ore

JHR 57: I-87 (2017) JOURNAL OF *0eerevowed opevaccers ural
doi: 10.3897/jhr.57. 12892 (MEE Hymenopter a

http://jhr.pensoft.net The Inerational Society of Hymenoptersts. RESEARCH

Eustochomorpha Girault, Neotriadomerus gen. n., and
Proarescon gen. n. (Hymenoptera, Mymaridae),
early extant lineages in evolution of the family

John T. Huber'

| Natural Resources Canada, clo Canadian National Collection of Insects, AAFC, K.W. Neatby Building, 960
Carling Avenue, Ottawa, ON, KIA 0C6, Canada

Corresponding author: John T: Huber (john.huber@agr.gc.ca)

Academic editor: Petr Jansta | Received 23 March 2017 | Accepted 8 May 2017 | Published 30 June 2017
http://zoobank.org/6201 DACE-9900-4A2F-92C9-D3014851100D

Citation: Huber JT (2017) Eustochomorpha Girault, Neotriadomerus gen. n., and Proarescon gen. n. (Hymenoptera,
Mymaridae), early extant lineages in evolution of the family. Journal of Hymenoptera Research 57: 1-87. https://doi.
org/10.3897/jhr.57.12892

Abstract

Eustochomorpha Girault, with one described species, E. haeckeli Girault, from Australia is redescribed.
Neotriadomerus Huber, gen. n., is described, together with seven new species, all from Australia: NV. bur-
welli Huber, sp. n., NV. crassus Huber, sp. n., NV. darlingi Huber, sp. n., NV. gloriosus Huber, sp. n., NV. lon-
giovipositor Huber, sp. n., NV. longissimus Huber, sp. n. (one of the largest species of Mymaridae), and WN.
powerae Huber, sp. n. Proarescon Huber, gen. n., is described for P primitivum (Huber), comb. n., trans-
ferred from Borneomymar Huber, and P similis Huber, sp. n., from Thailand. The previously unknown
male of Borneomymar madagascar Huber is described and the genus is redescribed from critical point dried
and slide mounted specimens. Triadomerini, stat. n., is proposed to include six genera: Borneomymar, Eu-
stochomorpha and Neotriadomerus, and the Cretaceous Carpenteriana Yoshimoto, Macalpinia Yoshimoto
and Triadomerus Yoshimoto. Aresconini is proposed to include five (possibly six) genera: Arescon Enock,
Kikiki Huber and Beardsley, Proarescon Huber and Tinkerbella Huber and Noyes, and the Cretaceous
Myanmymar Huber and, tentatively, also Enneagmus Yoshimoto. ‘The two tribes are proposed as being the
earliest lineages in Mymaridae, with Neotriadomerus and Triadomerus being sister genera to the remaining

extant and extinct genera, respectively.

Keywords
Taxonomy, Chalcidoidea, fairyfly, new genera, Australia, Thailand, earliest lineages

Copyright John T. Huber. 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.

2 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Table of contents

TiNGhOGLU GtiO tlt. due acc ccseaceg cele lee die ratte teat Rese att out Pann lane aul Dns sn lemnaulh wees Aes. 2

PSEC gs attested: ahacenco china Aha phen My ladle Pee Akal t Mee ty: diel e tase Ate EN tot 3

PRS UTiEG: nn cose seek ancatetervcctstmnneatticet hee boat tehde eit ae Geel Mat ot A ae ee he POS De 4

EUStOCMOMON pa Raita. IVS ak. Bove sane teenage wcest teecateE pus sceta= oo pa:seaiee vnnutdlg ooaeiean dene 4

EWGOVOMOLP IDA CHL GIRANU aS wcciviny iu SintetaElp cate dus eendip dah Ounelesrvnsae Sonedog ens 6

IN GOLT AONE VIDOE SO CI. Wsrecotle axe nist nehatoaiarnbisticlace chon sdetsnsdocentornttin ebesiche 14

INGOT TAA ONICTUS-DUTWCI MAUDEL, SP Ie auccdew A 2c sux sasepmencesnveons Pav eape reseed steed oevees 25

INERTIA OMErUS. Crassus TIUDEL, SSP: Als.i.0eixeessesscasestee cagncal. sagen eaten 2iystees chases. 28

I NCOLTIMAAOMETURAATIINGE MUD EIS > Waist. mtr anchsHesla ott ei bMos cased aeatle. 32

