New species and records of Evanioidea and Stephanoidea from New Caledonia (Hymenoptera)

. Three new species of aulacid wasps, Aulacus pascali sp. nov., Pristaulacus elveni sp. nov. and Pristaulacus villemantae sp. nov., and a new species of gasteruptiid wasp, Gasteruption jenningsi sp. nov., are described and ﬁ gured. Additionally, we update identi ﬁ cation keys to New Caledonia species of aulacids and gasteruptiids. We also provide new data on New Caledonian Evaniidae, and Stephanidae. from New Caledonia. Additionally, we provide new data on other Evanioidea and Stephanoidea from New Caledonia.


Introduction
2005) known from this territory. Additionally, a summary of the hymenopteran fauna of New Caledonia was provided by Jennings et al. (2013).
The family Aulacidae currently includes 310 species. Its diversity has greatly benefi ted from the recent descriptions of species, mostly from Africa and Asia (Smith 2001;Turrisi , 2007Turrisi , 2014Turrisi et al. 2009;Turrisi & Madl 2013;Jennings et al. 2018;. Even if previously known from three genera, aulacid wasps are now placed within two genera: Aulacus Jurine 1807, with 122 species, and Pristaulacus Kieff er 1900 (including the former Panaulix Benoit, 1984) with 188 species . Both genera are represented in all biogeographic regions, except Antarctica, with the genus Aulacus unknown from the Afrotropics (Turissi 2014 and included references). The monophyly of the family is well supported (Jennings & Austin 2000;Turrisi et al. 2009;Li et al. 2018). Extant species of aulacids are known to be koinobiont endoparasitoids of wood-boring larvae of Hymenoptera Linnaeus, 1758 and Coleoptera Linnaeus, 1758 (Deyrup 1984;Gauld & Hanson 1995;.
The family Gasteruptiidae comprises six extant genera (Crosskey 1962;Jennings & Austin 1997, 2002Macedo 2009;van Achterberg & Talebi 2014; but to date, many species, especially fossils, are placed as incertae sedis making it diffi cult to set up a species catalog. Extant gasteruptiids are placed in two extant gasteruptiid subfamilies. The subfamily Hyptiogastrinae Crosskey, 1953 has a restricted Gondwanan distribution and is currently known in Australian mainland, Tasmania, New Zealand and South America, as well as in New Caledonia, New Guinea, Fiji and Vanuatu (Jennings & Austin 1994a, 1994b, 1997, 2002. Among this subfamily, the genus Hyptiogaster Kieff er, 1903 is endemic to Australia, while Pseudofoenus Kieff er, 1902 displays an Australasian distribution (Jennings & Austin 2002). The subfamily Gasteruptiinae Ashmead, 1890, mainly represented by the genus Gasteruption Latreille, 1796, displays a worldwide distribution while other genera are found only in South America (van Achterberg & Talebi 2014;Macedo 2009Macedo , 2011Zhao et al. 2012). As for all other extant families composing the Evanioidea, the monophyly of Gasteruptiidae is well supported (Jennings & Austin 2002;Macedo 2009;Li et al. 2018;Parslow et al. 2020). Adults of extant species of gasteruptiids are free-living insects feeding on nectar and pollen ) while predator-inquiline, which mean that their larvae feed on the larval food of solitary bees, after consuming the egg or larva of the bee or solitary wasps (Malyshev 1968;Parslow et al. 2020;. The family Evaniidae is common, nearly cosmopolitan, and includes approximately 580 extant species in 21 genera, but its diversity is clearly underestimated (Deans 2005;Mullins et al. 2012). Interestingly, their larvae are considered as predators of cockroach eggs in oothecae (Huben 1995). However, the association between evaniid larvae and their prey is much more reminiscent of parasitoid behavior.
The crown wasp superfamily, Stephanoidea, is easily distinguishable from most of the other Hymenoptera in having tubercles on the vertex (Goulet & Huber 1963). The number of extant Stephanidae has been greatly increased since the last catalog of Aguiar (2004) and is now represented by 345 species (Hong et al. 2011) occurring mainly in subtropical and tropical forests (Vilhelmsen 1997;van Achterberg 2002). Species of Stephanidae develop as solitary idiobiont ectoparasitoids of larvae of wood-boring insect (Aguiar 2004).
Herein we describe a new species of Aulacus, two new species of Pristaulacus and a new species of Gasteruption from New Caledonia. Additionally, we provide new data on other Evanioidea and Stephanoidea from New Caledonia.

