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A Taxonomic Review of the Indian Species of the Genus Eumenes Latreille

Written By Sunday, April 17, 2022 Add Comment Edit

© The Authors, published by EDP Sciences, 2020

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1 Introduction

The subfamily Eumeninae is cosmopolitan and the largest subfamily in the Vespidae, consisting of more than iii,500 described species in 210 genera [i, two, Table 1]. Most of eumenine wasps are solitary even though they get together around a suitable nesting site and a few genera are known to be communal nesters [ii, 3]; some genera, such every bit Orancistrocerus and Calligaster, are subsocial. Many eumenine wasps construct nests with soil/mud mixed with relatively modest amount of wasps' oral secretion or saliva, but Calligaster species use plant materials, and Epsilon is known to brand their nests with institute-origin resin. Others use pre-existing cavities as their nesting places, and some are burrowers in the soil or wood, in which cell partitions or closing plugs are usually made of soil/mud or only sometimes plant cloth such as in Zethus [2, 4, 5].

Eumenine wasps are predaceous on terrestrial insects, hunting mainly lepidopterous caterpillars or sometimes coleopterous larvae equally food for their larvae ([half dozen, vii]; Iwata, 1971 in [iii]), and thus they are more than or less at college positions in a food web of terrestrial arthropods. They likewise visit flowers for nectar as their own free energy source and thus are considered as potential pollinators of many plants [five, 6, vii].

The Indonesian Archipelago, consisting of about 17,500 islands spreading from Sumatera in the west to New Republic of guinea in the eastward, is known for the mega biodiversity. In the archipelago, 4 (Eumeninae, Stenogastrinae, Polistinae and Vespinae) of the six vespid subfamilies occur, of which the Eumeninae are the most various, with a total of 226 species in 51 genera so far recorded in the archipelago. The species recorded from Sarawak, Sabah, Brunei, Timor Leste and Papua New Guinea were included in the full biodiversity of Eumeninae from Indonesian Archipelago, since the countries' edge may take no biological meaning in the distribution of species.

Compared with the other 3 vespid subfamilies, to which the species belonging are all social, the taxonomy of the Eumeninae, in general, is still much more poorly studied. Of the eumenine wasps known from the archipelago, 16 genera are amongst those treated as "potter wasps with a petiolate metasoma", which are characterized by having the offset metasomal tergum at least twice as long as its apical width. These eumenine wasps with a petiolate metasoma were originally described in the genus Eumenes Latreille, 1802, genera in the and then-chosen "Zethinae" and in the genus Labus.

Inside the Eumeninae, the wasps with a petiolate metasoma distributed in the Indo-Australian Region have been more than or less well studied compared with those with a non-petiolate metasoma, but their generic affinities and the concepts of some species yet remain unestablished. In the present report, the taxonomy and distribution (or biogeography) of potter wasps with a petiolate metasoma occurring in the Indonesian Archipelago are reviewed.

two Material and Methods

2.1 Specimens examined

Examination of the specimens was carried out to study the morphological characters and clarify the distributional record, including determining the color/marking pattern inside local populations. The specimens examined in the nowadays report were those having deposited in the collections of the Museum Zoologicum Bogoriense, Cibinong, Bogor, Indonesia (MZB), the Natural History Collection at Ibaraki University, Mito, Nippon (IUNH), the Systematic Entomology Institute, Hokkaido Academy Museum, Sapporo, Nippon (SEIHU), the Natural Resources Inventory Center, National Establish for AgroEnvironmental Sciences, Tsukuba, Japan (NRIC), the National Museum of Nature and Science, Tsukuba, Japan (NSMT), and Plant of Ecology and Biological Resources, Hanoi, Vietnam (IEBR). In improver to those specimens in the collections, we have collected many specimens in various places in the Indonesian Archipelago, which were pinned and dried and deposited in the MZB and IUNH.

two.2 Literature review

A literature review was conducted to study the taxonomic history of Eumeninae at the genus, species and subspecies levels. Placements of the genera into the tribes are based on Hermes et al. [2], and the use of the "Labus group" is based on the authors' opinion, since the taxonomic handling has not been carried out to clarify their phylogenetic position.