NCOP IAAONIEN US GIOLIOSU OMA DEL SPY. Txpucessiiantaneonesensontaerbunes yt ovetinn wt uenictixs 40

Neotriadomerus longiovipositor Huber, sp. 1. ......scssesseeseceeeseeseseseeseeeseeseens 43

Neotriadomerus longissimus Huber, sp. 0. .....eescssesssessessecsesseceseseseeseseaeeseees 51

INCOLTIMLONIENUS DOWENACR MAD EIS ES D2 IN wgeue sxe tuesank Wes eoonee suoeteaapovesinneveeteniel te 53

INGO ETI OMERUS SD 2 V seuuu cia cnvtonsiy tere enc vetidah Moa Ne teeter erceete ae Ene tneenaeees 60

TN CO TVTABONICRUESD A Da nats cases eiSoad souract nl ORs pts cate Btathog pire d Me ipa sons eh eR pene eeht te 60

EN LOGIC TOV USSD SAL alts alas Gan OM it tes ds Spe Mie dean oi ach lia bonleoys fle 60

IN COEPIAAOTIETUS SDs SR te tae cen 4 on At ie ae oars ge cece a Gack omeazes 62

PRO MPCCOMATUDEL: POM tai asta tense ter tind Peete cametaede Meteo coat seen du cabo boruios taentats 66

P FOGFENCON PIIMBIE VIS TEL LIDEL Vy COMAD: Me desea cosnnsicseniceebevaadienieandtedebanletosbalta’ 68

PPO ATERCOTT SUITEZ LESAN CG BIS daar galrsatow ecco Be pap odes dat teen's de Bae chops taaetaao 69

DOPRLOR NIT NDAD Cth acct eR unss Boon vitesse MANaM a ATA olLdesupdacelagtacsTrotdgius vhae ray We

Borneomymar macdgascar TUDE... .ccbsctabvssceasnncdoedsboveshbonsesoacnoeronntisnnnnssntas 81

ribs litac Oreri ny Staten. § <meta, noe. kaka Pees eRe eatin a eee 82

See Pe SCO Tt a: 8 See «or Pee on ES cert Rt Nee, Rc On eM re Ol cee 83

APS CUSS ONE hh tere ee ae ee coche ee atten bot, Stee tee aeleea st eciaeth Me eres. nett 84

ABOUTS On seen. Pecan tare aes et eee ee Ree eee i So et oes be. 85

ACK OWlEd Gene ES evs: sat. sgncses Aseatassng tavssson RamesUTe Raat one Tee CURE A cig esata ateatnah decades 85

PRETENCE Srey ee enn Nek Peet ft ce Mick A ey maces ai Noe Mates seme Cys ce hi Mr Ruane Aaa oe 86
Introduction

Eustochomorpha Girault (Hymenoptera: Mymaridae) includes a single species, E. hae-
ckeli Girault (Girault 1915). Annecke and Doutt (1961) included Eustochomorpha in
a world key, basing its placement in the key only on Girault'S, succinct but accurate
original description; they did not locate the type. Dahms (1984) provided notes on
the holotype and only known specimen but otherwise the genus remained unknown.
Based mainly on two specimens collected in Western Australia Eustochomorpha is re-
described. Specimens of an undescribed genus, apparently related to Eustochomorpha,
were found in various collections and are described. A third genus is proposed for Bor-
neomymar primitivum Huber (Huber 2002). A key is given to these genera and their

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... a

included species. Several other genera are included because they may well be closely
related. An attempt to place the Cretaceous fossils of Mymaridae in context with the
extant fauna is made.

