leiden (abbreviated as NNM), but also on those in the islands, one of the centers of divergence of this genus, following museums and institutions: has never been studied thoroughly. American Museum of Natural History, New York It had been the intension of the late Prof. Dr. J. van (AMNH); der Vecht “to discuss the species of this area in a future Australian National Insect Collection, CSIRO, paper” (van der Vecht, 1962: 4). I started, jointly with Canberra (ANIC); him, the taxonomic study of the New Guinean Bernice P. Bishop Museum, Honolulu (BISH); Ropalidia nearly 20 years ago. However, the study was California Academy of Science, San Francisco interrupted half way through due to his illness and death (CAS); (in 1992) (see van Achterberg, 1992). Recently I had Hope Entomological Collection, University Mu- several opportunities to visit the Nationaal Natuur- seum, Oxford (OUM); historisch Museum, Leiden, including one-year stay at Institut Royal des Sciences Naturelles de Belgique, the museum in 1996, and reexamined a large part of the Brussels (IRSNB); materials that we studied before. Museum of Comparative Zoology, Harvard Uni- At the beginning of the preparation of this versity, Cambridge, Massachusetts (MCZ); taxonomic revision, I intended to publish it as a joint Muséum d’Histoire Naturelle, Genève (MHN); paper with the late van der Vecht. However, I decided Museum National d’Histoire Naturelle, Paris later to take responsibility for all the nomenclatural (MNHN); changes, because many conclusions presented here are Museum Zoologicum Bogoriense, Bogor (MZB); very different from those on which we agreed before Natural History Laboratory, Ibaraki University, and/or the late van der Vecht reached. Most of these Mito (IUNH); discrepancies between my conclusions given in this The Natural History Museum, London (BMNH); paper and those we previously reached are due to the South Australian Museum, Adelaide (SAM); following three reasons. First, when we started this Zoölogisch Museum, Amsterdam (ZMA).

Ropalidia of New Guinea 33

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

The New Guinea mainland is divided into the under a stereoscopic dissecting microscope with an following four sectors, mainly for the convenience in ocular micrometer. When the terminal sterna and male reference (Fig. 1): “North-Western (abbreviated as NW) genitalia were examined they were dissected out, New Guinea” is the part of Papua (formerly Irian Jaya) briefly cleared in KOH and mounted on glass slide in north of the central mountains, including the Vogel Kop glycerin to observe under a stereoscopic dissecting Peninsula; “South-Western (SW) New Guinea”, the part microscope and/or a compound microscope. Termi- of Papua south of the central mountains; “North-Eastern nology of the male genitalia follows Kojima (1999c). (NE) New Guinea”, the part of Papua New Guinea The following morphometric characters and views north of the central mountains; and “South-Eastern (SE) of observation are often ill defined in the literature. In New Guinea”, the part of Papua New Guinea south of the present study they are defined as follows (Figs. 2– the central mountains, including Owen Stanley Range. 7): width of clypeus (CW in Fig. 2) is measured as a The specimens examined are listed according to this distance between the uppermost points where clypeus subdivision of the mainland and the names of adjacent touches eyes; height of clypeus (CH in Fig. 2) is islands. When exact localities are not indicated on the measured as a distance from the dorsal emargination to labels or are not recognized, the specimens are listed the apex; distance between inner eye margins at vertex under “New Guinea”. (OV in Fig. 2) refers to the distance between them at The name-bearing types (holotypes and lectotypes) the level of anterior ocellus in frontal view of head, and of the previously described species are listed with the that at clypeus (in most case equivalent to CW in Fig. complete information on the attached labels; for some 2) refers to the distance at the level where inner eye species, such information is available elsewhere margins approach each other most closely; inter- (Kojima and Carpenter, 1997; Kojima and van antennal (AA in Fig. 2) and antennocular (AO in Fig. 2) Achterberg, 1997), but is repeated here for distances are measured as horizontal distances at the convenience. When label data are quoted, a slash (/) level of middle of antennal socket; widths of eye (EW denotes the beginning of a new line on the label; in Fig. 3) and gena (GW in Fig. 3) are measured as the however, I failed to record the line breaks for labels of maximum for each in strictly lateral view of the head; the specimens in the OUM and BMNH, and their data distance between posterior ocelli (POD in Fig. 5) and are given without indicating the line breaks. My notes distance between posterior ocellus and inner eye margin on the label data and citations are given in brackets. (OOD in Fig. 5) are measured in dorsal view of head; length of scape (SL in Fig. 4) is measured excluding the Methods radicle; length of first metasomal tergum (T1L in Fig. 6) was measured in profile as a distance between the The adult morphology except for the terminal sterna posterior end of the basal slit for the reception of the and male genitalia was observed on pinned specimens 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 widths of first and second metasomal segments refer to (or segment) (T2L in Fig. 6) was measured in profile as their maximum widths in dorsal view; height of second a distance from the bottom of the basal depression or metasomal segment (MT2H in Fig. 6) is measured as “neck” to the posterodorsal margin of the tergum; the maximum height in lateral view; length of posterior

Ropalidia of New Guinea 35 widened part of first metasomal sternum (PL in Fig. 7) are paralectotypes, but I failed to locate the last is measured as a distance between the hypothetical line specimen. on the level where ventral margins of the first tergum Icaria bispinosa Meade-Waldo, 1912: Meade-Waldo approach each other most closely and the hypothetical (1912: 449) described this species based on four line connecting the posterolateral ends of the sternum, females from Mimika River collected in July and and its width (PW in Fig. 7) refers to the maximum August 1910. Cheesman (1952: 9) treated this posterior width. The “dorsal view of first metasomal taxon as a subspecies of Ropalidia fasciata (Smith, tergum” (DV in Fig. 6) refers to the view perpendicular 1859) and listed the specimens she examined as to the dorsal face of the posterior widened part of the follows: “1♀ (holotype), 1♀ (paratype), Mimika tergum, and “anterodorsal view of first metasomal River, 1911 (A. F. R. Wollaston)”. Again her tergum” (ADV in Fig. 6) is perpendicular to the first statement cannot be regarded as the designation of metasomal sternum. the lectotype. I could find only one specimen of Only major references are listed in this paper, and Meade-Waldo’s syntypes, and here designate it as other references can be found in a catalog of ropalidiine the lectotype of I. bispinosa: female (BMNH), species (Kojima and Carpenter, 1997; or on a web-site labeled “Holotype” (BM type label), “New Guinea ~ http://www.sci.ibaraki.ac.jp/ jkrte/wasp/ropa/top.html). Mimika R. A. F. R. Wollaston, 1911-229 Aug. For each original citation, the name of the taxon, author 1910”, “Icaria (Icariastrum) bispinosa sp. n., Type, name, and year of publication are followed by the ♀ , G. Meade-Waldo det.”, “fasciata Sm. v. sex(es), if stated, and the quotation of the type locality. bispinosa M. W., L. E. Cheesman det. 1950”, and In cases in which I did not examine the types, the type “B.M. TYPE HYM 18.851”. depositories are also given. Icaria torrida Smith, 1863: Smith (1863: 42) described Species are arranged according to the species- I. torrida from “Ceram” without giving any groups based on an on-going cladistic analysis of the indications for the number of specimens he genus Ropalidia (Kojima and Carpenter, unpublished). examined. The OUM houses two females labeled “Cer” [= Ceram] (circular) and “Icaria torrida Designation of the lectotypes Smith”, that are obviously from the type series. One of them is here designated as the lectotype of I. In any studies of New Guinean Ropalidia before torrida, and the other specimen (metasoma detached Bequaert (1932), the holotypes were not designated and and glued on the circular label) is a paralectotype. the numbers of specimens examined were rarely mentioned. Kojima and Carpenter (1997) designated Summary of synonymies the lectotypes of 16 New Guinean taxa described by Smith (1859, 1860, 1863), Cameron (1906a, b, 1911, Synonymies that are newly proposed in the present 1913) and Cheesman (1952). The lectotypes of the paper and those that were previously published and are following three taxa are herewith designated and confirmed in the present study are summarized below. labeled so. Ropalidia maculiventris Guérin-Méneville, 1831: = Icaria wollastoni Meade-Waldo, 1912: Meade-Waldo Icaria sexmaculata Cameron, 1911, NEW (1912: 449) described this species based on four SYNONYMY; = Ropalidia pratti Cheesman, 1952, females from Mimika River collected in July 1910. NEW SYNONYMY; = Ropalidia maculiventris Cheesman (1952: 7) inadvertently referred to one of humboldti Cheesman, 1952, NEW SYNONYMY; = them as “holotype”, stating “1♀ (holotype), 3♀♀, Ropalidia mondoensis Cheesman, 1952, NEW S.W. New Guinea, Mimika R., 1911 (A. F. R. SYNONYMY. Wollaston)”. Without specifying an individual Ropalidia wollastoni (Meade-Waldo, 1912): = Icaria specimen by its label data and/or characters, her sericea Cameron, 1911, non 1905, synonymized by statement cannot be regarded as the designation of Kojima and Carpenter (1997: 60). the lectotype (Article 74.3 in the International Code Ropalidia australis (de Saussure, 1853): = Icaria of Zoological Nomenclature). The following bispinosa Meade-Waldo, 1912, NEW SYNONY- specimen is herewith designated as the lectotype of I. MY; = Ropalidia fasciata erratica Cheesman, 1952, wollastoni: female (BMNH), labeled “Holotype” NEW SYNONYMY. (BM type label), “New Guinea, Mimika R., A. F. R. Ropalidia conservator (Smith, 1860): = Polybia Wollaston, 1911-229 July 1910”, “Icaria limatula Smith, 1863, synonymized by van der (Icariastrum) wollastoni sp. n., Type, ♀, G. Meade- Vecht (1966: 6); = Icaria parvimaculata Cameron, Waldo det.”, and “B.M. TYPE HYM 18.838”. I 1911, non 1907, synonymized by van der Vecht examined two other specimens in the BMNH with (1934: viii); = Icaria insularis Cameron, 1911, the same collection data as the lectotype and they synonymized by van der Vecht (1934: viii); =

36 J. Kojima

Ropalidia conservator goodfellowi Cheesman, 1952, Ropalidia lefebvrei (Le Guillou, 1841): = ? Icaria synonymized by Richards (1978: 59); = Icaria reactionalis Saussure, 1853. waigeuensis Cameron, 1913, synonymized by Ropalidia deminutiva Cheesman, 1952: = Ropalidia Meade-Waldo and Morley (1914: 405). integra Cheesman, 1952, NEW SYNONYMY. Ropalidia dispila (Cameron, 1913): = Ropalidia Ropalidia impetuosa (Smith, 1860): = Icaria fasciola Richards, 1978 (replacement name for spilocephala Cameron, 1906, synonymized under R. Icaria fasciata Smith, 1859, non Eumenes fasciata gregaria by van der Vecht (1941a: 104); = Icaria Fabricius, 1804), NEW SYNONYMY. cohni du Buysson, 1909, synonymized under R. Ropalidia jaculator (Smith, 1863): = Ropalidia ivorina gregaria by van der Vecht (1941a: 104); = Cheesman, 1952, NEW SYNONYMY. Ropalidia gregaria inquieta Cheesman, 1952, Ropalidia novaeguineae (von Schulthess, 1913): = synonymized under R. gregaria spilocephala by Polybia novaeguineae du Buysson, 1913, synony- Richards (1978: 86); = Ropalidia gregaria var. mized by van der Vecht (1966: 6). tolerans Cheesman, 1952, synonymized under R. Ropalidia semihyalinata (Meade-Waldo, 1912): = gregaria spilocephala by Richards (1978: 86). Ropalidia novaeguineae insolens Cheesman, 1952, Designating the lectotype of Icaria gregaria de NEW SYNONYMY. Saussure, 1854, I (Kojima, 2001: 5) showed that Ropalidia brunnea (Smith, 1859): = Polistratus “Ropalidia gregaria” of authors, including van der cariniscutis Cameron, 1906, synonymized by van Vecht (1941a) and Richards (1978), is not the true der Vecht (1934: viii); = Ropalidia wudai R. gregaria and adopted R. impetuosa for “R. Cheesman, 1952, NEW SYNONYMY; = Ropalidia gregaria” of authors; van der Vecht (1941a: 104) wudai trullissima Cheesman, 1952, NEW synonymized Icaria impetuosa under R. gregaria. SYNONYMY. Ropalidia gracilis (Smith, 1859): = Icaria deceptor Ropalidia conspicua (Smith, 1863): = Odynerus Smith, 1863, synonymized by Cheesman (1952: 18). mysolicus Dalla Torre, 1889 (unnecessary Ropalidia marginata jucunda (Cameron, 1898): = replacement name for Odynerus conspicuus Smith, Icaria pruinosa Cameron, 1906, synonymized by 1863); = Ropalidia conspicua alleni Cheesman, van der Vecht (1941a: 104); = Ropalidia canaria 1952, synonymized by Kojima and Carpenter (1997: Cheesman, 1952, NEW SYNONYMY. 16) without indicating that their treatment was a new synonymy [Richards (1978: 57) treated alleni Misidentifications of New Guinean as a junior synonym of R. brunnea (Smith, 1859), Ropalidia in the literature possibly erroneously taking “conspicua alleni” as “cariniscutis alleni”; see also Kojima and Carpenter, The following misidentifications of New Guinean 1997: 16]. Ropalidia published in the literature were proved during Ropalidia nigra (Smith, 1859): = Icaria morosa Smith, the course of the present study. 1863, synonymized by Cheesman (1952: 10); = Cameron (1906a: 63) recorded a female from Icaria spilostoma Cameron, 1906, synonymized by Manokwari as probably “Icaria irritata Smith”. van der Vecht (1934: viii); = Odynerus sariensis The specimen is in the ZMA and belongs to R. Cameron, 1906, synonymized by van der Vecht conservator (Smith). (1934: viii); = Odynerus confraternus Cameron, Cameron (1913: 76) recorded “Icaria irritata Smith” 1911, synonymized by van der Vecht (1934: viii); = from Waigeo, stating that “A male taken in January Ancistrocerus catharinae Cameron, 1913, synon- is probably this species . . .” This specimen is in the ymized by van der Vecht (1934: viii); = Ropalidia NNM and actually the female of R. australis (de domestica Chessman, 1952, NEW SYNONYMY; = Saussure). Ropalidia sepicana Richards, 1978, NEW SYN- Von Schulthess (1932: 39) recorded “Polistes ONYMY. tasmaniensis de Saussure” from Sakoemi. The Ropalidia festina (Smith, 1865): = Icaria zonata specimen is in the IRSNB and is a female of R. Cameron, 1906, synonymized by Meade-Waldo conservator (Smith). (1911: 100). Cheesman (1952: 11) recorded two females, each from Ropalidia albobalteata (Cameron, 1906): = Polybia Japen and Waigeo, under “Ropalidia irritata”. I papuana Cameron, 1913, REVISED SYNONYMY; located the female from Japen in the BMNH and = ? Icaria (Icariola) fluviatilis Meade-Waldo, 1912; this specimen is certainly not R. irritata, but = ? Ropalidia fluviatilis petulans Cheesman, 1952. probably an undescribed species; details will be Ropalidia catharinae (Cameron, 1913): = ? Ropalidia given in the second part of the present series. fluviatilis petulans Cheesman, 1952. Kojima and Spradbery (1987): “Ropalidia leopoldi” (p. 606) is not R. leopoldi and an undescribed species,

