Research Article |
Corresponding author: Akira Shimizu ( aquilashimizu7@gmail.com ) Academic editor: Maksim Proshchalykin
© 2021 Akira Shimizu, James P. Pitts, Jin Yoshimura, Raymond Wahis.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Shimizu A, Pitts JP, Yoshimura J, Wahis R (2021) New genus and species of Pompilinae spider wasps from the Oriental Region (Hymenoptera, Pompilidae). In: Proshchalykin MYu, Gokhman VE (Eds) Hymenoptera studies through space and time: A collection of papers dedicated to the 75th anniversary of Arkady S. Lelej. Journal of Hymenoptera Research 84: 339-359. https://doi.org/10.3897/jhr.84.68810
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The new genus Erythropompilus Shimizu & Pitts, gen. nov. from the Oriental Region (Pompilidae, Pompilinae) is described, based on the new species E. malaysiensis Pitts & Shimizu, sp. nov. from Malaysia. Two other new species of this genus, E. thailandensis Pitts & Shimizu, sp. nov. from Thailand and E. taiwanensis Pitts & Shimizu, sp. nov. from Taiwan, are also described. A key to the species is provided, and the taxonomic position of the genus is discussed.
Erythropompilus, Malaysia, Pompilidae, Pompilinae, Taiwan, taxonomy, Thailand
In the course of studying the systematics of spider wasps (Hymenoptera: Pompilidae) globally and more specifically from the Oriental and Australasian regions, we recently examined specimens from Malaysia, Thailand, and Taiwan belonging to the subfamily Pompilinae. From these regions, we found species, specimens of which were collected in Malaise traps and yellow-pan traps, of an enigmatic and unknown genus. A comparison of this genus with other genera in the Old World was performed, and we concluded that the specimens belong to an undescribed genus. Through detailed examination of the specimens, we also concluded that those specimens from the three regions belong to three separate undescribed species.
In this paper, we create a new genus, Erythropompilus Shimizu & Pitts, gen. nov., based on the new species E. malaysiensis Pitts & Shimizu, sp. nov. from Malaysia (type species). We also describe two other new species of the genus, E. thailandensis Pitts & Shimizu, sp. nov. from Thailand and E. taiwanensis Pitts & Shimizu, sp. nov. from Taiwan, we provide a key to the species, and we discuss the taxonomic position of the genus at the subfamily level.
Holotypes were photographed with a digital camera (Nikon Coolpix 4500 and MDC lens equipped with a stereo microscope Leitz TS and a transmitted light microscope Leitz Dialux). Photographs were stacked by using CombineZM (
The terminology for general morphology, including the wing veins and cells, follows
FW fore wing;
Fl1, Fl2, Fl3 … the first, second, third, … flagellomeres;
HW hind wing;
LID lower interocular distance;
MID middle interocular distance;
OOD ocello-ocular distance;
POD postocellar distance;
S1, S2, S3 ... the first, second, third, ... metasomal sterna;
SGP subgenital plate;
SMC1, SMC2, SMC3 the first, second and third submarginal cells of the fore wing;
T1, T2, T3 ... the first, second, third, ... metasomal terga;
TFD transfacial distance (= head width);
UID upper interocular distance.
The names of institutions in which specimens examined are deposited are abbreviated as follows:
QSBG Queen Sirikit Botanic Gardens, Chiang Mai, Thailand.
Erythropompilus malaysiensis Pitts & Shimizu, sp. nov. (here designated).
This genus differs from other genera of the subfamily Pompilinae by the following combination of features: Both sexes: setae on body sparse; clypeus small, as broad as or slightly narrower than LID (Figs
Erythropompilus malaysiensis Pitts & Shimizu, sp. nov., holotype, female A head in anterior view B head and pronotum in dorsal view C head in lateral view D head, pronotum and mesonotum in posterodorsal view (arrow, occipital carina) E scutellum, metanotum, metapostnotum and propodeum in dorsal view F whole body in lateral view G left fore tarsus in dorsal view H left fore and hind wings I mandible in anterior view J labio-maxillary complex in posterior view (left maxilla removed; arrow, preapical membranous area of prementum) K head and mesosoma in lateral view. Scale bars: 0.3 mm (C, J); 0.5 mm (G, I); 1 mm (A, B, D–F, H, K).
