Natural History Bulletin of Ibaraki University 5: 1–22, 9 February 2001

Taxonomic notes on types of the social wasp tribe Ropalidiini (Insecta: Hymenoptera; Vespidae, Polistinae) housed mainly in the Muséum d’Histoire Naturelle, Genève

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

Abstract Notes on types of the social wasp tribe Ropalidiini housed mainly in the Muséum d’Histoire Naturelle, Genève are provided. Lectotypes of the following species are designated: Icaria aethiopica du Buysson, 1906, I. antennata de Saussure, 1890, I. cohni du Buysson, 1909, I. gregaria de Saussure, 1854, I. guttatipennis de Saussure, 1853, Ropalidia montana Carl, 1934, Icaria plebeja de Saussure, 1863, non 1862, and Belonogaster pictus de Saussure, 1900. A new species in Belonogaster is described based on the specimens in von Schulthess’ collection in the Eidgennössische Technische Hochshule, Zürich. Nomenclatural changes include: synonymy of Ropalidia variegata interrupta van der Vecht, 1941, under R. gregaria (de Saussure), NEW SYNONYMY; application of R. impetuosa (Smith, 1860) to “R. gregaria” of authors, REVISED STATUS, and synonymies of Icaria spilocephala Cameron, 1906, I. cohni du Buysson, 1909, Ropalidia gregaria inquieta Cheesman, 1952, and R. gregaria var. tolerans Cheesman, 1952, under R. impetuosa, REVISED SYNONYMIES; synonymy of Polistes bioculata Fabricius, 1804, under Ropalidia fasciata (Fabricius, 1804), NEW SYNONYMY; synonymy of Icaria plebeja de Saussure, 1863, non 1862, with Ropalidia plebeiana Richards, 1978, NEW SYNONYMY; synonymy of Icaria duchaussoyi Gribodo, 1896, under Ropalidia clavata (de Saussure, 1853), NEW SYNONYMY; treating Belonogaster malagassa de Saussure, 1900 and B. pomicolor de Saussure, 1900 as valid species, REVISED STATUS; and replacement of B. pictus de Saussure, 1900, non Kohl, 1894, with B. confusa NEW REPLACEMENT NAME. Key words Hymenoptera, Vespidae, Polistinae, Ropalidiini, lectotype designation, new species

Introduction Richards (1982) did not specify any paralectotypes and even failed to designate the “lectotype” for Belono- In Kojima and Carpenter’s (1997) recent catalog of gaster brevipetiolata de Saussure, 1890, and did not species in the social wasp tribe Ropalidiini that includes label the “lectotype” of B. apicalis de Saussure, 1900 the four Old World polistine genera (Ropalidia Guérin- (see also Carpenter, 1999). Types of Ropalidia (= Méneville, 1831, Parapolybia de Saussure, 1854, Icaria de Saussure, 1853) in the MNH are as yet poorly Belonogaster de Saussure, 1854, and Polybioides du studied (Kojima et al., 1998; Carpenter, 1999). Buysson, 1913), the Muséum d’Histoire Naturelle, During my two-month stay in the MHN in 1999, I Genève (abbreviated as “MHN”) is listed as one of the compiled a list of ropalidiine types in the museum. In major institutions that house primary types, including the course of this research I located syntypes of four syntypes, of ropalidiine species group taxa; 33 species species, for which the depository has long been (23 in Ropalidia, ten in Belonogaster). Most of them unrecognized, and thus exact taxonomic status of these (31 species) were described by de Saussure (1853-58, taxa has been in question. As mentioned in “Methods”, 1890, 1900), whose collection was deposited in the in the course of the present study I had chances to MHN (Bedot, 1906; Horn et al., 1990). examine specimens in the Eidgennössische Technische Richards (1982) and Hensen and Blommers (1987) Hochshule, Zürich (“ETH”) and the Muséum National studied most of the Belonogaster types in the MHN, d’Histoire Naturelle, Paris (“MNHN”). The present and designated the lectotypes for six and three species, paper primarily aims to designate the lectotypes for the respectively. However, they provided neither details of taxa for which I examined most of syntypes, in order to label data nor characteristics that allow us to easily define the concept of each taxon. Detailed taxonomic recognize the name-bearing types (holotype and/or notes are also provided for the taxa for which I reached lectotype); such information should be provided when different taxonomic opinions from those of previous designating the name-bearing types according to authors. A new species in the genus Belonogaster is Recommendation 74C in the International Code of described based on specimens in the ETH. Although Zoological Nomenclature (International Commission on the MHN and MNHN house types of most of the Zoological Nomenclature, 2000). Furthermore, Malagasy species of Ropalidia, they will be treated in a 2 J. Kojima separate paper on the taxonomic revision of Malagasy relationships among species for Belonogaster. For each fauna of Ropalidia. species group taxon, the pagination of the original description is cited, followed by the sex of the Methods specimen(s) if stated. The locality given in the original description is then quoted. When label data are quoted, It is well known that H. de Saussure made his a slash (/) denotes the beginning of a new line on the taxonomic work in a close relationship with the label. Some specimens in the MHN have labels Museum Paris (= MNHN), and in the later stage of his specific to the museum; each label consists of two research life R. du Buysson took care of a large part of columns by three rows: the datum in the upper left cell de Saussure’s collection of social wasps. Consequent indicates the specimen number (or code); upper right results were that not a few specimens on which de cell is for the locality in a lower category (such as town, Saussure’s descriptions were based were retained by de etc.) if available; middle left cell is for the year when Saussure and/or exchanged between the MNHN and de the specimen was housed in the museum; middle right Saussure. In the present paper, the specimens in the cell shows the locality in a higher category (such as MNHN are also noted for some species described by de country or region); and the bottom row (two cells) gives Saussure and du Buysson. Another person who had a the collector (or donator) of the specimen. The five close relationship with de Saussure as well as the MHN data given on the MHN label are separated by colons was A. von Schulthess, whose collection is now in the (:). “MP” is an abbreviation used here for “MUSEUM ETH. A part of de Saussure’s material is in the ETH PARIS”, and the MNHN (co-) type labels are indicated (Carpenter, 1999) and the specimens in this institute are with “MP (co-) type label”. My comments and also referred to. In addition to these three collections, additions to the label data as well as citations are given the following museums and/or institutions are referred in brackets. to: In the descriptions of species several morphometric Australian National Insect Collection, CSIRO, characters are used. The parts measured are defined as Canberra (ANIC); follows: width of clypeus was measured as a distance Museo di Zoologia Sistematica, Università di between the uppermost points where clypeus touches Torino, Torino (MIZT; the specimens in this eyes; height of clypeus was measured as a distance museum were not seen); from the bottom of dorsal emargination to the apex; Museo Civico di Storia Naturale, “Giacomo Doria”, distance between inner eye margins at the vertex refers Genova (MCGD; not seen); to the distance between them at the level of the anterior Museo Civico di Storia Naturale, Venice (MCSN; ocellus in frontal view of head, and distance between not seen); inner eye margins at clypeus refers to the distance at the Museum of Comparative Zoology, Harvard Uni- level where inner eye margins approach each other versity, Cambridge (MCZ); most closely; widths of eye and gena were measured as Nationaal Natuurhistorisch Museum, Leiden the maximum for each in a strictly lateral view of the (NNM); head; length of the first metasomal tergum was The Natural History Museum, London (BMNH); measured in profile as a distance between the posterior Naturhistorika Riksmuseet, Stockholm (NR; not end of the basal slit for the reception of the suspensory seen); ligament of the mesosoma and the posterodorsal end of Zoologisches Museum der Humboldt-Universität, the tergum; widths of the first and second metasomal Berlin (ZMH; not seen); segments refer to their maximum widths in dorsal view. Zoologisches Sammlung München (ZSM; not seen); In the genus Ropalidia, the female antenna has 12 Zoologische Sammlung, Naturhistorisches Museum, articles and the male antenna has 13 articles as in the all Wien (ZSNM; not seen). the other polistine wasps except for Belonogaster and Judging from the handwritings a large part of Polybioides, in which the female antenna has 11 articles determination labels of de Saussure’s types not only in and the male antenna 12. the MNHN but also in the MHN was prepared by du Buysson. The specimens of social wasps (also other aculeates) in the MHN were rearranged in the current Genus Ropalidia Guérin-Méneville, 1831 style during the period while J. Carl and/or C. Ferrière The type materials in the MHN here treated are all were curating the hymenopteran collection in the in the box “Vespidae 18” unless noted. museum; the original arrangement by de Saussure seemed to be retained for most specimens originating aethiopica (du Buysson) from de Saussure’s collection. Icaria aethiopica du Buysson, 1906: 110, female, male – Species group taxa are listed alphabetically in “Ethiopie méridionale: Dirouhoria”. Ropalidia, while according to the arguments on the

Taxonomic notes on Ropalidiini 3

One male and one female in the MNHN that are labeled with the MP type label in addition to “MP/ÉTHIOPIE MÉRID/DIROUHORIA/Maurice de Rothschild/1905” and “Icaria/aethiopica/♂ type Buyss. [or type ♀ Buyss.]/R. DU BUYSSON det. 1909.” are from the type series. They obviously belong to the same species; the male is here designated as the lectotype, and the female as a paralectotype. The structure of the paralectotype female is similar to the types of Icaria guttatipennis de Saussure, 1853, and those of I. politica de Saussure, 1854, which are later referred to. In order to establish their exact taxonomic status, it is necessary to make thorough comparison of the type materials of the following Figs. 1–3. Left antenna of the lectotype male of Icaria Ropalidia species from continental Africa, preferably antennata de Saussure. 1, frontal view; 2, apical with reference to new materials collected from various articles; 3, apical articles, showing tyloids. Scale lines: 0.5 mm. localities: aethiopica (du Buysson, 1906); aethiopica bimaculata Giordani Soika, 1981 (holotype in the ZSM; Kojima and Carpenter (1997: 8) indicated “Monaco” as “Delagoabay / 620 94” and a determination label the depository without listing the ZSM in the list of “Icaria / antennata / Sss [possibly in von Schulthess’ museums and/or institutions they referred to); africana handwriting]”, indicating the same origin as the (Cameron, 1910) (type probably in the NR; Bequaert female and the nest in the MHN, thus is not a syntype. (1918: 338) treated this taxon as a probable synonym of The male in the MHN is here designated as the R. aethiopica); cariniscutis (Cameron, 1910, non 1906) lectotype. (type in the BMNH; junior secondary homonym of Bequaert (1918: 338) treated Icaria ambigua Ropalidia cariniscutis (Cameron, 1906); Meade-Waldo Gribodo, 1894 as a probable synonym of R. capensis (1913: 53) treated this taxon as a synonym of I. (de Saussure, 1862), and Richards (1978: 58), without distigma Gerstaecker); cincta (Lepeletier, 1836) any comment, listed them as synonyms. Under (current type depository unknown); distigma “ambigua” in the MHN, there are three females and a (Gerstaecker, 1857) (type possibly in the ZMH); male together with two nests, all of which are labeled irrequieta (Kohl, 1907) (type possibly in the ZSNM); “Rikatla / Delagoa / (Junod)” and von Schulthess’ nobilis (Gerstaecker, 1857) (type possibly in the ZMH); identification label as “ambigua”. The MHN houses politica (de Saussure, 1854) (see later section in this also two females and two males with the same locality paper); tomentosa (Gerstaecker, 1857) (type probably in label as the above-mentioned specimens. All of them the ZMH); and unidentata Giordani Soika, 1981 are indicated to be from nest “69” and with von (holotype in the ZSM; Kojima and Carpenter (1997: 57) Schulthess’ identification label as “antennata”; one indicated “Monaco” as the depository). female with identification label “I. antennata / = I. ambigua / det. Schulthess 92”. The males doubtless antennata (de Saussure) belong to R. antennata, and then both R. antennata and Icaria antennata de Saussure, 1890: 136, footnote, female, R. capensis (de Saussure, 1862) are certainly good male – “côte de Mozambique”. species: comparison of a R. antennata female and the R. The female and male were described in the footnote capensis holotype (NNM; labeled “♀”, a circle with and de Saussure did not give any indication for the “Pr. b. Sp.”, “Mus Leiden”, a blue square with “typus” number of specimens examined nor depository. Under printed on it, “Typus / Ropalidia / capensis /♀ Sauss”) “antennata” in the MNH, there are one female, one showed that the female gena of R. capensis is distinctly male and a nest. The female and the nest have the same proportionately wider (about 0.75 times as wide as eye label, specific to the MHN, “620:[blank]:[18]94: in profile) than in R. antennata (0.63-0.67), and that the Delagoa:ach. Dayrolle” and are certainly of later origin, first metasomal segment tended to be proportionately while the male, having a small gold square and de shorter in R. antennata than in R. capensis. A brief Saussure’s determination label “Icaria / antennata / ♂ examination of specimens placed under “ambigua” in Sss. [in his handwriting on a pale blue-green label]”, is the MNHN showed that they included at least three doubtless from the type series. The ETH houses one species; two of them seemed to be R. antennata and R. female marked as a cotype, which Carpenter (1999: 34) capensis. referred to as “may well be a syntype”. The specimen I elsewhere (Kojima, 1999b) described the male is, however, labeled with “Delagoa / Dayrolle”, antennae and genitalia of two mainland African species