Ropalidia of New Guinea 37

which will be treated in the second part of this Head in frontal view about 1.2 times as wide as series; “Ropalidia loriana” (p. 607) is R. high; in dorsal view little more than twice as wide as longipetiolata (Cameron) as I have already pointed long, strongly narrowed behind eyes, strongly out elsewhere (Kojima, 1998: 218); “Ropalidia sp. emarginate posteriorly (Fig. 8), slightly wider than possibly R. irritata (Cameron, 1863)” [error in mesosoma including tegulae. Gena in lateral view author name; this should be “Smith”] (p. 603) is R. widest at level of mid-height of eye (about 0.65 times as irritata. wide as eye), then gradually narrowing ventrally; Spradbery and Kojima (1989): “Ropalidia kurandae” occipital carina complete, smoothly and weakly curved. (p. 639) is an undescribed species, which will be Ocelli proportionally larger than most of other treated in the second part of this series; “Ropalidia Ropalidia species; posterior ocelli slightly closer to leopoldi” (p. 649) and “Ropalidia loriana” (p. 633) each other than to anterior ocellus; distance between are the materials from the same colonies as those in posterior ocelli about 0.65× their diameter, and about Kojima and Spradbery (1987); and “sp. close to R. 0.3× distance between posterior ocellus and inner eye bispinosa” (p. 644) is R. australis (de Saussure). margin; space between posterior ocelli flat (Fig. 9). Kojima and Kojima (1994): “Ropalidia leopoldi” (p. Inner eye margins weakly converging ventrally, about 52) is the material from the same colony as in 1.1 times further apart at vertex than at clypeus. Kojima and Spradbery (1987) and Spradbery and Clypeus rather convex, pointed apically, slightly wider Kojima (1989). 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; Taxonomy of the Ropalidia loriana-group third article slightly more than twice as long as its apical width, about as long as fourth and fifth articles The R. loriana-group includes three species, R. combined; tenth and eleventh articles widest in flagellar loriana (du Buysson, 1909), R. longipetiolata articles, each distinctly wider than long, and about 1.3 (Cameron, 1911) and a species described in this paper. times as wide as apical width of third article; terminal They are, so far known, endemic to the New Guinea article bullet-shaped, about 1.5 times as long as its basal mainland. Richards (1978) placed R. loriana and R. width (Fig. 10). Mandible rather stout, not twisted; longipetiolata in the subgenus “Icarielia” only based on teeth not elongate. weak sculpture, but these species are very different Mesosoma not thick; in dorsal view, nearly 1.5 from the other species that he placed in “Icarielia” (see times as long as wide. Pronotum in dorsal view with Kojima, 1996, 1997b). The diagnostic characters (not lateral margins diverging posteriorly, barely convex; all of them is apomorphic) of the R. loriana-group are: pronotal carina raised into fine lamella, more strongly body surface barely sculptured; mid-femur slender raised at humeral angles, not sinuate laterally. Scutum throughout its length (Fig. 33); propodeal valvula not convex, nearly 1.1 times as long as its width between carinate basally; first metasomal segment elongate and tegulae. Scutellum convex (Fig. 11); lateral margins of slender (Figs. 12, 13, 26, 27, 36–38); tergum and disc not carinate, barely diverging posteriorly. Disc of sternum of second metasomal segment not fused, but metanotum convex, with no lateral marginal carina, overlapping (Fig. 36). slightly produced posteromedially into rounded corner. Mesepisternum without carina; secondary entrance of Ropalidia longipetiolata (Cameron) first spiracle raised, followed posteriorly by depression, (Figs. 8–21) which is margined dorsally by fine carina. Posterior Icaria longipetiolata Cameron, 1911: 189, female – face of propodeum convex, without median furrow or “Lorentz”. — van der Vecht, 1941a: 105 (subfamily depression; in posterodorsal view, lateral sides of Polybiinae). — Kojima and Carpenter, 1997: 4 propodeum converging posteriorly in nearly straight (designation of lectotype). lines; propodeal orifice round above, about twice as Ropalidia longipetiolata; Bequaert, 1932: 50 (“probablement long as wide; propodeal valvula rather large, with un Parapolybia”). — Kojima, 1996: 328. broadly rounded apex (Fig. 12), in profile covering ? Ropalidia loriana (du Buysson); Richards, 1978: 58 (in ventral half of propodeal teeth. subgenus Icarielia), 128 (key). First metasomal segment slender, about 3.5 times as Ropalidia loriana; Kojima and Spradbery, 1987: 607, figs. long as wide, raised at spiracles; in dorsal view with 37–41 (larva). — Spradbery and Kojima, 1989: 633 (nest; basal parallel-sided part about three times as long as colony population). — Spradbery, 1991: 351 (colony wide, weakly widening posteriorly from level slightly population). anterior to spiracles, slightly constricted near apical Female — Body length (head + mesosoma + first margin, and then weakly widening again at apex, apical two metasomal segments) 9–10 mm; forewing length width about twice as wide as width of basal parallel- 9.5–10 mm.

38 J. Kojima sided part (Fig. 13); in lateral view dorsal surface very Specimens examined — NW New Guinea: 1♂ slightly arising posteriorly from posterior margin of (NNM), Vogelkop, Kebar Val., W of Manokwari, 550 basal slit for reception of mesosomal suspensory m, 4-31.i.1962, S. Quate & L. Quate; 1♀ (NNM), Neth. ligament, then weakly and smoothly convex in posterior Ind.-Amer. New Guinea Exp., Mountain slopes above one-third (Fig. 12); ventral margins of tergum weakly Bernhard Camp, 750 m, 27.iii.1938, L. J. Toxopeus. approaching each other before posterior divergence; SW New Guinea: ♀ (ZMA, lectotype designated by posterior margin of sternum weakly emarginate Kojima and Carpenter (1997: 4)), labeled “Z. Nieuw medially. Second metasomal segment flattened Guinea/Lorentz 1909-10/Noord rivier/IX. 09”, “Nest dorsoventrally, about as long as wide, about 1.2 times B?”, “longipetiolata [in Cameron’s handwriting]”, as long as high; in dorsal view spiracle weakly “Holotype”, “Icaria/longipetiolata/ ♀ Cam. (1911) protruding, smoothly expanding posteriorly in anterior /HoloTYPE/sec. J. v. d. Vecht 33 [in van der Vecht’s half, then slightly narrowing towards apex; thyridium handwriting” and “Ropalidia ♀ /longipetiolata/(Cam.) distinct, elongate oval on tergum and line on sternum. /det. J. v. d. Vecht 1933”. NE New Guinea: 1♀ Terminal sternum with basal suboval membranous area (NNM), Torricelli Mts., Sugoitei Vill., 900 m, 6- armed with sparse, short hairs (Fig. 14) (in most 9.ii.1959, W. W. Brandt. SE New Guinea: 24♀ (12♀ Ropalidia species, this area is in peculiar bifurcate in NNM, 9♀ in IUNH, 3♀ in AMNH), from a colony, shape; see van der Vecht, 1968). Central District, Baruanumu, Musgrave R., 19.v.1973, Body shining, covered with fine tomentum except in J. P. Spradbery; 1♀ (NNM), Fly R., Olsobip, 400-600 apical half of clypeus, and sparse short, erect, silvery m, viii.1969, J. & M. Sedlacek. hairs. Head and mesosoma with sparse, shallow, ill- defined punctures; metasoma unpunctured. Ropalidia loriana (du Buysson) Color brown to dark brown; first metasomal (Figs. 22–27) segment (often with brown markings on posterior Polybia loriana du Buysson, 1909b: 314, female – “Nouvelle widened part of tergum), and basal part of second Guinée S. E. Moroka 1300 m” segment, yellow; specimens from Musgrave River with Parapolybia loriana; von Schulthess, 1913: 153 (key), 158, upper half of clypeus, base of antennal scape, ventral pl. 11 fig. 5 (female). part of ocular sinus, gena (partly), yellowish brown, and Ropalidia loriana; van der Vecht, 1966: 6. most of legs (except tarsi), scutellum, metanotum, Female — Body length (head + mesosoma + first propodeum, yellow. two metasomal segments) about 10 mm; forewing Male — Body length (head + mesosoma + first two length about 10 mm. metasomal segments) 9 mm; forewing length 9.5 mm. Structure and coloration as in female of R. Structure as in female except following characters. longipetiolata, but different in the following characters. Head relatively thinner, about 1.2 times as wide as high, Head in dorsal view with lateral sides behind eyes about 2.3 times as wide as long; posterior emargination hardly convex, more strongly convergent posteriorly in dorsal view shallower. Clypeus smaller, nearly 1.1 (Fig. 22); gena in profile proportionally narrower, times as wide as high. Inner eye margins more strongly slightly more than half as wide as eye Space between convergent below, about 1.25 times further apart at posterior ocelli with slightly raised, short, longitudinal vertex than at clypeus. Eye more strongly swollen. carina (Figs. 23, 24). Scutellum less convex (Fig. 25 vs. Gena narrow, in lateral view about 0.3 times as wide as 11 for longipetiolata); posterior face of propodeum eye. Antennal scape about 3.7 times as long as its slightly more convex (Fig. 26 vs. 12 for longipetiolata). apical width; third article nearly 2.5 times as long as its First metasomal tergum slightly more slender; in profile apical width, slightly longer than fourth and fifth dorsal margin nearly parallel with its ventral margin in articles combined; terminal article about 1.8 times as anterior half, smoothly and weakly convex in posterior long as wide; fourth to terminal articles with slightly half (Fig. 26); in dorsal view nearly parallel-sided in raised tyloids (oval shaped rather than of narrow ridge). anterior half, and weakly swollen posteriorly from just Terminal sterna and genitalia as in Figs. 17-21. in front of spiracle, then weakly constricted subapically Parameral spine without hairs; Parameral Spine Index (Fig. 27). Punctures shallower and sparser; scutellum of Charnley (1973): (length of projection beyond and metanotum barely punctured. Body dark, blackish lobe)/(total length of paramere and spine) × 100 = 14. brown; antennal scape slightly paler; posterodorsal Volsella as in most Ropalidia species (see Kojima, margin of pronotum sometimes with narrow band 1999c). Penis valves about 1.2 times as long as basal slightly paler; extreme apex of mesosoma, sometimes apodeme. area around propodeal spiracle, first metasomal Color as in females from Musgrave River except segment (sometimes posterior widened part of the brown scutellum. tergum dark brown), base of second metasomal

Ropalidia of New Guinea 39 segment, yellow to orange-yellow. Wings semihyaline, slightly infuscate along anterior margin. Male — Not known.

40 J. Kojima

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.

Specimens examined — NW New Guinea: 7♀ W. W. Brandt. SE New Guinea: ♀ (MNHN, holotype; (NNM, 1 ♀ in IUNH), Neth. Ind.-American New see also Carpenter, 1999: 36), labeled “N. GUINEA S. Guinea Exped., Rattan Camp [3°30’S, 139°09’E], L. J. E./Moroka 1300 m/LORIA VII-XI 93”, “MUSEUM Toxopeus (1 ♀ : 1,150 m, 12.i.1939; 3 ♀ : 1,150 m, PARIS/G. Gestro 1909”, “TYPE”, “Polybia/Loriana ii.1939; 2 ♀ : 1,200 m, 4.iii.1939; 1 ♀ : 1,100 m, Buyss./type/R. DU BUYSSON det 1909”. iii.1939); 1♀(NNM), Neth. Ind.-American New Guinea Exped. Araucaria Camp [3°30’S, 139°11’E], 800 m, Ropalidia principalis new species iii.1939, L. J. Toxopeus; 1♀ (NNM), Star Mts., Sibil (Figs. 28–41) Val., 1,245 m, 18.x-8.xi.1961, L. W. Quate; 1 ♀ (NNM), NE, Swart Val., Karubaka 1,300 m, 7.xi.1958, ? Ropalidia longipetiolata; Richards, 1978: 58 (in subgenus J. L. Gressitt. NE New Guinea: 1♀ (BISH), Torricelli Icarielia; list), 128 (key). Mts., Nengian Vill. [3°22’S, 142°05’E], 17-24.xi.1958,

Ropalidia of New Guinea 41

Female — Body length (head + mesosoma + first mid-height of eye, little more than 0.7 times as wide as two metasomal segments) 11–12.5 mm; forewing eye, gradually narrowing ventrally in ventral half; length 10–11 mm. occipital carina complete, smoothly and weakly curved Head in frontal view about 1.2 times as wide as high (Fig. 29). Ocelli large, arranged close to each other in (Fig. 28); in dorsal view slightly more than twice as nearly equilateral triangle; anterior ocellus slightly wide as long, distinctly wider than mesosoma including larger than posterior ones; distance between posterior tegulae, deeply emarginate posteriorly, with lateral ocelli about 0.6× their diameter and about 0.25× sides behind eyes convex, and strongly converging distance to inner eye margin; interocellar space flat. posteriorly (Fig. 30); in profile, gena widest at level of 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. punctures on scutum more or less defined, with central 28). Clypeus weakly convex, pointed apically, nearly postules; mesepisternum posteriorly with ill-defined 1.1 times as wide as high. Antenna weakly thickened sparse punctures, anteriorly unpunctured, border apically; scape slightly curved, slightly more than 4 between them ill-defined; mesopleura and sides of times as long as its apical width; third article rather propodeum with scattered, shallow punctures; short, slightly more than twice as long as its own apical metasoma barely punctured. Body with sparse silvery width, about as long as fourth and fifth articles tomentum, which is dense on upper half of clypeus, combined; fourth and fifth articles each about as long as sides of mesosoma and second to sixth metasomal wide; tenth article widest among flagellar articles, about segments. 1.2 times as wide as long, and nearly 1.2 times as wide Body pale ferruginous; antennal flagellum fuscous; as fourth article; terminal article bullet-shaped, nearly mandible brown, except dark brown basal margin and 1.5 times as long as its basal width (Fig. 31). Mandible teeth; vertex often dark brown; first metasomal tergum rather stout, not twisted; dorsal tooth truncate apically, dark brown in apical one-third, with ill-defined pale- remaining three teeth pointed apically and subequal in ferruginous apical band; second to sixth metasomal length. segments dark brow to blackish brown. All tarsi and Mesosoma not thick, in dorsal view about 1.7 times hind tibia brown to dark brown. Wings slightly as long as wide. Pronotum in dorsal view broadly and infuscate, more strongly infuscate along anterior weakly rounded anteriorly, with lateral sides diverging margin. posteriorly in nearly straight lines; ventral corner Male — Body length (head + mesosoma + first and gradually narrowed; pronotal carina fine, thinly raised second metasomal segments): about 11 mm; forewing throughout its length, hardly sinuate on lateral sides of length about 10 mm. pronotum. Scutum convex, with fine, very shallowly Structure and color as in female except: head more depressed line in anterior one-third to half. Disc of transverse, nearly 1.3 times as wide as high (Fig. 39); scutellum trapezoid, strongly convex (Fig. 32), with inner eye margins more strongly converging below, convex, margined laterally by neither carinae nor nearly 1.25 times further apart at vertex than at clypeus; truncation. Posterior face of propodeum weakly convex clypeus smaller, about as wide as high, covered with (Fig. 36), without median depression or furrow, slightly rather dense silvery tomentum; gena proportionally depressed only just in front of orifice, smoothly curved narrower, in profile little more than 0.3 times as wide as down into lateral faces; propodeum in posterodorsal eye (Fig. 40); antennal scape shorter, about 3.5 times as view with lateral sides converging posteriorly in nearly long as its own apical width; flagellum more slender; straight lines; valvula large, rounded, without basal third antennal article slightly shorter than fourth and carina, in profile covering most of propodeal teeth (Fig. fifth articles combined; terminal article bullet-shaped, 36); orifice rounded above, about twice as long as wide about 1.5 times as long as its own basal width (Fig. 41); (Fig. 35). Mid femur with short fringe beneath, with fourth to terminal articles with slightly raised, ill- sparse outstanding fine setae. defined tyloids. Metasoma slender. First segment elongate, 3.7-3.9 times as long as wide; in dorsal view, with basal Holotype — ♀ (NNM), from a colony, NW New parallel-sided part about three times as long as wide, Guinea “Neth. Ind.-American/New Guinea Expedit weakly thickening posteriorly from level slightly /Bernhard Camp [3°29’S, 139°13’E] B 100 m/9.iv.1939 anterior to spiracle, widest well before apex, then L. J. Toxopeus”. slightly narrowed, and very weakly widened again near Paratypes — 1♂ (allotype) + 112♀1♂ (NNM, 4♀ apex, widest part about 2.2 times as wide as base (Fig. in IUNH, 1♀in AMNH and 4♀in MZB; 1♀in the MZB 37); in profile dorsal face very weakly arising in has an additional label “Ropalidia/loriana/♀det. v. d. straight line from posterior end of basal slit, becoming V.”), same data as the holotype. weakly and smoothly curved near level of spiracle (Fig. Other specimens examined — NW New Guinea: 36); ventral margins of tergum weakly approaching 7♀ (MZB; 1♀in BMNH, identified as “longipetiolata” each other before posterior divergence; posterior by the late van der Vecht), same data as the holotype; 4 widened part of first sternum about as long as wide, ♀ (NNM), Maffin Bay [1°57’S, 138°51’E], 25.v.1944, with posterior margin emarginate medially (Fig. 38). E. S. Ross; 3 ♀ (NNM), Hollandia [= Jayapura], Second segment shortly petiolate basally, about as wide vi.1937, W. Stüber; 10♀ (NNM, 1♀ in BMNH), Mts. as long (Fig. 37), about 1.1 times as wide as high; Bewani, iii.1937, W. Stüber; 1 ♀ (NNM), N. N. articulation of tergum hardly separated from “neck”. Guinean Exp., Mamberamo, Alb.-Bivak, 1926, W. Clypeus except apical one-third, front, pronotum, Docters v. Leeuwen (with a label “Parapolybia loriana scutellum, and anterior half of metanotum with sparse, det. Schulthess 93”); 1 ♀ (AMNH), Maffin Bay, shallow punctures; vertex and gena barely punctured;