Repositories
Specimens examined for the present study are deposited in the following collection: MNHN = Muséum national d'histoire naturelle, Paris, France

Specimen examination
Specimens were examined using an AMSCOPE and a Leica MZ APO stereo microscopes equipped with a micrometer, at several magnifi cation planes. Series of photographs were taken at diff erent focal planes using a Canon EOS 5D mark II camera mounted on a Leica MZ APO stereo microscope or an Olympus TOUGH TG-5. These series of photographs were stacked using Helicon Focus. The fi gures were processed with Adobe Illustrator and Photoshop software.

Terminology and abbreviations
The general terminology follows  and Goulet & Huber (1993). The sculpture terminology follows Harris (1979

Etymology
The specifi c epithet is a patronym honoring Olivier Pascal, manager of the terrestrial part of the "Our Planet Reviewed" expeditions. The species epithet is to be treated as a noun in the genitive case. . Head, mesosoma and coxae II and III, ovipositor valves black; hind legs tarsomeres, tibia III, propleuron, metasoma and ovipositor brown to dark brown; palpi, pronotum, pro-coxa, trochanter white; mandibles, scape, pedicel, femora, pro-and mesotibia, pro-and mesotarsomeres yellow; fl agellomeres yellow basally, gradually becoming dark brown toward apex; wings hyaline with brown veins. H ( Fig. 1A-C). In full-face view, 1.23 × as wide as long; mandibles visible but closed [tooth number not visible]; clypeus slightly convex, with small blunt projecting tooth medially; malar space 0.58 × as long as scape, margin sinuate; antenna with 14 antennomeres, scape 1.60 × as long as wide and slightly longer than pedicel and as long as F-I, F-I slightly longer than F-II, both more than twice as long as wide; fl agellum thickening toward apex, apical fl agellomere longer than preceding one; subantennal groove shallow and indistinct; frons without lateral medial carina above torulus; vertex with ocelli arranged in small isosceles triangle, mid-ocellus separated from lateral ocelli by its diameter, lateral ocelli separated  1D). Propleuron ventrolateral carina pronounced; pronotum without angular process, covered with tiny, erect and dense pilosity; mesoscutum in lateral view slightly angular anterodorsally, notauli conspicuous, parapsidal lines present; metapostnotum narrow with posterior margin strongly curved; mesepimeron broad; propodeum straight under metasomal insertion; protrochantellus present; tibial spurs formula 1-2-2 with mid-and hind-tibial spurs equal and similar in shape; hind coxa with ovipositor guide on inner surface, somewhat distal, transverse and somewhat oblique; hind femur 0.68 × as long as hind tibia; TS 1 3.20 × as long as TS 2; TS 2 1.25 × as long as TS 3; TS 3 1.60 × as long as TS 4; TS 4 0.55 × as long as TS 5; hind tarsal claw 0.35 × as long as TS 5; tarsal claws simple. F (Fig. 1F). Pterostigma twice as wide as costal cell; marginal cell with Rs abscissae curved; sub-marginal cell with vein 2-Rs as long as 1-Rs, vein 2-Rs + M shorter than 2-Rs; veins 2r-m and 3r-m largely nebulous; discal cell rectangular, vein 1-M and 1m-cu parallel; second discal cell elongate with vein 2m-cu ending at mid-length of 3-M vein; sub-basal cell thin, 2.6 × as wide as discal cell. H (Fig. 1F). Venation reduced, Sc + R tubular, C and A nebulous, with 3 hamuli, second and third closer to each other. M (Fig. 1E). Clavate, slightly longer than mesosoma; T-I about 3.00 × as long as T-II; T-II to T-VI of similar length slightly shortening toward apex.

Distribution
It is known only from its type locality, Katalupaik (Fig. 12A).