4 J. Kojima under “antennata” and “capensis” based on the nest; it was certainly R. artifex with its second specimens in the MCZ identified by Bequaert, but metasomal segment typical to the species (Fig. 4), and neither are R. antennata (Figs. 1-3; see also Kojima’s van der Vecht’s (1941) interpretation was confirmed. (1999b) figs. 15 and 16). My “antennata” is presumably R. capensis and possible candidates for my brazzai (du Buysson) “capensis” are the two species described by Giordani Icaria brazzai du Buysson, 1906: 110, female, male – Soika (novissima in 1944: the holotype in the MCSN; “Congo, Lékéti”. crassipunctata in 1981: the holotype in the ZSM). Bequaert (1918: 339) listed brazzai as a possible Detailed taxonomic study including the comparison synonym of R. guttatipennis; this was rejected by among the types of I. antennata, I. capensis, I. ambigua Giordani Soika (1977: 128), who mentioned that “My (probably in the MCGD), and the two species described examination of the type of this species in the Paris by Giordani Soika is required to understand the museum . . . not at all the same as guttatipennis”. mainland African fauna of Ropalidia. However, du Buysson did not indicated the number of specimens that he examined, but indicated only the artifex (de Saussure) locality and depository. In the MNHN there are three Icaria variegata Smith; de Saussure, 1853, in de Saussure, females that have the same collection label 1853-58: 25, female – “Java”. “MP/CONGO/Lékéti/de BRAZZA 1886”. They are Icaria artifex de Saussure, 1854, in de Saussure, 1853-58: equally considered as from the type series. Giordani 236. Soika’s (1977) statement is regarded as the designation De Saussure (1853-58: 25) described the female of of the lectotype by inadvertently referring to the only this species under “Icaria variegata Smith”, then, specimen that posses the MP type label and du having come to the conclusion that the species was not Buysson’s determination label “Icaria/Brazzai identical with variegata of Smith, gave it the name Buyss./type/R. DU BUYSSON det 1907” among the artifex (1853-58: 236). As van der Vecht (1941: 134) three (Article 74.6 in the International Code of pointed out, this species had not been recognized before Zoological Nomenclature). Carpenter (1999: 34), him since 1854 and the species recorded under artifex following Giordani Soika (1977), referred to the by various authors were either R. stigma or R. specimen as “the holotype”. Now the other two mathematica (see also Kojima and Carpenter, 1997). specimens are paralectotypes. However, neither van der Vecht (1941, 1962) nor any The species is easily distinguished not only from other authors have seen the type material of this species. other African species but also from any other Ropalidia The original description indicated the MHN and species by the combination of the following characters: Spinola collection (now deposited in MIZT) as body nearly entirely black; body surface strongly depository, but Casolari and Casolari Moreno (1978, sculptured (punctures as well as striae on the 1979) did not list “artifex” or “variegata” under Icaria propodeum); eye densely haired; propodeum with in the Spinola collection. I also failed to find any adult strongly raised, paired longitudinal carinae, running in specimens of R. artifex in the MHN; only the nest parallel from the posterolateral corners of the disc of illustrated in de Saussure (1853-58, pl. IV fig. 3a) was metanotum to the posterior margin of propodeum. located. I took out a fully chitinized pupa from the cabeti (de Saussure) Icaria cabeti de Saussure, 1853, in de Saussure 1853-58: 26, pl. 4 fig. 2, pl. 5 fig. 2, female – “La Tasmanie”. Ropalidia romandi cabeti; Richards, 1978: 54, 59, 63, 65, 110; Kojima, 1999a: 375 (syn. of R. romandi). De Saussure (1853-58: 26) described Icaria cabeti from “La Tasmanie” and noted “très commune”; type depositories indicated are “Musée de Paris, ma collection et celle de M. Spinola”. Richards (1978: 111) designated one of the ten females indicated as “type” in the MNHN as the lectotype. He also stated “there are also four more females from Tasmania (Verreaux)” but did not refer to their status. Aside from Fig. 4. Lateral view of the first and second the lectotype, I could locate only 12 females in the metasomal segments of a fully chitinized pupa MNHN, each with a label “MP/TASMANIE/VER- of Ropalidia artifex female, taken out from the REAUX 4-46” and a purple circle with “4/46” on its nest illustrated in de Saussure (1853-58). Scale underside. All of them are presumably from the line: 1 mm.

Taxonomic notes on Ropalidiini 5 original series and are paralectotypes. I have not seen “Transvaal” to localities; I was unable to find any the collection in the MIZT nor did Carpenter (1999) specimens identified as “clavata” by Bequaert in the refer to cabeti. In the MHN are 13 females under MCZ collection. In any case, the holotype of I. clavata “cabeti Sauss./Australie” (beneath it is an older label “I. was never referred to until Carpenter (1999: 34), who Cabeti, Sauss./Tasmanie ♀ Mr Verreaux”). Of them mentioned the second label as “an illegible label”. three females are labeled “Tasm.” and are certainly from the type series, thus are now paralectotypes. Ropalidia clavata (de Saussure) I elsewhere (Kojima, 1999a) mentioned that, “at the Icaria clavata de Saussure, 1853, in de Saussure, 1853- time when Verreaux visited Australia . . . southeastern 58: 40. part of Australian Continent was often called as Icaria duchaussoyi Gribodo, 1896: 13, female – “Nova “Tasmania” . . .” However, this was probably not Caledonia”. NEW SYNONYMY. correct; the part was called “New Hollandia” or “New Icaria marginata var. duchaussoyi; von Schulthess, South Wales” at that time. An alternative interpretation 1915: 48. could be that Hobert in Tasmania was the large port, Ropalidia clavata; Bequaert, 1918: 340. where insect specimens collected in Australian Ropalidia marginata duchaussoyi; van der Vecht, 1941: Continent could have been traded, and some or a large 104, 124. number of these specimens would have been labeled as Ropalidia duchaussoyi; Richards, 1978: 58. if they were from “Tasmania” or “Van Diemen’s Land” (J. Cardale, personal communication). cohni (du Buysson) Icaria cohni du Buysson, 1909a: 306, male – “Bougainville”. clavata (de Saussure) Van der Vecht (1941: 108) synonymized this Icaria clavata de Saussure, 1853, in de Saussure, 1853-58: 40, species under R. gregaria (de Saussure), and Richards female – “Le Cap de Bonne-Espérance”. (1978: 86) treated this taxon as a junior synonym of R. In the MNHN, there is the holotype that is labeled gregaria spilocephala (Cameron). As discussed in the with “MP/Cap de B. Espérance/COLL BOSC 1828”, next section, however, R. gregaria is a different species “Cinerca [? illegible]/ne Cap. B. Spe”, “I. clavata from that of authors. /Sauss/♀ Cap B. Sp. [in de Saussure’s handwriting on Under cohni in the MNHN, there are two males; one blue label]” and “TYPE”. The specimen is doubtless labeled “MP/I. Bougainville/(leg. Cohn)/Alfken 1909”, conspecific with R. duchaussoyi (Gribodo, 1896) the MP type label and “Icaria Cohni Buyss./types/R. described from New Caledonia. The MNHN houses a DU BUYSSON det. 1909”, and the other with the same specimen under duchaussoyi that is marked as type and collection label but “(Cohn leg.)” instead of “(leg. with the labels “MP / N. Caledonie / 1904 /A. Cohn)”. Although Carpenter (1999: 34) expressed elle Duchaussoy”, “N Caledonie” and “Icaria /Duchaus- some doubt on the type status of the second specimen, soyi / Grib. 1896 [in handwriting]”. Carpenter (1999: this is doubtless from the type series as du Buysson’s 35) doubted its type status, stating “Gribodo did not determination label for the first specimen indicates specify a type depository, however other types of “types” (in pleural form). The first specimen with the species described by him are in the MCGD.” Even if MP type label is here designated as the lectotype, and the specimen does not have type status, R. duchaussoyi the other is a paralectotype. is the only Ropalidia species so far recorded from New Caledonia and endemic to the islands. The second label gregaria de Saussure for the holotype of Icaria clavata is partly illegible and I could neither recognize the exact spelling of the first Icaria gregaria de Saussure, 1854, in de Saussure, 1853-58: word, but the label shows that the specimen came from 236, female – “La Nouvelle Hollande”. somewhere other than Cap de Bonne-Espérance (= De Saussure (1853-58: 236) indicated “Collect. de Cape Town). The original locality indicated in de M[onsieur Bernard] de Romand” as the type depository. Saussure’s original description is presumably based on The first reviser of this species was himself (de incorrect labeling. Here I synonymize Icaria duchaus- Saussure, 1862: 137), stating that the specimens soyi under Ropalidia clavata. examined in 1862 were in his collection. Since then the Smith (1857: 97) and Dalla Torre (1894: 118; 1904: name “gregaria” did not appear in any taxonomic 73) listed I. clavata in their catalogs possibly without works for nearly 80 years until van der Vecht (1941); seeing any specimens. Gribodo (1894: 125, 126) exceptions are catalogs by Dalla Torre (1894 and 1904) compared his Icaria ambigua with I. clavata, and that listed Icaria gregaria as a valid species without Bingham (1903: 48) listed a female from “Durban” seeing specimens, and Tillyard (1926) who used under I. clavata. I wonder, however, if their “clavata” “Ropalidia gregaria” to describe a biological aspect of was correctly identified. Bequaert (1918: 340) added an Australian paper wasp. Van der Vecht (1941)