3 Results and Discussion

3.1 Taxonomic history

Hermes et al. [2] performed a comprehensive cladistic analysis of the Eumeninae and corroborated a natural classification of Eumeninae at the tribal level to recover the Eumenini, Odynerini and Zethini, which had been previously bachelor. They also clearly showed that the potter wasps with a petiolate metasoma do not course a monophyletic group, but they are placed in all the 3 monophyletic tribes, namely Zethini, Odynerini and Eumenini, suggesting that the shape of the get-go metasomal segment is a weak character due to its high heterogeneity (see also [viii]). Yet, their report did non include several Oriental and/or Afrotropical genera with a petiolate metasoma, such as Calligaster de Saussure, 1852, Labus de Saussure, 1867 and Cyrtolabulus van der Vecht, 1969, and their phylogenetic positions have non still been discussed.

Many species originally described in the genus Eumenes have been transferred to several genera equally the results of taxonomic works, including the establishment of new genera, past subsequent authors. The most important taxonomic studies on the Oriental Eumeninae with a petiolate metasoma are van der Vecht [9,10,11,12,13] and Giordani Soika [14,15], which resulted in a proliferation of generic names in the Eumeninae: Omicroides Giordani Soika, 1935, Pseumenes Giordani Soika, 1935, Nortozumia van der Vecht, 1937, Coeleumenes van der Vecht, 1963 and Cyrtolabulus van der Vecht, 1969. These taxonomic treatments to split the preexisting genera were then continued by Giordani Soika [xvi] and Gusenleitner and Gusenleitner [17], in which Phimenes Giordani Soika, 1992 and Flavoleptus Giordani Soika, 1992, and Norteumenes Gusenleitner and Gusenleitner, 2013 were proposed.

The first tape of potter wasps under Zethus from the Indonesian Archipelago was Cameron [eighteen], who described Z. quadridentatus and Z. varipunctatus from Borneo. The other Oriental genus in the so-called "Zethini", Calligaster de Saussure, 1852, was described based on C. cyanoptera de Saussure, 1852 from Coffee, only since de Saussure (1855) treated Calligaster as a division of Zethus its taxonomic position had been ambiguously interpreted until Bequaert [19] definitely reinstated its generic status.

The Labus grouping is an Oriental and Ethiopian eumenine fauna. Ii genera are recognized, Labus from the Oriental region and Cyrtolabulus from Indo-Ethiopian region. The genus Labus was described past de Saussure (1867) based on L. spiniger de Saussure, 1867 from Java, and since then received many interpretations, as it differs in many characters from its related genera [8, 20]. Yet, Bequaert [20] discussed the generic characters and established skillful diagnosis based on Ethiopian species. Van der Vecht [9] newly described another three species from Java, having contributed to a better agreement of the Labus animate being in the Indonesian Archipelago. The genus Cyrtolabus was described by van der Vecht [13] to accommodate ii species from the Indian subcontinent; subsequently Cyrtolabulus van der Vecht, 1969 was established as a replacement name for Cyrtolabus van der Vecht, 1963.

iii.2 Diversity and biogeography

Potter wasps with a petiolate metasoma in the Indonesian Archipelago consisting of 131 species-group taxa (80 species and 51 subspecies; Table one). The diversity of a petiolate metasomal genera in the archipelago consisted of widely distributed species (33) and endemic species (47). Sumatra, Java, Bali and Borneo are more than likely representing Oriental element with less endemic species occurs; Sulawesi, Lesser Sunda Islands and Moluccas species more or less consisting mainly with the occurrence of endemic species; New Guinea species probably underrate, equally they may less explore areas (Fig. one).

The tribe Eumenini includes four genera known to occur in the Indonesian Archipelago: Eumenes Latreille, 1802, Delta de Saussure, 1855, Phimenes Giordani Soika, 1992 and Omicroides Giordani Soika, 1935. Delta, Eumenes and Phimenes take widely distributed genera, while Omicroides is representing Oriental element. In this tribe, a full of 38 species in iv genera with a petiolate metasoma are recognized from the Indonesian Archipelago. They are ix species in the genus Delta; 20 species in the genus Eumenes; the species in the monotypic genus Omicroides; and eight species in the genus Phimenes.