Ropalidia of New Guinea 43

25.xi.[19]44, E. S. Ross. NE New Guinea: 1♀ times as wide as eye. Space between posterior (NNM), Lae, v.1945, W. Harden. ocelli flat (Fig. 9). First metasomal tergum in profile weakly and gradually swollen posteriorly Etymology — The loriana-group, to which the from its base, then more strongly swollen near present species belongs, is the basal clade in Ropalidia spiracle (Fig. 12) ...... R. longipetiolata (Cameron) (Kojima and Carpenter, unpublished), thus the group can be considered as the first (principalis) group in the genus. Taxonomic notes mainly on types Remarks — This species can be distinguished from of the species so far described the previous two species by the characters given the following key. The present species seems to have been Ropalidia maculiventris Guérin-Méneville identified as “longipetiolata” or “loriana”. For and its closely related species example, the late van der Vecht identified a female in Ropalidia maculiventris Guérin-Méneville, 1831 the MZB as “loriana” and the specimens in the NNM and its very closely allied species appear to be widely as “longipetiolata”. Furthermore, he identified R. distributed and rather common in New Guinea and longipetiolata as “loriana” (see Spradbery and Kojima, some neighboring islands. It shows a remarkable 1989). He probably sent misidentified specimens to O. variation not only in color pattern, but also in the degree W. Richards, and Richards’ (1978: 128) key to species of development of the posterolateral angles of the of “subgenus Icarielia” seems to be based on those propodeum, which are called “propodeal apophyses”. specimens. Specimens from various localities have been described as different species and/or subspecies, and at this Key to the species in the R. loriana-group moment seven species and/or subspecies are recognized (females, but may be applicable to males) (see also Kojima and Carpenter, 1997): R. maculiventris, R. maculiventris pratti Cheesman, R. 1. Larger species; body length (head + mesosoma + first humboldti Cheesman, R. mondoensis Cheesman, R. two metasomal segments) 11–12.5 mm, forewing sexmaculata (Cameron), R. fulvopruinosa (Cameron), length 10.5–11 mm. Dorsal surface of first and R. wollastoni (Meade-Waldo). The variability of metasomal tergum in profile weakly and smoothly the first five “species” and/or “subspecies”, as well as arising from posterior margin of basal slit for the occurrence of various transitional forms, indicate reception of mesosomal suspensory ligament, and that they should be considered as a single diagnosable broadly and smoothly curved down to posterior species. The remaining two forms are treated as margin in posterior half (Fig. 36). Interocellar space species, as they are diagnosable with stable characters flat. Punctures on head and pronotum slightly even only in color pattern. deeper, those on scutum more or less defined and with central postules. Head yellowish brown or pale ferruginous ...... R. principalis new species Ropalidia maculiventris Guérin-Méneville – Smaller species; body length (head + mesosoma + Ropalidia maculiventris Guérin-Méneville, 1831: pl. 9 fig. 8 – first two metasomal segments) 9–10 mm, forewing “Dory” [= Manokwari, New Guinea] (holotype in the length 9.5–10 mm. First metasomal tergum in Museo Civico di Storia Naturale, “Giacomo Doria”, profile more distinctly swollen dorsally near level of Genova); 1838: 266 (Rhopalidia [!]). — Ducke, 1913: spiracle rather than evenly swollen posteriorly from 331 (Rhopalidia [!]; correct name of Icaria maculipennis posterior margin of basal slit for reception of [!] de Saussure). — Bequaert, 1932: 50. — van der Vecht, mesosomal suspensory ligament. Punctures on head 1941b: 28. — Guiglia, 1948: 177 (Rhopalidia [!]; type: a and mesosoma very sparse, shallow and ill defined. damaged specimen from “Austr. Voy. Coquille” ex coll. Head brown to dark brown...... 2 Guérin-Méneville in MCG). — Cheesman, 1952: 2, 4. Icaria maculiventris; de Saussure, 1853-58: 23. 2. Head in dorsal view with lateral sides behind eyes Icaria sexmaculata Cameron, 1911: 188, female – “Bivak barely convex (Fig. 22); gena in profile narrower, Island”. NEW SYNONYMY. — Kojima and Carpenter, slightly more than half as wide as eye. Space 1997: 5 (designation of lectotype). between posterior ocelli slightly raised into short, Ropalidia sexmaculata; Bequaert, 1932: 51. — van der Vecht, longitudinal carina (Figs. 23, 24). First metasomal 1941b: 28. tergum in profile with dorsal and ventral margins Ropalidia maculiventris humboldti Cheesman, 1952: 5, more or less parallel in anterior half, then rather female – “Hollandia”. NEW SYNONYMY. abruptly swollen dorsally just behind spiracle (Fig. Ropalidia pratti Cheesman, 1952: 2, 6, female, male – 26) ...... R. loriana (du Buysson) “Papua, Kokoda”. NEW SYNONYMY. — Richards, – Head in dorsal view with lateral sides behind eyes 1978: 56 (in subgenus Ropalidia). weakly convex (Fig. 8); gena in profile about 0.65

44 J. Kojima

Ropalidia mondoensis Cheesman, 1952: 2, 6, female – Ropalidia fulvopruinosa; Bequaert, 1932: 50. — Richards, “Papua, Mafulu, 4000 ft”. NEW SYNONYMY. — 1978: 54 (nest), 56 (list), 63 (key), 67, figs. 12, 25, 33, 41 Richards, 1978: 56 (in subgenus Ropalidia). (in subgenus Ropalidia; redescription). Ropalidia humboldti; Richards, 1978: 56 (in subgenus Ropalidia). This species seems to be distributed in the southern Ropalidia maculiventris pratti; Kojima and Spradbery, 1987: coast of New Guinea, from southern coast of Vogel 601, figs. 1-7 (larva) — Spradbery, 1991: 351, 354 Kop Peninsula and Aru to the Fly River, and in Cape (colony population). York Peninsula in Australia. I examined, in addition to the holotype female The late J. van der Vecht, having examined the (ZMA: second to sixth metasomal segments lacking, as holotype of Ropalidia maculiventris, suggested me to mentioned in the original description), labeled “Nieuw treat all of I. sexmaculata, R. pratti, R. maculiventris Guinea-/Expeditie 1909 / [this slash is written on the humboldti and R. mondoensis as subspecies of R. label, not indicating line break] 5/Etna-baai [3°58’S, maculiventris. I examined the name-bearing types of 134°40’E; NW New Guinea]” “Ropalidia [in van der all of these taxa except for R. maculiventris. All of Vecht’s handwriting] / Leionotus / fulvopruinosus / them were so different in the marking pattern, body size Cam. Type /New Guinea [in Cameron’s handwriting] and developmental degree of propodeal apophyses that /Holotype ♀ sec. J. v. d. Vecht ‘33 [in van der Vecht’s they all could be treated as belonging to different handwriting]”, specimens from several localities in species. However, examination of many additional New Guinea and Australia (they will be referred to in specimens, as mentioned in the second part of this the second part of this series), and the most reliable series, showed that these differences are intergraded character distinguishing R. fulvopruinosa from R. and concluded to follow the late van der Vecht’s view; maculiventris is the combination of develop-mental however, these taxa are all synonymized under R. degree of the propodeal apophyses and yellow spots on maculiventris in stead of treating them as subspecies. them. In R. fulvopruinosa, the apophyses are never Types examined — NW New Guinea: ♀ (BMNH, well developed, and the yellow spots are always present holotype of Ropalidia maculiventris humboldti on them. On the other hand, in R. maculiventris, the Cheesman), labeled “Holotype” (BM type label), apophyses are usually well developed and yellow spots “Dutch New Guinea, Humboldt Bay, Hollandia, on them are found only when the apophyses are iv.1936, L. E. Cheesman B.M.1936-271”, “L. E. produced into sharp teeth. Cheesman det. 1948, maculiventris ssp. humboldti Chees. Type”, and “B.M. TYPE HYM 18.832”. SW Ropalidia wollastoni (Meade-Waldo) New Guinea: ♀ (ZMA, lectotype of Icaria sexmaculata Cameron, designated by Kojima and Icaria wollastoni Meade-Waldo, 1912: 448 (key), 449, female (in subgenus Icariastrum) – “Mimika River”. Carpenter (1997: 5)), labeled “Z. Nieuw Guinea Icaria sericea Cameron, 1911: 188, female – “Bivak Island” /Lorentz 1909-10/Bivak Eiland [= Bivak I.; 5°00’S, (lectotype Amsterdam). Junior primary homonym of I. 138°39’E]/II-’10 [last line vertically written]”, “6- sericea Cameron, 1905. — Kojima and Carpenter, 1997: maculata [in Cameron’s handwriting]”, “Icaria ♀ 5, 60 (designation of lectotype; syn. of Ropalidia /sexmaculata Cam/Holotype by design./J. v. d. Vecht wollastoni). 1933 [in van der Vecht’s handwriting]”, “Ropalidia Ropalidia wollastoni; Bequaert, 1932: 51. — van der Vecht, /sexmaculata (Cam.)/det. J. v. d. Vecht ’34 [in van der 1941b: 28. — Cheesman, 1952: 2, 6 (redescription of the Vecht’s handwriting]” and “Holotype”; 1 ♀ (ZAM, type). paralectotype of I. sexmaculata), Lorentz 1909-10, Beside the color pattern, such as wings very Noord Rivier, ix.[19]09. SE New Guinea: ♀ (BMNH, strongly infuscate, colored blue-black rather than dark holotype of R. pratti Cheesman), labeled “Holotype” brown, I could not find any characters that distinguish R. (BM type label), “Papua: Kokoda 1200 ft. vi.1933, L. wollastoni from R. maculiventris or R. fulvopruinosa. E. Cheesman B.M. 1933-427”, “pratti Chees. Type, L. E. Cheesman det. 195”, and “B.M. TYPE HYM Types examined — SW New Guinea: 3 ♀ 18.839”; ♀ (BMNH, holotype of R. mondoensis (BMNH, lectotype and paralectotypes of Icaria Cheesman), labeled “Holotype” (BM type label), wollastoni Meade-Waldo), Mimika R., vii.1910, A. F. “Papua, Mafulu 4000ft, i.1934, L. E. Cheesman B.M. R. Wollaston [for the label data, see the section 1934-321”, “mondoensis Ch. Type L. E. Cheesman det. “Designation of the lectotypes” in this paper]; ♀ 195”, and “B.M. TYPE HYM 18.833”. (ZMA, lectotype of Icaria sericea Cameron, 1911, designated by Kojima and Carpenter (1997: 5)), labeled Ropalidia fulvopruinosa (Cameron) “Z. Niuew Guinea/Lorentz 1909-10/Bivak Eiland I 10”, “sericea [in Cameron’s handwriting]”, “Icaria ♀ sericea Odynerus (Leionotus) fulvopruinosa Cameron, 1906b: 225, Cam 1911/Holotype by design. J. v. d. Vecht 1933 [in female – “Etna Bay”. van der Vecht’s handwriting]”, “= wollastoni M. Waldo

Ropalidia of New Guinea 45

1912 [in van der Vecht’s handwriting]”, and Icaria australis de Saussure, 1853, in de Saussure, 1853-58: “Ropalidia/wollastoni (M. Waldo)/= sericea (Cam.) 24, pl. 34 fig. 5, female – “Dorey. Nouvelle-Guinée”. 1911/nec. 1906/det. J. v. d. Vecht 1934”; 1♀ (ZMA; Icaria irritata Smith; Cameron, 1913: 76 [misidentification]. Kojima and Carpenter (1997: 5) stated that the Icaria bispinosa Meade-Waldo, 1912: 448 (key), 449, female lectotype of I. sericea was “probably unique”, but this (in subgenus Icariastrum) – “Mimika River”. NEW SYNONYMY. specimen is evidently from the type series and is a Ropalidia australis; Bequaert, 1932: 50. paralectotype of I. sericea), labeled “Z. Nieuw Ropalidia bispinosa; Bequaert, 1932: 50. — Richards, 1978: Guinea/Lorentz 1909-10/Bivak Eiland IX 09”. 56 (in subgenus Ropalidia). Ropalidia fasciata bispinosa; Cheesman, 1952: 2, 9. Ropalidia fasciata erratica Cheesman, 1952: 2, 9, female – Other species in the R. maculiventris-group “Cyclops Mt.” NEW SYNONYMY. Ropalidia bicolor (Smith) Ropalidia erratica; Richards, 1978: 56 (in subgenus Ropalidia). Icaria bicolor Smith, 1865: 90, female – “New Guinea”. Ropalidia bicolor; Bequaert, 1932: 50. — Cheesman, 1952: 2, In the MNHN, under “Icaria australia” there are 7 (redescription of the type). — Richards, 1978: 56 (in three females and a male: a female labeled “MUS. subgenus Ropalidia). PARIS/Dorey [= Manokwari]/Dumont d’Urv. 1841”, The OUM houses a single female labeled circle with “Dorey/d Urville”, “TYPE” and “Holotype” (circular), “N” [= New Guinea] (circular), “Icaria/australis/type Sauss. [in du Buysson’s hand- and “Icaria bicolor Sm.” This is presumably the writing]/H. DE SAUSSURE DET. [in print]”, that is holotype of I. bicolor Smith, although Smith (1865: 90) evidently the holotype of Icaria australis (see also Carpenter, 1999: 34). A male labeled “MUS. PARIS did not give any indication for the number of specimens lle he examined. Smith (1871) did not mention this /N Guinie/allemande/J. Bequaert 1909”, “Finschhafen species in his catalog of the aculeata wasps in India and [NE New Guinea]/Duitsch Nieuw/Guinea” and “Icaria the Eastern Islands while he listed all the other species /australis /Sauss./R. DU BUYSSON det 1909”, and two of his New Guinean Icaria. females (but they are actually R. irritata (Smith)), labeled “MUS. PARIS/NOUV. GUINÉE/I. SALA- WATI [NW New Guinea]/RAFFRAY & MAINDRON Ropalidia irritata (Smith) /1878” are apparently of later origin. Icaria irritata Smith, 1863: 42, female – “Mysol”. Cameron (1906a: 62) and Meade-Waldo (1912: Ropalidia irritata; Bequaert, 1932: 51. — Cheesman, 1952: 2, 448) recorded I. australis from “Moaif” and “Mimika 9 (redescription of type). — Richards, 1978: 56 (in River”, respectively, but the holotype of I. australis had subgenus Ropalidia). not long been referred to until Carpenter (1999). As the The OUM houses two females: one is labeled “M”, identification labels for R. dispila (Cameron, 1913) “Icaria irritata Smith”, “Icaria irritata Sm.”, “Type specimens show, the late van der Vecht regarded that R. locality = Misool, this true type is labeled “M” the other dispila was a synonym of R. australis (see the label data specimen is from “N” = New Guinea v. d. Vecht, 1958 for the specimens listed under R. dispila). Actually, [in van der Vecht’s handwriting]”; the other is labeled however, they are different species. “Type Mysol” “N” (circular), “Icaria irritata Sm.” As When he described I. bispinosa, Meade-Waldo suggested in van der Vecht’s label, the former is the (1912: 450) stated that “This species comes very near I. true type (and the holotype) of Icaria irritata because spilostoma, Cam. . . .”; however, I. spilostoma is a “M” of Smith’s label indicates “Mysol” [= Misool]. synonym of R. nigra (Smith, 1859), which is very different from R. australis in structure. Meade-Waldo’s Ropalidia incurva Cheesman remarks are based only on the coloration. Cheesman (1952) treated Meade-Waldo’s (1912) Ropalidia incurva Cheesman, 1952: 2, 7, female, male - “bispinosa” as a subspecies of R. fasciata (Smith, 1859) “Papua, Mondo, 4000 ft”. and described a new subspecies, R. fasciata erratica. The holotype female in the BMNH is labeled She (p. 8) examined and redescribed the holotype of “Holotype” (BM type label), “Papua: Mondo [8°35’S, Icaria fasciata Smith, 1859, and referred to the 147°18’E; SE New Guinea] 5000 ft. ii.1934, L. E. differences only in marking pattern to treat bispinosa as Cheesman B.M.”, “L. E. Cheesman det. 1948, incurva a subspecies of R. fasciata, and in the structure of Chees.” and “B.M. TYPE HYM 18.1033”. Cheesman scutellum and metanotum as well as marking pattern to (1952: 8) listed also a male from the same locality as describe R. fasciata erratica. She, however, overlooked the holotype, but I failed to locate this male specimen. the facts that the gena and the first metasomal tergum of the types of I. bispinosa and R. fasciata erratica are Ropalidia australis (de Saussure) distinctly proportionally wider than those of the type of

46 J. Kojima

I. fasciata. I could not find any differences among the 10”, “Mevr. de Beaufort leg”, “Holotype”, “Icaria [in holotypes of I. australis, I. bispinosa and R. fasciata Cameron’ handwriting]/dispila [in Cameron’s hand- erratica to treat them as different species. writing] ♀ [in van der Vecht’s handwriting]/Cam. Type [in Cameron’s handwriting]/Holotype sec. J. v. d. Other specimens examined — Waigeo: 1♀ Vecht ’33 [in van der Vecht’s handwriting]”. Aru: ♀ (NNM), “Waigeoe”, 5.i.[19]10, Mevr. de Beaufort (also (OUM, holotype of Icaria fasciata Smith, 1859), labeled “Icaria irritata Sm. ♂ !, det. Cam. [in labeled “Type” (BM type label), “Icaria fasciata Smith Cameron’s handwriting]” and “Ropalidia bispinosa [in Smith’s handwriting]” (blue); 12♀ (NNM, IUNH), (M.W.) var.? ♀, det. J. v. d. Vecht, 34 [in van der labeled “Ins. Aru” and “Museum Leiden, Ropalidia Vecht’s handwriting]”). NW New Guinea: ♀ australis Sauss. ♀ , det. J. v. d. Vecht, ‘33”; 1 ♀ (BMNH, holotype of Ropalidia fasciata erratica (NNM), labeled “Icaria fasciata Smith, Ins. Aru” and Cheesman), labeled “Holotype” (BM type label), “Museum Leiden, Ropalidia australis Sauss. ♀, det. J. “Dutch New Guinea, Cyclops Mts. Camp. 1: 1200 ft. v. d. Vecht, ‘33”; 3♀ (NNM), Rosemberg leg. [1♀ 22.v.1936 L. E. Cheesman B.M.1936-271”, “fasciata with a label “Ropalidia australis Sauss., fasciata Sm., Sm. erratica Ch., Type, L. E. Cheesman det., 1951”, dispila Cam.”, and 1♀ with a label “Ropalidia australis and “B.M. TYPE HYM 18.1038”. SW New Guinea: (Sauss.), J. v. d. Vecht”]; 10♀ (NNM), labeled “Ins. ♀ (BMNH, lectotype of Icaria (Icariastrum) bispinosa Aru, Rosemberg” and “Museum Leiden, Ropalidia Meade-Waldo) from Mimika River, labeled as shown in australis Sauss. ♀, det. J. v. d. Vecht, ‘33”. the section “Designation of the lectotypes”.