Erythropompilus malaysiensis Pitts & Shimizu, sp. nov., paratype, male (from Pasoh Forest Res., Negri S., Malaysia) A head in anterior view B head and pronotum in dorsal view C whole body in lateral view D left fore and hind wings E left flagellomeres 3–5 in medial view F metasomal sternum 6 and subgenital plate in ventral view G genitalia in dorsal view H genitalia in ventral view view I head and mesosoma in lateral view. Scale bars: 0.25 mm (F–H); 0.5 mm (E); 1 mm (A–D, I).
Of these features, the very sharply grooved outer orbit in both sexes, placoid sensilla on the flagellomeres, a pair of large, medially directed lateral hooks on S6, the very short, peg-like paramere, and the very broad and sinuate parapenial lobe in the male are unique to this genus. Our preliminary phylogenomics strongly support the close relationship of the genus with Tachypompilus Ashmead, 1902 and Ctenagenia Saussure, 1892.
Female. Greater part of body and legs with silvery-white pubescence densely. Vertex along inner orbit, labrum, and fore coxa posterobasally with sparse fine setae; S2–S6 with short erect setae, those on S6 dense.
Clypeus slightly convex with lateral margin strongly slanted, apicolateral corner broadly rounded (Figs
Pronotum with dorsum steeply sloping anteriorly, merging into declivity, latter being short and almost vertical (Figs
Fore tibia with several short spines internally. Fore tibial spur pale yellow. Fore tarsus lacking tarsal comb, without spines interiorly and exteriorly (Fig.
Erythropompilus taiwanensis Pitts & Shimizu, sp. nov., holotype, female A head in anterior view B head, pronotum, and scutum in dorsal view C scutellum, metanotum, metapostnotum and propodeum in dorsal view D whole body in lateral view E left fore tarsus in dorsal view F left fore and hind wings G head and mesosoma in lateral view H right hind tibia in dorsal view. Scale bars: 1 mm.
FW inner fascia broad on both sides of second abscissae of veins Rs and M (basal vein) and crossvein cu-a (Figs
S2 without transverse groove or depression. S6 not compressed laterally without median carina.
Male. Antenna much longer than head and mesosoma combined (Figs
Erythropompilus taiwanensis Pitts & Shimizu, sp. nov., paratype, male (from Kaohsiung, Luluei Forest Sta., 750 m, Taiwan) A head in anterior view B head and pronotum in dorsal view C whole body in lateral view D left fore and hind wings E left flagellomeres 2–4 in medial view F metasomal sternum 6 and subgenital plate in ventral view G genitalia in dorsal view H genitalia in ventral view. Scale bar: 0.25 mm (F–H); 0.5 mm (A, B, E); 1 mm (C, D).
The Oriental Region (Thailand, Malaysia and Taiwan).
The generic name is derived from the type species, of which the female mesosoma is orange-red for most part. The name is considered masculine.
Female
1 | Thorax (mesosoma except propodeum) dark in coloration, almost black (Fig. |
E. taiwanensis Pitts & Shimizu, sp. nov. |
– | Thorax mostly orange-red (Figs |
2 |
2 | Gena, in dorsal view, very thin, abruptly receding posteriorly (Fig. |
E. malaysiensis Pitts & Shimizu, sp. nov. |
– | Gena, in dorsal view, not very thin, roundly receding posteriorly (Fig. |
E. thailandensis Pitts & Shimizu, sp. nov. |
Male
1 | Frons, in profile, spherically convex (Fig. |
E. taiwanensis Pitts & Shimizu, sp. nov. |
– | Frons, in profile, gently convex (Figs |
2 |
2 | Thorax usually dark rufous (Fig. |
E. malaysiensis Pitts & Shimizu, sp. nov. |
– | Thorax mostly orange-red (Fig. |
E. thailandensis Pitts & Shimizu, sp. nov. |
Holotype
, ♀: ‘Pasoh Forest Res. Negri S., Malaysia X.20.79 sec. for. P. & M. Becker’ (
Both sexes: clypeus as broad as LID (Figs
Holotype female. Length: Body 6.5 mm; forewing 5.7 mm. Head 1.3 × as broad as long. MID 0.55 × head width. UID:MID:LID = 7.4:10:8.0. POL:OOL = 1: 0.62. Clypeus 2.4 × as broad as long. Gena, in profile, 0.2 × eye width. Scape:Fl1:Fl2 = 7.6:10:8.9; Fl1 3.7 × as long as thick, 0.86 × as long as UID. Longer spur of hind tibia 0.69 × hind tarsomere 1. FW marginal cell removed from wing tip by 0.47 × its own length. SMC2:SMC3 = 1:1.5 on vein M, 1:0.43 on vein Rs. SMC2 narrowed on vein Rs by 1.0 × its length on vein M, receiving crossvein 1m-cu at basal 0.57. SMC3 narrowed on vein Rs by 0.29 × its length on vein M, receiving crossvein 2m-cu at basal 0.45, removed from outer wing margin by 1.5 × its own length.
