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Taxonomic notes on the genus Rhynchium Spinola (Hymenoptera: Vespidae: Eumeninae) from Vietnam, with a first checklist of the species worldwide

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The genus Rhynchium Spinola, 1806, is studied in Vietnam. Two species are recorded, namely, R. brunneum brunneum (Fabricius, 1793) and R. haemorrhoidale haemorrhoidale (Fabricius, 1775). A key to Vietnamese species of the genus is presented along with new distributional records. Twenty-one species and 36 subspecies are listed in a first checklist of species of the genus. New synonymy proposed in the present study is R. brunneum ceylonicum Giordani Soika, 1994, under the nominotypical subspecies R. b. brunneum (Fabricius, 1793).
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Ecol. Mont., 7, 2016, 530-537
ISSN 2336-9744 (online) | ISSN 2337-0173 (print)
The journal is available on line at www.biotaxa.org/em
Taxonomic notes on the genus Rhynchium Spinola (Hymenoptera:
Vespidae: Eumeninae) from Vietnam, with a first checklist of the
species worldwide
PHONG HUY PHAM1, P. GIRISH KUMAR2
1 Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet
Road, Hanoi-100000, Vietnam; E-mail: phong.wasp@gmail.com
2 Zoological Survey of India, Western Ghat Regional Centre, Kozhikode-673006, Kerala, India;
E-mail: kpgiris@gmail.com
Corresponding author: Phong Huy Pham
Received 6 September 2016 Accepted 12 November 2016 Published online 5 December 2016.
Abstract
The genus Rhynchium Spinola, 1806, is studied in Vietnam. Two species are recorded, namely, R. brunneum brunneum
(Fabricius, 1793) and R. haemorrhoidale haemorrhoidale (Fabricius, 1775). A key to Vietnamese species of the genus
is presented along with new distributional records. Twenty-one species and 36 subspecies are listed in a first checklist
of species of the genus. New synonymy proposed in the present study is R. brunneum ceylonicum Giordani Soika, 1994,
under the nominotypical subspecies R. b. brunneum (Fabricius, 1793).
Key words: distribution, new record, Oriental region, synonymy, Vietnamese species.
Introduction
The genus Rhynchium belonging to subfamily Eumeninae, named by Spinola (1806) with the type species
Rygchium europaeum Spinola, 1806, is an Old World genus, comprising relatively large-size species mainly
distributed in the Oriental Region. The main characters to distinguish wasps of this genus from wasps of
other genera are the scutellum and posterior part of mesoscutum smooth and at most finely and sparsely
punctuate. After referring to data from a website (http://www.eol/pages/31233), Kumar & Sharma (2013)
reckoned there were 47 species with several subspecies in the genus Rhynchium. Siddiqui et al. (2015) stated
that “the genus Rhynchium includes slightly more than 20 species and many subspecies, distributed
throughout the Old World”. Hence, the number of the Rhynchium species has not yet been obvious to date.
There have been two species in the genus Rhynchium recorded from Vietnam so far. The first Rhynchium
species recorded was R. brunneum (Fabricius, 1793) by Giordani Soika (1986). After eight years (1994), he
mentioned this as a nominotypical subspecies R. brunneum brunneum. The second species recorded for
Vietnam was R. haemorrhoidale haemorrhoidale (Fabricius, 1775) by Nguyen et al. (2015). In present
paper, taxonomic notes on the genus are mentioned along with a first checklist of species of the genus.
Research Article
PHAM & KUMAR
Ecol. Mont., 7, 2016, 530-537 531
Materials and Methods
The materials examined in the present paper are housed in the collections of the Institute of Ecology and
Biological Resources (IEBR) and of Zoological Survey of India, India (NZC).
The morphological and colour characters were observed and described on pinned and dried
specimens under a stereoscopic microscope. Photographic images were taken by using a Leica
Stereomicroscope with LAS software version 3.6.0. Provinces and cities are alphabetically listed in the text
as distribution of species in the genus.
Results and Discussion
Genus Rhynchium Spinola
Rygchium Spinola, 1806. Ins. Ligur., 1: 84, genus (incorrect original spelling of Rhynchium Spinola).
Rhynchium Spinola, 1806, Ins. Ligus., 1:84, genus, emendation of Rygchium validated by Opinion 747 (ICZN, 1965).