Ropalidia dispila (Cameron) Ropalidia mimikae (Meade Waldo) Icaria fasciata Smith, 1859: 167, female – “Aru”. Junior Icaria mimikae Meade-Waldo, 1912: 448 (key), 450, female secondary homonym of Ropalidia fasciata (Fabricius, (in subgenus Icariastrum) – “Mimika River”. 1804). Ropalidia mimikae; Bequaert, 1932: 50. — Cheesman, 1952: Icaria dispila Cameron, 1913: 77, female – “Waigeu”. 2, 9 (redescription of the type). Ropalidia dispila; Bequaert, 1932: 50. Although Icaria mimikae is the only species for Ropalidia fasciata; Bequaert, 1932: 50. — Cheesman, 1952: which Meade-Waldo (1912) did not give the number of 2, 8 (redescription of the type). specimens he examined, a female in the BMNH labeled Ropalidia fasciola Richards, 1978: 56. (in subgenus Ropalidia). Replacement name for Icaria fasciata Smith, “Holotype” (BM type label), “New Guinea Mimika R. 1859, non Eumenes fasciata Fabricius, 1804 [Richards [SW New Guinea], A. F. R. Wollaston, 1911-229 aug. sated “Icaria fasciata Smith, 1858]. NEW SYNONYMY. 1910”, “Icaria (Icariastrum) mimikae sp. n., Type, ♀, G. Meade-Waldo det.”, “Ropalidia mimikae Meade- Although Smith (1859) does not give any indication Waldo, L. E. Cheesman det., 1951”, “B.M. TYPE for the number of specimens, the specimen in the OUM HYM 18.852”, and “BM 79” (red) is evidently the is presumably a unique specimen on which his holotype of I. mimikae. description of Icaria fasciata was based, and thus the This specimen is the only known specimen for R. holotype. The specimen in the ZMA marked as type of mimikae; this species could be very rare. Icaria dispila is evidently the holotype as Cameron (1913: 77) mentioned that “The 2nd and following segments are absent.” Ropalidia conservator (Smith) Judging from the identification labels by the late van Icaria conservator Smith, 1860: 130, female – “Dory”. der Vecht, he regarded that both Icaria fasciata and I. Polybia limatula Smith, 1863: 43, female – “Mysol”. — dispila were the synonyms of Ropalidia australis; he, Meade-Waldo, 1911: 100 (syn. of Icaria conservator however, undoubtedly did not examine the holotype of Smith)). — van der Vecht, 1966: 6 (syn. of Ropalidia conservator (Smith)). — Kojima and Carpenter, 1997: 5 Icaria australis in the MNHN, and thus recognized R. (designation of lectotype). bispinosa Meade-Waldo as a good species (see the Icaria irritata Smith; Cameron, 1906a: 63 [misidentification]. identification label for a specimen from Waigeo listed Icaria parvimaculata Cameron, 1911: 189, female – “Bivak in the section for R. australis). I did not find any Island”. Junior primary homonym of I. parvimaculata differences between the holotypes of I. fasciata and I. Cameron, 1907. — van der Vecht, 1934: viii (syn. of dispila to reject the late van der Vecht’s opinion that the Ropalidia conservator (Smith)). — Kojima and two taxa belong to the same species. Here I propose to Carpenter, 1997: 5 (designation of lectotype). synonymize R. fasciola Richards under R. dispila Icaria insularis Cameron, 1911: 188, female – “Bivak Island”. (Cameron). — van der Vecht, 1934: viii (syn. of R. conservator (Smith)). Specimens examined — Waigeo: 1♀ (ZMA, Parapolybia limatula; von Schulthess, 1913: 164 (syn. of holotype of Icaria dispila Cameron; second to sixth Icaria conservator). metasomal segments lacking), labeled “Waigeoe 5. 1.

Ropalidia of New Guinea 47

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, vertically written]”, “parvimaculata [in Cameron’s 1934: viii. — Cheesman, 1952: 2, 15 (redescription). — handwriting]” and “Icaria ♀ /parvimaculata Cam Richards, 1978: 54, 59, 130 (in subgenus Icarielia). /Holotype sec. J. v. d. Vecht ’33 [in van der Vecht’s Ropalidia insularis; Bequaert, 1932: 50. handwriting]”; 1♀ (ZMA, paralectotype of Icaria Ropalidia parvimaculata; Bequaert, 1932: 50. parvimaculata; damaged specimen), labeled “Z. Nieuw Polistes tasmaniensis de Saussure; von Schulthess, 1932: 39 Guinea/Lorentz 1909-10/Bivak Eiland/II-’10 [last line [misidentification]. Ropalidia conservator goodfellowi Cheesman, 1952: 2, 16, written vertically]”, “parvimaculata [in Cameron’s female – “S. Dutch New Guinea, Mimika R.”. handwriting]” and “Icaria ♀ Cam. /parvimaculata Ropalidia conservator waigeuensis; Cheesman, 1952: 2, 16. /Paratype [in van der Vecht’s handwriting]”; ♀ (ZMA, Ropalidia conservator form goodfellowi; Richards, 1978: 130. holotype of Icaria insularis Cameron), labeled Ropalidia waigeuensis; Richards, 1978: 59 (syn. of R. “Holotype”, “Z. Nieuw Guinea/Lorentz 1909-10/Heurel conservator (Smith)). Bivak/XI-09/750 m [last two lines are written Ropalidia conservator form waigeuensis; Richards, 1978: vertically]”, “insularis [in Cameron’s handwriting]”, 130. “Icaria ♀/insularis Cam./HOLOTYPE/designated by/J. In addition to the specimens listed below, I v. d. Vecht 1933 [in van der Vecht’s handwriting]”; ♀ examined many specimens from various localities (they (BMNH, holotype of Ropalidia conservator good- will be referred to in the second part of this series); fellowi Cheesman), labeled “Type” (BM type label), these specimens showed so large variations in the “New Guinea Mimika R. A. F. R. Wollaston 1911-229”, marking pattern and the shape of the first metasomal “L. E. Cheesman det. 1948 conservator Sm. ssp. segment that the differences among the types of above goodfellowi Chees.” and “B. M. TYPE HYM 18.850”; listed taxa are intergraded. 2♀ (BMNH, paratypes of R. conservator goodfellowi), Mimika R., vii.1910, A. F. R. Wollaston. Specimens examined — Waigeo: ♀ (ZMA, holotype of Icaria waigeuensis Cameron), with labels Ropalidia novaeguineae (von Schulthess) “Holotype”, “Waigeoe 31 XII 09”, “Mevr. de Beaufort leg”, and a label on which are Cameron’s handwriting Parapolybia novaeguineae “Saussure nom. mus.” von “Icaria/waigionensis/Cam. Type [in Cameron’s hand- Schulthess, 1913: 154 (key), 162 (description based on writing]” and van der Vecht’s handwriting “= Icaria notes received from Meade-Waldo) – “Neu-Guinea”. Polybia novae-guineae du Buysson, 1913: 297, female – /conservator/Smith”. Misool: ♀ (OUM, lectotype of locality not stated, but evidently from New Guinea. Polybia limatula Smith, designated by Kojima and Junior secondary homonym of Ropalidia novaeguineae Carpenter (1997: 5)), labeled “M” [= Mysol], “Polybia (von Schulthess). — van der Vecht, 1966: 6 (syn. of limatula Smith” and “Lectotype desig. J. v. d. Vecht Ropalidia novaeguineae (von Schulthess)). Aug. 1958”; 1♀ (OUM; Kojima and Carpenter (1997) Ropalidia novae guineae [!]; Cheesman, 1952: 3, 23 did not refer to this specimen when they designated the (redescription of a type of “Polybia novaeguineae du lectotype of Polybia limatula, but this specimen is Buysson”). evidently from the type series of P. limatula and is now Ropalidia novaeguineae; van der Vecht, 1966: 6. — Richards, a paralectotype), labeled “M” and “Polybia limatula 1978: 59 (in subgenus Icarielia; listed as R. novaeguineae Smith”. NW New Guinea: ♀ (BMNH, holotype of (von Schulthess)), 129 (R. novaeguineae (R. du Buysson) Icaria conservator Smith), labeled “Type” (BM type in a key to species of the subgenus Icarielia). label), “This must be regarded as the holotype of Ic. Based on the notes received from Meade-Waldo, conservator Sm. no other specimens from Dory being von Schulthess (1913, April) described Parapolybia preserved in Oxford! J. v. d. Vecht, 1934 [in van der novaeguineae, indicating “Neu-Guinea (Type Museum Vecht’s handwriting]”, “conservator Smith”, “Dor” [= London)”. Meade-Waldo’s notes was evidently based Dory] (circular), “Smith coll. press by Mrs. Farren on a female in the BMNH (second to terminal White 99-303”, “B.M. TYPE HYM 18.849”, and metasomal segments are separately attached to a small “Holotype Icaria conservator Smith, 1860, Teste C. piece of card board), that is labeled “Type H. T” O’Toole, 1993”; 1♀ (IRSNB), Sakoemi [probably near (circled with red), “New Guin.”, “Type. F. Sm. coll. Triton Bay], with an identification label as “Polistes 79.22”, “Polybia novae guineae Sauss. Type”, tasmaniensis” by von Schulthess; 1♀ (ZMA), labeled “Saussure’s name in ms. R. du Buysson det. Bull. ent. “Manokwari 27 mei [19]03”, “Icaria irritata Sm. Sec. Soc. Fr. p. 297 (1913) G. Meade-Waldo det.” and Cam. New Guinea [in Cameron’s handwriting]”. SW “B.M. TYPE HYM 18.876”. Thus this specimen is the New Guinea: ♀ (ZMA, lectotype of Icaria holotype of Parapolybia novaeguineae von Schulthess. parvimaculata Cameron, designated by Kojima and In July of the same year, du Buysson (1913) Carpenter (1997: 5)), with labels “Holotype [in van der described Polybia novaeguineae, mentioning “. . . n. sp., Vecht’s handwriting]”, “Z. Nieuw Guinea/Lorentz in mus. SMITH 1879. . . . Cet insecte est unique et il 1909-10/Heurel Bivak/XI 09/750 m [last two lines are porte de la main de SMITH l’étiquette: «Polybia Novae-

Ropalidia of New Guinea 49

Guineae Sauss» . . .” A female in the OUM labeled E. Cheesman B. M. 1933-427”, “ivorina Chees. L. E. “New Guinea” (circular), “Polybia Novae Guineae Cheesman det. 1951” and “BM TYPE HYM 18.1036”, Sauss.”, “Polybia novae-guineae Saussure et Smith, and a paratype (BMNH) of R. ivorina from the same type, det. 1912, R. du Buysson”, “Coll. Smith, 1879” is locality as the holotype, and found no specific evidently the holotype of Polybia novaeguineae du differences among them. I herewith propose to Buysson. There are no distinct differences between the synonymize R. ivorina under R. jaculator. holotypes of Parapolybia novaeguineae von Schulthess and Polybia novaeguineae du Buysson that suggest they Ropalidia semihyalinata (Meade-Waldo) belong to different species; van der Vecht’s (1966: 6) synonymy is confirmed. Icaria semihyalinata Meade-Waldo, 1912: 449 (key), 451, female (in subgenus Icariola) – “Mimika River”. Cheesman (1952: 23) mentioned “novae guineae du Ropalidia semihyalinata; Bequaert, 1932: 51. — Cheesman, Buyss., 1913 . . . Named by Saussure (Polybia novae 1952: 3, 24 (semihyalineata [!]; redescription). — guineae) but not described. Smith’s Collection. Richards, 1978: 56 (in subgenus Ropalidia). Collected by A. P. R. Wallace on Salawatti Is., W. New Ropalidia novaeguineae insolens Cheesman, 1952: 3, 24, Guinea. Type, B.M. . . . a redescription follows. . . . 1 female – “Japen, Mt. Baduri, 1000 ft.”. NEW ♀ (holotype), Smith’s collection, New Guinea, 1 ♀ , SYNONYMY. — Kojima and Carpenter, 1997: 6 labelled 62:91, which signifies 1862, Salawatti (designation of lectotype). (Accession Register).” The specimen that she Ropalidia insolens; Richards, 1978: 58 (in subgenus considered as the holotype of “novaeguineae du Icarielia), 129 (key). Buysson, 1913” is evidently the holotype of Meade-Waldo (1912: 451) clearly indicated that he Parapolybia novaeguineae von Schulthess, 1913. examined a single female for his Icaria semihyalinata Furthermore, neither von Schulthess (1913) nor du and the holotype in the BMNH is labeled “Holotype” Buysson (1913) referred to the exact locality or the (BM type label), “New Guinea, Mimika R. [SW New collector; Cheesman (1952) seemed to refer to the label Guinea], A. F. R. Wollaston, 1911-229 July 1910”, datum for the specimen labeled “62:91” as if it were for “Icaria (Icariola) semihyalinata sp. n. Type, G. Meade- the holotype of Parapolybia novaeguineae. Waldo det.”, and “B.M. TYPE HYM 18.874”. Richards (1978: 59, 129) placed this species in the I failed to locate the specimens from the type series subgenus “Icarielia”, but the species may be keyed to of Ropalidia novaeguineae insolens other than the the “subgenus Ropalidia” with his key to the subgenera lectotype designated by Kojima and Carpenter (1997: 6; of Ropalidia (p. 51). erroneously mentioning “Ropalidia insolens Cheesman, 1952”), which is labeled “Type” (BM type label), Ropalidia jaculator (Smith) “Dutch New Guinea, Japen Is., Mt. Baduri, 1000 ft., Odynerus fallax Smith, 1863: 40, female – “Mysol”. viii.1938, L. E. Cheesman, B.M. 1938-593”, “L. E. Odynerus jaculator Smith, 1871: 377. Replacement name for Cheesman det. 1948, insolens Chees.”, and “B.M. O. fallax Smith, 1863, non 1861. TYPE HYM 18.875”. A headless female (BMNH) Odynerus tertius Dalla Torre, 1889: 125. Unnecessary identified as insolens by Cheesman is from “Humboldt replacement name for Odynerus fallax Smith, 1863, non Bay”, the locality listed by Cheesman (1952: 24). 1861 and de Saussure, 1852. However, the collection date of this specimen is “vi Ropalidia ivorina Cheesman, 1952: 3, 25, female – “Papua, 1937”, while Cheesman mentioned “Humboldt Bay, ii Kokoda, 1200 ft.” NEW SYNONYMY. — Richards, 1936”, thus this specimen is not a paralectotype. 1978: 58, (in subgenus “Icarielia”), 129 (key to species in I could not find any specific differences among the the subgenus “Icarielia”). holotype of I. semihyalinata, the lectotype of R. Ropalidia jaculator; Kojima and Carpenter, 1997: 6. novaeguineae insolens and a female identified as Since Smith (1871) replaced his Odynerus fallax “insolens” by Cheesman; R. novaeguineae insolens is a with Odynerus jaculator this species had not long been synonymy of R. semihyalinata. On the other hand, they referred to until Kojima and Carpenter (1997: 6); are different from the types of R. novaeguineae in the exceptions are Dalla Torre (1889, 1894, 1904), but he shape of the first metasomal segment (details will be certainly did not see any specimens of this species. The given in the second part of this series). holotype of Odynerus fallax that Kojima and Carpenter Cheesman (1952) described R. novaeguineae (1997: 6) examined is in the OUM and labeled “M” [= insolens in the “Group VIII”, which included also R. Mysol] (circular), “Odynerus fallax Smith”, “Icaria novaeguineae, R. semihyalinata and R. ivorina. tertia DT”, and “Type 091 Hope Dept. Oxford”. I Richards (1978) raised insolens to the species rank and compared this specimen with the holotype of Ropalidia placed it in “subgenus Icarielia”, in which he placed ivorina (BMNH), labeled “Type” (BM type label), also R. novaeguineae and R. ivorina. On the other hand, “Papua: Kokoda [SE New Guinea] 1200 ft VI 1933 L.