The type series. Female. (Measurements of the holotype are given in parentheses.) Length: Body 5.6–7.7 (6.5) mm; forewing 5.4–7.1 (5.7) mm. Body and legs dark rufous to black, except for mesosoma mostly orange-red (Fig.
Pubescence on body and legs mostly silvery, but that on following sericeous: upper frons, vertex, mesosoma dorsally (except for propodeum posteriorly), upper meso- and metapleura, metasomal terga anteriorly and posteriorly, metasomal sterna posteriorly, fore tibia and tarsus, and mid and hind tarsi.
Head 1.3–1.4 (1.3) × as broad as long. Vertex not convex between eye tops (Fig.
Pronotum with posterior margin subangulate medially (Fig.
Mid tibia with five to eight spines externally and two or three spines internally. Hind tibia with six to eight spines externally and five to seven spines internally. Longer spur of hind tibia 0.63–0.71 (0.69) × hind tarsomere 1.
FW and HW as shown in Fig.
Male. Body 4.6–6.2 mm; forewing 4.2–5.6 mm. Body and legs black (Fig.
Head 1.3 × as broad as long. Vertex, in frontal view, raised in ocellar area above level of eye tops (Fig.
Pronotum, in dorsal view, distinctly narrowing anteriorly (Fig.
Mid tibia with five or six spines externally and two or three short spines internally. Hind tibia with five to eight spines externally and four or five spines internally. Longer spur of hind tibia 0.65–0.83 × hind tarsomere 1.
Fore and hind wings as shown in Fig.
SGP somewhat concave ventrally. Genitalia (Fig.
The species name is derived from its locality, i.e., Malaysia.
Holotype
, ♀: ‘Wushe. Taiwan 1150m. V-15-83 Henry Townes’ (
Both sexes: clypeus narrower than LID (Figs
Holotype female. Length: Body 6.0 mm; forewing 5.9 mm. Body and legs black (Fig.
Head 1.4 × as broad as long. Vertex, in frontal view, almost transversely straight but slightly raised in ocellar area (Fig.
Pronotum gently and arcuately emarginate posteriorly (Fig.
Mid tibia with four spines externally and one spine internally; hind tibia with five or six spines externally and five spines internally (Fig.
FW and HW as shown in Fig.
Male. Based on the only paratype. Body 3.8 mm; forewing 3.6 mm. Body and legs reddish brown (Fig.
Head 1.3 × as broad as long. Vertex distinctly convex between eye tops (Fig.
Metapostnotum very short, 0.06 × as long as metanotum at midline.
FW and HW as shown in Fig.
Mid tibia with very small spines dorsally, three spines externally and two spines internally. Hind tibia with spines longer than those on mid tibia, five or six spines externally and three or four spines internally. Longer spur of hind tibia 0.65 × hind tarsomere 1.
SGP somewhat concave beneath. Genitalia (Fig.
The species name is derived from its locality, i.e., Taiwan.