Type species: Rygchium europeaum Spinola, 1806 [= Vespa oculata Fabricius, 1781], by monotypy.
Rhynchium Billberg, 1820, Enum. Ins.: 109. Emendation of Rychium (!) Spinola.
Rhynchium Sturm, 1829, Verz. Ins. Nurnberg: 12. Emendation.
Rhygchium de Saussure, 1853, Ét. Fam. Vesp. 1: xxxi, 276. Emendation.
Rhynchuium (!) de Saussure, 1863, Mém. Soc. Phy. Hist. Nat. Genéve 17: 242.
Eurrhynchium Dalla Torre, 1904, Gen. Ins. 19: 33. New name.
Rygohium (!) Willink, 1982, Bol. Ac. Nac. Sci. 55: 195.
Diagnosis: Palpal formula 6:4; scutellum and posterior part of mesoscutum smooth, at most finely and
sparsely punctuate; metanotum depressed medially and bluntly projecting laterally, as seen from above and
in front distinctly concave; propodeum with valvula small and never rectangular, submarginal carina not
projecting; tegula not evenly rounded posteriorly, emarginate adjoining parategula; tegula shorter than
parategula posteriorly; midtibia with one spur; mid femora of male emarginated at base; forewing with
second cubital cell not petiolate; third cubital cell further away from the apex of the radial cell; parastigma of
fore wing more than half as long as the stigma; metasomal segment I sessile, with tergite without transversal
ridge or carina, segment I with width more than half that of II, much less than twice as long as wide.
Key to species of the genus Rhynchium in Vietnam
1 Metasomal segment I and II usually entirely black, rarely with a thin reddish brown band apically of
first or second or both segments (Fig. 6). .......... R. haemorrhoidale haemorrhoidale (Fabricius, 1775)
- Metasomal segment I and II black at basal half (Figs. 1-2); Mandible long with apical tooth sharp,
acute (Figs. 3-4); Central frons with pubescences dense (Fig. 5); Metasomal segment I and
metasomal segment II with a broad yellowish brown band apically (Figs. 1-2). ....................................
.............................................................................................. R. brunneum brunneum (Fabricius, 1793)
1. R. brunneum brunneum (Fabricius, 1793)
(Figs. 1-5)
Vespa brunnea Fabricius, 1793, Entomol. Syst., 2: 264, Syntype, sex not mentioned, “Tranquebariae, India
(Universitetets Zoologiske Museum, Copenhagen, Denmark)”.
Odynerus brunneus Latreille, 1806, Gen. Crust. Ins., 1: pl. 14 fig. 3.
Rhynchium brunneum Spinola, 1808, Ins. Ligur, 2: 189.
Rhynchium haemorrhoidale var. brunneum de Saussure, 1862, Stettin. Ent. Ztg., 23: 190.
Rhynchium quinquecinctum var. brunneum Dalla Torre, 1894, Cat. Hym., 9: 45 (cat.).
Rhynchium brunneum brunneum: Gusenleitner, 2006, Linzer biol. Beitr., 38(1): 692 (Sikkim, Meghalaya, West
Bengal).
TAXONOMIC NOTES ON THE GENUS RHYNCHIUM FROM VIETNAM
532
Material examined: VIETNAM: Bac Ninh: 1 , Tu Son, v.2010, P.H. Pham. Hai Phong: 1 , Vinh Bao,
25.vi.2013, P.H. Pham. Hoa Binh: 2 + 1 , Mai Chau, Mai Chau, 3.vi.2012, P.H. Pham. Hung Yen: 1
+ 1 , Trieu Duong, Tien Lu, 25.vi.2013, P.H. Pham. Nam Dinh: 1 + 1 , Xuan Thuy National Park,
Xuan Thuy, 22.vi.2013, P.H. Pham. Quang Ninh: 1 + 3 , Hong Thai Tay, Dong Trieu, 26.vi.2013, P.H.
Pham. Thai Binh: 2 + 2 , Hong Minh, Hung Ha, 23-24.vi.2013; 2 , Hung Nhan, Hung Ha, 25.vi.2013,
P.H. Pham; 1 , Dong Minh, Tien Hai, 20°2023’’N, 106°3589’’E, alt. 0 m, 21.vi.2013, H.P. Pham. Thanh
Hoa: 2 , Sam Son Town, 20-21.vi.2016, P.H. Pham. SRI LANKA: Paradeniya: 5 , 2.vi.1910, name of
collector unknown, NZC Regd.Nos.13752/H3 to 13756/H3.