6 J. Kojima extensively revised the taxonomy of Ropalidia in Indo- (Lepeletier, 1836) under “ferruginea”, that lacks in the Australian region, and synonymized Icaria cohni du metasoma and is labeled “Icaria/ferruginea/Fab [in de Buysson, 1909, I. impetuosa Smith, 1860, and I. Saussure’s handwriting]”, indicating that de Saussure’s spilocephala Cameron, 1906, under Ropalidia gregaria. “ferruginea” is R. marginata (see also Kojima and Richards (1978) returned R. impetuosa to a valid Carpenter, 1997: 37). Further, there is a female of species without any comments and treated spilocephala “gregaria” of authors labeled “amboina” under “Icaria as a subspecies of R. gregaria. I (Kojima, 1999a) marginata” in the same box in the MHN; the specimen synonymized I. spilocephala again under R. gregaria. might have been seen by de Saussure and this seems to The name “gregaria” has been applied for more than 50 be his “marginata” worker. This interpretation is also years since van der Vecht (1941) to the wasp that can supported by de Saussure’s (1853-58: 238) description be separated with certainty from R. fasciata (Fabricius, of Icaria picta (= R. fasciata): “Formes de l’I. 1804) only in the male but shows completely allopatric ferruginea, mais plus petite . . . ressemble beaucoup à distribution with the latter (see Kojima and Carpenter, l’I. marginata . . .” 1997). However, no one has referred to the types of I added a red label “Lectotype/Icaria gregaria/de Icaria gregaria. Saussure, 1854/desig. by J. Kojima, 2000” to one of the In the MHN, under “gregaria Saussure/Australia” three specimens to designate it as the lectotype of Icaria (beneath it is an older label “I. gregaria, Sauss/Nouv. gregaria (it lacks the left antenna, but is still in the best Hollande”) are three females labeled “N. Holl. [in condition among the three specimens); other two print]”. They are conspecific and I reached, based on specimens were labeled as paralectotypes. This the following observations, the conclusion that they are consequently resulted in the synonymy of Ropalidia all or partly of syntypes of Icaria gregaria – they variegata interrupta van der Vecht, 1941 under R. belong to a different species from the wasps having gregaria and the application of R. impetuosa to “R. been called “R. gregaria”. (1) As Richards (1978) gregaria” of authors, as discussed below. pointed out, de Saussure received the material from de Ropalidia gregaria (de Saussure) Romand, and according to Horn et al. (1990) hymenopteran specimens in de Romand’s collection Icaria gregaria de Saussure, 1854, in de Saussure, were deposited in the MHN through de Saussure, and 1853-58: 236. thus the types of Icaria gregaria may well be in the Ropalidia variegata interrupta van der Vecht, 1941: MHN. (2) Richards (1978: 86) stated that the form 158, female – “Thursday Island” (holotype in the from Northern Territory of Australia that he called “R. MCZ, examined). NEW SYNONYMY gregaria gregaria” “is the form that agrees best with de Ropalidia interrupta interrupta; Richards, 1978: 89. Saussure’s original description [of Icaria gregaria]”, Ropalidia interrupta; Kojima, 1999a: 373. but there are some distinct differences between Below is the redescription of the lectotype, with specimens of “gregaria” of authors from Northern variations in paralectotypes indicated in brackets. Territory and de Saussure’s descriptions (1853-1858 Body length (head + mesosoma + first two meta- and 1862). In the original description, de Saussure somal segments), about 7.5 mm; forewing length, about (1853-58) stated “Petite taille de l’I. socialistica”, while 5.7 mm. “gregaria” of authors is as long as or even longer than Head rather large and thick, about 1.2 times as wide R. socialistica (de Saussure, 1853). “R. gregaria” of as mesosoma excluding tegulae, a little more than 1.1 authors has the discs of scutellum and metanotum and times as wide as high (Fig. 5); in dorsal view about 1.8 paired longitudinal bands on the posterior face of times as wide as long, hardly emarginate posteriorly, propodeum all colored yellow, which is not mentioned with lateral sides behind eyes weakly convex and in de Saussure (1853-58, 1862). Furthermore, the slightly convergent posteriorly (Fig. 7). Inner eye second metasomal segment is obliquely cut off margins convergent ventrally; distance between them at posteriorly in “gregaria” of authors and is colored vertex about 1.3 times as long as at clypeus. Clypeus ferruginous in the Northern Territory specimens of convex, transverse, about 1.3 times as wide as high. “gregaria” of authors, while de Saussure’s (1862) Mandibles not twisted, with dorsal tooth truncate description says that the second metasomal segment is apically. Vertex wide, slightly convex; area among “tronqué verticalement à l’extrémité”, and “noir” in ocelli hardly raised; smoothly but distinctly sloping 1853-58 and “brun ou noirâtre” in 1862. The three down to occipital carina. Ocelli arranged as follows: specimens in the MNH agree with de Saussure’s distance between posterior ocelli about 1.3 times larger original and 1862 descriptions. (3) There is no doubt than distance between anterior and posterior ocelli, and that de Saussure (1853-58: 237 and 1862: 139) about 2.4 times as long as diameter of posterior ocellus, identified “gregaria” of authors as “Icaria marginata”. which is slightly smaller than anterior ocellus; posterior In the MHN there is a female of R. marginata ocelli separated about 0.75 times as far from each other

Taxonomic notes on Ropalidiini 7 as from inner eye margin. Antenna strongly clavate Disc of scutellum convex, but median part nearly flat, apically, thickest at tenth article, where it is about 1.5 margined laterally by weak truncation, with fine times as wide as apical width of third article (Fig. 8); median, impressed line. Disc of metanotum nearly flat scape about 3.5 times as long (excluding radicle) as medially, margined laterally with carinae; posterior wide at apex; third article a little more than 1.5 times as margin nearly straight. Propodeum with wide median long as wide at apex, slightly shorter than length of concavity on posterior face; median furrow distinct; in fourth and fifth articles combined, each of which is profile, posterior face slightly convex; propodeal orifice distinctly wider than long; tenth and eleventh article rounded above, about twice as long as wide (Fig. 10); nearly twice as wide as long; terminal segment bullet- valvula large, semicircular in outline. shaped, nearly as long as wide at base. Gena rather Metasoma as in Figs. 11 and 12; first tergum about developed, in lateral view about 0.8 times as wide as 1.4 times as long (measured in profile from posterior eye (Fig. 6), weakly depressed posteroventrally; margin of basal slit to posterodorsal margin) as wide occipital carina complete, fine entirely, in profile of (maximum width in dorsal view) [this segment is more head smoothly curved. or less elongate in “gregaria” of authors while de Pronotum in dorsal view broadly rounded anteriorly, Saussure’s (1862) description states “assez court”]; with lateral sides hardly diverging posteriorly, slightly ventral margins of tergum closely approaching to each convex or nearly straight; pronotal carina complete, other before posterior divergence; posterior margin of produced anteriorly in thin lamella at humeral angles, first sternum weakly emarginate; articulation of second where it is distinctly sinuate backward (Fig. 9). Scutum tergum separated from “neck” by sharp truncation about as long as wide between tegulae, weakly convex. laterally and blunt truncation medially; second segment

Figs. 5–12. Lectotype female of Icaria gregaria de Saussure. 5–7, head (5, frontal view; 6, lateral view; 7, dorsal view); 8, right antenna; 9, lateral view of pronotum; 10, propodeal orifice; 11–12, first two metasomal segments (11, lateral view; 12, dorsal view). Scale line: 1 mm, but 0.5 mm for Fig. 10.

8 J. Kojima in profile vertically cut off posteriorly; posterior margin metasomal segments with yellow apical band; all coxa of second segment widely depressed into lamella. with yellow spot. Wings hyaline; infuscate antero- Body covered with dense appressed silvery apically. tomentum and suberect hairs; hairs in apical part of clypeus sparser but longer; posterior semioval area of I elsewhere (Kojima, 1999a) synonymized R. metanotum polished, without hairs; anteroventral part mutabilis mutabilis Richards, 1978, and R. mutabilis of mesepisternum without distinct hairs. Punctures on torresiana Richards, 1978, under R. interrupta and R. body surface dense, with their interspaces less than their flavinoda van der Vecht, 1941, respectively. However, diameters; punctures on clypeus shallow, sparse and ill- recent re-examinations of the holotypes, paratypes and defined, but larger than those on front; punctures on additional specimens in the ANIC made me to doubt vertex slightly sparser than on front and gena; posterior these synonymies. The taxonomic reconsideration of polished area of metanotum without punctures, the R. variegata-group, which includes R. gregaria, R. occupying medially in half to two-third length of variegata flavinoda, R. mutabilis mutabilis, R. mutabilis metanotum; anteroventral part of mesepisternum torresiana, R. variegata variegata (Smith, 1852), R. unpunctured, its posterior margin bordering with variegata dichroma van der Vecht, 1941, R. jacobsoni coarsely punctured posterodorsal area well defined, but (du Buysson, 1908) R. pendula (Smith, 1857) and R. not carinate; metapleura sparsely punctured dorsally, revolutionalis (de Saussure, 1853), will be published hardly punctured ventrally; propodeum barely elsewhere. punctured in anterolateral part and ventral part of Two names may be applicable for the species of posterior concavity, with oblique striae together with “gregaria” of authors; namely, R. bioculata (Fabricius, coarse punctures between striae on posterolateral face; 1804) and R. impetuosa (Smith, 1860). Although I dorsal part of posterior concavity of propodeum with have not seen the holotype of Polistes bioculata punctures and weak transverse striae. Anterior face of deposited in the Zoologisk Museum in Kobenhaven, Dr. first metasomal tergum without punctures, and its R. Meier kindly sent me photographs of the holotype of posterior widened part with large but shallow P. bioculata and informed me its label data (with the punctures; second tergum with large, rather dense following three labels “biocula/ta [in Fabricius punctures, each of which is ill-defined posteriorly; handwriting]”, a standard Kobenhaven red type label, punctures on second sternum similar to those on and “Ropalidia ♀/fasciata (F.)/=bioculata (F.)/det J. v. tergum, but smaller. d. Vecht 1963”). Richards (1978: 86) stated that the Body ferruginous with yellow (darkened possibly female holotype of P. bioculata “. . . is in bad condition due to bad preservation during the transportation from [both eyes and right antenna lacking, and with some Australia to Europe) marks and black (or blackish molds] and might be either R. gregaria or R. fasciata”, brown) parts as follows: clypeus with large obscure which would be the reason why Richards proposed to yellow spots laterally [wide u-shaped band along apical ignore “bioculata”. However, I do not regard this as a margin yellow]; mandible with a yellow spot at sufficient reasoning to ignore any names. Fabricius dorsoproximal corner [absent], teeth darker margined (1804: 278) described bioculata from “nova Cambria”, with blackish brown; spot occupying the bottom of which could be somewhere in eastern part of Australia, inner eye margin and spot between antennal sockets distribution range of “gregaria” of authors, but not of R. yellow; ill-defined spot above each antennal socket and fasciata. Nevertheless, the following facts suggest that area among ocelli black; ill-defined small spot at Fabricius’ locality was incorrect, and lead me to bottom of gena yellow [absent]; antennal scape and propose to synonymize Polistes bioculata under second article yellow below, fourth to terminal articles Ropalidia fasciata (Fabricius, 1804), NEW yellowish beneath, brown to dark brown above; narrow SYNONYMY, and to adopt R. impetuosa for “R. band along pronotal carina yellow [reduced]; margins gregaria” of authors. of scutum nearly black [scutum entirely darker than The photographs clearly showed that, as Richards pronotum]; anterior and posterior margins and anterior stated, the holotype of P. bioculata is, in its structural half of median line of scutellum black; metanotum characters, either “R. gregaria” of authors or R. fasciata. black in anterior margin, with ill-defined, dark-yellow The specimen has its black marking on the head spots laterally [yellow spots hardly recognizable]; restricted to the area among ocelli, and the pronotum tegula yellow, with outer large ferruginous spot; largely yellow dorsally and the second metasomal metapleura and ventral margin of propodeum darker; tergum with a pair of large lateral yellow markings near propodeal teeth yellowish brown; second metasomal base. The marking pattern is rarely found in “R. segment darker than first tergum, blackish brown, with gregaria” of authors; I have never seen such a marking narrow, yellow pre-apical band [first tergum also with a pattern in any specimens of “R. gregaria” of authors trace of yellow apical band]; third to terminal from Australia. Instead, this marking pattern is, as far