Methods

This study is based on about 35 specimens on card-mounts, point-mounts and slides.
Slides of the specimens were prepared as described in Huber (2015). Morphological
terms used in the descriptions are mostly as defined in Gibson (1997) and Huber
(2012, 2015). All measurements of specimens are given in micrometers and, depend-
ing on specimen length and preparation, taken with an ocular micrometer attached to
a Nikon compound microscope at 200x magnification or a Leitz-Wetzlar binocular
microscope at 64x and 160x magnification. Because of rounding errors, it appears
that the ratios are slightly incorrect compared to those calculated using the absolute
measurements (micrometers) but, in fact, those calculated from the ocular microm-
eter readings before converting to micrometers are more accurate, so are given in the
descriptions. Photographs of slide mounts were taken with a ProgRes C14?"* digi-
tal camera attached to a Nikon Eclipse E800 compound microscope, and a selection
of the resulting layers combined electronically in Zerene Stacker™. Card- or point
mounted specimens were photographed with a Leica DFC 500 camera mounted on a
Leica Z16 APO Zoom macroscope or on the Nikon compound microscope. Abbre-
viations used in the descriptions are: fl for funicle segment, gt for gastral tergum and
mps for multiporous plate sensilla. Because the number and position of setae and the
type of sculpture on species within a given genus of Mymaridae is usually quite similar
the sculpture and setation are described in detail under the single species in Eustocho-
morpha, the single new species under one of the new genera and, to avoid repetition,
under the genus name for the new genus containing seven new species. The number
and position of setae is described for one side of the body only. So when observing a
specimen in dorsal view the total number of setae on a particular structure is twice
that described. The setae of appendages (antenna, mouthparts, wings, legs, genitalia)
are usually excluded because they are numerous and often similar; only obviously dif-
ferent ones are mentioned, where informative. In general, the position of body setae,
particularly on the mesosoma, is fairly stable and consistent within and even among
genera, though setal shape, length and thickness may vary. Specimens are deposited in
the following institutions:

ANIC Australian National Insect Collection, Canberra, Australian Capital Terri-
tory, Australia;

CAS California Academy of Sciences, San Francisco, California, USA;

CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa,
Ontario, Canada;

NZAC New Zealand Arthropod Collection, Auckland, New Zealand;

4 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

QMBA Queensland Museum, Queensland, Brisbane, Queensland, Australia;

QDPC Queensland Primary Industries Insect Collection, Brisbane, Queensland,
Australia;

QSBG =e Queen Sirikit Botanic Garden, Chiang Mai, Chiang Mai, Thailand;

ROM _ Royal Ontario Museum, Toronto, Ontario, Canada;

UCRC Entomology Research Museum, University of California, Riverside, Cali-
fornia, USA.

Results

Eustochomorpha Girault, 1915
Figs 1-31

Type species. Eustochomorpha haeckeli Girault, by monotypy.

Diagnosis. Female. Antenna with funicle 8-segmented and clava 2-segmented
(Figs 3, 7, 14). Venation about 0.9x fore wing length. Postmarginal vein distinct, uni-
formly thick and about 0.7x marginal vein length, with at least 2 short, socketed setae
(Fig. 8, apical one indicated by arrow, Fig. 20c). Hypochaeta possibly present (Fig.
20a), originating on the parastigma instead of on the costal cell. Ovipositor strongly
extending posteriorly beyond gastral apex but not extending anteriorly under meso-
soma (Figs 2, 4, 28, 30).

Description. Female. Body ~1280-1290 um in length, excluding section of
ovipositor exserted beyond apex of hypopygium. Colour. Generally brown (Fig. 4),
with a few metallic reflections (an artefact of lighting?) on gaster. Gaster at base and
most parts of legs lighter brown to white. Wings hyaline, with venation light brown.
Head. Head about 1.7x as wide as long, 1.3x as wide as high and 1.4x as high as long; in
lateral view with anterior surface convex, receding strongly from ventral margin of eye
to mouth opening, and bulging medially compared to laterally (at preorbital groove);
posterior surface slightly, evenly convex (Fig. 4). Face about as wide as high (Fig. 5),
without subantennal grooves; preorbital groove ventral to torulus straight then, more
ventrally, slightly curving medially to ventrolateral margin of mouth opening. Torulus
in slight triangular depression about 2.0x as high as torulus width and separated by
1.5x torulus width from transverse trabecula (Fig. 5). Vertex in lateral view (Fig. 11)
sloping obliquely to transverse trabecula to form an obtuse angle with face, and vertex
posteriorly merging smoothly into occiput medially but separated laterally from occi-
put by short transverse vertexal suture. Ocellar triangle raised slightly above level of rest
of vertex, the mid ocellus almost vertical, the lateral ocelli oblique, facing posterolat-
erally; ocelli with POL about 2.0x LOL and about 1.26 OOL. Transverse trabecula
separated at each apex from supraorbital trabecula (Fig. 5); preorbital trabecula (ap-
parently absent because not sclerotized) short, extending to inner orbit at about dorsal
margin of torulus (Fig. 5); supraorbital trabecula in 2 subequal sections, the posterior
sections slightly converging and the sutures outside posterior sections continuing onto