Distribution: Vietnam: Bac Giang, Bac Ninh (new record), Cao Bang, Dien Bien, Gia Lai, Ha Giang, Ha
Noi, Ha Tinh, Hai Duong, Hai Phong (new record), Hoa Binh, Hung Yen (new record), Kon Tum, Lai
Chau, Lam Dong, Lao Cai, Nam Dinh, Nghe An, Quang Nam, Quang Ninh, Quang Tri, Thai Binh, Thai
Nguyen, Thanh Hoa, Thua Thien Hue, Vinh Phuc. Elsewhere: India, Seychelles, Bangladesh, Cambodia,
China, Guam, Indonesia, Laos, Malaysia, Marianas, Myanmar, New Britain, Pakistan, Palau, China, Taiwan,
Thailand.
Remarks: There are three subspecies recorded from the Oriental Region namely, R. brunneum brunneum
(Fabricius, 1793), R. brunneum ceylonicum Giordani Soika, 1994 and R. brunneum maladivum Gusenleitner,
2003. Out of these three subspecies, the nominotypical subspecies namely R. b. brunneum is widely
distributed in the Oriental Region. R. b. ceylonicum is recorded from Sri Lanka and R. b. maladivum
recorded from the Maldives (Kumar & Sharma, 2013). We have examined specimens collected from Sri
Lanka. These specimens are of the subspecies R. b. ceylonicum. This subspecies is distinguished from R. b.
brunneum by colour pattern. In R. b. brunneum the body is marked with black and reddish brown
colouration, whereas R. b. ceylonicum the body is marked with black and yellowish brown colouration.
Among specimens collected in Vietnam, there are 5 specimens that they have a slight difference in colour
from the other specimens [1 , Mai Chau, Hoa Binh Province; 2 , Hung Nhan, Hung Ha, Thai Binh
Province; 1 + 1 , Hong Thai Tay, Dong Trieu, Quang Ninh Province]. Their colour pattern is very close
to that of the subspecies R. b. ceylonicum, and these specimens can represent the formally named color form
R. b. ceylonicum. Thus we propose to synonymize R. b. ceylonicum under the nominotypical subspecies R. b.
brunneum.
The black marks on the body of R. b. brunneum are highly variable. On mesoscutum: A small
triangular mark in front can be extended to become a large mark at the centre, and a narrow transverse line
along the apex can be also extended to become a large line at the central part. Therefore two marks can join
at the centre. On Metasoma: from basal third to basal two thirds of metasomal tergite I and II, metasomal
sternite II from almost to entirely black, metasomal sternite III-VI in females and metasomal sternite III-VII
in males from basal third to basal two thirds. Marks on metasoma are also from light to dark black
colouration.
A female collected at Dong Minh commune, Tien Hai district, Thai Binh province has the mandible
different from that of all of the other specimens of R. brunneum and also of the other Rhynchium species
(Figs. 3-4). This means that this specimen can be of a new species. But at present time only a specimen of
the female is available it is arranged under the nominotypical species R. brunneum and the future studies
should include it.
2. Rhynchium haemorrhoidale haemorrhoidale (Fabricius, 1775)
(Fig. 6)
Vespa haemorrhoidalis Fabricius, 1775, Syst. Entomol.: 366, Type ♂, “ad Cap. b. Spei” (British Museum of Natural
History).
Odynerus dimidiatus Guèrin, 1834, in Bèlanger, Voyage Indes-Orient. Zool.: 503, pl. 4 fig. 4, “Côte du Coromandel”.
Rygchium haemorrhoidale de Saussure, 1852, Ét. Fam. Vesp., 1: 109 (, ).
Odynerus haemorrhoidalis Bequaert, 1918, Bull. Am. Mus. Nat. Hist., 39: 300 (occurrence in Cape Province very
doubtful).
Rhynchium haemorrhoidale Bingham, 1897, Fauna Brit. India, Hym., 1: 353 (key), 354, , (India; Burma; Ceylon;
Tenasserim; Java; Cape of Good Hope).