The tribe Odynerini is consist of eight genera known to occur in the Indonesian Archipelago: Coeleumenes van der Vecht, 1963, Ectopioglossa Perkins, 1912, Flavoleptus Giordani Soika, 1992, Norteumenes Gusenleitner & Gusenleitner, 2013, Nortozumia van der Vecht, 1937, Pareumenes de Saussure, 1855, Pseudozumia de Saussure, 1875 and Pseumenes Giordani Soika, 1935. Flavoleptus, Norteumenes, Nortozumia and Pseudozumia are representing the Oriental genera in the Odynerini; Coeleumenes, Ectopioglossa, Pareumenes and Pseumenes are widely distributed genera and occur forth the archipelago. In the Odynerini, the following 26 species in eight genera are recognized in the Indonesian archipelago: three species in the genus Pareumenes, 3 species in the genus Pseumenes, six species in the genus Coeleumenes, seven species in the genus Ectopioglossa, two species in the genus Nortozumia, 3 species in the genus Pseudozumia, and each of the monotypic genera Flavoleptus and Norteumenes.

The tribe Zethini consists of three genera known to occur in the Indonesian Archipelago: Calligaster de Saussure, 1852, Elimus de Saussure, 1852 and Zethus Fabricius, 1804. In the Zethini, eight species in three genera are recognized in the Indonesian Archipelago; namely, iii species in the genus Calligaster, four species in the genus Zethus and one species in the genus Elimus.

The Labus group is consisting but single genus known to occur in the Indonesian Archipelago, representing by eight species and distributed from Sumatra in the w to the Lesser Sunda Islands in the east. In the genus Labus, which may class, together with the Indo-Ethiopian genus Cyrtolabulus, an independent tribe, viii species are recognized in the Indonesian Archipelago.

With extremely loftier biodiversity characterized by a high level of endemism and circuitous geological history, the Indonesian Archipelago, the islands between continental Southeast Asia and Australia, has long attracted the attention of biologists from biogeographical and evolutionary (in terms of speciation) points of view. From the geological perspective, the Indonesian Archipelago is comprised of two major components, namely the Sunda Shelf component (Borneo, Sumatra, Java and Bali), where the fauna consists of mainly Oriental elements; and the Sahul Shelf component (New Republic of guinea and its adjacent islands, including Aru Islands), where the animal is comprised of Papua-Australian elements.

In the fauna of potter wasps with a petiolate metasoma of the Indonesian Archipelago, genera such as Calligaster, Zethus, Flavoleptus, Norteumenes, Nortozumia, Pseudozumia, Omicroides and Labus are the Oriental elements (Figs two, 3), while Elimus is the Papua- Australian element (Fig. 2). Sumatra, Java, Bali and Kalimantan, as a part of Sunda Shelf/Oriental component, accept mainly Oriental eumenine fauna, including several area-specific species of the Oriental genera (Fig. iv). New Guinean animal is comprised of Papua-Australian species of the widely distributed genera, together with widely distributed species and species endemic to New Republic of guinea (Fig. 4).

The islands lying betwixt the 2 shelves form a detail area, known as Wallacea, which is considered every bit the harbour of endemic species, as well as those on potter wasps with a petiolate metasoma (Fig. 4). The creature of potter wasps with a petiolate metasoma in the Wallacea is constituted mainly with the area-specific species in the rather widely distributed genera, such as Eumenes, Delta, Phimenes, Coeleumenes, Ectopioglossa, Pareumenes, and Pseumenes; in the western part of Wallacea, namely Sulawesi and Lesser Sunda Islands, they are represented mainly past widely distributed genera, together with Oriental genera Calligaster, Zethus and Oriental genera. Labus (Fig. 2); in the eastern role of Wallacea, namely the Moluccas (excluding Aru Islands), they are constituted mainly by area endemic species of widely distributed Occurrences of four Papua-Australian species in the Moluccas show that the area is the transition zone betwixt Oriental and Australasian faunae (Fig. 4).

thumbnail Fig. i

Total species and number of endemic species of potter wasps with a petiolate metasoma in Indonesian Archipelago.