Taxonomic notes on Ropalidiini 9 as I know, common in the populations of R. fasciata Carl (1930: 22-23) briefly described nests of a from Okinawa, Taiwan and mainland China. Ropalidia wasp from India, referring it as “Icaria montana n. sp.” If Article 12 in the International Code Ropalidia impetuosa (Smith) of Zoological Nomenclature is strictly applied, this act Icaria impetuosa Smith, 1860: 131, female – “Batchian; is regarded as the description of a new species, with the Amboyna” (lectotype in the OUM, examined). — nests he described as types (Article 72.5.1). However, van der Vecht, 1941: 104 (syn. of R. gregaria). because Carl (1930: 23) mentioned “c’est sous ce nom Ropalidia impetuosa: Bequaert, 1932: 51. — Richards, que nous décrirons prochainement l’Insecte”, I propose 1978: 58 (listed as a good species). to take his later description of the wasp (Carl, 1934) as Icaria spilocephala Cameron, 1906: 230, female – the original description of Ropalidia montana, as “Etna Bay” (lectotype in the ZMA, examined). — Kojima and Carpenter (1997) did. van der Vecht, 1941: 104 (syn. of R. gregaria). Under “montana” in the MHN are five females Ropalidia gregaria spilocephala; Richards, 1978: 53, labeled “Indes/Coonoor/J. Carl [in handwriting]” that 62, 86. — Kojima, 1999a: 371 (syn. of R. gregaria, are undoubtedly the specimens that his measurements erroneously marked “Syn. nov.”). of body size were based on; he stated “6 mm - 7 mm, 5” Icaria cohni du Buysson, 1909a: 306, male – (p. 676). These specimens are accompanied with a “Bougainville” (MHNH). — van der Vecht, 1941: label “voix nombraux specimens/dans Boire de doubles 104 (syn. of R. gregaria). /(Vespidae n° 17)”. A glass tube containing a large Ropalidia gregaria inquieta Cheesman, 1952: 3, 20, number of wasps (apparently more than 100 wasps) is female, male – “Papua, . . . Mafulu, 4000 ft.” in the box for the “duplicate” or “extra” specimens, but (lectotype female in the BMNH, examined). — I wonder if Carl used these specimens as the bases for Richards, 1978: 86 (syn. of R. gregaria spilo- his description. Another five pinned females were in cephala). the other box for the “duplicate” specimens; three of Ropalidia gregaria var. tolerans Cheesman, 1952: 3, them are labeled “Voy. Carl et Escher / Inde 21, female – “Cyclops Mts. Sobron” [New Guinea] mèridionale / Coonoor / Nilgiris 23-XII-26” and the (lectotype in the BMNH, examined). — Richards, remaining two specimens are without labels. They 1978: 86 (syn. of R. gregaria spilocephala). presumably constitute the type series. I transferred Ropalidia gregaria of authors, listed in Kojima and them to the box “Vespidae 18” and placed together with Carpenter (1997). the above-mentioned five females. The ETH houses three females and a male marked as cotypes. The guttatipennis (de Saussure) females are labeled also with “S. India /Nilgiris/(Carl)” Icaria guttatipennis de Saussure, 1853, in de Saussure, 1853- and “I. montana/Carl det.” and are probably from the 58: 40, pl. 5 fig. 8, female – “Le Sénégal”. type series, while the male that has a collection label “Nilgiris/S. India/Carl, 1929” and a determination label In the MNHN, there are four females marked with of von Schulthess is probably not from the type series. the MP type label: two females have also a label “MP One of the five females labeled “. . . Coonoor . . .” in /SÉNÉGAL/GUÉRIN” and a circle with “Senegal the MHN was labeled as the lectotype “Lectotype /Guerin”, and one of them has an additional long, blue /Ropalidia montana/Carl, 1934 /desig. J. Kojima, label on which is written “I. guttatipennis/Sauss. [in de 2000”, and the remaining 12 females are paralectotypes. Saussure handwriting]”; the other two females have I failed to locate any males of R. montana or R. additional labels “MP/ SÉNÉGAL/ROBERT 1836” and artifex that he examined for their genitalia. Judging a blue circle with “1[or 7?]60/36” on its underside. As from his drawings and the fact that many specimens of pointed out by Carpenter (1999: 35) all of them are R. fasciata are placed under “artifex” in the MNH, presumably from the type series. Under “guttatipennis” Carl’s “artifex” may well be R. fasciata (see also in the MHN is one females labeled with a small silver Kojima, 1999b; note a proportionately shorter penis square and a label written in hand as “Seneg”, that is valve in Carl’s drawing), although Kojima and also presumably from the type series although de Carpenter (1997) listed Carl’s “artifex” under R. stigma Saussure indicated only the MNHN as depository. The (Smith, 1858). specimen with de Saussure’s determination label in the MNHN is here designated as the lectotype, and the revolutionalis (de Saussure) other four specimens are paralectotypes. Icaria revolutionalis de Saussure, 1853, in de Saussure, 1853- montana Carl 58: 29, pl. 5 fig. 7, female – “La Nouvelle Hollande ou la Tasmanie”. Ropalidia montana Carl, 1934: 675, figs. 1-6, 8-13, female, male – “le versant méridional de Nilgiris . . . Coonoor De Saussure (1853-58: 29) described this species [India]”. from “La Nouvelle-Hollande ou la Tasmanie” and listed

10 J. Kojima the MNHN as depository. Although de Saussure (p. 30) socialistica (de Saussure) referred to a single female that was uncertain for its Icaria socialistica de Saussure, 1853, in de Saussure, 1853- affiliation, he did not give any indication for the 58: 27, pl. 4 fig. 6, female – “La Tasmanie.” number of specimens on which his description of Icaria revolutionalis was based. Richards (1978: 91) stated De Saussure (1853-58: 27) described this species that “The holotype female at MNHN, Paris, is labelled from “La Tasmanie” and indicated the MNHN and his ‘Australie (Verreaux) 2-47’. . .”, which should be collection as depository. Richards (1978: 81) considered as the designation of the lectotype by designated one of the three females marked as type in inadvertently stating it as “the holotype” (Article 74.6 the MNHN as the lectotype. In addition to the MP type in the International Code of Zoological Nomenclature label and designation label in Richards’ handwriting, (ICZN, 2000)). The specimen is labeled also with a the lectotype has labels of “MP/TASMANIE purple circle with “2/47” on its underside, /VERREAUX 4-46”, a purple circle with “4/46” on its “revolutio/nalis [in pencil, possibly in de Saussure’s underside, “TYPE/HOLOTYPE” [in handwriting on handwriting]” and “Icaria/revolutionalis/type Sauss. [in red label], and “Icaria/socialistica/types Sauss. [in du du Buysson’s handwriting]/H. DE SAUSSURE DET Buysson’s handwriting]/H. DE SAUSSURE DET [in [in print]”. This is the only specimen of R. press]”. In addition to another two females that revolutionalis that I could find in the MNHN and de Richards referred to, four females (three are headless) Saussure indicated only the MNHN as depository. with the same collection label are placed together with Nevertheless the following specimens in the MHN the lectotype. They are also presumably from the type placed under “revolutionalis/Sauss./ Australie” (beneath series and are now paralectotypes. In the MHN, there it is an older label “I. revolutionalis/Nouv. Holl. Sauss. are five females under “socialistica/Sauss./Australie” /Mr Verreaux”) may well constitute type series and are (beneath it is an older label “I. socialistica, thus paralectotypes: one female labeled “N. H.”; one Sss./Tasmanie”) that are labeled with only a piece of female with a small silver square and a label silver square but are also presumably from the type “Rockhampton/N. Holl.”; two females with the same series as de Saussure indicated his collection as small silver square. depository; I labeled them as paralectotypes. plebeja (de Saussure) Genus Belonogaster de Saussure, 1854 Icaria plebeja de Saussure, 1863: 235, female – “Nouvelle- Hollande”. The specimens of Belonogaster in the MHN are in the box “Vespidae 16”. De Saussure used “Icaria plebeja” twice for quite different species, once in 1862 for the species from brevipetiolata de Saussure Celebes (= Sulawesi) (the holotype is housed in the Belonogaster brevipetiolatus de Saussure, 1890: 89 (key), 98, NNM), and in 1863 for the species from Australia. pl. 4 fig. 1, female, male – “Madagascar. – Fianarantsoa Richards (1978: 75) described Ropalidia plebeiana as a (pays des Betsileo) et environs d’Andrangoloka (à la new concept, listing “Probably Icaria plebeja de limite est de la province d’Imerina) . . . Còte sud . . . Saussure, 1863” and stating “I have not seen the type Nosibé”. material [of I. plebeja de Saussure, 1863, non 1862] but In the MHN, there are 11 females and nine males I am following the usual interpretation of the species.” under “brevipetiolatus”, including a male labeled In the MHN, there are four females and a nest: one “MADAGASCAR/Fianarantsoa”, and “Belonogaster female without any label; three females and a nest /brevipetiolatus/de Saussure/Lectotype ♂ /Richards labeled with “N. Holl” (the three females also with a 1979 [in Richards’ handwriting]”. Richards (1982: small silver square; the nest with an additional label “N 101), however, did not publish the designation of the H” in handwriting). De Saussure (1863: 235) described lectotype; the specimen was later designated as the a nest, with which this nest well agrees; then the four lectotype by Carpenter (1999: 30) and has his red females are highly probably from the type series of I. designation label “LECTOTYPE/Belonogaster/brevi- plebeja de Saussure, 1863. Here I designate a female petiolata ♂ /(Saussure 1891)/des. J. M. Carpenter, with additional labels “Ropalidia ♀ /plebeiana 1991”. Paralectotypes designated by Carpenter have Rich./det. J. v. d. Vecht 1975” and “MS-name!” as the been also properly labeled. lectotype and added a red label “Lectotype/Icaria Hensen and Blommers (1987: 16) mentioned that plebeja/de Saussure, 1863, non 1862 /desig. by J. “Most specimens examined by Richards belong to the Kojima, 2000”. The synonymy of Icaria plebeja de related species B. erythrocephala . . . The ♂ Saussure, 1863, non 1962, with R. plebeiana Richards, identified . . . by Richards . . . as B. brevipetiolata 1978, is now established. belongs to a third species, B. betsileo . . .” However,