PHAM & KUMAR
Ecol. Mont., 7, 2016, 530-537 533
Figures 1-6. 1-5 - Rhynchium brunneum brunneum (Fabricius, 1793): 1-2 - Female - Habitus; 3-4 - Mandible; 5 - Male
- Head front view; 6 - Rhynchium haemorrhoidale haemorrhoidale (Fabricius, 1775): Female - Habitus.
TAXONOMIC NOTES ON THE GENUS RHYNCHIUM FROM VIETNAM
534
Distribution: Vietnam: Kien Giang. Elsewhere: Cambodia, India, Indonesia, Malaysia, Myanmar, New
Guinea, Singapore, Sri Lanka, China, Thailand.
Remarks: Giordani Soika (1994) recorded this species occurring in Vietnam, but did not give its exact
locality. This species is presently recorded from only one province in Vietnam (Kien Giang).
A checklist of species of the genus Rhynchium Spinola, 1806
1. R. acromum Giordani Soika, 1952
2. R. annuliferum (Boisduval, 1835)
3. R. atrissimum van der Vecht, 1968
4. R. atrum de Saussure, 1852
4.1. R. atrum atrum de Saussure, 1852
4.2. R. atrum jacobsoni Giordani Soika, 1994
4.3. R. atrum palawanense van der Vecht, 1968
5. R. bathyxanthum van der Vecht, 1963
6. R. brunneum (Fabricius, 1793)
6.1. R. brunneum brunneum (Fabricius, 1793)
6.2. R. brunneum maladivum Gusenleitner, 2003
7. R. carnaticum (Fabricius, 1798)
8. R. claripenne Giordani Soika, 1994
9. R. cyanopterum de Saussure, 1852
9.1. R. cyanopterum cyanopterum de Saussure, 1852
9.2. R. cyanopterum assabense Gioadani Soika, 1987
9.3. R. cyanopterum somaliense Giordani Soika, 1987
10. R. haemorrhoidale (Fabricius, 1775)
10.1. R. haemorrhoidale haemorrhoidale (Fabricius, 1775)
10.2. R. haemorrhoidale andamanicum Kumar & Sharma, 2013
10.3. R. haemorrhoidale dohertyi Schulz, 1905
10.4. R. haemorrhoidale sanguneum de Saussure, 1852
10.5. R. haemorrhoidale tahitense (de Saussure, 1867)
10.6. R. haemorrhoidale umeroatrum Gusenleitner, 1991
11. R. magnificum Smith, 1869
12. R. marginellum (Fabricius, 1793)
12.1. R. marginellum marginellum (Fabricius, 1793)
12.2. R. marginellum congicum (Bequaert, 1918)
12.3. R. marginellum sabulosum de Saussure, 1855
12.4. R. marginellum somalicum Magretti, 1899
13. R. medium (Maindron, 1882)
14. R. mirabile de Saussure, 1855
14.1. R. mirabile mirabile de Saussure, 1852
14.2. R. mirabile biakense van der Vecht, 1960
14.3. R. mirabile expolitum van der Vecht, 1960
15. R. nigrosericeum Giordani Soika, 1989
16. R. oculatum (Fabricius, 1781)
16.1. R. oculatum oculatum (Fabricius, 1781)
16.2. R. oculatum adenense Giordani Soika, 1952
16.3. R. oculatum distinguendum du Buysson, 1913
16.4. R. oculatum hebraeum Giordani Soika, 1952
16.5. R. oculatum leviscutis (Cameron, 1908)
16.6. R. oculatum lefebvrei Lepeletier, 1841
17. R. quinquecinctum (Fabricius, 1787)
17.1. R. quinquecinctum quinquecinctum (Fabricius, 1787)
17.2. R. quinquecinctum murotai Tano, 1983
17.3. R. quinquecinctum nambui Yamane, 1983
17.4. R. quinquecinctum fukaii Cameron, 1911
PHAM & KUMAR
Ecol. Mont., 7, 2016, 530-537 535
17.5. R. quequincinctum samurayi Giordani Soika, 1973.
17.6. R. quinquecinctum tahitense de Saussure, 1867
18. R. rubropictum Smith, 1861
18.1. R. rubropictum rubropictum Smith, 1861
18.2. R. rubropictum nigriventre Giordani Soika, 1994
18.3. R. rubropictum tenimberense Giordani Soika, 1994
19. R. salomonis Meade-Waldo, 1910
20. R. superbum de Saussure, 1852
21. R. vittatum du Buysson, 1909
Acknowledgements
The authors are grateful to Dr. K. Venkataraman, former Director, Dr. Kailash Chandra, Director, and Dr. Gaurav
Sharma, former Officer-in-Charge, Hymenoptera Section, Zoological Survey of India, for providing facilities and
encouragements. First author also thanks The Centre for Science and Technology of the Non-Aligned and Other
Developing Countries (NAM S&T Centre), Government of India, for granting the scholarship for the research training
course at Zoological Survey of India. The authors also extend thanks to two reviewers, Dr. James M. Carpenter,
Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th street, New York,
NY 10024, U.S.A. and Dr. Tingjing Li, Institute of Entomology & Molecular Biology, College of Life Sciences,
Chongqing Normal University, Chongqing, 401331, P. R. China, for their valuable comments and suggestions on the
manuscript.