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 5

occiput as the occipital grooves (short lines each appearing as posterior extension of
supraorbital suture and strongly converging towards occipital foramen) (Fig. 6) separ-
ating occiput medially from temple dorsolaterally. Occiput separated from vertex by
vertexal suture, this only present laterally behind lateral ocelli (Fig. 12). Eye large with
numerous (about 100) small facets, in lateral view slightly higher than wide (10:9)
and clearly separated dorsally from back of head (temple about 0.3x eye width). Ocu-
lar apodeme fairly long, crooked, needle-like. Malar sulcus absent (Fig. 11). Gena at
level of ventral margin of eye wider than malar space. Occiput entire, not separated
by transverse groove into upper and lower portions (Fig. 6a). Mouthparts. Labrum
with 2? setae; mandible with 3 teeth. Antenna. Scape about 2.6x as long as wide, with
radicle distinct from rest of scape and about 0.25x total scape length (Fig. 13); pedi-
cel about 2.1x as long as wide, almost as wide and about 0.7x as long as entire scape;
funicle 8-segmented; clava 2-segmented (Figs 3, 7, 14), 1.08x as wide as apical funicle
segment and 0.32—0.36x as long as entire funicle. Mesosoma. Mesosoma about 2.2x
as long as wide, 2.3x as long as high and 1.1x as wide as high. Pronotum entire (Figs 9,
12, 15), in dorsal view clearly visible, medially about 0.3x as long as mesoscutum; col-
lar almost horizontal, with a shallow median longitudinal impression and prominent
angular shoulders; neck shorter than collar, posteriorly strongly sloping up to collar; in
lateral view, pronotum rectangular, slightly overlapping anterior margin of mesoscu-
tum, with lateral surface merging smoothly into dorsal surface, with a shallow, oblique
impression for femur. Spiracle (Figs 17, 18) apparently slightly stalked, at posterolat-
eral angle of pronotum in a slight depression and facing dorsolaterally. Propleura near
anterior apex not quite abutting, then gap widening slightly more anteriorly. Proster-
num rhomboidal, incompletely divided medially by faint longitudinal groove poster-
iorly. Mesoscutum about 1.7x as long as scutellum, in dorsal view with shallow, thin,
slightly diverging notauli apparently absent posteriorly (Figs 9, 15, 16), in lateral view
mesoscutum (Fig. 4) almost flat except anteriorly. Scutellum slightly wider than long;
anterior scutellum about 3x as long as narrowly trapezoidal, transverse frenum and
separated from it by a shallow, curved frenal depression (Figs 9, 16); campaniform
sensilla small, slightly closer to lateral margins of anterior scutellum than to each other,
with apex of large, cone-shaped fenestra projecting anteriorly between them (Fig. 9).
Axilla slightly advanced, the transscutal articulation laterally forming an obtuse an-
gle with median section (Fig. 16); axillula long and narrow, separated from anterior
scutellum by posteriorly recurved axillular groove; mesophragma widely convex pos-
teriorly, extending to posterior apex of propodeum. Prepectus probably spindle-shaped
(Fig. 17, partly hidden by fore leg); mesopleuron somewhat rectangular, with shallow
depression separating mesepisternum from mesepimeron (Fig. 17). Metanotum with
distinct lens-shaped dorsellum (only visible Fig. 9, a cleared, slide mount) and lateral
panel length (toward hind wing articulation) about half length of dorsellum (Fig. 16).
Metapleuron triangular, with a narrow anterior extension almost to base of hind wing,
the margin at junction with mesopleuron deeply concave and posterior margin straight
and vertical. Propodeum with a complete longitudinal groove medially, in lateral view
weakly sloping, almost in same plane as dorsellum, in dorsal view with faint median