50 J. Kojima

Richards (1978) placed R. semihyalinata in “subgenus Ropalidia brunnea; Bequaert, 1932: 51. — van der Vecht, Ropalidia”. 1934: viii. — Cheesman, 1952: 2, 14 (redescription of the type) [error: year of publication, 1864]. — Richards, 1978: 57 (in subgenus Polistratus; list; syn.: cariniscutis Species in the R. bidens-group (Cameron), alleni Cheesman) [alleni as syn., apparently by error; see also Kojima and Carpenter, 1997: 16], 121 Ropalidia bidens van der Vecht (key). Ropalidia bidens van der Vecht, 1996, in van der Vecht and Ropalidia brunnea var. cariniscutis; van der Vecht, 1934: viii. Kojima, 1996: 99, female, male – “N. W. New Guinea, Ropalidia cariniscutis; Bequaert, 1932: 51. — Richards, Klamono Oilfds [1°08’S, 131°30’E].” 1978: 121 (syn. of R. brunnea (Smith)). Ropalidia wudai Cheesman, 1952: 2, 14, female – “Papua, The holotype female (NNM) is kept in the type Kokoda, 1200 ft.” NEW SYNONYMY. — Richards, collection separately from the ordinary specimens, and 1978: 57 (in subgenus Polistratus). labeled “NW New Guinea/Klamono Oilfds/18-24.viii. Ropalidia wudai trullissima Cheesman, 1952: 2, 15, female – 1948/M. A. Lieftinck”. The paratype male and female “Dutch New Guinea, Cyclops Mts., 930 ft.” NEW in the BISH are labeled as follows: male: “NEW SYNONYMY. — Richards, 1978: 57 (in subgenus GUINEA: NETH./Bodem, 100 m, 11 km/SE of Polistratus). Oerberfaren/July 7-17, 1959” and “T. C. Maa/Collector The OUM houses a female that was referred to by /BISHOP”; and female paratype: “NEW GUINEA: Cheesman (1952: 14) and is labeled “Paratype O. U. NETH/Boden, 100 m, 11 km/SE of Oerberfaren/July 7- M.”, “M”, “N. G. Ins.” and “Icaria brunnea Sm”. “M” 17, 1959/T. C. Maa” is written in Smith’s handwriting and indicates that the specimen was from “Mysol” [= Misool]; thus the Ropalidia clypeata Kojima specimen is not from the type series of I. brunnea. A Ropalidia clypeata Kojima, 1996, in van der Vecht and female in the BMNH labeled “Aru” is evidently the Kojima, 1996: 103, male, female – “New Guinea, Sivipi holotype of I. brunnea. near Sasambata, Popondetta subdist.” Cameron (1906a: 59) did not mention the sex and The holotype male (NNM) is kept in the type the number of specimens when he described Polistratus collection separately from the ordinary specimens, and cariniscutis. A female in the ZMA is certainly the labeled “Sivipi near Sasambota/Popondetta Subdist. unique specimen as Kojima and Carpenter (1997: 12) /Papua 1 Nov. 1963/D. K. McAlpine”, “Austr. Mus. mentioned. Collection” and “Museum Leiden/ex. collectie/J. v. d. In addition to the holotypes of I. brunnea and P. Vecht”. I failed to locate the lost female paratypes. 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 Species in the R. nigra-group differences in the marking pattern that may diagnose any of them. I synonymize R. wudai and R. wudai Ropalidia bambusae Richards trullissima under R. brunnea. Ropalidia bambusae Richards, 1978: 54 (nest), 57 (list), 65 Types examined —NW New Guinea: ♀ (ZMA, (larva in key), 122 (key), 123, female, male, nest (in subgenus Polistratus) – “New Guinea: Morobe District, holotype of Polistratus cariniscutis Cameron), labeled Wau”. “Manokuwari 2 mei 03”, “Ropalidia brunnea (Sm.) var. cariniscutis Cam. ♀/det J. v. d. Vecht ’34 [in van der This species is certainly a valid species. Vecht’s handwriting]” and “Polistratus cariniscutis Cam. Types examined — NE New Guinea: ♀ (BMNH, Type/New Guinea [in Cameron’s handwriting] holotype), labeled “Holotype” (BM type label), “NG42 /Holotype by designation J. v. d. Vecht 1933 [in van der nest, New Guinea: Morobe district, Wau, 5.i.1973, Vecht’s handwriting]”; ♀ (BMNH, holotype of O.W. Richards”, and “Ropalidia bambusae Richards ♀, Ropalidia wudai trullissima Cheesman), labeled Holotype”; 1 ♂ (NNM, paratype), same data as the “Holotype” (BM type label), “Dutch New Guinea, holotype; 1 ♀ (NNM, paratype), nest NG6, Wau, Cyclops Mts. Sabron 930 ft. iv.1936, L. E. Cheesman 20.xii.1973, O. W. Richards. B.M. 1936-271”, “L. E. Cheesman det. 1949, wudai Chess. ssp. trullissima Chess.”, and “B.M. TYPE HYM Ropalidia brunnea (Smith) 18.861”; 1♀ (BMNH, paratype of R. w. trullissima), same data as the holotype of R. w. trullissima. Aru: ♀ Icaria brunnea Smith, 1859: 167, female – “Aru”. (BMNH, holotype of Icaria brunnea Smith), labeled Polistratus cariniscutis Cameron, 1906a: 59, [female] – “Holotype” (BM type label), “Aru” (blue, circular), “Manokwari”. — van der Vecht, 1934: viii (syn. of R. brunnea (Smith)). “Icaria brunnea Smith” (blue), “Smith coll. press by Mrs. Farren White 99-303”, and “B.M.TYPE HYM

Ropalidia of New Guinea 51

18.863”. SE New Guinea: ♀ (BMNH, holotype of Ropalidia melania Richards Ropalidia wudai Cheesman), labeled “Holotype” (BM Ropalidia melania Richards, 1978: 55 (nest), 57 (list), 65 type label), “Papua: Kokoda 1200 ft. viii.1933, L. E. (larva in key), 122 (key), 126, female, nest (in subgenus Cheesman B.M. 1933-427”, “L. E. Cheesman det. Polistratus) – “New Guinea: Morobe District, Wau”. 1948, brunnea Sm. wudai Chess.”, and “B.M. TYPE This species certainly a valid species, distinguished HYM 18.862”; 1♀ (BMNH, paratype of R. wudai), from other species in the R. nigra-group by a wide same collection data as the holotype of R. wudai. median impression at the occipital margin as noted in the original description by Richards (1978). Ropalidia conspicua (Smith) Types examined — NE New Guinea: ♀ (BMNH, Odynerus conspicuus Smith, 1863: 40, female – “Mysol” holotype), labeled “Holotype” (BM type label), “Nest Odynerus mysolicus Dalla Torre, 1889: 124. Replacement NG13, New Guinea, Morobe district, Wau, 24.xii.1972, name for Odynerus conspicuus Smith, 1863, non de O.W. Richards”, “Ropalidia melania Richards, ♀ , Saussure. [Unnecessary replacement, incorrectly ♀ considering de Saussure, 1870, as de Saussure, 1857]. holotype”; 1 (paratype, NNM), same collection data Ropalidia conspicua; Cheesman, 1952: 2, 13 (redescription of as the holotype. the type). Ropalidia conspicua alleni Cheesman, 1952: 2, 13, female – Ropalidia nigra (Smith) “Mysol”. — Kojima and Carpenter, 1997: 16 (syn. of R. Icaria nigra Smith, 1859: 167, female – “Aru”. — Meade- conspicua, without indicating a new synonymy). Waldo and Morley, 1914: 406 (syn.: Ancistrocerus Ropalidia alleni; Richards, 1978: 57 (syn. of R. brunnea catharinae (Cam.) as var.). — Kojima and Carpenter, (Smith) [this treatment was apparently based on 1997: 4 (designation of lectotype). erroneously taking “conspicua alleni” as “cariniscutis Icaria morosa Smith, 1863: 42, female – “Waigiou”. — alleni”]). Cheesman, 1952: 10 (syn. of Ropalidia nigra (Smith)). Ropalidia mysolica; Richards, 1978: 57 (in subgenus Icaria spilostoma Cameron, 1906a: 62, female, male – Polistratus; Odynerus conspicuus as a synonym). “Manokwari”. — van der Vecht, 1934: viii (syn. of Carpenter (1999: 34) referred to the holotype of Ropalidia nigra (Smith)). — Cheesman, 1952: 10 (syn. of Odynerus conspicuus Smith, stating “In the OUM are Ropalidia nigra (Smith)). — Kojima and Carpenter, 1997: 5 (designation of lectotype). two females marked as types, labeled with type Odynerus sariensis Cameron, 1906a: 63, female – “Siari”. — numbers 097 and 098. The first has no locality label, van der Vecht, 1934: viii (syn. of Ropalidia nigra and the determination label states, “By mistake (Smith)). — Kojima and Carpenter, 1997: 5 (designation described/as an Odynerus.” Number 098 is from “New of lectotype). Guinea”, and has a labed [sic] . . . stating, “Not a type!” Odynerus confraternus Cameron, 1911: 192, female – As the locality is incorrect, number 098 is a pseudotype. “Lorentz River”. — van der Vecht, 1934: viii (syn. of Number 097 is therefore apparently the holotype.” I Ropalidia nigra (Smith)). — Kojima and Carpenter, 1997: examined both specimens and number 097 is labeled 5 (designation of lectotype). “Holotype O.U.M” (circled with red), “M” [= Mysol] Icaria confraternus; Meade-Waldo and Morley, 1914: 506. (circular label), “Holotype”, “Icaria conspicuus by Ancistrocerus catharinae Cameron, 1913: 78, female – mistake described as an Odynerus” (blue label), and “Waigeu”. — Meade-Waldo and Morley, 1914: 406 (var. of Icaria nigra). — van der Vecht, 1934: viii (syn. of “Type 097 Hope Dept. Oxford”, and number 098 is Ropalidia nigra (Smith)). — Kojima and Carpenter, 1997: with labels “New Guinea” (circular), “Odynerus 4 (designation of lectotype). conspicuus Smith” (blue), “Icaria”, “Not a type!” and Icaria nigra var. catharinae; Meade-Waldo and Morley, “Type 098 Hope Dept. Oxford”. Carpenter overlooked 1914: 406. the locality label for number 097 specimen, which Ropalidia confraterna; Bequaert, 1932: 50. definitely supports his interpretation. Ropalidia nigra; Bequaert, 1932: 50. — van der Vecht, 1934: Although Cheesman (1952: 13) stated that the viii (syns.: Icaria spilostoma Cameron, Odynerus holotype of R. conspicua alleni was “labeled by Smith sariensis Cameron, Odynerus confraternus Cameron, “Odynerus conspicuus type, Mysol Island”, this speci- Ancistrocerus catharinae Cameron). — Cheesman, 1952: men in the BMNH is actually labeled “Type” (BM type 2, 10 (redescription of the type female, description of label), “S” (circular), “Smith coll. press by Mrs. Farren male; syns.: Icaria morosa Smith, Icaria spilostoma Cameron; distribution [erroneously recorded from White 99-303”, “Odynerus conspicuus Smith, New Makassar in Celebes]). — Richards, 1978: 57 (in Guinea, Mysool”, “conspicua Sm. ssp. alleni Chees., L. subgenus Polistratus; syn.: spilostoma (Cameron)), 121 E. Cheesman det. 1950”, and “B.M.Type HYM 18.860”. (key). Cheesman (1952) overlooked the locality label “S” [= Ropalidia spilostoma; Bequaert, 1932: 51. Salwatty = Salwati]; NW New Guinea]. Furthermore, Ropalidia morosa; Bequaert, 1932: 51. the fourth label does not include “type”. Ropalidia domestica Cheesman, 1952: 2, 12, female, male – “Papua, Kokoda”. NEW SYNONYMY. — Richards,

52 J. Kojima

1978: 54 (nest), 57 (in subgenus Polistratus), 122 Kojima and Carpenter (1997: 4)), labeled “Waigeoe/4. (holotype stated to be female; redescription). 1. 10”, “Mevr de/Beaufort/Leg” and “Icaria [in van der Ropalidia sepicana Richards, 1978: 57 (list), 122 (key), 127, Vecht’s handwriting]/Ancistrocerus [in Cameron’s figs. 22, 27, female, male (in subgenus Polistratus) – handwriting]/Beauforti/Cam. Type [in Cameron’s hand- “New Guinea: Sepik district, Sepik River, Kondanggei”. writing]/Holotype sec. J. v. d. Vecht ’33 [in van der NEW SYNONYMY. Vecht’s handwriting]”. NW New Guinea: ♀ (BMNH, I examined many specimens from various localities lectotype of Icaria spilostoma Cameron, designated by in addition to the types of all the six species here treated Kojima and Carpenter (1997: 5)), labeled “Manokwari, as synonyms of R. nigra (Smith) and that of Icaria 23 mei 03”, “Icaria spilostoma Cam. Type [in nigra. Despite the fact that some of these taxa can be Cameron’s handwriting]”, “B.M. TYPE HYM 18.873a” diagnosed by minor characters when only the type and “Ropalidia spilostoma (Cam.) Lectotype [in specimens are concerned, additional specimens showed Richards’ hand-writing]”; 2♂ (ZMA, paralectotype of that they are intergraded as given in the second part of Icaria spilostoma), Manokwari, 2.v.[19]03; 1♂ (ZMA, this series. Van der Vecht (1934: viii) synonymized paralectotype of Icaria spilostoma), Manokwari, Cameron’s four species under R. nigra (smith), and 15.v.[19]03; 1 ♀ (ZMA, lectotype of Odynerus Cheesman (1952: 10) synonymized Smith’s species I. sariensis Cameron, designated by Kojima and morosa under R. nigra. They have been, since then, Carpenter (1997: 5)), labeled “Holotype”, “Sari [= treated as the synonyms of R. nigra (Smith). I agree Siari; 1°58’S, 131°58’E]/12. II 03” and “Odynerus with their treatments. Cheesman (1952: 12), however, /Sariensis/Cam. Type/New Guinea [in Cameron’s described Ropalidia domestica in comparison with R. handwriting]/Holotype sec. J. v. d. Vecht [in van der nigra; besides minor differences in the marking pattern, Vecht’s handwriting]”. Aru: ♀ (OUM, lectotype of a distinct difference recognized in her descriptions is Icaria nigra Smith, designated by Kojima and “width of 3rd [antennal flagellar article] 1/3 of 9th” (p. Carpenter (1997: 4)), labeled “Aru” (circular), “Type” 10) for the type of I. nigra and “3rd segment width 2/3 (BM type label), “Icaria nigra Smith”, “Icaria nigra that of 9th” (p. 12) for R. domestica. The types Sm.” and “Lectotype v. d. Vecht desig. 1958”; 1♀ (lectotype and paralectotype) of I. nigra, however, do (OUM; Kojima and Carpenter (1997: 4), when they not have antennae so strongly dilated apically. designated the lectotype of Icaria nigra, stated Richards (1978: 57) listed only I. spilostoma as a “probably unique”, but this specimen is presumably synonym of R. nigra and regarded R. domestica and his from the type series and is a paralectotype of I. nigra), species R. sepicana as good species; he did not refer to labeled “I. Aru” (circular), “Icaria nigra Isl. Aru”, the other species of Cameron and Smith that had been “Smith coll. press by Mrs. Farren White 99-303” and synonymized by van der Vecht (1934) and Cheesman “Icaria nigra Sm”. SW New Guinea: ♀ (ZMA, (1952), respectively. The grounds on which Richards lectotype of Odynerus confraternus Cameron, recognized R. domestica and R. sepicana as good designated by Kojima and Carpenter (1997: 5)), labeled species are given in his key to species of the “subgenus “Z. Nieuw Guinea/Lorentz 1909-10/Noord Rivier IX. Polistratus” (Richards, 1978: 121-122). In the couplet 09”, “HOLOTYPE”, “Odynerus confraternus Cam. 3, he mentioned that R. nigra has the eyes “bare”, the Type [in Cameron’s handwriting; “Odynerus” is lined widened part of the first metasomal tergum “4 times as on it with pencil and “Icaria” in van der Vecht’s wide as long” and “half as wide as tergite 2”, and the handwriting is above it]/HOLOTYPE ♀ SEC J. v. d. scutum “entirely dull”. On the other hand, he stated Vecht 1933 [in van der Vecht’s handwriting]”. NE that R. domestica and R. sepicana have their eyes New Guinea: ♀ (ANIC, holotype of Ropalidia “usually with a few hairs”, and the widened part of their sepicana Richards), labeled “NEW GUINEA first metasomal tergum is “5 times as wide as long”. /Kandanggei,/Sepik River/29 Feb. 1964/D. H. Colless” However, the types (lectotype and paralectotype) of and “Ropalidia / sepicana / ♀ Rich. / holotype [in Icaria nigra Smith have eyes possessing sparse hairs. Richards’ handwriting]”; 1♀ (NNM, paratype of R. Furthermore, their first metasomal tergum is wider than sepicana), same collection data as the holotype of R. half the width of the second tergum. Richards’ sepicana. SE New Guinea: ♂ (BMNH, holotype of measurements of the parts of metasomal segments are Ropalidia domestica Chessman), labeled “Holotype” so poorly defined that nobody can repeat his (BM type label), “Papua: Kokoda 1200 ft. iv.1933, L. measurements. E. Cheesman B.M. 1933-427”, “L. E. Cheesman det. Types examined — Waigeo: ♀ (OUM, holotype 1949, domestica Chess. ♂”, and “B.M. TYPE HYM of Icaria morosa Smith), labeled “Wag” [= Waigiou] 18.1037”; 2 ♀ (BMNH, paratypes of R. domestica), (circular), “Icaria morosa Smith” and “= nigra Smith, same collection data as the holotype of R. domestica; 1 L. E. Cheesman, 1949”; ♀ (ZMA, lectotype ♀ (BMNH, paratype of R. domestica), same collection Ancistrocerus catharinae Cameron, designated by data as the holotype of R. domestica but vi-vii.1933.