Holotype
, ♀: ‘THAILAND Nakhon Nayok Khao Yai NP near Training Center 2 14°24.515'N, 101°22.432'E 750m Pan traps 5–6.ii.2007 Wirat Sukho leg. T2235’ (QSBG). Paratypes: Thailand: 1♀, same data as that of holotype (QSBG); 1♀, same locality, nature trail in secondary forest, 14°24.522'N, 101°22.434'E, 750 m, Malaise trap, 5.vii–12.viii.2006, Pong Sandao leg., T399 (QSBG); 1♂, same locality, behind headmasters house, 14°24.781'N, 101°22.689'E, 770 m, Malaise trap, 5–12.vii.2006, Pong Sandao leg., T144 (QSBG); 2♂, same locality and collector, Malaise trap, 19–26.vii.2006, 150 (QSBG); 1♀, same locality, Lum Ta Kong View Point, 14°25.82'N, 101°23.754'E, 744 m, Malaise trap, 5–12.iv.2007, Pong Sandao leg., T2123 (QSBG); 1♂, same locality, 12–19.iv.2007, Wirat Sukho leg., T2126 (QSBG); 1♀, same locality, 12–19.iv.2007, Wirat Sukho leg., T2126 (QSBG); 1♂, same locality, moist evergreen forest near the reservoir, 14°27.119'N, 101°21.482'E, 699 m, Malaise trap, 19–26.xi.2006, Wirat Sookkho leg., T997 (QSBG); 1♀, Nakhon Ratchasima, Khao Yai NP, Cobra zone near fire protection office, 14°28.524'N, 101°22.928'E, 757 m, Malaise trap, 19–26.vi.2007, Pong Sandao leg., T2227 (QSBG); 1♀, Tang Prov., Nayong, Khaochong, 75 m, 7°33.068'N, 99°47.369'E, 2005 (QSBG); 1♀2♂, Trang Prov., near Nam Tok, Tan Prov. Khoa Chong Mt., 140 m, 7°32'15"N, 99°47'35"E, vii.2005, D. Lohman (QSBG); 3♀1♂, same locality, date and collector, 7°32'15"N, 99°47'36"E (QSBG); 1♀2♂, same locality, date and collector, 7°32'16"N, 99°47'36"E (QSBG); 1♂, Trang Prov., Nayong Khaochong Lab, Malaise trap, 75 m, 7°33.38'N, 47°47.369'E, 20–22.vi.2005 (QSBG); 1♀, Khonkaen, Phu Pha Man NP, teak plantation, 16°43.241'N, 101°59.580'E 265 m, Malaise trap, 6–13.vi.2006, Vinai Boonma leg., T3 (
Both sexes: mid and hind femora with small spines set in pits apicodorsally. Female: gena, in dorsal view, thin but roundly receding posteriorly (Fig.
Erythropompilus thailandensis Pitts & Shimizu, sp. nov., holotype, female A head in anterior view B head and pronotum in dorsal view C scutellum, metanotum, metapostnotum, and propodeum in dorsal view D whole body in lateral view E left fore and hind wings F left maxilla in lateral view G labio-maxillary complex in posterior view (arrow, preapical membranous area of prementum) H head and mesosoma in lateral view I mid and hind tibiae in dorsal view. Scale bars: 0.3 mm (F, G); 1 mm (A–E, H, I).
Holotype female. Length: Body 7.1 mm; forewing 6.6 mm. Head 1.4 × as broad as long. MID 0.57 × head width. UID:MID:LID = 7.6:10:8.1. POL:OOL = 1:0.72. Clypeus 2.5 × as broad as long. Gena, in profile, 0.3 × eye width. Scape:Fl1:Fl2 = 7.5:10:8.6; Fl1 4.4 × as long as thick, 0.86 × as long as UID. Longer spur of hind tibia 0.68 × hind tarsomere I. FW marginal cell removed from wing tip by 0.53 × its own length. SMC2:SMC3 = 1:1.6 on vein M, 1:0.70 on vein Rs. SMC2 narrowed on vein Rs by 0.90 × its length on vein M, receiving crossvein 1m-cu at basal 0.57. SMC3 narrowed on vein Rs by 0.40 × its length on vein M, receiving crossvein 2m-cu at basal 0.51, removed from outer wing margin by 1.5 × its own length.
The type series. Female. (Measurements of the holotype are given in parentheses.) Length: Body 5.1–9.0 (7.1) mm; forewing 4.7–8.0 (6.6) mm. Body and legs black, except for mesosoma mostly orange-red (Fig.
Pubescence on dorsal part of mesosoma except for blackish part of propodeum, T1–5 medially and metasomal sterna posteriorly sericeous.
Head 1.3–1.4 (1.4) × as broad as long. Vertex slightly convex between eye tops (Fig.
Erythropompilus thailandensis Pitts & Shimizu, sp. nov., paratype, male (from Phu Pha Man NP Teak plantation, Khonkaen, Thailand) A head in anterior view B head and pronotum in dorsal view C whole body in lateral view D left fore and hind wings E right flagellomeres 2–4 in medial view F metasomal sternum 6 in ventral view G subgenital plate in ventral view H genitalia in dorsal view I genitalia in ventral view J head and mesosoma in lateral view. Scale bars: 0.25 m.
Pronotum with posterior margin gently arcuate or subangulate medially (Fig.
Mid tibia with about five spines externally, two or three spines internally (Fig.
FW and HW as shown in Fig.
Male. Body 4.1–7.9 mm; forewing 4.1–7.6 mm. Body and legs black, but pronotum except anteriorly, meso- and metanota, metapostnotum, and meso- and metapleura except ventrally orange-red (Fig.
Head 1.3–1.5 × as broad as long. Vertex, in frontal view, distinctly convex above level of eye tops (Fig.