6 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

longitudinal groove. Propodeal spiracle facing dorsolaterally in a slight depression and
separated from anterior margin of propodeum by about its own diameter. Wings. Fore
wing (Figs 4, 8, 19) wide, with completely transparent membrane sparsely scattered
with minute microtrichia. Venation complete; submarginal vein with 1 subbasal seta
and about 16 bullae, the most apical one at level of proximal macrochaeta (Fig. 8);
parastigma, 0.5x submarginal vein length; marginal vein present, its length (from distal
macrochaeta) about 2.0x length of parastigma; stigmal vein short but distinct, curving
away from wing margin (Fig. 20) then closely paralleling it, with 4 apical campaniform
sensilla (Fig. 21b); postmarginal vein present, about 0.8x as long and as thick as mar-
ginal vein, with 4 microchaetae along anterior margin; hypochaeta absent (Fig. 8) but
one similar to it originating on parastigma (Fig. 20a); proximal campaniform sensillum
near posterior margin of marginal vein just beyond apical macrochaeta (far removed
from the 4 campaniform sensilla on stigmal vein). Hind wing (Fig. 8) with membrane
not extending to base of wing, relatively wide medially, tapering gradually almost to
a point apically. Fore wing membrane with one irregular row of microtrichiae basally
and two or three distally. Legs. Profemur and metafemur narrow (Fig. 17), metafemur
widest, about 2x as wide as mesofemur. Tarsi 5-segmented. Calcar (moveable protibial
spur) apparently without setae along outer margin, and with the inner tine about 0.5x
as long as outer tine. Middle and hind legs with tarsomere 1 shorter than tarsomere
2. Metasoma. About 2.2x as long as wide, 2.3x as long as high and 0.9x as wide as
high; its length, excluding exserted part of ovipositor, about 2.0x that of mesosoma
(Fig. 4). Petiole ring-like, about 0.3x as long as wide. Gastral terga apparently with
fine longitudinal wrinkles except gt; gt, shorter than remaining terga and apparently
et, the longest tergum, gt, apically acutely triangular. Cercus a distinctly raised, longer
than wide, peg-like plate, with 4 setae (Figs 22, 25). Gt, with spiracle minute (appar-
ently absent). Hypopygium extending posteriorly well beyond level of cercus (Figs 4,
23, 25). Ovipositor sheath exserted beyond gastral apex by about one third of its total
length and abruptly upturned apically, apparently without subapical seta (Figs 22, 24).

Male. Unknown.

Hosts and habitat. Hosts are unknown. The holotype was collected in forest

(Girault 1915).

Eustochomorpha haeckeli Girault
Figs 1-31

Eustochomorpha haeckeli Girault 1915: 156 (original description); Annecke and Doutt,
1961: 4, 14 (key, diagnosis); Dahms, 1984: 675 (holotype data); Lin et al., 2007:
33 (diagnosis in part, figs 129-131).

Type material. Holotype female (QMBA) on slide labelled as shown (Fig. 1). The
holotype (Fig. 2) is in uncleared in Canada balsam under 1 coverslip fragment be-
tween two other fragments and is poorly mounted more or less dorsoventrally with legs

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 7p

Figures 1-3. Eustochomorpha haeckeli, holotype female. | type slide, specimen under middle cover slip
2 body + legs 3 antenna. Scale bar = 100 um.

mostly folded up against body; left antenna with clava missing; right antenna (Fig. 3)
detached beyond pedicel and positioned next to left (very faint) hind wing; other hind
wing detached and its membranous portion hidden under head; right fore wing folded
over and partly hidden by legs. Except for estimated body length, measurements of the
holotype are not given in the species description below (my measurements were inac-
curate when I borrowed the specimen).

Diagnosis. Female. Among extant genera, E. /aeckeli is distinguished from the
first new genus described below, having a 2-segmented clava (3-segmented in the first
new genus), and the ovipositor strongly exserted posteriorly beyond apex of gaster and
not extending anteriorly under the mesosoma. It differs from the second new genus
described below by the postmarginal vein shorter than the marginal vein (postmar-
ginal vein longer than marginal vein in the second new genus), hypochaeta absent
(hypochaeta present in the second new genus), and it differs from Borneomymar by its
2-segmented clava (1-segmented in Borneomymar).

8 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

i
-
¥
: F
:
q

Figures 4-7. Eustochomorpha haeckeli, female. 4 habitus (Australia, WA, Stirling Range National Park)
5 head, anterior; 6a head, posterior 6b tentorium 7 antenna. Scale bar for 4 = 500 um; 5=7 = 100 pm.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 9

fA igdoale ”
cee ie o

2 F va
es So cae a 25)

ZLE ff [PPP

SASYYO yy SS ahs. =
anak hex ete vy a ay : ' We Ata fi sacsscie as . uhreN wh AOE Bhs

Ne

a ;
4

x

Wij

pe

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|

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fenestra a

Figures 8-10. Eustochomorpha haeckeli, female. 8, wings 9 mesosoma, dorsal 10 metasoma, dorsal. Scale
bar = 200 pm.