Taxonomic notes on Ropalidiini 11 they did not refer to any syntypes in the MHN. The of Belonogaster in Madagascar and provided a key to lectotype and all the paralectotypes in the MNH are species. The holotype of B. apicalis may not be highly likely conspecific and Hensen and Blommer’s properly keyed; because it has the fore coxa with dense “brevipetiolata” is the true B. brevipetiolata. white pubescence and rather dense, black setae slightly longer than the white pubescence, the user may stop at apicalis de Saussure couplet 9, in which they mentioned “Fore coxa with Belonogaster apicalis de Saussure, 1900: 207 (key), 208, white pubescence, rarely with a few longer dark setae” female – “Madagascar”. to lead to B. apicalis. They doubted that the two males Richards (1982: 105) designated a female in the in the NNM examined by Richards (1982) belonged to MHN as the lectotype of B. apicalis, but he did not this species. I examined these two males and agreed label it (see also Carpenter, 1999: 30). A female in the with Hensen and Blommers based on the following MHN labeled “MADAGASC. [in print on blue label]”, observations. (1) The two male specimens showed “Belonog./apicalis/♀ Sss. [in probably de Saussure’s distinct sexual differences not only in the characters handwriting]”, and “bonnne éspéce/du Buysson [in du usually representing sexual dimorphism in Belono- Buysson’s handwriting]” is undoubtedly the specimen gaster (such as coloration and structure in the head), but that Richards referred to. Although de Saussure did not also in the coloration of wings and metasoma, and in give any indication for the number of specimens he the shape of the first and second metasomal segments, examined, du Buysson’s (1909b: 234) statement “coll. all of which are usually not distinctly different between H. de Saussure, Musée de Genève, type” indicates that sexes in Belonogaster. (2) The ETH houses two males the specimen is the holotype of B. apicalis, and I added that well agree with the type female of B. apicalis in the a label indicating that the specimen is the holotype of B. coloration of wings and metasoma as well as in the apicalis. shape of the first two metasomal segments. They have Hensen and Blommers (1987) revised the taxonomy the antennae and terminal tarsal segments distinctly

Figs. 13–21. Male of Belonogaster apicalis de Saussure. 13–14, head (13, frontal view; 14, lateral view); 15, right antenna; 16–17, apical part of left antenna of the same specimen as of Fig. 15; 18, apical part of right antenna of another specimen; 19, tarsal segments of hind leg; 20–21, first two metasomal segments (20, lateral view; 21, dorsal view). Scale lines: 1 mm, but 0.5 mm for Figs. 16–19.

12 J. Kojima different from the description and drawings for the male 15. Forewings dark gray, with the tip light red-brown in of Richards’ “apicalis”. Thus I concluded that these both sex; mesosoma entirely black; last article of two males are the true males of B. apicalis. The two male antenna weakly curved, hardly narrowed males in the NNM that Richards (1982) referred to as apically with rounded apex (Figs. 15–18); apical the males of “B. apicalis” seem to be different species segment of male fore tarsus hardly dilated apically, from each other, but their exact affiliations were not with truncate apical margin (Fig. 19)……...…………… determined...... apicalis de Saussure The male of B. apicalis is described below based on — Forewings uniformly yellow to brown; mesosoma the two males in the ETH; both labeled “Rogez/O. never entirely black...... 16 Madagasc.” and “ ♂ ” (one with an additional identification label “?apicalis/det. Schulthess 93”; and bicolor de Saussure the other with “?apicalis/sec. coll. Guep [last word illegible]”). Belonogaster bicolor de Saussure, 1900: 207 (key), 208, female – “Madagascar”. Male — Body length (head + mesosoma + first two Richards (1982: 104) designated the lectotype, metasomal segments) about 18 mm; forewing length which is in the MHN and labeled “madgasc/Madagasc about 17.5 mm. [in print on a blue label]”, “ne sont que des/variations Head in frontal view about 1.1 times as wide as high de colorie/du B. prasinus/De Saussure/Du Buysson [in (Fig. 13). Inner eye margins hardly converging below; du Buysson’s handwriting]”, “bicolor, Sss./ ♀ distance between them at vertex only slightly longer Madagascar [in handwriting]”, and “Belonogaster than that at clypeus. Clypeus slightly concave below, /bicolor Sauss./Lectotype ♀ /Richards, 1979 [in higher than wide, rather strongly produced below, with Richards’ handwriting]”. However, Richards did not angled apex. Gena narrow, in profile about half as wide refer to other syntypes. A female in the MHN labeled as eye (Fig. 14). Antennal scape similar to that of “Madagascar/F Sikora [in print on blue label]” and female, about 4.5 times as long as wide at apex (Fig. “bicolor/ Sss. ♀ [possibly in de Saussure’s handwriting, 15); third to terminal articles with tyloids of on blue label]” is certainly from the type series; the longitudinal carina; third article about three times as specimen is now a paralectotype. long as wide, about 1.5 times as long as fourth article; The type of B. bicolor stops at couplet 11 in the key each of fourth and fifth articles about twice as long as of Hensen and Blommers (1987), who synonymized B. wide; tenth and eleventh articles nearly as wide as malagassus under B. bicolor. However, the two taxa preceding article, each about twice as long as wide; certainly belong to different species as noted in the next terminal article about 0.7 times as wide as eleventh section, and the key of Hensen and Blommers was article, about 3.2 times as long as wide at base, weakly undoubtedly based on the lectotype (and possibly curved and bluntly rounded apically (Figs. 16–18). paralectotypes) of B. malagassa that they observed. Mesosoma and metasoma (Figs. 20, 21) as in the female. malagassa de Saussure Body black; gena tinged with blackish green; front Belonogaster malagassus de Saussure, 1900: 207 (key), 210, below level of eye emargination, clypeus and mandible female – “Madagascar”. pale ivory, irregularly tinged with pale leaf-green; wide median band on clypeus and front among dorsal Hensen and Blommers (1987) designated the emargination of clypeus black; mandibular teeth brown. lectotype and synonymized B. bicolor under B. Antenna ferruginous; scape blackish brown, beneath malagassa. However, their decision was certainly with whitish band tinged with pale leaf-green; second to based on the description of B. bicolor by Richards sixth articles dark above. First metasomal segment and (1982: 105) but not on examination of the lectotype of basal part of second segment dark green; first tergum B. bicolor. Belonogaster bicolor is different from B. posteriorly tinged with black; second (except basal part) malagassa in several characters as noted below, and to terminal metasomal segments blackish brown to both should be treated as good species. There are four black, each apically with ill-defined, dark brown band. females of B. malagassa in the MHN as Hensen and Legs black; tarsi dark ferruginous; fore coxa sometimes Blommers (1987) mentioned, and all of them are tinged with blackish leaf-green. Wings as in the properly labeled to indicate their type status: the ♀ female; dark gray, with apical one-fourth of fore wing lectotype is labeled “Belonog./malagassus/ Sss. [in de semitransparent, colored light reddish brown. Saussure’s handwriting]”, “Madagascar/F Sikora [in print on blue-green label]”, “LECTOTYPE ♀ Couplet 15 of Hensen and Blommers’ (1987) key is /Belonogaster/Malagassa Sss./R. Hensen 1986” [red ♀ modified to summarize the male characters. label], and “Belonogaster/ /bicolor Sss./Det. R. Hensen/1986”.

Taxonomic notes on Ropalidiini 13

The most distinct structural difference between B. long as wide at apex) than in B. bicolor (Fig. 25: less bicolor and B. malagassa is the condition of the than 3 times as long as wide at apex), but the condition posterior face of the propodeum; namely, it is only in the paralectotype of B. bicolor (Fig. 26) approaches weakly convex and with a distinctly impressed line in at that in the lectotype of B. malagassa. The tarsi of the least the posterior two-thirds in B. malagassa (Fig. 22). hind leg (especially first to third tarsal segments) are On the other hand, in B. bicolor the posterior face of the more slender in B. malagassa (Fig. 27) than in B. propodeum is more strongly convex and lacks the bicolor (Fig. 28). impressed median line although a dark line, which may Couplet 11 of Hensen and Blommer’s (1987) key is look impressed, is present along its entire length (Fig. modified as follows to distinguish between B. 23); Richards (1982: 105) stated “impressed line strong, malagassa and B. bicolor. complete”. Body color is also distinctly different between the two species: the mesosoma of B. 11. Propodeum with impressed median line at least in malagassa is entirely dark ferruginous, while in B. posterior two-thirds or with dark median line in bicolor the anterior part of the mesosoma (pronotum, nearly its entire length...... 11’ scutum, mesepisternum except posterior margin) is — Propodeum without such impressed or dark median black and its posterior part is colored with ferruginous line...... 12 to pale ferruginous; Richards (1982: 105) stated 11’. Posterior face of propodeum relatively weakly “Mesosoma light ferruginous, proepisternum convex, with impressed median line, which is ferruginous”. Further, the following differences were evanescent anteriorly (Fig. 22). Mesosoma entirely recognized between the lectotypes of the two species, ferruginous to dark ferruginous………………………. but these differences could hardly serve as the … ...... malagassa de Saussure characters diagnosing the two species when further — Posterior face of propodeum convex, with dark, but additional materials are examined. The antenna of B. not impressed median line in nearly its entire length malagassa has the flagellum relatively more slender (Fig. 23). Mesosoma black anteriorly (pronotum, (Fig. 24: second antennal article more than 3 times as scutum, and mesepisternum except posterior margin), ferruginous or light ferruginous posteriorly. …...... bicolor de Saussure

eumenoides de Saussure Belonogaster eumenoides de Saussure, 1890: 89 (key), 94, pl. 17 fig. 12, female – “Madagascar . . . aux environs d’Andrangoloaka”. The lectotype designated by Richards (1982: 110) is labeled “Madagascar/Andrengoloka”, “eumenoides /Saussure ♂ [probably in du Buysson’s handwriting]” and has a lectotype designation label in Richards’ handwriting. Another five females with the same collection label and du Buysson’s (probably) determination label (all of them are also labeled with Hensen’s determination label; and one of them with an additional label “Belonogaster/eumenoides/♀ Sss [in de Saussure’s handwriting on a larger label]”) are certainly from the original series and are paralectotypes. Another female with the same collection label but a determination label “Belonogaster eumenoides/♂ S [in de Saussure’s handwriting]” may well be from the type series: the presence of a male with the same collection label and a label “Belonogaster/eumenoides/♀ Sss [in de Saussure’s handwriting]” suggests that the determination labels by de Saussure might have been

Figs. 22–28. Females of Belonogaster malagassa de swapped by accident in the course of research by Saussure (22, 24, 27) and B. bicolor de Saussure someone else. Another four females that are labeled (23, 25–26, 28). 22–23, propodeum, dorsolateral “Madagasc/Annanarive” are also presumably from the view; 24–26, left antenna; 27–28, tarsus of hind leg. original type series as de Saussure indicated “Région Scale lines: 1 mm. centrale (province d’Imerina)” as a locality. Five