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Islands, Japan (Hymenoptera: Eumenidae). Memoirs of Kagoshima University Research Center for
South Pacific, 4(1), 119132.
... The phylogenetic relationships within the Rhynchium genera group have been discussed in a previous study based on morphological features and mitochondrial genomes [15], but uncertainties and conflicting classification schemes remain among genera and subgenera and on the taxonomic status of certain species. Firstly, the taxonomic status of Anterhynchium (Dirhynchium) is controversial. ...
... In one clade, a monophyletic Anterhynchium is the sister to a clade comprising Dirhynchium and Rhynchium. Anterhynchium, Dirhynchium, and Rhynchium form a monophyletic lineage that shares a common feature: propodeum and metanotum at different levels in lateral view [5,[8][9][10][11]14,15]. This supports the elevation of Dirhynchium to a valid genus, as it is proposed based on a mitogenomic analysis [16] and corroborated by morphological differences. ...
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Simple Summary The eumenine wasps of the genus Rhynchium Spinola, 1806; Allorhynchium van der Vecht, 1963; Anterhynchium de Saussure, 1863; and Pararrhynchium de Saussure, 1855 are related and sometimes hard to tell apart from each other. In this study, we first reconstructed the phylogenetic relationships of these genera based on universal single-copy orthologs and ultraconserved elements extracted from 10 newly sequenced low-coverage whole genomes. The results showed that Allorhynchium and Lissodynerus are distinct from the other four taxa. The genus Rhynchium was recovered as monophyletic, whereas Anterhynchium was recovered as paraphyletic, with Anterhynchium (Dirhynchium) as a sister to Rhynchium and hence deserving a separate genus Dirhynchium; and within the genus Pararrhynchium, P. septemfasciatus feanus and P. venkataramani were separated, not clustered on a branch. It is suggested that the genus Lissodynerus should be restituted as a valid genus, not a synonym of the Pararrhynchium. The results are consistent with previous eumenine mitochondrial genome phylogenetic analyses. This paper confirms the feasibility of low-coverage whole genome eumenine wasp phylogenetics and provides a reference for subsequent research in Eumeninae. Abstract The subfamily Eumeninae is a large group of fierce predatory insects that prey mainly on the larvae of Lepidoptera pests. Because of the highly similar morphologies of the genus Rhynchium and its related genera in the subfamily, including Rhynchium Spinola, Allorhynchium van der Vecht, Anterhynchium de Saussure, Pararrhynchium de Saussure, it is essential to delineate their relationships. A previous phylogenetic analysis based on mitochondrial genomes suggested the inconsistent relationships of these genera under traditional classification based on morphological characters. In this study, we first used single-copy orthologs [USCO] and ultraconserved elements [UCE] extracted from 10 newly sequenced low-coverage whole genomes to resolve the phylogenetic relationships of the above genera. The newly sequenced genomes are 152.99 Mb to 211.49 Mb in size with high completeness (BUSCO complete: 91.5–95.6%) and G + C content (36.31–38.76%). Based on extracted 5811 USCOs and 2312 UCEs, the phylogenetic relationships of Rhynchium and its related genera were: ((Allorhynchium + Lissodynerus) + (Pararrhynchium + (Anterhynchium + (Dirhynchium + Rhynchium)))), which was consistent with the mitochondrial genome results. The results supported the genus Rhynchium as monophyletic, whereas Anterhynchium was recovered as paraphyletic, with Anterhynchium (Dirhynchium) as a sister to Rhynchium and hence deserving generic status; In addition, in the genus Pararrhynchium, P. septemfasciatus feanus and P. venkataramani were separated, not clustered on a branch, just as P. septemfasciatus feanus was not together with P. striatum based on mitochondrial genomes. Since Lissodynerus septemfasciatus, the type species of the genus Lissodynerus, was transferred to Pararrhynchium, it is considered that the genus Lissodynerus should be restituted as a valid genus, not a synonym of Pararrhynchium.