Remarks
Aulacus pascali sp. nov. can be distinguished from A. coracinus Jennings, Austin & Stevens, 2004 by the reduced hind wing venation (R + Rs, M + Cu, Cu, r-m and 2-M absent), the pronotum white and the appendages mostly yellow, and its smaller size (3.18 mm vs 11.5 mm), from A. emineo Jennings, Austin & Stevens, 2004 by the presence of the transverse ovipositor guide on the inner surface of the hind coxa and from A. burwelli Jennings, Austin & Stevens, 2004 by its coloration (body of A. pascali sp. nov. mostly black vs body of A. burwelli black with extensive orange coloration on mesosoma and metasoma), the metasoma more clavate and its smaller size (3.18 mm vs 4.1-5.9 mm).

Distribution
Aulacus burwelli is known from the type locality, Table Unio road , and from Katalupaik (Fig. 12B).

Remarks
During the 2017 "Our Planet Reviewed" expedition, 11 females were collected in Malaise traps at Katalupaik, 110 km north-west of the type locality (Fig. 12B). These females range from 4.1 to 5.9 mm in total length. Based on these new specimens, the variability of color pattern of A. burwelli is: pronotum from completely yellow to yellow with the dorsal half brown; median lobe of mesoscutum from orange with a basal and transverse black stripe to orange with the basal half almost completely black; lateral lobes of mesoscutum from black to orange with a median black stripe; scutellum from black to black with two lateral orange spots; coxa III from yellow with base black to almost completely black with a variable amount of orange anteriorly and posteriorly; metasoma from largely orange with variable amount of black, especially dorsally, to almost black with some orange laterally; wings hyaline with the apex slightly infuscated; venation brown with pterostigma lighter medially.

Diagnosis
Light orange species with few black or dark brown markings on head, mesoscutum and metasoma. Head 1.35 × as wide as long; antenna with scape 1.58 × as long as pedicel; F-II more than twice as long as F-I. Dorsal face of propodeum in lateral view strongly concave basally then convex, with single transversal carina on convexity.

Etymology
The specifi c epithet refers to the fi rst name of Thibault Ramage's son, Elven. The species epithet is to be treated as a noun in the genitive case.

Female holotype C
. Light orange species; mandibles apex, pedicels and fl agellae, large spot around ocelli reaching dorsal margin of eyes, transversal stripe on posterior margin of mesoscutum and hind tarsomeres black; apical ⅔ rd of hind tibiae, sub-basal (sometimes absent) and sub-apical rounded spot on T-I, median rounded spot on T-II to T-VI and ovipositor brown. Fore wing with membrane yellow basally, hyaline in its apical half and infuscated apically; veins brown. Hind wing with membrane yellow basally, hyaline in its apical half and slightly infuscated apically; veins brown. H (Fig. 4A, C). In full-face view, 1.35 × as wide as long, vertex convex; mandibles tridentate; clypeus slightly convex, with small blunt projected tooth medially; malar space as long as scape, margin sinuate; antenna 14-segmented, scape 1.58 × as long as wide, 1.58 × as long as pedicel, and as long as F-I, F-I slightly longer than F-II, both more than twice as long as wide; fl agellomeres III-XI shortening gradually toward apex; apical fl agellomere slightly longer than preceding one; frons without lateral medial carina above torulus; vertex with ocelli arranged in small isosceles triangle, mid-ocellus separated from lateral ocelli by its diameter, lateral ocelli separated by nearly 3.00 × their diameter and by twice their diameter from eye margin; posterior margin of head conspicuously concave in dorsal view; occipital carina present. M (Fig. 4B). Pronotum without angular process; mesoscutum in lateral view slightly angular anterodorsally, notauli conspicuous and parapsidal lines present; small transverse depression between axillae, mesoscutum and scutellum; metapostnotum narrow; mesepimeron elongate; dorsal face of propodeum in lateral view strongly concave basally becoming convex distally, with transversal carina separating medially dorsal convex and concave surface, and carina surrounding metasomal insertion directed downward to hind coxa, posterior face of propodeum slightly concave under metasomal insertion; protrochantellus present; tibial spurs formula 1-2-2, protibial spurs bifi d apically, with midand hind-tibial spurs equal and similar in shape; hind coxa with ovipositor guide on inner surface, somewhat distal, transverse and somewhat oblique; hind femur 0.63 × as long as hind tibia; TS 1 2.80 × as long as TS 2; TS 2 1.60 × as long as TS 3; TS 3 2.50 × as long as TS 4; TS 4 0.50 × as long as TS 5; hind tarsal claw 0.50 × as long as TS 5; tarsal claw with two sub-apical teeth equidistant and equal in length, apical tooth as long as preceding ones.  4D). Elongate, longer than mesosoma; T-I more than 3.00 × as long as T-II; T-II to T-VI of similar length.