14 J. Kojima females and a male collected by Sikora are all labeled them at vertex about 1.2 times as long as that at as “var.” and are not paralectotypes. clypeus. Clypeus nearly flat, slightly higher than wide, weakly produced below, and with bluntly angled apex. Gena narrow, in profile nearly half as wide as eye (Fig. 40); malar space rather short, about as long as diameter of antennal socket. Mandible normal. Ocelli arranged in equilateral triangle and close to each other; distance between posterior ocelli about 0.7 times as long as their diameter, and nearly half distance between posterior ocelli and inner eye margin. Antenna slender, curled apically (Fig. 41); tyloids in longitudinal slightly raised carina (Fig. 42), present in apical half to one-third of third article and on fourth to terminal articles, weak on terminal article; scape short, very weakly swollen medially, about three times as long as wide; third article long, about 4.7 times as long as wide, nearly 0.8 times as long as fourth and fifth articles combined; fourth and fifth articles each 2.5-3.5 times as long as wide; last three articles distinctly narrower than preceding articles; tenth and eleventh articles each about three times as long as wide; terminal article nearly five times as long as wide at base, curved, tapering into sharply pointed apex in apical one-third (Figs. 43, 44). Scutum weakly convex, about as long as wide, with median carina in anterior half. Scutellum weakly Figs. 29–34. Belonogaster eumenoides de Saussure. convex, with median carina in anterior half. 29–32, lectotype female; 33, paralectotype Propodeum weakly and evenly convex, with median, male; 34, male indicated as “var.” 29, slightly depressed line, which is obliterated anteriorly. scutellum, metanotum and propodeum, lateral Apical segment of fore tarsus hardly dilated apically view; 30, first and second metasomal segments, (Fig. 45). lateral view; 31, first metasomal tergum, dorsal First metasomal segment about ten times as long as view; 32, second metasomal tergum, dorsal wide (Figs. 46, 47); in dorsal view weakly widened view; 33–34, apical part of right antenna. Scale lines: 1 mm. posteriorly to just behind spiracle, then nearly parallel- sided, where the width is about 1.5 of basal width; in profile slightly thickening apically, with dorsal surface Von Schulthess’ collection in the ETH houses a slightly and smoothly curved. Second segment male identified as “eumenoides” by von Schulthess and gradually widening posteriorly after basal stalk, with another male from Madagascar, which are conspecific posterior width about five times as wide as basal width and certainly belong to an undescribed species of and about 3.5 times as wide as first tergum; basal stalk Belonogaster. A female also identified as “eume- about 1.75 times as long as wide. noides” by von Schulthess agrees with these two males Body covered with dense, appressed, white in the characters that usually do not show sexual tomentum; tomentum sparser and shorter on metasoma dimorphism in Belonogaster. A new species is than on other body parts; vertex and mesosoma (except described below based on these three specimens. metapleura) with dense, long, dark brown hairs, protruding from tomentum; ventral face of first Belonogaster schulthessi new species metasomal segment with similar, but much sparser, hairs; fore coxa with the same dark brown hairs on (Figs. 36–47) anterior face. Vertex and mesosoma covered with Male — Body length (head + mesosoma + first two dense, microscopic punctures, some of which are metasomal segments) about 15 mm; forewing length slightly larger; mesepisternum posteroventrally covered about 13.5 mm. with punctures similar to those on scutum, antero- Head about as wide as mesosoma excluding tegulae, ventrally smooth with scattered shallow punctures, with in frontal view about 1.1 times as wide as high (Fig. ill-defined border between punctured and smooth areas; 39), slightly more than twice as wide as long. Inner eye metapleura smooth, with scattered shallow punctures. margins weakly converging ventrally; distance between Metanotum hardly punctured.

Taxonomic notes on Ropalidiini 15

Head colored ivory yellow; clypeus with wide propodeum dark leaf-green. Legs dark leaf-green; fore median longitudinal band colored pale brown; coxa ivory yellow; tarsi brown to dark brown. Wings mandibular teeth margined with brown; front above hyaline, colored pale brown. level of center of eye emargination and vertex dark leaf- Female — Body length (head + mesosoma + first green, extending laterally on gena; antenna ferruginous, two metasomal segments) about 16 mm; fore wing with scape paler and tinged with pale leaf-green; length about 15 mm. flagellum darker above. Pronotum dark leaf-green, Structure similar to male, but head different as with paired narrow bands on anterolateral margins follows. Head about 1.05 times as wide as high (Fig. [absent in the paratype]; scutum nearly black [dark 36), slightly less than twice as wide as long; inner eye brown in the paratype]; scutellum and metanotum margins less convergent below, distance between them nearly black, tinged with dark blackish green, with ill- at vertex about 1.1 times as long as that at clypeus; defined pale-brown band on each posterior margin; clypeus strongly produced below, nearly 1.1 times as mesepisternum dark leaf-green, with ill-defined pale- high as wide, with sharply angled apex, bearing sparse, brown scrobal spot and anteroventral spot [in the dark-brown setae, without tomentum in apical produced paratype, mesepisternum pale ferruginous, with ill- part; gena in profile nearly 0.6 times as wide as eye (Fig. defined green spot near dorsal margin]; metapleura and 37); malar space long, longer than diameter of antennal

Figs. 35–47. Belonogaster eumenoides de Saussure (35) and B. schulthessi new species (36–47). 35–38, female; 39–47, male. 36–37, 39–40, head (36, 39, frontal view; 37, 40, lateral view); 35, 38, 41, left antenna; 42, first two flagellar articles, showing tyloids; 43–44, apical part of left antenna in different views; 45, tarsal segments of hind leg; 46–47, first two metasomal segments (46, lateral view; 47, dorsal view). Scale lines: 1 mm, but 0.5 mm for Figs. 42–45.

16 J. Kojima socket. Antenna slender (Fig. 38); scape about 4.4 times as long as wide at apex; third article about 3.5 times as long as wide at apex, about as long as length of fourth and fifth articles combined; fourth article about 1.4 times as long as wide; fifth article slightly longer than wide; antennal flagellum thickest at ninth article, which is about 1.35 times as wide as long and slightly wider than apical width of fifth article; terminal article bullet shaped, about 1.3 times as long as wide at base. Head including mandibles pale ferruginous, tinged with pale leaf-green; apical margin of clypeus and teeth of mandible brown; front above level of lower margins of antennal sockets and vertex blackish green or dark brown tinged with blackish green. Antenna blackish brown; scape tinged with blackish green, paler beneath; flagellum reddish brown beneath. Coloration of mesosoma and metasoma as in the male, but second to terminal metasomal segments darker; fore coxa pale leaf-green. Holotype — male, labeled “Rogez/O. Madagasc”, “♂”, and “eumenoides/det. Schulthess 93” (deposited in the ETH). I added a label “Holotype/Belonogaster schulthessi Kojima, 2001”. Paratypes — 1 male, labeled “Mada-/gascar” (deposited in the ETH); 1 female, labeled “Mada- /gascar”, “♀” and “eumenoides/det. Schulthess 93”. Figs. 48–53. Belonogaster ornata de Saussure. Etymology — The specific name is after A. von 48–52, female; 53, male. 48–49, head (48, Schulthess. lateral view; 49, dorsal view); 50, scutellum and metanotum, lateral view; 51–52, first two ornata de Saussure metasomal segments (51, lateral view; 52, dorsal view); 53, terminal articles of antenna. Belonogaster ornatus de Saussure, 1900: 207 (key), 209, Scale lines: 1 mm, but 0.5 mm for Fig. 53. female, male – “Madagascar.” Under “eumenoides Sauss./var./(= ornatus Sauss.) Buysson’s handwriting]”. The two specimens are /Madagascar” in the MHN, there are seven males and doubtless conspecific and I here designate the first three females clearly labeled by Hensen to indicate the specimen (with the label “. . . F Sikora”) as the lectotype lectotype and paralectotypes. The lectotype male is of B. pictus de Saussure. It seems certain that de labeled “MADAGASC.”, “ornatus/Sauss. ♂ [probably Saussure considered that the two specimens constituted in du Buysson’s handwriting]”, and “ne sont qu’une a good species (namely pictus) when he produced the demi=/gamme du coloris du/B. eumenoides Sss./Du key, and that later he regarded them as a variety of B. Buysson [in du Buysson’s handwriting]”. hildebrandti. In fact, his description of B. hildebrandti in 1900 includes only the description for “var.”, with pictus de Saussure which the two females well agree. The species is a Belonogaster pictus de Saussure, 1900: 207 (key), female – good species and I propose Belonogaster confusa as its “Madagascar” replacement name. This species, a junior homonym of B. pictus Kohl, The lectotype of B. pictus de Saussure is described 1894 (currently treated as a junior synonym of B. grisea below under a new replacement name, Belonogaster (Fabricius, 1775)), was described only in the key and its confusa; variations for the paralectotype are given in affiliation has long been in question. In the MHN there brackets. are two females under “hildebrandti/Sauss./var. Belonogaster confusa new replacement name /Madagascar”, that are labeled “B. pictus/♀ Sss. [in de (Figs. 54–60) Saussure’s handwriting]”. One of them is labeled also with “Madagascar/F Sikora” and “Hildebrandti/♀ var Belonogaster pictus de Saussure, 1900, non Kohl, 1894. Sauss. [in probably de Saussure’s handwriting on a Female — Body length (head + mesosoma + first large label]”, and the other has labels “Madagascar two metasomal segments) about 14 mm [13 mm]; fore- /Sikora” and “Hildebrandti/Sauss. var:/ ♀ [in du wing length about 14 mm [13 mm].

Taxonomic notes on Ropalidiini 17

yellow. First metasomal segment nearly black, irregularly tinged with green; apical semitransparent lamella brown. Second to terminal segments dark brown; second segment basally tinged with blackish- green; following markings ivory-yellow: large irregular lateral spots and paired small posteromedian spots on second tergum, irregular lateral bands and paired posteromedian spots on third and fourth terga, large irregular spot on fifth tergum, small lateral spots on sixth tergum, paired large irregular spots on second to fifth sterna, and most visible part of sixth sternum. Legs dark brown to black; trochanters, basal margins of femora and tibiae yellow. Wings infuscate (hyaline in ornata). Etymology — The specific name refers to a situation in which de Saussure may have been when he worked with this species.

pomicolor de Saussure Belonogaster pomicolor de Saussure, 1900: 207 (key), 209, female, male – “Madagascar.”