Mid tibia with four to six spines externally and two to six spines internally. Hind tibia with four or eight spines externally and four to six spines internally. Longer spur of hind tibia 0.67–0.82 × hind tarsomere 1.
FW and HW as shown in Fig.
SGP flattened ventrally with several long setae laterally near middle (Fig.
The species name is derived from its locality, i.e., Thailand.
In recent molecular phylogenetic analyses, the monophyly of the subfamily Pompilinae was well supported, more so than the other large subfamily Pepsinae (
The mid and hind femora have one or several small spines set in grooves or pits apicodorsally (
In practice, however, this characteristic is found not solely in Pompilinae but also in Ceropalinae (Ceropales Latreille, 1796), Notocyphinae (Notocyphus Smith, 1855), and some Pepsinae (e.g., Minotocyphus Banks, 1934, Calopompilus Ashmead, 1900, Austrosalius Turner, 1917, Dolichocurgus Haupt, 1937, Mimocurgus Haupt, 1937, Alococurgus Haupt, 1937, Pachycurgus Haupt, 1937, Eremocurgus Haupt, 1937, Pompilocalus Roig-Alsina, 1989, Hypoferreola Ashmead, 1902, Aimatocare Roig-Alsina, 1989, Anacyphonyx Banks, 1946, and Cordyloscelis Arnold, 1935). In older classifications, Minotocyphus was placed in Notocyphinae (
FW vein CuA1 is deflected posteriorly at the base (
This feature is also found in Ceropalinae (Irenangelus Schulz, 1906, and Ceropales) and some Pepsinae (e.g., Minotocyphus, Dolichocurgus, Pachycurgus, Plagicurgus Roig-Alsina, 1982, Anacyphonyx, Pepsis Fabricius, 1804, Hemipepsis Dahlbom, 1843, Cyphononyx Dahlbom, 1845 and Entypus Dahlbom, 1844). Thus, the feature is not always characteristic of Pompilinae.
Spines at the apex of the hind tibia are of unequal length, more or less splayed out and irregularly spaced (
This is a good feature for Pompilinae. The condition is, however, sometimes subtle. For example, in some parasitoid genera, such as Allochares Banks, 1917 and Eoferreola Arnold, 1935 (Pompilinae), the spines are short and a few, and their splayed-out condition is indistinct. Nevertheless, in these genera, the spines are of unequal thickness, as in the other genera of Pompilinae. Thus, the differing thickness of the apical spines on the hind tibia is also available for the definition of this subfamily.
A cluster of the HW basal hamuli are strongly proximal to the separation of vein C from vein Sc+R+Rs (
This characteristic is not applicable to a few Pompilinae, e.g., Priochilus Banks, 1944, Braunilla Wasbauer & Kimsey, 2019 (= Balboana Banks, 1944), and Parabatozonus Yasumatsu, 1936, in which the cluster of basal hamuli are at or distal to the separation of vein C from vein Sc+R+Rs, as in most Pepsinae. In contrast, in some Pepsinae, e.g., Psoropempula Evans, 1974, Trichocurgus Haupt, 1937 and some Priocnemis Schiødte, 1837 (e.g., P. pusilla (Schiødte, 1837)), the cluster of the basal hamuli are strongly proximal to the separation of vein C from vein Sc+R+Rs.
The prementum has a preapical circular, or heart- or spade-shaped membranous area (
This feature is applicable to almost all members of Pompilinae, although the verification of the condition is difficult. To examine the degree of the sclerotization of the apical prementum, in many cases, the labio-maxillary complex has to be removed from a specimen after softening the specimen with water vapor.
Erythropompilus possesses all the above features, and, thus, this genus should be treated as a member of Pompilinae. We are currently conducting molecular phylogenetic analyses using ultra-conserved elements, and the preliminary phylogenomics highly support this subfamilial position of the genus and its close relationship with Tachypompilus and Ctenagenia.
We thank Dr. A. S. Lelej (Federal Scientific Center of the East Asia Terrestrial Biodiversity, FEB RAS, Vladivostok, Russia) for his help in various ways in the course of our studies on spider wasps. We also thank G. Broad (Natural History Museum, London, UK) for the examination of various specimens from the world, and A. Harris (Otago Museum, Dunedin, New Zealand), R. A. Cambra (University of Panama, Panama City, Panama) and two anonymous reviewers for critically reviewing the manuscript and making important suggestions. This research was supported by the Utah Agricultural Experiment Station, Utah State University, and approved as journal paper number 9473.