S
. Head, pronotum, sides of mesosoma densely puncticulate (except small transverse depression between axillae and anterior median triangle on mesoscutum). Mesoscutum (except anterior median triangle and posterior margin), axillae, scutellum with transversal ridges. Lateral depression of metanotum, metapostnotum with longitudinal ridges. Propodeum basally and medially with 3 or 4 longitudinal ridges, laterally with 3 ridges directed toward metasomal insertion and under propodeal declivity with longitudinal and oblique ridges.

P
. Small, erect, dense on all the body, sparse and less conspicuous on metasoma, absent on mesoscutum anterior median triangle and T-I and T-II.

Distribution
It is known only from its type locality, Katalupaik (Fig. 12A).

Remarks
Pristaulacus elveni sp. nov. can easily be distinguished from P. villemantae sp. nov. by its yellow coloration, the stouter mesosoma, and the dorsal face of propodeum more concave basally and with the presence of a single transversal ridge on posterior half (dorsal face of propodeum shallowly concave basally and with several transverse ridges on posterior half in P. villemantae sp. nov.).

Etymology
The specifi c epithet is a patronym honoring Dr Claire Villemant (MNHN), who greatly contributes to the knowledge of Hymenoptera, especially of Ichneumonidae. The species epithet is to be treated as a noun in the genitive case. Female holotype C . Mainly black species; base of mandibles, fore and mid legs, hind protrochantellus and base of tibiae, metasomal spiracle, lateral small transverse stripe at T-I mid-length, ovipositor dark-orange. Fore wing with membrane slightly yellow basally, hyaline in its apical half conspicuously infuscated apically; veins black. Hind wing with membrane hyaline; veins black.  (Fig. 5A, C). In full-face view, 1.20 × as wide as long, vertex slightly convex; mandibles tridentate; clypeus slightly convex, with small blunt projected tooth medially; malar space as long as scape, margin sinuate; antenna 14-segmented, scape about 1.30 × as long as wide, 2 × as long as pedicel; F-I 1.26 × as long as scape, F-II twice as long as F-I, both more than twice as long as wide; fl agellomeres III-XI shortening gradually toward apex; apical fl agellomere slightly longer than preceding one; frons without lateral medial carina above torulus; vertex with ocelli arranged in isosceles triangle, mid-ocellus separated from lateral ocelli by slightly less its diameter, lateral ocelli separated by slightly more than their diameter and by 1.50 × their diameter from eye margin; posterior margin of head slightly concave in dorsal view; occipital carina present. M (Fig. 5B). Propleuron ventrolateral carina present; pronotum without angular process; mesoscutum in lateral view slightly angular anterodorsally, notauli conspicuous and parapsidal lines present; small transverse depression between axillae, mesoscutum and scutellum; metapostnotum wide; mesepimeron elongate; propodeum in lateral view shallowly concave basally becoming convex distally, with transversal carina separating medially dorsal convex surface and carina surrounding metasomal insertion and directed downward to hind coxa, posterior face of propodeum concave under metasomal insertion; protrochantellus present; tibial spurs formula 1-2-2, protibial spurs bifi d apically, with midand hind-tibial spurs equal and similar in shaped; hind coxa with ovipositor guide on inner surface, somewhat distal, transverse and somewhat oblique; hind femur 0.65 × as long as hind tibia; TS 1 2.88 × as long as TS 2; TS 2 1.80 × as long as TS 3; TS 3 2.50 × as long as TS 4; TS 4 0.40 × as long as TS 5; hind tarsal claw 0.40 × as long as TS 5; tarsal claw with two equidistant sub-apical teeth, basal tooth shorter than second one; apical tooth longer than preceding ones. F (Fig. 5E). Vein 2-Rs + M very short; second discal cell elongate; vein 2r-m almost absent except for slight node on medial vein; vein 3r-m tubular in anterior ⁄ th and posterior ⁄ th , remainder nebulous. H (Fig. 5E). With 2 hamuli, venation reduced in Sc + R, r-m, M + Cu, A; beginning of Cu and 2-M present; Sc + R and 2-M tubular, others nebulous. M (Fig. 5D). Elongate, longer than mesosoma; T-I more than 3.00 × as long as T-II; T-II to T-VI of similar length.