Under “eumenoides Sauss./var./(= pomicolor Sauss) /Madagascar” in the MHN, there are six females that are designated as the lectotype and paralectotypes by

Figs. 54–60. Belonogaster confusa new replacement Hensen and Blommers (1987). The lectotype is labeled name (lectotype female of Belonogaster pictus “Madagasc”, “Belonogaster/pomicolor/Sss. [in de de Saussure). 54–55, head (54, frontal view; 55, Saussure’s handwriting]”, “ne sont qu’une demi = lateral view); 56, scutellum and metanotum, /gamme du coloris du/B. eumenoides Sss./Du Buysson lateral view; 57, 59, first metasomal segment [in du Buysson’s handwriting]”, “LECTOTYPE (57, lateral view; 59, dorsal view); 58, second /Belonogaster ♀/pomicolor Sss./R. Hensen 1986” and and third metasomal segments, lateral view; 60, Hensen’s determination label as B. ornata. The second metasomal tergum, dorsal view. Scale remaining five females (one without metasoma) labeled lines: 1 mm. “Madagasc” or Madagascar/F Sikora” are clearly labeled as paralectotypes by Hensen. These females are certainly conspecific. Morphology similar to that of B. ornata, but Hensen and Blommers (1987: 25) stated that “It is distinctly different from it in the scutellum more possible that B. pomicolor is a distinct species, but as strongly convex (Fig. 56 vs. Fig. 50 for B. ornata) and long as the male is not known we prefer to leave it in in the first metasomal tergum in both dorsal and lateral the synonym of B. ornata”, but they synonymized B. views with posterior part little more distinctly swollen pomicolor under B. ornata for the first time; Richards (Figs. 57 and 59; hardly swollen in ornata, Figs. 51 and (1982: 110) listed all of B. pomicolor, B. ornata and B. 52). malagassa as probable synonyms of B. eumenoides, but Head blackish green, with following part yellow: he failed to locate the type materials of these species. I mandible except brown teeth, clypeus except large, ill- do not agree with the treatment by Hensen and defined dorsal greenish spot, ill-defined wide band Blommers. Actually there are following differences in along inner eye margin, and gena. Antenna the females between B. pomicolor and B. ornata: (1) ferruginous; scape darker above. Pronotum blackish gena in B. ornata (in profile, 0.4-0.45 times as wide as green, with narrow band along posterodorsal margin eye; Fig. 48) proportionately narrower than in B. and ventral corner colored yellow; scutum dark brown, pomicolor (0.5-0.6 times as wide as eye; Fig. 61), and irregularly tinged with black; scutellum and metanotum in dorsal view head behind eyes (= gena) slightly more yellow, except for black anterior margin and median strongly narrowed posteriorly in B. ornata (Fig. 49) line of scutellum; mesepisternum and metapleura brown, than in B. pomicolor (Fig. 62); (2) metanotum more with ill-defined yellow markings; propodeum black, strongly convex in B. pomicolor (Fig. 63) than in B. slightly green-tinged, with paired yellow spots above ornata (Fig. 50); (3) first metasomal tergum in dorsal propodeal orifice, and anterolateral parts of propodeum view more or less distinctly narrowed posteriorly near

18 J. Kojima

Figs. 61–72. Belonogaster pomicolor de Saussure. 61–65, female; 66–72, male. 61–62, 66, head (61, lateral view; 62, dorsal view; 66, frontal view); 63, scutellum and metanotum, lateral view; 64–65, 71–72, first two metasomal segments (64, 71, lateral view; 65, 72, dorsal view); 67, left antenna; 68–69, apical articles of left antenna in different views; 70, tarsal segments of hind leg. Scale lines: 1 mm, but 0. 5 mm for Figs. 68–70. apical margin in B. ornata (Fig. 52), while hardly Male — Body length (head + mesosoma + first two constricted in B. pomicolor (Fig. 72), and the segment metasomal segments) about 14.5 mm; forewing length in lateral view slightly thicker in B. pomicolor (Fig. 71) about 12 mm. than in B. ornata (Fig. 51). Structure and coloration similar to the female, but head different in the following characters (female A male in the ETH, labeled “Rogez/O. Madagasc”, characteristics are given in brackets). “♂” and “eumenoides/v. pomicolor/sec. coll Guep [last Head in frontal view about 1.25 [1.05-1.1] times as word illegible]” certainly belongs to B. pomicolor, and wide as high (Fig. 66); inner eye margins strongly has the antennae distinctly different from those of B. convergent below, distance between them at vertex ornata, as described below. about 1.25 [1.15] times as long as that at clypeus;

Taxonomic notes on Ropalidiini 19 clypeus smaller than in female, flat [convex], weakly article about 3.5 times as long as wide at apex; each produced below into bluntly angled apex [strongly of ninth and tenth articles slightly wider than long; produced below], covered with dense white tomentum terminal article about 1.5 times as long as wide at [tomentum sparser]; gena narrow, in profile about 0.4 base. Male terminal antennal article sharply pointed [0.5-0.6] times as wide as eye. Antenna slender (Fig. apically (Figs. 41–44)...... schulthessi new species 67); scape short and thick, swollen medially, about 2.5 24. First metasomal segment in dorsal view nearly times as long as wide; third article about 4 times as long parallel-sided posteriorly (Fig. 65), in profile only as wide, about 1.5 times as long as fourth article; fourth weakly thickening posteriorly (Fig. 64). Wings and fifth articles in nearly same length, each about 2.7 infuscate. Male terminal antennal article long, times as long as wide; last three articles distinctly curved, tapering apically, with pointed apex (Figs. thinner than preceding article; each of tenth and 68, 69) ...... pomicolor de Saussure eleventh articles more than 2.5 times as long as wide; — First metasomal segment in dorsal view weakly terminal article elongate, about five times as long as constricted near posterior margin (Figs. 52, 59); in wide at base, slightly flattened, weakly curved and profile thickening posteriorly towards well behind pointed apically (Figs. 68, 69; in B. ornatus weakly spiracle, then narrowing towards posterior margin widened apically, with rounded apex as in Fig. 53). (Figs. 51, 57). Wings hyaline or semihyaline...... 25 Body yellow or ivory yellow; median part of clypeus and front among dorsal emargination of clypeus 25. First metasomal tergum in profile weakly swollen pale ferruginous; front above level of eye emargination behind spiracle (Fig. 57); second tergum relatively and vertex brown, area behind ocelli tinged with dark slender, maximum width about four times as wide leaf-green; dorsal face of pronotum, scutum (except ill- as basal width; basal stalk proportionately longer defined median and lateral bands colored brownish (Fig. 60)...... confusa new replacement name yellow), scutellum, metanotum and posterior face of — First metasomal tergum in profile smoothly and propodeum leaf-green. First metasomal segment dark weakly thickening, not swollen (Fig. 51); second leaf-green, with posterior ill-defined narrow band tergum proportionately wider, maximum width colored ferruginous; second segment largely leaf-green, about five times as wide as basal width; basal stalk posteriorly irregularly tinged with yellow; remaining shorter (Fig. 52)...... ornata de Saussure metasomal segments brown or reddish brown. Legs yellow; hind coxa and femur tinged with leaf-green; all longistylus de Saussure tarsi ferruginous. Belonogaster longistylus de Saussure, 1890: 89 (key), 97, pl. The last couplet (couplet 22) of Hensen and 17 fig. 13, female – “Madagascar”. Blommer’s (1987) key is modified and another three The lectotype designated by Richards (1982: 103) is couplets are added to distinguish the above five species. in the MHN and labeled “Belonogaster/longistylus 22. Eyes with sparse, but conspicuous hairs; first /Sauss. ♀ [probably in du Buysson’s handwriting]” and metasomal segment in dorsal view more or less with Richards’ designation label. Another female under parallel-sided posteriorly (Figs. 31, 47). Metasoma longistylus is conspecific with the lectotype and labeled without leaf-green and/or yellow markings...... 23 “Madagascar/Elliot” and “Belonogaster/longistylus/ ♀ — Eyes without hairs, or if present, very short and Sss.” [in de Saussure’s handwriting] that is not a indistinct; first metasomal segment in dorsal view syntype as de Saussure definitely mentioned Hildbrandt nearly parallel-sided or narrowed posteriorly. and Sikora as collectors. Metasoma with leaf-green and/or yellow markings... Both Richards (1982) and Hensen and Blommers …...... 24 (1987) treated B. longistylus as a synonym of B. madecassa (de Saussure, 1853). 23. Body ferruginous to dark ferruginous. Female antenna relatively short (Fig. 35); third article about 2.5 times as long as wide at apex; ninth and tenth prasina de Saussure article distinctly wider than long; terminal article Belonogaster prasinus de Saussure, 1890: 89 (key), 92, pl. 19 distinctly less than 1.5 times as long as wide. Male fig. 5, female, male – “Madagascar . . . aux environs de terminal antennal article not narrowed or slightly Fort-Dauphin”. dilated apically, with rounded apex (Figs. 33, 34)... The lectotype designated by Richards (1982: 107) …...... eumenoides de Saussure is in the MHN and labeled “Madagascar/Elliot”, — Body blackish green; head and mesosoma partly “prasinus/Sss. ♂ [probably in du Buysson’s hand- ferruginous and/or pale yellow; second and writing, green label]”, and “Belonogaster/prasinus/de subsequent metasomal segments largely dark brown. Saussure/ Lectotype [with red underline] ♂/Richards Female antenna relatively long (Fig. 38); third 1979 [in Richards’ handwriting]”.

20 J. Kojima

In the MHN, under “prasina” are four males with Under “dubia Kohl” in the MHN are two females. the same collection label (one with the same One of them is labeled “Spec. ? [in handwriting]”, “B. determination label as the lectotype; another with a saevus/Sss./♀ Cap. [in de Saussure’s handwriting]” and label “Belonogaster/prasinus/♂ Sss [in de Saussure’s “? Belonogaster/dubia Kohl/ ♀ var. [in Richards’ handwriting]”), a female labeled “MADAGASC”, handwriting]” and matches in general the original “612” and “Belonog./prasinus/var. Sss. ♀ [in de description of B. saeva. However, the specimen is Saussure’s handwriting]”, and a female labeled much smaller (about 27.5 mm in the total body length) “Madagascar/F Sikora” and the same determination than the size range de Saussure’s reported (32-34 mm). label as the lectotype. All these specimens, except for Further, “Cap.” is not tropical Africa, and thus the the female labeled as “var.” and the last specimen, are specimen is probably not a syntype. Richards (1982) highly probably from the original series and are did not refer to this specimen; de Saussure’s paralectotypes. As Carpenter (1999: 31) pointed out, de determination label is folded and might have been Saussure did not mention “Sikora” as a collector. overlooked by Richards. The specimen keys to B. grisea (Fabricius, 1804) in Richards (1982). brunnea nigriclava Richards Another female labeled “Lourenco-Marquès/Dr G. Belonogaster brunnea nigriclava Richards, 1982: 38, 43 Andeoud.” and “Belonogaster/dubius Kohl/♀ [in du (key), 66, figs. 26-27, male, female – holotype male Buysson’s handwriting]” is certainly B. dubia, but (BMNH) from “Uganda: Mt Kokanjero, SW. Mt Elgon, agrees well with de Saussure’s description of “var. ♀” 6400 ft”. of B. saeva and its total body length is about 32.5 mm. A female labeled “Namanga, Kenya/13.2.58 – P. I have at this moment no exact opinion about the Strinati/et V. Aellen”, “Belonogaster/sp. (prés freyi affiliations of “B. saeva” of du Buysson (1909b: 242) dB.)/Ch. Ferrière det. mais pattes noires”, and “B. and Richards (1982). Richards’ description does not distinguenda / ♀ Kohl / spp. / nigriclava Rich. [in match in several points with de Saussure’s original Richards’ handwriting]” in the MHN is undoubtedly a description of B. saeva; for example, Richards (1982: paratype, although he (p. 66) indicated “MHN, Genoa” 61) stated that the wings are “usually blackish brown, as the depository for this specimen, undoubtedly by an sometimes paler” and “dark or black” in the key (p. 41), error in typesetting. while de Saussure’s original description says “alis Bequaert (1918: 329) synonymized B. distinguendus ferruginescentibus, apice griseis” – this is the state Kohl under B. brunnea. Richards (1982: 64) designated mentioned for B. grisea in Richards (1982: 88). the lectotype of B. distinguendus and marked as “Syn. n.”, overlooking Bequaert (see also Kojima and Carpenter, 1997: 72). Genus Polybioides du Buysson, 1913 raphigastra (de Saussure) pusilloides Richards Polybia raphigastra de Saussure, 1854, in de Saussure, 1853- Belonogaster pusilloides Richards, 1982: 37, 43 (key), 80, fig. 58: 204, female – “L’Amérique? Etiqueté par erreur de 54, female, male – holotype female (BMNH) from Pulo-Pinang (Asie)”. “Uganda: top of escarpment, east of Batiaba, L. Albert, Under “raphigastra/Sauss/Sumatra” in the box 3200 ft”. “Vespidae 14” in the MHN, there is a female that is A female labeled “630/78 Congo/ach. Deyrolle labeled with a small silver square, “Pulo/Pinang” and leg.”, “Congo/630/78”, and “Belonogaster/pusilloides/ “Parapolybia/raphigastra/Saus [possibly in von ♀ Rich” [in Richards’ handwriting] in the MHN is Schulthess’ handwriting]”. The specimen is in bad obviously a paratype although Richards indicated condition, lacking wings except for the left hind wing. “MNHN, Paris” as depository for this specimen. De Saussure indicated the Spinola collection as the depository, and Casolari and Casolari Moreno (1979: freyi du Buysson 98; for abbreviations, see Casolari and Casolari Moreno, Belonogaster freyi du Buysson, 1909: 217 (key), 255, female 1978) listed one specimen under “Polybia rhafigastra – “Afrique australe”. [sic]” in the Spinola collection. Van der Vecht (1966: A female in the MHN labeled “B. freyi D.B./type 13) referred to the specimen in the MIZT as “holotype [underlined with red]/afr. austr. [in du Buysson’s ♀ leg. D. Westermann, Coll. Spinola, MT”. The MHN handwriting]” is undoubtedly the holotype. I added a specimen is presumably from the original series red label indicating it as the holotype. retained by de Saussure. Then van der Vecht’s statement is regarded as the lectotype designation and saeva de Saussure the specimen in the MHN is now a paralectotype. Belonogaster saevus de Saussure, 1890: 92, footnote, female – “l’Afrique tropicale”.