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Interactions between Strepsiptera (Pseudoxenos Saunders, 1872) and solitary wasps (Eumeninae) are recorded for the first time in Brazil for Pachodynerus grandis Willink & Roig-Alsina, 1998. An updated worldwide checklist of the host species of Eumeninae for Pseudoxenos is provided.
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A total of 45 species belonging to 26 genera of the solitary wasp subfamily Eumeninae are listed, including 13 species and six genera that are newly recorded for Vietnam. The references for distribution in Vietnam are provided. Ectopioglossa keiseri nigra Giordani Soika is synonymized under Ectopioglossa keiseri van der Vecht; the male of this species is also described for the first time.
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A total of 45 species belonging to 26 genera of the solitary wasp subfamily Eumeninae are listed, including 13 species and six genera that are newly recorded for Vietnam. The references for distribution in Vietnam are provided. Ectopioglossa keiseri nigra Giordani Soika is synonymized under Ectopioglossa keiseri van der Vecht; the male of this species is also described for the first time.
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Six genera and nine species of Eumeninae (Eumenes punctatus de Saussure, E. papillarius (Christ), Delta dimidiatipenne(de Saussure), D. p. pyriforme (Fabricius), D. esuriens (Fabricius), Rhynchium brunneum (Fabricius), Anterhynchium ab-dominale (Illiger), Antepipona sibilans (Cameron), Allorhynchium argentatum (Fabricius)) , two genera and six species of Polistinae (Polistes wattii (Cameron), P. olivaceus (De Geer), P. rothneyi carletoni van der Vecht, P. indicus Stolfa), Ropalidia brevita Das & Gupta, R. cyathiformis (Fabricius), and one genus and four species of Vespinae (Vespa basalis Smith, V, orientalis Linnaeus, V, tropica (Linnaeus), V. velutina Lepeletier de Saint Fargeau) are recorded from Pothwar region. Eumenes punctatus and E. papillarius are newly recorded from Pakistan. An illustrated keys to subfamilies, genera and species are given
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Four new species and one subspecies of Eumenidae collected in the Oriental region are reported: Stenodynerus incurvitus nova spec, 5 from Taiwan, Parancistrocerus siamensis nova spec, $>, from Thailand, Rhynchium brunneum maladivum nova. ssp. cs <J from the Maledives, Allorhynchium violaceipenne nova spec, 5, from Philipp. Islands. The differences between Zethus varipunctatus (CAMERON) and Zethus nigerrimus GUSENLEITNER are presented.
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It has generally been accepted that the Philippine Islands are inhabited by only a single species of the Eumenid genus Rhynchium. In Miss Baltazar's valuable catalogue (1966) it is listed as a subspecies, atrum Saussure, of R. haemorrhoidale (Fabr.). However, when in 1963 I examined an extensive collection of Eumenidae from these islands in the U.S. National Museum, Washington ), the available material of this genus proved to consist of two very similar, though readily separable, species. The question thus arose, to which of these should be applied the name atrum Saussure. This species was described from "Les Indes Orientales, les Iles de la Sonde, la Nouvelle Guinée [Warou]. (Musée de Paris)". Unfortunately I did not find any specimens of this certainly rather heterogeneous series in the Paris Museum, and I therefore had to rely on a comparison of my material with the original description. It appeared that this description is almost certainly based on one or more specimens from the Philippine Islands, for it applies better to one of my two Philippine species than to any other of a great number of forms from various parts of the Oriental region. In order to establish the identity of R. atrum with certainty, however, I have considered it desirable to designate a neotype for this species. The material discussed in this paper is preserved in the following collections. AMNH = American Museum of Natural History, New York, U.S.A. BISH = Bernice P. Bishop Museum, Honolulu, Hawaii, U.S.A. BPI = Bureau of Plant Industry, Manila, Philippines. CNHM = Chicago Natural History Museum, Chicago, Ill., U.S.A.