Table 1

An updated listing of the Potter Wasps (Hymenoptera: Vespidae, Eumeninae) with a Petiolate Metasoma from Republic of indonesia Archipelago (based and modified from Nugroho et al. [21]).

thumbnail Fig. two

Map showing the collection records of the Labus group and Zethini at the generic level in the Indonesian Archipelago. (a) Wallace's line. (b) Lydekker's line
thumbnail Fig. 3

Map showing the drove records of Oriental genera of Odynerini in the Indonesian Archipelago. (a) Wallace's line. (b) Lydekker's line
thumbnail Fig. four

Number of species of the potter wasps with a petiolate metasoma in the Indonesian Archipelago. Sumatra, Coffee, Bali and Borneo are function of Sunda Shelf; Wallacea consists of Lesser Sunda, Sulawesi and Moluccas; New Guinea is Sahul Shelf component, and including Aru Islands
thumbnail Fig. 5

Marking blueprint and distribution of the subspecies Phimenes flavopictus and Phimenes arcuatus [modified from van der Vecht (1959)]: 1—16: Phimenes flavopictus. (i) P. f. flavopictus. (2) P. f. continentalis. (3) P. f. kalimantenus. (4) P. f. formosanus. (v) P. f. andamanicus. (half-dozen) P. f nicobaricus. (7) P. f. simalurensis. (8) P. f. umbripennis. (9) P. f telonus. (10) P. f maidli. (11) P. f. engganensis. (12) P. f. blanchardi. (thirteen) P. f. baweanus. (14) P. f. timorensis. (fifteen) P. f. dammae. (16) P. f. aidrytus. 17—24: Phimenes arcuatus. (17) P. a. arcuatus. (xviii) P. a. praslinius. (19) P. a. buruanus. (20) P. a. obiensis. (21) P. a. lyratus. (22) P. a. transilis. (23) P. a. amboinensis. (24) P. a. muruensis

thumbnail Fig. half dozen

Map showing the parallel distribution of marking pattern in four species: (one) Phimenes flavopictus, (two) Pareumenes quadrispinosus, (3) Pseumenes depressus, (iv) Coeleumenes impavidus

3.3 Color variation and marking pattern

Species that are widely distributed generally show wide range of variation in the marker patterns, such as Phimenes flavopictus, P. arcuatus, Pareumenes quadrispinosus, Pseumenes depressus and Coeleumenes impavidus. The marking patterns of two closely related polymorphic species, Oriental Phimenes flavopictus and Papuan P. arcuatus, are remarkable (Fig. 5). Their mark patterns consist of those of the "mainland populations (or subspecies)" that inhabit a wide range of geographical areas and those of "peripheral populations (or subspecies)" inhabiting adjacent islands (van der Vecht, 1961). The mainland subspecies of P. flavopictus, flavopictus in continental Asia (Fig. five [1]), continentalis occurring in continental Asia and Sumatra (Fig. five [2]) and formosanus in Taiwan (Fig. 5 [iv]), are more or less extensively marked with yellow, and occur sympatrically in some areas. The Bornean subspecies, kalimantenus (Fig. 5 [three]), has the marking pattern similar to those of these mainland subspecies but its yellowish markings are constantly narrower or reduced. The peripheral subspecies of P. flavopictus inhabiting modest islands from the Andaman Islands, through the group of islands off the westward coast of Sumatra and Java, to Lesser Sunda Islands and Tanimbar, and also the Philippines, are characterized by the island(s)-specific marking patterns. In some subspecies, mosaic marking patterns are recognized, that is, in some body parts xanthous markings are more extensive than those in mainland subspecies and in other torso parts the yellow markings are reduced or even absent-minded (Fig. 5 [5-7, 9, ten]). In other subspecies, yellow markings are much reduced, or even the body is nearly entirely black (Fig. 5 [12, thirteen, 16]), while others accept the marker pattern with yellow marking only slightly reduced and/or only slightly extensive than those of the mainland subspecies (Fig. 5 [viii, 11, 14, fifteen]).

The mainland subspecies of P. arcuatus, arcuatus (Fig. 5 [17]), has the marking pattern like to those of the mainland subspecies of P. flavopictus, and occurs in New Guinea (including Aru Islands) and North Queensland of Commonwealth of australia, and also in Kei Islands, where P. flavopictus dammae [Fig. v (15)] with the marking pattern like to P. a. arcuatus occurs. The peripheral subspecies of P. arcuatus are found in the Moluccas and adjacent islands of New Guinea [Fig. 5 (18-24)]. They evidence, as the peripheral subspecies of P. flavopictus, the island(s)-specific mark patterns, which are more often than not characterized by orange markings instead of yellowish markings [Fig. v (18-21)] or reduction of yellow markings [Fig. 5 (22)]. Subspecies ambonensis is similar to mainland subspecies, simply the markings on the metasoma are slightly darker [Fig. 5 (23)].