Taxonomic notes on Ropalidiini 21

Acknowledgments Dalla Torre, K. W. 1904. Vespidae. Genera Insectorum 19: 1–108. I thank J. M. Carpenter for critical reading of the Fabricius, J. C. F. 1804. Systema Piezatorum. Brunsvigae. manuscript, and I. Löbl, B. Merz, G. Cuccodoro and staff Giordani Soika, A. 1944. Contributo alla conoscenza dei members of the MHN for their help during my stay in the vespidi solitari e sociali della regione etiopica. Atti Ist. MHN. My thanks are also due to C. van Achterberg, A. Ven. Sci., Lett. Arti Cl. Sci. Mat. Nat. 103: 149–178. Müller, S. Bieri, C. Villemont and M. Lachais for their help in Giordani Soika, A. 1977. Results from the Danish Expedition research in the NNM, ETH and MNHN, and to R. Meier for to the Cameroons 1949-1950. 33. Hymenoptera, Vespidae the photographs and information on the holotype of Polistes and Eumenidae (Insecta). Steenstrupia 4: 125–129. bioculata. The present study was mainly supported by the Giordani Soika, A. 1981. Notulae vespidologicae XLII. Nuovi Swiss Academy of Science and Japan Society for the vespidi della regione Afrotropicale. Boll. Soc. Entomol. Promotion of Science and partly by a Grant-in-Aid (no. Ital. 113: 177–175. 11833001) from the Japan Society for the Promotion of Gribodo, G. 1896. Hymenopterorum novorum diagnoses Science. proecursoriae. Misc. Entomol. 4: 12–14. Hensen, R. V. and L. H. M. Blommers. 1987. Revision of the References Malagasy species of Belonogaster Saussure (Hymenoptera; Vespidae). Tijdschr. Entomol. 130: 11– Bedot, M. 1906. Henri de Saussure Notice biographique. Rev. 31. Suisse Zool. 14: 1–32. Horn, W., I. Kahle, G. Friese and R. Gaedike. 1990. Bequaert, J. 1918. A revision of the Vespidae of the Belgian Collectiones Entomologicae. Akademie der Landwirt- Congo based on the collection of the American Museum schafswissenschaften Deutschen Demokratischen Repu- Congo expedition, with a list of Ethiopian diplopterous blik, Berlin. wasps. Bull. Am. Mus. Nat. Hist. 39: 1–384. International Commission on Zoological Nomenclature, 2000 Bequaert, J. 1932. Aculeata (Vespidae) genre Ropalidia. Pp. (published in 1999). International Code of Zoological 47–51. In Résultats scientifiques du voyage aux Indes Nomenclature. Fourth Edition. orientales Néerlandaises de LL. RR. le Prince et la Kojima, J. 1999a. Taxonomic notes on Australian Ropalidia Princesse Léopold de Belgique. Vol. 4 (5). Mém. Mus. R. Guérin-Méneville, 1831 (Hymenoptera: Vespidae, Hist. Nat. Belg. Hors Sér. Polistinae). Entomol. Sci. 2: 367–377. Bingham, 1903. On the Hymenoptera collected by Mr. W. L. Kojima, J. 1999b. Male genitalia and antennae in an Old Distant in the Transvaal, South Africa, with descriptions World paper wasp genus Ropalidia Guérin-Méneville, of supposed new species. Ann. Mag. Nat. Hist. (7) 12: 1831 (Insecta: Hymenoptera; Vespidae, Polistinae). Nat. 46–69. Hist. Bull. Ibaraki Univ. 3: 51–68. Buysson, R. du. 1906. Hyménoptères nouveaux. Rev. Kojima, J. and J. M. Carpenter. 1997. Catalog of species in Entomol. 25: 103–112. the polistine tribe Ropalidiini (Hymenoptera: Vespidae). Buysson, R. du. 1909a. Deux Hyménptères nouveaux Am. Mus. Novitat. 3199: 1–96. d’Océanie. Bull. Soc. Entomol. France 1909: 305–306. Kojima, J., J. W. Wenzel and J. M. Carpenter. 1998. The Buysson, R. du. 1909b. Monographie des Vespides du genre types of Icaria galimatia de Saussure, 1853, and I. Belonogaster. Ann. Soc. Entomol. France 1909: 199–270, ignobilis de Saussure, 1890, with establishment of their pl. 2–8. synonymy (Hymenoptera: Vespidae, Polistinae). Entomol. Cameron, P. 1906. Hymenoptera of the Dutch Expedition to Sci. 1: 393–401. New Guinea in 1904 and 1905. Part. 1: Thynnidae, Meade-Waldo, G. 1913. New species of Diploptera in the Scolidae, Pompilidae, Sphegidae and Vespidae. Tijdschr. collection of the British Museum. Part IV. Ann. Mag. Nat. Entomol. 49: 221–233. Hist. (8) 11: 44–54. Carl, J. 1930. Dans les massifs montagneux de l’Inde Richards, O. W. 1978. The Australian social wasps méridionale. Mémoires du Gulobe 69: 1–153, 32 pls. (Hymenoptera: Vespidae). Aust. J. Zool. Suppl. Ser. 61: Carl, J. 1934. Ropalidia montana n. sp. et son nid. Un type 1–132. nouveau d’architecture vespienne. Rev. Suisse Zool. 41: Richards, O. W. 1982. A revision of the genus Belonogaster 675–691. de Saussure (Hymenoptera: Vespidae). Bull. Br. Mus. Carpenter, J. M. 1999. Taxonomic notes on paper wasps (Nat. Hist.) Entomol. 44 (2): 31–114. (Hymenoptera: Vespidae; Polistinae). Am. Mus. Novitat. Saussure, H. de. 1853-1858. Études sur la famille des 3259: 1–44. Véspides. Vol. 2. Monographie des Guêpes sociales, ou Casolari, C. and R. Casolari Moreno. 1978. Catalogo della a de la tribe de Vespiens. Masson, Paris, and Kessmann, collezione imenotterologica de Massimiliano Spinola. 1 Genève. Parte. Boll. Mus. Zool. Univ. Torino (5): 27–74. Saussure, H. de. 1862. Sur divers Vespides Asiatiques et Casolari, C. and R. Carolari Moreno. 1978 Catalogo della a Africains du Musée de Leyden. Stettin. Entomol. Ztg. 23: collezione imenotterologica de Massimiliano Spinola. 3 129–141. Parte. Boll. Mus. Zool. Univ. Torino (4): 91–130. Saussure, H. de. 1863. Mélanges Hyménoptérologiques II Cheesman, L. E. 1952. Ropalidia of Papuasia. Ann. Mag. Nat. Vespides. Mem. Soc. Phys. Hist. Nat. Genève 17: 171– Hist. (12) 5: 1–26, 4 pls. 245. Dalla Torre, K. W. 1894. Vespidae (Diploptera). Catalogus Hymenopterorum 9: 1–181.

22 J. Kojima

Saussure, H. de. 1890. Volume 20. Histoire Naturelle des Hyménoptères. In A. Grandidier, Histoire Physique, Vecht, J. van der. 1941. The Indo-Australian species of the Naturelle et Politique de Madagascar. genus Ropalidia (= Icaria) (Hym., Vespidae) (first part). Saussure, H. de. 1900. Wissenschaftliche Ergebnisse der Treubia 18: 103–190 Reisen in Madagaskar und Ost-Afrika in der Jahren 1889- Vecht, J. van der. 1962. The Indo-Australian species of the 1895 von Dr. A. Voelzkow. Hymenoptera. Vespidae. genus Ropalidia (Icaria) (Hymenoptera, Vespidae) Abh. Senckenb. Natureforsch. Ges. 26 (2): 203–240. (second part). Zool. Verh., Leiden 57: 1–72, 8 pls. Schulthess, A. von. 1915. Hymenopteren von Neu-Caledonien Vecht, J. van der. 1966. The east-Asiatic and Indo-Australian und den Loyalty-Inseln. Pp. 45–53. In F. Sarasin and J. species of Polybioides Buysson and Parapolybia Saussure Roux (eds.), Nova Caledonia, Zoologie 2, 1(3). (Hym., Vespidae). Zool. Verh., Leiden 82: 1–42, 3 pls. Smith, F. 1857. Catalogue of Hymenopterous Insects in the Collection of the British Museum. Part 5. Vespidae. Smith, F. 1860 (1861). Catalogue of Hymenopterous insects collected by Mr. A. R. Wallace in the Islands of Bachian, Received 3 August 2000; Kaisar, Amboyna, Gilolo, and at Dory in New Guinea. J. revised 5 October 2000; Proc. Linn. Soc. Zool. 5: 93–143. accepted 7 October 2000. Tillyard, R. J. 1926. The Insects of Australia and New Zealand. Angus and Robertson, Ltd., Sydney.