Ropalidia of New Guinea 53

Ropalidia leopoldi and its related species Kokoda. It is a very different species and agrees better (or species in the R. flavopicta-group) with the description. I have transferred the type label to this specimen”. However, the type label was not Ropalidia leopoldi Bequaert transferred, but Richards merely added his own labels Ropalidia leopoldi Bequaert, 1932: 47, female - “Nouvelle- to the specimens: one of the two females is labeled Guinée hollandaise. Lac d’Angi-Gita” [= Anggi Gita; “Type” (BM type label), “Papua, Kokoda, 1200 ft. IX. 1°23’S, 133°58’E]. — Richards, 1978: 58 (in subgenus 1933, L. E. Cheesman, B.M. 1934-321”, “L. E. Icarielia; list), 129 (key). Cheesman det. 1948, integra Chees.”, “B.M. TYPE Icaria spec.; von Schulthess, 1932: 40 (Nouvelle-Guineé, Lac HYM 18.1035” “BM ‘79” (red), “This is not type of de Angi-Gita). integra OWR, 1974 [in Richards’ handwriting]”; and This species is included in a key to species of the other is with “Paratype” (circled with yellow), “subgenus Icarielia” by Richards (1978: 129). As “Papua: Kokoda, 1200 ft. viii.1933., L. E. Cheesman, discussed in detail in the second part of this series, in B.M. 1933-427”, “L. E. Cheesman det. 1948, integra New Guinea there could be several species of this Chees.”, “♂”, “This is the real type of integra OWR species-group. I failed to locate any specimens of R. ‘74, ♀ ”. There is no grounds to accept Richards’ leopoldi that Richards (1978) used to produce the key. interpretation even if the specimen labeled “Paratype” Types examined — NW New Guinea: ♀ (IRSNB, were to agree better with the original description than holotype), labeled “Lac d’Angi-Gita/(Nouvelle Guinée) the specimen labeled “Type” does. Only one fact that /10-III-1929/Prince Léopold”, “Ropalidia/leopoldi J. may support Richards’ interpretation is that Cheesman Beq./Holotype [in red]/Dr. J.C.Bequaert det., 1932 [in (1952: 23) listed the material as “1♀ (holotype), 1♂ Bequaert’s (?) handwriting except the printed last (paratype) . . . viii., ix. 1933”. If this means that “♀ and line]”, and “cf. Mem. Mu. Hist. Nat. Belg. (hors serie) ♂ was collected in August and September, IV 5 (1932) p.47, fig. 1, A-D”; 40 ♀ (paratypes, respectively”, the holotype must be labeled “viii. 1933”. IRSNB; 2 ♀ in MCZ), same collection data as the The specimen labeled “paratype” was collected in “viii. holotype. 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 Ropalidia deminutiva Cheesman (1952) listed the dates of collection chronologically. Ropalidia deminutiva Cheesman, 1952: 3, 23, female – “S. of The holotype of R. integra should be the specimen Humboldt Bay, inland 16 miles, Mt. Nomo”. — Richards, collected in “ix.1933” and a specimen labeled “♂” is 1978: 54 (diminutiva [!]; nest), 58 (in subgenus Icarielia), the paratype. The holotype agrees well in structure 129 (key). with the holotype of R. deminutiva Cheesman, and R. Ropalidia integra Cheesman, 1952: 3, 22, female, “male” – integra is synonymized under R. deminutiva. The “Papua, Kokoda, 1200 ft”. NEW SYNONYMY. — paratype of R. integra is not, as Richards pointed out, Richards, 1978: 58 (in subgenus Icarielia), 129. conspecific with the holotype of R. integra, but its Richards (1978: 129) noted that “All the specimens definitive affiliation could not be determined because [of deminutiva] . . . labelled Bougainville, Mt. Nomo. I its metasoma is missing. Richards’ (1978: 128) “R. cannot trace Mt Nomo, but I suspect that Bougainville integra” in his key to species of “subgenus Icarielia” is an error, possibly for Mt. Bougainville . . . in West was based on the paratype of R. integra, which lacks Irian.” I examined the holotype and a paratype of R. the metasoma. 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 Species in the R. lefebvrei-group “L. E. Cheesman B.M. 1936-271”; the holotype is also Ropalidia lefebvrei (Le Guillou) with “Type” (BM type label), “Ropalidia diminutiva TYPE [in Cheesman’s (?) handwriting] Det. L. E. Polistes lefebvrei Le Guillou, 1841a: 325, female – “Triton- Bay”; 1841b: 322. Cheesman 195”, “B.M. TYPE HYM 18.1034”; and the ?Icaria lefebvrei; de Saussure, 1853-58: 241 (unidentified paratype is with additional labels “cotype”, “Ropalidia species). deminutiva Chees. Det. L. E. Cheesman”. Richards Icaria lefebvrei; Smith, 1857: 99 [Kojima and Carpenter (1978) misread the locality label. (1997: 33) listed the pagination as “95” by error]. Richards (1978: 129) referred to the types of R. Ropalidia lefebvrei; Bequaert, 1932: 50. integra Cheesman and stated that “The specimen in the ? Icaria reactionalis de Saussure, 1853, in de Saussure, 1853- BMNH labeled type is a female from Kokoda [SE New 58: 28, female – “La Nouvelle-Guinée” (coll. Guérin- Guinea] but is, I think, a specimen of R. deminutiva Méneville, current depository unknown). Cheesman. The paratype, though labelled male, is ? Ropalidia reactionalis; Bequaert, 1932: 51. — Cheesman, actually a female without a gaster, also coming from 1952: 3, 22 (group of festina; female).

54 J. Kojima

Le Guillou published the name Polistes lefebvrei Méneville were not deposited in the MHN (see Horn et twice in 1841. Although the exact dates of these two al., 1990). I failed to locate the type(s) of I. publications are not known, I take Le Guillou (1841a) reactionalis. as the original description of P. lefebvrei; in Le Guillou (1841b: 322) the author name of P. lefebvrei is clearly Ropalidia gemmea Cheesman indicated as “Le Guillou” (see also Kojima and Ropalidia gemmea Cheesman, 1952: 3, 22, female, male – Carpenter, 1997: 33; Carpenter, 1999: 36). In the “Papua, Owen Stanley Mts., Kokoda, 1200 ft.” — MNHN, there is a female labeled “MUSEUM Richards, 1978: 58 (in subgenus Icarielia), 129 (key). PARIS/Triton bay [3°45’S, 134°10’E; NW New Guinea]/LE GUILLOU”, “114”, “Type”, “314 - I examined: the holotype (in the BMNH) labeled Polistes ne 18/Lefebvrei - Leg./Triton bay:” and “Ann. “Papua: Kokoda [SE New Guinea], 1200 ft., ix.1933, Soc. Ent. 1841/n. 322/no. 26/par le 8 Le Guillou”, that L. E. Cheesman, BM 1934-321” “L. E. Cheesman det. is evidently the holotype of P. lefebvrei (see also 1950, TYPE, gemmea Chees.” and “B.M. TYPE HYM Carpenter, 1999: 36). 18.1032”; a paratype female (BMNH) from the same De Saussure (1853-58: 241) referred to Icaria locality as the holotype; and a headless paratype lefebvrei (Le Guillou) as an unidentified species, and (NNM) labeled “Para-/type” (BM type label), “PAPUA: this is the only citation of I. lefebvrei by de Saussure. Kokoda. /1,200ft. vi.1933/L. E. Cheesman. /B.M.1933- Thus, de Saussure should not have seen the type of 427”, “Museum Leiden/ex. coll. J. v. d. Vecht”, “L. E. Polistes lefebvrei. Du Buysson (1911: 229) referred to Cheesman det. 1950/gemmea Chees”. They are the holotype of Polistes lefebvrei, but he never cited presumably conspecific, and I concluded that R. Icaria reactionalis de Saussure, 1853. Although gemmea is a valid species. Bequaert (1932: 50 and 51) cited both species simply listing them as species so far recorded from New Ropalidia catharinae (Cameron) Guinea, it seems highly probable that he did not see the Icaria catharinae Cameron, 1913: 77, female – “Waigeu”. — types of either taxa. Cheesman (1952: 21) cited I. Meade-Waldo and Morley, 1914: 406. — van der Vecht, reactionalis when she redescribed Icaria festina Smith, 1934: viii. 1864, as a subspecies of Ropalidia reactionalis (de ? Ropalidia catharinae; Bequaert, 1932: 50. Saussure), but she did not examine the type(s) of I. ?Ropalidia fluviatilis petulans Cheesman, 1952: 18, female – reactionalis. A female in the MHN labeled with a blue “Japen Is.” circle with “voyage/d’Cap/d’Urville [probably in de Cameron (1913) described two “catharinae” of New Saussure’s handwriting]”, “Ropalidia/spec. [in van der Guinean Ropalidia, respectively in Icaria and Vecht’s handwriting]/det. J. v. d. Vecht 1967” and Ancistrocerus. Meade-Waldo and Morley (1914: 406) “Ropalidia lefebvrei /comp. w / HOLOTYPE / det. J. treated Ancistrocerus catharinae as a variety of Icaria Kojima 1999” is R. lefebvrei and agrees, in general, nigra Smith, and stated “All the species published with with the original description of Icaria reactionalis. the specific name “catharinae” in Cameron’s paper . . . This specimen, however, has the scutum colored bear labels with the specific name “beauforti” on the ferruginous with a pair of yellow longitudinal bands type-specimens”, without referring to the taxonomic and the propodeum is entirely yellow, while de status of Icaria catharinae. Bequaert (1932) listed “R. Saussure (1853-58: 28) stated for I. reactionalis that catharinae (Cameron, 1913)” as a species distributed in “Corselet jaune; métathorax [= propodeum] New Guinea, but he did not indicate which ferrugineux, avec deux lignes jaunes longitudinales”. “catharinae” he referred to. Van der Vecht (1934: viii) In the original description of I. reactionalis, the body stated “Icaria catharinae . . . is niet synoniem met . . . parts are referred to in an unusual order; that is, de Ancistrocerus catharinae . . . ; of deze sort reeds Saussure (1853-58: 28) mentioned “Corselet . . . vroeger beschreven was, hoopt Spr. nog nader te métathorax [= propo-deum] . . . écusson [= scutellum]”. onderzoeken” [Icaria catharinae is not a synonym of This fact may suggest that “métathorax” in the Ancistrocerus catharinae; the speaker [= the late van description of I. reactionalis could be an erroneous der Vecht] wish to make further investigation if this writing in the manuscript or erroneous typesetting for species was already described previously]. “mésothorax”. Even if this interpretation is correct, this In the ZMA is a female marked as “Holotype” [red specimen in the MHN could not be regarded as the type label, detached metasoma glued], with labels “Waigeoe of I. reactionalis. De Saussure (1853-58: 29) [= Waigeo] 29 XII 09”, “Mevr. de Beaufort leg” and mentioned “La Nouvelle-Guinée” as the locality and “Icaria/Catharinae/Beauforti/Cam. type/HoloTYPE sec. “Collection de M. Guérin-Méneville” as the depository: J. vdV. ’33 [in Cameron’s handwriting, but the specimen does not have a locality label and “Catharinae” and the last line are in van der Vecht’s hymenopteran specimens in the collection of Guérin- handwriting]”, that is evidently the holotype of Icaria