S
. Head, mesosoma (except small transverse depression between axillae and anterior median triangle on mesoscutum) densely and slightly puncticulate. Frons and vertex with shallow ridges. Mesoscutum (except anterior median triangle and posterior margin), axillae, scutellum, dorsum of propodeum, dorsal face of mid and hind coxae with transversal ridges. Lateral depression of metanotum with longitudinal ridges. Metapostnotum, lateral and posterior face of propodeum reticulate. Tergite of metasoma micro-reticulate, reticulation becoming stronger toward apex.

P
. Small, erect, dense on all the body, sparse and less conspicuous on metasoma, absent on mesoscutum anterior median triangle and T-I and T-II.

Distribution
It is known only from its type locality, Katalupaik (Fig. 12A).

Remarks
Pristaulacus villemantae sp. nov. can easily be distinguished from P. elveni sp. nov. by its black coloration, the more elongate mesosoma, and the dorsal face of propodeum less concave basally and with the presence of several transversal ridges on the posterior half (dorsal face of propodeum strongly concave basally and with a single transverse ridge on the posterior half in P. elveni sp. nov.).   Balhoff et al. 2013: appendix 2).

Remarks
This female agrees with the description of S. deercreeki except for its size (7.1 mm vs 5.3-6.6 mm for the types) and the presence of the median carina of lower face. We are reluctant to describe a new species based only on the latter criteria, additional specimens from various parts of New Caledonia as well as DNA barcoding may solve this problem.

Distribution
Species known only from New Caledonia. See additional data in Balhoff et al. (2013: appendix 2).

Remarks
Except for their body lengths (6.2 and 7.8 mm vs 6.5-6.8 mm for the types), these two females agree with the description of S. irwini.

Remarks
The unidentifi ed males of Szepligetella collected during the 2016 and 2017 "Our Planet Reviewed" expeditions are listed here. Based on the identifi cation key and diagnoses in Balhoff et al. (2013) these males belong to either S. irwini or S. levipetiolata. The only criteria to separate the males of these species is the presence or absence of long setae. The males examined show a great variability in head pilosity, as well as size (BL from 3.9 to 7.4 mm) or coloration of front leg (from totally black to tibia and tarsomeres orange). We prefer here to not assign these males to either S. irwini or S. levipetiolata. The use of DNA barcoding would be of great help to determine males of Szepligetella and redefi ne the species delimitations.

Diagnosis
Mainly black species, large sub-basal ring on hind tibia and large sub-apical ring on ovipositor white. Head somewhat elongate, 1.13 × as long as wide; scape 1.77 × as long as pedicel; F-I 1.30 × as long as scape, F-II 0.90 × as long as F-I. Hind coxa punctulate, 3.67 × as long as wide. Fore wing with fi rst discal cell absent.