Every bit mentioned in a higher place, their marking patterns oftentimes consist of those of the "mainland populations (or subspecies)" that inhabit a broad range of geographical areas and those of "peripheral populations (or subspecies)" inhabiting next islands; the mainland subspecies more or less testify similar markings with simply slightly reduced and/or only a slightly extensive variation and occurs sympatrically in some areas, while the peripheral subspecies usually characterized past the isle(s)-specific marker patterns (Fig. 5). Sympatric occurrences of forms with quite unlike color patterns in widely distributed species may invite us to make a further intensive study to constitute their taxonomic condition, namely whether they are variations within a given area or unlike species.

Trunk color markings in aculeate wasps are commonly establish to exist contrasted patterns of bright and night (usually black) color as a alarm signal to their predators, despite the fact that the degree of darkness drastically varies amid local populations of a given species. The color variations in the polymorphic and widely distributed eumenine species with a petiolate metasoma in the Indonesian Archipelago show mosaic pattern of distribution merely they practise not change along geographical clines, and thus such the color variations have been more probable driven by aposematic pressure level rather than by abiotic factors. Sympatric occurrences of virtually the same marking pattern in four species (Phimenes flovopictus, Pareumenes quadrispinosus, Pseumenes depressus, and Coeleumenes impavidus), probably take resulted from the Mullerian mimicry, and are recognized in the continental Asia, the Philippines, Borneo and Java (Fig. 6).

The present study was supported by the Japan Society for the Promotion of Science (JSPS). We are thankful to Dr. Masahiro Ohara (Systematic Entomology Institute, Hokkaido University Museum), Dr. Akihiko Shinohara (National Museum of Nature and Science, Tsukuba, Japan), Dr. Shinichi Yoshimatsu (Natural Resources Inventory Eye, National Institute for AgroEnvironmental Sciences, Tsukuba, Nippon) and Dr. Nguyen Thi Phuong Lien (Institute of Environmental and Biological Research, Hanoi, Vietnam) for arranging loans of the specimens.

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All Tables

Tabular array 1

An updated list of the Potter Wasps (Hymenoptera: Vespidae, Eumeninae) with a Petiolate Metasoma from Indonesia Archipelago (based and modified from Nugroho et al. [21]).

All Figures

thumbnail Fig. 1

Total species and number of endemic species of potter wasps with a petiolate metasoma in Indonesian Archipelago.
In the text
thumbnail Fig. 2

Map showing the collection records of the Labus grouping and Zethini at the generic level in the Indonesian Archipelago. (a) Wallace's line. (b) Lydekker's line
In the text
thumbnail Fig. 3

Map showing the collection records of Oriental genera of Odynerini in the Indonesian Archipelago. (a) Wallace's line. (b) Lydekker'south line
In the text
thumbnail Fig. 4

Number of species of the potter wasps with a petiolate metasoma in the Indonesian Archipelago. Sumatra, Java, Bali and Borneo are part of Sunda Shelf; Wallacea consists of Lesser Sunda, Sulawesi and Moluccas; New Republic of guinea is Sahul Shelf component, and including Aru Islands
In the text
thumbnail Fig. five

Marking pattern and distribution of the subspecies Phimenes flavopictus and Phimenes arcuatus [modified from van der Vecht (1959)]: ane—16: Phimenes flavopictus. (one) P. f. flavopictus. (ii) P. f. continentalis. (3) P. f. kalimantenus. (4) P. f. formosanus. (5) P. f. andamanicus. (vi) P. f nicobaricus. (7) P. f. simalurensis. (8) P. f. umbripennis. (9) P. f telonus. (10) P. f maidli. (11) P. f. engganensis. (12) P. f. blanchardi. (13) P. f. baweanus. (14) P. f. timorensis. (15) P. f. dammae. (sixteen) P. f. aidrytus. 17—24: Phimenes arcuatus. (17) P. a. arcuatus. (18) P. a. praslinius. (19) P. a. buruanus. (20) P. a. obiensis. (21) P. a. lyratus. (22) P. a. transilis. (23) P. a. amboinensis. (24) P. a. muruensis

In the text
thumbnail Fig. 6

Map showing the parallel distribution of marker pattern in four species: (1) Phimenes flavopictus, (ii) Pareumenes quadrispinosus, (three) Pseumenes depressus, (four) Coeleumenes impavidus

In the text

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