Ropalidia of New Guinea 55 catharinae. I examined the types of almost all taxa the types of Icaria festina and I. zonata. The late van described from New Guinea and its adjacent islands, der Vecht distributed at least to the BMNH the and I concluded that R. catharinae (= Icaria specimens of an undescribed species (this will be catharinae) is a valid species. treated in the second part of this series) with his Cheesman (1952: 18) described R. fluviatilis identification label as “R. festina”; Richards could use petulans based on three females from Japen Island. In those specimens when he produced the key. the drawer in the BMNH that contains the two paratypes of R. fluviatilis petulans is a label “Type Ropalidia albobalteata (Cameron) missing”; I searched the holotype in vain. The both paratypes are labeled “Dutch New Guinea, Japan Is. Mt. Polistes albobalteatus Cameron, 1906a: 61, female – Baduri 1000 ft. viii 1938 L. E. Cheesman BM. 1938- “Manokwari”. — van der Vecht, 1934: viii (syn. of R. 593” and “L. E. Cheesman det. 1949 fluviatilis M. W. festina (Smith)). ? Icaria fluviatilis Meade-Waldo, 1912: 449 (key), 451, ssp. petulans Chees”. These two specimens apparently female (in subgenus Icariola) – “Mimika River”. belong to different species; one agrees well with the Polybia papuana Cameron, 1913: 77, female – “Waigeu”. holotype of Icaria catharinae, and the other with the REVISED SYNONYMY. — van der Vecht, 1934: viii holotype of Polistes albobalteatus Cameron, 1906. (syn. of R. festina (Smith)); 1966: 6. — Kojima and Unless the holotype of R. fluviatilis petulans is Carpenter, 1997: 6 (designation of lectotype). eventually found, the exact taxonomic status of this ? Ropalidia fluviatilis; Bequaert, 1932: 51. — Cheesman, taxon cannot be determined. 1952: 2, 17 (redescription; distr.), 26 (fluvialis [!]). — Richards, 1978: 58 (in subgenus Icariola; list). Ropalidia festina (Smith) ? Ropalidia fluviatilis petulans Cheesman, 1952: 2 (petulaus [!]), 18, female – “Japen Is., Mt. Baduri, 1000 ft.” Icaria festina Smith, 1865: 90, female – “New Guinea”. Ropalidia papuana; van der Vecht, 1966: 6. Icaria zonata Cameron, 1906a: 61, female – “Moaif”. — ? Ropalidia petulans; Richards, 1978: 58 (in subgenus Meade-Waldo, 1911: 100 (syn. of I. festina). — van der Icariola). Vecht, 1934: viii (syn. of R. festina (Smith)). — Kojima Ropalidia albobalteata: Kojima and van Achterberg, 1997: and Carpenter, 1997: 5 (designation of lectotype). 166 (“they [albobalteata and festina] differ at species Ropalidia festina; Bequaert, 1932: 51. — van der Vecht, level”). 1934: viii. Ropalidia reactionalis festina; Cheesman, 1952: 3, 21 Cameron (1906a: 61) stated “3. Polistes (redescription of the type). albobalteatus sp. nov. Manokwari, February. One Ropalidia zonata; Cheesman, 1952: 3, 22. — Richards, 1978: example”, thus a female in the ZMA labeled “Mano- 58 (in subgenus Icarielia), 129 (key). kwari [NW New Guinea]/14-28 II 03”, “Polistes ? Ropalidia festina; Richards, 1978: 58 (in subgenus Icarielia; /albobalteatus/Cam. Type/New Guinea [in Cameron’s list), 129 (key). handwriting] /♀Holotype sec. J. v. d. Vecht ‘33 [in van Although Smith (1865) did not give any indication der Vecht’s handwriting]”, “Ropalidia/festina (Sm.)/var. for the number of specimens, a female in the OUM albobalteata (Cam.)/det. J. v. d. Vecht 34 [in van der labeled “Holotype” (BM type label), “N” [= New Vecht’s handwriting]” and “Holotype” is certainly the Guinea] (circular) and “Icaria festina Smith” is holotype of P. albobalteatus. While van der Vecht presumably the holotype of Icaria festina Smith, 1865. (1934: viii) synonymized P. albobalteatus under R. I compared this specimen with the lectotype and festina (Smith), Kojima and van Achterberg (1997: paralectotypes of Icaria zonata Cameron, 1906 (the 166) noted that they are different species. As detailed lectotype designated by Kojima and Carpenter (1997: 5) in the second part of this series, comparison of the is in the ZMA and labeled “Holotype”, “Moaif [prob. holotypes of I. festina and P. albobalteatus together NW New Guinea]/26 VI - 4 VII”, “Icaria /zonata/Cam. with the examination of additional specimens led me to Type/New Guinea [in Cameron’s handwriting]/ ♀ conclude that they are different species. HOLOTYPE/Sec. J. v. d. Vecht 33 [in van der Vecht’s Van der Vecht (1934: viii) synonymized Polybia handwriting]” and “Ropalidia zonata ♀ (Cam.) det J. v. papuana Cameron, 1913, under R. festina, but later he d. Vecht 1974”), and I confirmed the synonymy of I. (1966: 6) transferred it to the genus Ropalidia and zonata under R. festina proposed by Meade-Waldo treated it as a valid species. I did not find any specific (1911: 100) and van der Vecht (1934: viii). differences between the holotype of P. albobalteatus Richards (1978) treated both species as valid species and the lectotype of P. papuana (ZMA, designated by and included them in his key to species of the Kojima and Carpenter (1997: 6), and labeled “subgenus Icarielia” (p. 129), stating that the first “Holotype”, “Waigeoe [= Waigeo]/25 XII 09”, “Mevr. metasomal tergum is “about as long as mid femur” in R. de Beaufort leg” and “Polybia/papuana/Cam. Type [in festina and “clearly shorter than mid femur, wider” in Cameron’s handwriting]/Holotype ♀ sec J. v. d. Vecht R. zonata. However, I wonder if Richards examined

56 J. Kojima

33 [in van der Vecht’s handwriting]”) and propose to (de Saussure)). — Kojima and Carpenter, 1997: 5 synonymize P. papuana under R. albobalteata. (designation of lectotype). Meade-Waldo (1912: 452) described Icaria Icaria cohni du Buysson, 1909b: 306, male – “Bougainville”. (Icariola) fluviatilis based on a single female from — van der Vecht, 1941a: 104 (syn. of R. gregaria (de “Mimika River”. The drawer in the BMNH, in which Saussure)). — Richards, 1978: 86 (kohni [!]; syn. of R. gregaria spilocephala (Cameron)). the holotype should be, contains a note “headless type Ropalidia impetuosa; Bequaert, 1932: 51. — Richards, 1978: loaned to v.d. Vecht”. When I visited the NNM in 1999, 58 (in subgenus Icariola), 86 (possible syn.: Ropalidia I located the holotype of I. fluviatilis among the gregaria spilocephala (Cameron)). specimens brought from the late van der Vecht’s house Ropalidia gregaria; van der Vecht, 1941a: 104 (syn.: Icaria after his death [the specimen is tentatively kept at the cohni du Buysson, I. impetuosa Smith, I. spilocephala IUNH and I will return it to the BMNH on my Cameron), 111 (key), 149 (distribution [including responsibility]; the specimen is headless and labeled subspecies spilocephala]). — Cheesman, 1952: 3, 20 “Type” [BM type label]; “New Guinea./Mimika R./A. (male). — Spradbery, 1991: 349 (colony population). F. R. Wollaston. /1911-229./aug 1910” [in print but last Ropalidia cohni; Bequaert, 1932: 51. line in handwriting]; “B. M. TYPE/HYM./18.859”; Ropalidia spilocephala; Bequaert, 1932: 51. “BM ’79”, and “Icaria/(Icariola) /fluviatilis sp. n./Type Ropalidia gregaria inquieta Cheesman, 1952: 3, 20, female, male – “Papua, . . . Mafulu, 4000 ft.” — Richards, 1978: /G. Meade-Waldo det”. Although I could not find any 86 (designation of lectotype [error in locality: Mafalu]; differences between the holotypes of Polistes albobalte- syn. of R. gregaria spilocephala). atus and Icaria fluviatilis to consider them as different Ropalidia gregaria var. tolerans Cheesman, 1952: 3 (as species, headlessness of the I. fluviatilis holotype subspecies), 21 (as var.), female – “Cyclops Mts. Sabron”. prevents me from making a definitive conclusion. — Richards, 1978: 86 (designation of lectotype; syn. of R. The BMNH houses two females identified as gregaria spilocephala). “fluviatilis” by Cheesman; they could be the specimens Ropalidia gregaria gregaria; Richards, 1978: 53 (nest), 62 referred to in Cheesman (1952: 17), and one is from (key), 86 (in subgenus Icariola). — Spradbery, 1991: 351 Kokoda and the other is from Humboldt Bay. While (colony population). the specimen from Kokoda and one of the paratypes of Ropalidia gregaria spilocephala; Richards, 1978: 53, 62 R. fluviatilis petulans, as noted before, are conspecific (key), 86 (syns.: Icaria kohni [!] du Buysson, Ropalidia gregaria inquieta Cheesman, Ropalidia gregaria var. with the holotype of P. albobalteatus, the specimen tolerans Cheesman; possibly syn. of Icaria impetuosa from Humboldt Bay seems to me to be R. gemmea Smith; “I examined the holotype male [!] in the (because of the orientation of the head, I could not Zoological Museum, Amsterdam.”). — Spradbery and determine its definite specific affiliation). Kojima, 1989: 636 (nest; colony population). — Spradbery, 1991: 351 (colony population). Ropalidia bensoni Richards During my recent research in the MHN, I located Ropalidia bensoni Richards, 1978: 55 (nest), 59 (list), 65 the types of Icaria gregaria de Saussure, 1854. They (larva in key), 129 (key), 130, female, larva, nest (in are different species of “Ropalidia gregaria” of authors, subgenus Icarielia) – “New Guinea: Morobe district, and I proposed to apply R. impetuosa (Smith) to “R. Wau”. gregaria” of authors (see Kojima, 2001: 5). This species is undoubtedly a valid species. Specimens examined — Halmahera: 3♀ (NNM, 1 Types examined — NE New Guinea: ♀ (BMNH, ♀ in IUNH), between Gita & Payahe, Tayawi, holotype), labeled “Holotype” (BM type label), “Nest 4.iii.1995, R. de Vries; 4♀ (NNM), nr. Payahe, Tayawi, NG 3, New Guinea, Morobe district, Wau, 12-16. Dec. R. de Vries (2♀: 10.iii.1995; 2♀: 14.iii.1995); 1♀ 1972, O. W. Richards”, “Ropalidia bensoni Richards, (NNM), road Payahe-Weda, E. of Payahe, 15.iii.1995, ♀, Holotype [in Richards’ handwriting]”; 1♀ (NNM, R. de Vries. Bacan: ♀ (OUM, lectotype of Icaria paratype), same collection data as the holotype. 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. Species in the R. impetuosa-group J. Toxopeus; 1♀ (ZMA), Station 9, 18.v.1921, L. J. Ropalidia impetuosa (Smith) Txopeus. Ambon: 1♀ (OUM, paralectotype of I. impetuosa), labeled “Amb”; 1♀ (NNM), 10 km NE Icaria impetuosa Smith, 1860: 131, female – “Batchian; Pago, 200 m alt. 3°38’S, 128°16’E, 21.iii.1988, R. Amboyna”. — van der Vecht, 1941a: 104 (syn. of R. ♀ ♂ gregaria (de Saussure)). — Kojima and Carpenter, 1997: Hensen; 2 1 (NNM), Ambon City, 200 m alt., 4 (designation of lectotype). 3°41’S, 128°10’E, 20.iii.1988, R. Hensen. Seram: 1♀ Icaria spilocephala Cameron, 1906b: 230, female – “Etna (NNM), 15 km NNE of Tehoru, Hatumete, 50 m alt., Bay”. — van der Vecht, 1941a: 104 (syn. of R. gregaria 3°17’S, 129°36’E, 24.iii.1988, R. Hensen. NW New Guinea: ♀ (ZMA, lectotype of Icaria spilocephala

Ropalidia of New Guinea 57

Cameron, designated by Kojima and Carpenter (1997: BUYSSON det 1909”; 1♂ (MNHN, paralectotype of I. 5)), labeled “Nieuw Guinea/Expeditie 1904/[slash]5 cohni); 2♀ (SAM), A. H. Voyce. Solomon Islands: 1 /Etna baai”, “Icaria / spilocephala /Cam. Type / New ♀ (CAS), Tenaru River, Guadalcanal, i.[19]45, G. E. Guinea [in Cameron’s handwriting] /♀ Holotype sec. J. Bohart; 1 ♀ (NNM), Guadalcanal, Lunga Estate, on v. d. Vecht 1933 [in van der Vecht’s handwriting]” and nest, 9.vii.1933, H. T. Pagden; 9♀1♂ (NNM), Tulgi, “Holotype”; 1 ♀ (ZMA), Baliemvalley, 5 km W. H. T. Pagden (3♀, nest on Hibiscus, 1.vii.1933; 1♂, Wamena, 1,500 m, 23.x.1993, A. J. de Boer, A. L. M. 2.ix.1933; 3♀, 23.vii.1934; 3♀, on nest, 16.ix.1934); 1 Rutten & R. de Vos; ♀ (BMNH, lectotype of Ropalidia ♀ (ZMA), Guadalcanal, Honiara, xi.1967. gregaria inquieta Chessman, designated by Richards (1978: 86)), labeled “Lectotype” (BM type label), “Dutch New Guinea, Cyclops Mts., Sabron, 930 ft., Species in the R. marginata-group v.1936, L. E. Cheesman, B.M. 1936-271”, “Ropalidia Ropalidia marginata jucunda (Cameron) gregaria var. tolerans det. L. E. Cheesman”, “Ropalidia Icaria jucunda Cameron, 1898: 46, female – “New Guinea”. gregaria (Sauss.) var. tolerans Cheesm. Lectotype”. Icaria pruinosa Cameron, 1906b: 228, female – “Digoel”. — NE New Guinea: 3♀ 3 ♂ (BMNH), Madang Dist., van der Vecht, 1941a: 104 (syn. of R. marginata jucunda). Finisterre Mts., Damanti, 3,550 ft., 2-11.x.1964, M. E. —Kojima and Carpenter, 1997: 5 (designation of Bacchus; 2♀ (BMNH), Madang Dist., Finisterre Mts., lectotype). Moro C., 5,550 ft., 30.x.-15.xi.1964, M. E. Bacchus; 5 Ropalidia jucunda; Bequaert, 1932: 50. ♀ (CAS), Madang Province, Bundi, 5°45’S, 145°15’E, Ropalidia pruinosa; Bequaert, 1932: 51. 10-12.iii.1987, W. J. Pulawski; 8 ♀ (CAS), Madang Ropalidia marginata jucunda; van der Vecht, 1941a: 104, 124. Province, Simbai, 5°17’S, 144°26’E, W. J. Pulawski; (6 Ropalidia canaria Cheesman, 1952: 3, 19, male - “Papua, Kokoda”. NEW SYNONYMY. — Richards, 1978: 58 (in ♀: 25-26.iii.1987; 1♀: 27.v.1988; 1♀: 31.v.1988); 10 subgenus Icariola). ♀ 1 ♂ (CAS), Madang Province, Tep Tep, 5°55’S, 146°30’E, 1,900 m, M. Wasbauer (6♀: 27.xi.1987; 4♀ In the latest citations, four subspecies are recognized 1 ♂ : 29.xi.1987); 11 ♀ (CAS), Madang Province, in R. marginata (Lepeletier, 1836), based primarily on Karisokora, 7.5 air km W. Bundi, 5°44’S, 145°10’E, ca. color pattern: nominate subspecies, jucunda (Cameron, 1,800 m, 17.v.1988, E. Koima & P. Yomane; 2 ♀ 1898), rufitarsis van der Vecht, 1941, and sundaica van (CAS), Madang Province, Brahman Catholic Mission, der Vecht, 1941 (see Kojima and Carpenter, 1997). 6°45’S, 145°23’E, 21.x.1987, M. Wasbauer; 1 ♀ Although such differences in color pattern could be (SAM), Wareo, L. Wagner; 10 ♀ (SAM), Finsch intergraded among marginata, jucunda and sundaica, I Haven, L. Wagner; 1♀ (SAM), Komba, L. Wagner; 2 leave at this moment them as subspecies of R. ♀ 2 ♂ (IUNH), foot of Mt. Missim, Wau, Morobe, marginata. Their taxonomic status (color variation or 11.vii.1985, J. & K. Kojima; 1♀2♂ (IUNH), Wau, diagnosable species) can be discussed only after a 23.xii.1973, T. Okazawa; 5 ♀ 4 ♂ (IUNH), Bulolo detailed comparative study of the R. marginata-group, Watut, 21.xii.1973, T. Okazawa; 1 ♀ (IUNH), which includes also R. spatulata van der Vecht, 1962, R. Kundiawa, 27.xii.1973, T. Okazawa. SE New Guinea: brevita Das and Gupta, 1989, R. laticincta laticincta ♂ , (BMNH, lectotype of Ropalidia gregaria var. van der Vecht, 1962, R. laticincta floresiana van der tolerans Cheesman, designated by Richards (1978: Vecht, 1962, and R. clavata (de Saussure, 1853) [= 86)), labeled “Lectotype” (BM type label), “Papua, Icaria duchaaussoyi Gribodo, 1896; see Kojima, 2001: Mafulu, 4000 ft. xii.1933, L. E. Cheesman, B.M. 1934- 5]. 321”, “ ♂ ’, “gregaria Sauss. inquieta Ch. L. E. I examined the types of Icaria jucunda Cameron, I. Cheesman det. 195”, “Ropalidia gregaria (Sauss.) spp. pruinosa Cameron, and R. carinata Cheesman, and inquieta Cheesm. ♂ lectotype”; 2♀ (IUNH), Waitape, concluded that they all belong to the same taxa. 20.i.1974, T. Okazawa; 7♀1♂ (SAM), Mt. Lamington, Specimens examined — New Guinea: ♀ (OUM, 1,300-1,500 feet, C. T. McNamara; 1 ♂ (BMNH), holotype of Icaria jucunda Cameron), labeled “Icaria Laloki, 6.xii.1980, J. W. Ismay; 6 ♀ (CAS), Port jucunda Cam. Type, New Guinea [Cameron’s Moresby, W. J. Pulawski (2♀: 1-2.iv.1987; 1♀: 29- handwriting]”, “Ropalidia ferruginea (F.) ♀ var. 30.iii.1987; 3♀: 22-23.vi.1988); 1♀1♂ (CAS), Central jucunda (Cam.), det. J. v. d. Vecht”, “12”. SW New Province, Lake Iaraguma, 20 km NW Port Moresby, Guinea: ♀ (ZMA, lectotype of Icaria pruinosa 21.vi.1988, W. J. Pulawski. New Ireland: 1♀ Cameron, designated by Kojima and Carpenter (1997: (BMNH), Nama Kainai, xi.1934, J. L. Froggatt. 5)), labeled “Nieuw Guinea/Expeditie 1904/Digoel”, Bougainville: ♂ (MNHN, lectotype of Icaria cohni du “Icaria/pruinosa/Cam. Type/New Guinea [in Cameron’s Buysson, designated by Kojima (2001: 5)), labeled handwriting] /Holotype sec. J. v. d. Vecht 1933 [in van “MUSEUM PARIS/I. Bougainville/(leg. Cohn)/Alfken der Vecht’s handwriting], and “Ropalidia ferruginea 1909”, “TYPE”, “Icaria/Cohni Buyss. /types/R. DU (F.)/var. jucunda (Cam.)/det. J. v. d. Vecht”; 2 ♀