Etymology
The specifi c epithet is a patronym honoring Dr John Jennings (University of Adelaide), who greatly contributes to the knowledge of Hymenoptera, including those of New Caledonia, and is to be treated as a noun in the genitive case.  . Mainly black species; coxa I and tibia III dark brown; femur I brown; fore leg trochanter, base and apex of femur, inner face of tibia, TS 4-5, mid leg inner face of tibia, ovipositor orange-brown; mandible (except for apex) and mouthparts yellowish; fore leg base, outer face and apex of tibia, TS 1-3, mid leg base and apex of tibia, TS 1 (except for apex), large sub-basal ring on hind leg tibia, apex of sheath, large sub-apical ring on ovipositor white. Fore and hind wing with membrane hyaline and black veins. H ( Fig. 8A-C). In full-face view, 1.13 × as long as wide, frontal carina very shallow, vertex strongly convex; mandibles bidentate; clypeus strongly sinuate; malar space ⁄ th the width of the pedicel; antenna 14-segmented, scape about 1.90 × as long as wide, 1.77 × as long as pedicel; F-I 1.30 × as long as scape, F-II 0.90 × as long as F-I; fl agellomeres III-XI shortening gradually toward apex; apical fl agellomere slightly longer than preceding one; vertex with ocelli arranged in isosceles triangle, mid-ocellus separated from lateral ocelli by ⁄ th its diameter, lateral ocelli separated by slightly less than their diameter and by 1.73 × their diameter from eye margin; occipital carina narrow.

P
. White, minute, oblique and very dense on all body, longer on mandibules, clypeus, lower face and underside of head, propleuron, pronotum, meso-and metapleuron, upper part of propodeum and coxa I.

Distribution
It is known only from its type locality, Ouinné (Fig. 12C).

Remarks
Gasteruption jenningsi sp. nov. can be distinguished from the three species of Gasteruption from New Caledonia, G. lacoulee Jennings, Krogmann & Parslow, 2015, G. maquis Jennings, Krogmann & Parslow, 2015and G. sarramea Jennings, Krogmann & Parslow, 2015, by the absence of the fi rst discal cell of the fore wing, its elongated hind coxa, and the presence of a white large sub-basal ring on the hind tibia and a large sub-apical ring on the ovipositor.
The absence of the fi rst discal cell of the fore wing is a rare condition in the species of Gasteruption, and reported from G. subhamatum Pasteels, 1958(Zhao et al. 2012) known from Borneo and China and from G. tomanivi Parslow, Stevens & Schwarz, 2018from Fiji (Parslow et al. 2018). Among these two species of Gasteruption, G. jenningsi sp. nov. seems more related to G. tomanivi with which it shares the presence of white rings on the hind tibia and the ovipositor, as well as the black TS 1 on the hind leg. However, G. jenningsi sp. nov. can be distinguished from G. tomanivi in particular by the coloration, and the shape of the propleuron and the hind coxa.

Remarks
Previously, G. maquis was known only from the holotype male collected at the Pic du Grand Kaori. During the 2016 "Our Planet Reviewed" expedition, a female was collected at Ouinné, 40 km northwest of the type locality. This female diff ers from the type by: hind tibia with a sub-basal small cream ring and its size (BL: 17.0 mm vs 25.7 mm for the holotype). As the female of G. maquis was not known, an additional character for species diagnosis is the ovipositor orange, and sheaths black with tip white. We believe this female belongs to G. maquis and the diff erences identifi ed above to be intra-specifi c variations or sexual dimorphism.

Remarks
The eight specimens from Aoupinié diff er from Pseudofoenus ritae (Cheesman, 1936) by the presence of a frontal carina and of two medial teeth on the mandibles, and agree with the description and measurements of P. caledonicus. However, they diff er from P. caledonicus in their size and coloration, these specimens being smaller (females: 7.4-8.3 mm (vs 8.7-10.0 mm for caledonicus); males: 7.9-8.5 mm (vs 10.5 mm for caledonicus)) and brown with reddish parts (vs dark brown for caledonicus).

Distribution
Species known only from New Caledonia (Fig. 12F). See additional detail in Aguiar & Jennings (2005: 10).  Jennings, Austin & Stevens, 2004; red circle = type locality; blue circle = specimens concerned by this study. C. Distribution of Gasteruption jenningsi sp. nov.; red circle = type locality; blue circle = specimens concerned by this study. D. Distribution of Gasteruption maquis Jennings, Krogmann & Parslow, 2015; red circle = type locality; blue circle = specimens concerned by this study. E. Distribution of Pseudofoenus caledonicus Jennings & Austin, 2005; red circles = localities previously published; blue square = specimens concerned by this study. F. Distribution of Parastephanus mouensis Aguiar, 2005; red circle = type locality; blue circle = specimens concerned by this study.