58 J. Kojima

(NNM), Neth. N. G. Exp. 1959, Tanah Merah, 17 m alt., Icaria unicolor Smith, 1859: 168, female – “Key Island” [= 5.ix.1959. NE New Guinea: 33 ♀ (IUNH, 1 ♀ in Kai]. NNM), Kundiawa, 27.xii.1973, T. Okazawa. SE New Ropalidia unicolor; Bequaert, 1932: 51 (“probablement un Guinea: ♂ (BMNH, holotype of Ropalidia canaria Parapolybia”). — van der Vecht, 1962 (in subgenus Cheesman), labeled “Type” (BM type label), “Papua: Anthreneida) — Richards, 1978: 58 (in subgenus Icariola). Kokoda 1200 ft. vii.1933, L. E. Cheesman B.M. 1933- Ropalidia mathematica unicolor; van der Vecht, 1941a: 133. 427”, “ ♂ ”, “L. E. Cheesman det. 1948, canaria Chess.”, “B.M. TYPE HYM 18.840”, and “R. In the OUM is a female labeled “Kei” (circular), marginata jucunda (Cam.) det. J. v. d. Vecht ‘79”; 1♀1 “Icaria unicolor Smith” (blue label), “Icaria unicolor ♂ (NNM), Port Moresby. 1889, L. Loria; 2 ♀ 2 ♂ Sm.”, that is presumably the holotype of Icaria unicolor (CAS), Port Moresby, W. J. Pulawski (1♀1♂, 29- Smith, as noted by van der Vecht (1941a: 133). 30.iii.1987; 1 ♀ 1 ♂ , 1-2.iv.1987); 2 ♀ (NNM), Port Moresby, Boroko, 5.ix.1959, J. I. H. Sient Ivany; 9♀2 Species in the R. gracilis-group ♂ (NNM), Kapakapa, 1891, L. Loria; 2♀ (NNM), Popondetta, 8.ii.1962, A. Catley; 1♀ (NNM), Central Ropalidia gracilis (Smith) District, Rouna Falls, 24.vi.1973, J. P. Spradbery. Icaria gracilis Smith, 1859: 167, female – “Aru”. — Dalla Torre, 1904: 73 [error: year of publication, 1868]. Icaria deceptor Smith, 1863: 42, female – “Mysol”. Species in the R. stigma-group Ropalidia gracilis; Bequaert, 1932: 50 (gracillis [!]; Ropalidia mathematica torrida (Smith) “probablement un Parapolybia”). — Cheesman, 1952: 3, 18 (syn.: Icaria deceptor Smith, 1863; redescription of Icaria torrida Smith, 1863: 42, female – “Ceram”. type). — Richards, 1978: 58, 99 (in subgenus Icariola). Ropalidia torrida; Bequaert, 1932: 51. Ropalidia deceptor; Bequaert, 1932: 50. — Cheesman, 1952: Ropalidia mathematica torrida; van der Vecht, 1941a: 104, 18 (syn. of Ropalidia gracilis (Smith, 1859)) — Richards, 133 (distr.); 1962: 20 (in subgenus Anthreneida; distr.; 1978: 53 (nest), 58 (list), 61 (key), 99, figs. 20, 34 (in “Perhaps this form will eventually prove to be closer to R. subgenus Icariola). socialis or unicolor than to R. mathematica.”). The R. stigma-group is distributed from India to Smith (1859: 167) described Icaria gracilis from Australia, but no species in this species-group has been Aru, but he did not refer to it when he later (1863: 42) known from New Guinea mainland. This species-group described Icaria deceptor from “Mysol” (= Misool). includes torrida and unicolor, and still awaits an Cheesman (1952: 18) synonymized Icaria deceptor intensive taxonomic study. In the present paper, the under Ropalidia gracilis; she listed the holotype of only taxonomic status of torrida and unicolor follows those I. gracilis, and thus it is not clear whether her in the latest citations. Other taxa included in the R. synonymy was based on the comparison of the types of stigma-group are (their taxonomic status is according to I. gracilis and I. deceptor. Richards (1978: 99) the latest citations; see Kojima and Carpenter, 1997): R. examined the types of both I. gracilis and I. deceptor artifex artifex (Smith, 1854), R. a. fuscata van der and treated them as valid species. I examined the types Vecht, 1941, R. darwini Richards, 1978, R. elegantula of I. gracilis and I. deceptor; both are presumably the Richards, 1978, R. hongkongensis hongkongensis (de unique original specimens (the holotypes). The Saussure, 1954), R. h. juncta van der Vecht, 1941, R. holotype of I. gracilis differs from the holotype of I. mathematica mathematica (Smith, 1860), R. m. deceptor in the following characters as Richards (1978: binotata van der Vecht, 1941, R. m. nigroplagiata 99) noted: punctuation more closely and stronger; (Cameron, 1900), R. m. sumbaensis van der Vecht, posterior widened part of first metasomal tergum in 1962, R. nigrita Das and Gupta, 1989, R. rufocollaris dorsal view more parallel-sided (Fig. 43 vs. 45); second rufocollaris (Cameron, 1900), R. r. atrata, van der metasomal segment proportionately longer (Figs. 42 Vecht, 1941, R. santoshae Das and Gupta, 1989, R. and 43 vs. 44). The shape of the basal parallel-sided socialis socialis (de Saussure, 1862), R. s. trimaculata part of the first metasomal tergum is, however, not so van der Vecht, 1962, R. stigma stigma (Smith, 1858), R. different as Richards noted. Additional specimens from s. nigrolineata van der Vecht, 1962, R. s. rufa van der several localities in New Guinea show that the shape of Vecht, 1941, and R. thailandia Gusenleitner, 1994. the first metasomal tergum is so variable (Figs. 46–52) that the differences between the holotypes of I. gracilis Types examined — Seram: 2♀ (OUM, lectotype and I. deceptor are intergraded. The punctuation is also and paralectotype), labeled as given in the section so variable that this character is barely used to “Designation of the lectotypes”. distinguish the two taxa. Furthermore, the marking pattern does not diagnose them. I follow Chessman’s Ropalidia unicolor (Smith) (1952) synonymy.

Ropalidia of New Guinea 59

In Australia, R. gracilis and its closely related dorsal surface of the hind coax in R. gracilenta. I species, R. gracilenta Richards, 1978, are distributed; (Kojima, 1999a) found that the states of this character the former is distributed in northern part of Australia’s are variable in R. gracilenta, while the condition of the Queensland and the latter from northern part of propodeum is more reliable to distinguish the two Queensland to northern part of New South Wales species: posterior face of propodeum distinctly and (Kojima, 1999b). The characters shared by these two widely excavated and in lateral view slightly convex in species are the gena proportionately wider (in terms of R. gracilenta, while only shallowly depressed medially

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. the gena width/eye width ratio) in the male than in the and in profile slightly concave or nearly straight in R. female (Figs. 53, 54; see also Kojima, 1987) and the gracilis (Fig. 55). pronotum with ventral corner obliquely cut off (Figs. 55, Specimens of R. gracilis examined — New 56). Similar states in both characters are also found in Guinea: 1♀ (OUM), labeled “N” [= New Guinea] the R. rufoplagiata-group (Fig. 57; see also Kojima, (circular), “Icaria gracilis Smith” (blue), and “not a 1999a). Richards (1978: 61) stated that R. gracilenta type”; 1♀ (NNM), labeled “N”, “compared with type” could be distinguished from R. gracilis [as R. deceptor] and “R. deceptor”; 1 ♀ (BMNH), labeled “N” and by the presence of a strong angular projection on the “Icaria gracilis Smith”; 1♀ (BMNH), New Guinea.

60 J. Kojima

Waigeo: 1♀ (BMNH), Mt. Nok, vi.1938, L. E. tergum wide and not strongly depressed, in profile Cheesman. Misool: ♀ (OUM, holotype of Icaria length of lamella distinctly longer than depth of deceptor Smith), labeled “M” [= Mysol] (circular), depression (Fig. 60). Papua New Guinea………...… “Icaria deceptor Smith” and “= gracilis Smith, L. E. …...... R. cristata Kojima Cheesman, 1949”; 3♀ (NNM), Fokal, 0-75 m, 8.ix.- – Ocelli less close to each other, distance between 20.x.1948, M. A. Lieftinck. NW New Guinea: 1♀ posterior ocelli 1.7× or more their diameter; space (NNM), Torricelli Mts., 200-1000’, i.[19]39, E. L. between posterior ocelli with a few punctures (Fig. Cheesman; 1♀ (SAM), Torricelli Mts., 1000’, i.[19]39, 61); gena in profile smoothly widened ventrally to E. L. Cheesman; 1 ♀ (NNM), Humboldt Bay; 1 ♀ the level of mid of eye, then smoothly narrowed (NNM), Neth. Ind.-Amer. New Guinea Exp., Bernhard toward mandibular base, occipital carina broadly Camp, 50 m, 16.ix.1939, L. J. Toxopeus. Aru: ♀ and regularly curved (Fig. 62); posterior lamella of (OUM, holotype of Icaria gracilis Smith), labeled the first metasomal tergum narrow, strongly “Aru” (circular), “Icaria gracilis Smith” (blue label), depressed, in profile length of lamella nearly the “Type [in Richards’ (?) handwriting]”; 1 ♀ (NNM), same as depth of depression (Fig. 63). Northeastern Mus. Leiden Snellius Exp., Aroe Eilanden, Manoem- Australia...... R. turneri Richards baai en omgeving [= Manoem Bay and environs], 11- 44.x.1929. SW New Guinea: 2♀1♂ (NNM, IUNH), Specimens of R. cristata examined — NE New 11 km S. Bupul, 20 m, 7°39’S, 140°53’E, 9.iv.1988, R. Guinea: 1♀ (NNM, paratype), Wau, Morobe Province, Hensen. SE New Guinea: 1♀ (SAM), Mt. Lamington, 1250 m, 11.i.1950, J. Sedlacek; 9♀ (IUNH, paratypes), 1,300-1,500 feet, C. T. McNamara; 1 ♀ (BMNH), Wau, 23.xii.1973, nest 731223-01, T. Okazawa; 3♀ Kokoda, 1200 ft., ix.1933, L. E. Cheesman. Australia: (IUNH), Wau, 1.vii.1985, nest RPNG8501, J. Kojima; 2 ♂ (BMNH), North Queensland, R. C. L. Perkins 7 ♀ (IUNH), Wau, 12.vii.1985, nest RPNG8504, J. [identified as “deceptor”]. Kojima; 1 ♀ (IUNH), Wau, 14.vii.1985, nest RPNG8509, J. Kojima. SE New Guinea: ♀ (ANIC, holotype), labeled “PAPUA NEW GUINEA/Central Species in the R. rufoplagiata-group District/Ower’s Corner/28 August 1973/J. P. Sprad- Ropalidia cristata Kojima bery”; 3♀ (NNM, paratypes), same collection data as the holotype; 2♀ (NNM, paratype; BMNH), Ower’s Ropalidia cristata Spradbery, 1985: 17. Nomen nudum. ♂ Ropalidia cristata Kojima, 1989: 143, female, male, larva – Corner, 3.ix.1973, J. P. Spradbery; 2 (NNM, “PAPUA NEW GUINEA . . . Ower’s Corner, Sogeri, Central paratype; BMNH), Uberi, Kokoda Trail, 26.iii.1974, J. Province”. — Spradbery and Kojima, 1989: 633 (colony P. Spradbery; 1♀ (BMNH), Kokoda, 1,200 ft., v.1933, population; nest; ethology). — Spradbery, 1991: 350 L. E. Cheesman. (colony population). This species is closely related to R. turneri Richards, Acknowledgments 1978, distributed in north-eastern Australia. Among the My special thanks are due to the late J. van der Vecht, characters I (Kojima, 1989) remarked to distinguish R. who had encouraged me to make an intensive study of cristata from R. turneri, the state of the propodeal Ropalidia, one of the largest and most interesting polistine concavity, the shape of the clypeus and the condition of genera; he always offered me valuable suggestions on the the pronotal carina were revealed to be less reliable biology not only of the Ropalidia but also of the Vespidae in after examination of many additional specimens of R. general. I am very pleased to have a chance to publish some turneri (mainly from the Atherton Tableland, Queens- more of his work. I thank C. van Achterberg for his help in various ways during the present study, without which the land, Australia; deposited in the IUNH). The following present study would not have been accomplished. My one- key gives the diagnostic characters, including those year stay in the Nationaal Natuurhistorisch Museum in 1996 newly recognized, to distinguish between R. cristata was supported by the Nederlandse Organisatie voor and R. turneri. Wetenschappelijk Onderzoek (NWO) and Japan Society for the Promotion of Science (JSPS). My thanks are also due to 1. Ocelli relatively close to each other, distance my family for their patience allowing me to spend most time between posterior ocelli less than 1.5× their of the one-year stay in the Netherlands in taxonomic work. I diameter; space between posterior ocelli hardly also wish to express my thanks to J. M. Carpenter for critical punctured (Fig. 58); gena in profile widened reading of an earlier draft, and to the following persons who ventrally to level of about one-third from eye top, assisted me in examining the specimens and/or provided me then posterior margin of gena (or occipital carina) with the specimens for this study: A. D. Austin, B. Bregge, J. Cardale, J. M. Carpenter, J. Casewetts-Weulersse, M. C. Day, run down in nearly parallel to posterior margin of P. Dessart, S. Hartini, S. Heydon, W. Hogeness, T. eye, recurved anteriorly near the base of mandible Huddleston, S. Kahono, M. Lachaise, S. Lewise, I. Löbl, E. G. (Fig. 59); posterior lamella of the first metasomal Matthews, A. Menke, B. Merz, I. D. Naumann, G. M.

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Nishida, W. A. Noerdjito, D. Notton, L. Tarel, C. O’Toole, Carpenter, J. M. 1999. Taxonomic notes on paper wasps W. Pulowski, J. P. Spradbery, C. Taylor, C. Villemont, R. de (Hymenoptera: Vespidae; Polistinae). Am. Mus. Novitat. Vries and Sô. Yamane. Beside the financial support from the 3259: 1–44. NWO and JSPS, the present study was partly supported by Carpenter, J. M. and J. Kojima. 1997. Checklist of the species Grants-in-Aid from the JSPS (nos. 11833001 and 11691161). in the subfamily Vespinae (Insecta: Hymenoptera: Vespidae). Nat. Hist. Bull. Ibaraki Univ. 1: 51–92. References Charnley, H. W. 1973. The value of the propodeal orifice and the phallic capsule in vespid taxonomy (Hymenoptera, Achterberg, C. van. 1992. Obituary and bibliography of Vespidae). Bull. Buffalo Soc. Nat. Sci. 26: 1–79. Jacobus van der Vecht (1906-1992). Zool. Med., Leiden Cheesman, L. E. 1952. Ropalidia of Papuasia. Ann. Mag. 66: 295–302. Nat. Hist. (12) 5: 1–26, 4 pls. Bequaert, J. 1918. 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Nova Guinea, Résultats de l’Expedition ptera; Vespidae). Tijdschr. Ent. 130: 11–31. scientifique Néerlandaise à la Nouvelle Guinée en 1907 et Horn, W., I. Kahle, G. Friese and R. Goedike. 1990. 1909 sous les auspices de Dr. H. A. Lorentz, 9: 185–248. Collectiones Entomologicae. Akademie der Landwirt- Cameron, P. 1913. On the Hymenoptera (exclusive of the schafswissenschaften Deutschen Demokratischen Repu- Anthophola and Fromicida) collected by Mrs. de Beaufort blik, Berlin. in the Papuan islands of Waigeu and Saonek. Bijdr. Kojima, J. 1987. The male of Ropalidia gracilenta Richards, Dierk. 19: 75–86. with notes on females and a nest (Hymenoptera, Carpenter, J. M. 1991. Phylogenetic relationships and the Vespidae). J. Austr. Entomol. Soc. 26: 149–151. origin of social behavior in the Vespidae. Pp. 7–32. In Kojima, J. 1989. A new polistine species of Ropalidia Ross, K. G. and R. W. Matthews (eds.), The Social (Hymenoptera, Vespidae) from Papua New Guinea. Jpn. Biology of Wasps. 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Vecht, J. van der and J. Kojima. 1996. Two new species of the genus Ropalidia from New Guinea (Hymenoptera: Vespidae: Polistinae). Zool. Med., Leiden 70: 99–104. Received 1 January 2001; revised 16 January 2001; accepted 20 January 2001.