10 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Figures | 1-18. Eustochomorpha haeckeli, female. | | head + prothorax, lateral 12 head + prothorax, dorsal

13 base of antenna, lateral (right antenna) and ventral (left antenna) 14 apex of antenna 15 mesosoma,

dorsal 16 mesosoma, posterolaterodorsal 17 mesosoma, lateral 18 base of wings + mesosoma, lateral

(medial portion). Scale bar for 11, 12, 16 = 50 um; 13, 14, 18 = 20 um; 15, 17 = 100 um.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 11

, Sipe

distall

_

macroch aetay

distal aes
eerie proximal |

. f macrochaeta Ascent

/ | sl

— eS NS

a
WIT Giney — a :
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—~—_., Pryoorrisisizi- »

ovipositor

@

Figures 19-24. Eustochomorpha haeckeli, female. 19 wings 20a parastigma, ventral 20b base of mar-

ginal vein, ventral 20c apex of marginal vein, stigma + postmarginal vein, ventral 21a parastigma,
dorsal 21b stigma, dorsal 22 gaster apex showing cerci, posterolaterodorsal 23 gaster apex showing
cerci, dorsal 24 gaster apex, posterolaterodorsal. Scale bar for 19 = 200 um; 20a, 20b, 2b, 22-24 =
20 um; 20c, 21a = 50 um.

1 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

lower valve
Wien, A upper valve

ovipositor
sheath

SOS eee
a: <a

UPPCTAV AVE,

sheaths

/\

Po.
ji wy
fa r sz
BAA
:
=
> a

DZ

Figures 25-31. Eustochomorpha haeckeli, female. 25 gaster apex, lateral 26 ovipositor apex, dorsal
27 ovipositor apex, ventral 28 ovipositor + sheaths, lateral 29 apex of ovipositor + sheaths, lateral 30 ovi-
positor + sheaths, dorsal 31 apex of sheaths (one sheath folded over on itself). Scale bar for 25 = 50 um;
26, 27 = 5 um; 28, 30 = 100 um; 29, 31 = 20 um.

Description. Female. Body length ~1280 (critical point dried specimen, Stirling
Range National Park). Colour. Body mainly brown (Fig. 4). Ocellar triangle, mesothor-
ax and fine longitudinal line laterally on gaster darker brown. Base of gaster, especially
basal sterna, legs except metacoxa and metafemur and apical tarsomere of all legs, flagel-
lum and mouthparts except mandibles, and a faint transverse band at level of anterior
ocelli lighter brown to white. Head. Width 202. Face with weak reticulate sculpture me-
dially, stronger laterally (Fig. 5), with setae distributed as follows: 3 medial to torulus and
8 ventral to torulus, the 2 submedially above mouth margin the shortest. Vertex smooth

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 13

anterior to mid ocellus and reticulate posterior to mid ocellus (Fig. 5), with 2 setae on
anterior orbit lateral to transverse trabecula and 2 setae on dorsal orbit, in angle between
eye and posterior section of supraorbital trabecula; ocellar triangle with 2 short setae
lateral to mid ocellus and 1 short setae just posteromedial to lateral ocellus. Malar area
apparently with 3 setae. Gena and occiput laterally longitudinally reticulate (Fig. 6a),
the reticulations on occiput shallower and transverse medially; occiput with 2 short setae
submedially above occipital foramen, about 9 setae sublaterally and laterally; tentorium
with two very short dorsal arms (Fig. 6b). Antenna. Fl —fl, respectively with 0, 1, 2, 2, 2,
3, 3, 3 mps; clava 1 with 3 (4?) mps on first segment and 5 (6?) mps on second segment
(Figs 3, 7). Length/width measurements (n=1): scape 70/27, pedicel 55/26, fl, 33/18, f1,
32/20, fl, 39/25, fl, 40/25, fl, 44/25, fl, 42/25, fl 42/24, fl, 42/26, clava 112/31 (first
segment 42, second segment 69). Mesosoma. Width 200 and length 415. Pronotum
with reticulate sculpture, with 3 short setae along posterior margin and 3 towards anter-
ior margin. Propleuron faintly, longitudinally reticulate, with 2 seta about midway be-
tween anterior and posterior apices. Prosternum smooth, with 1 seta submedially almost
at anterior margin and 1 setae laterally midway between anterior and posterior margins.
Mesoscutum faintly longitudinally reticulate, with 1 setae along inner margin midway
between anterior and posterior apices of notaulus, and 1 setae at lateral angle of side lobe.
Anterior scutellum smooth, with 1 setae on lateral margins level with campaniform sen-
sillum; frenum faintly reticulate. Axilla reticulate, and with 1 seta at anteromedian angle
and 1 dorsally on longitudinally reticulate lateral panel; axillula smooth. Metanotum
smooth medially, faintly reticulate laterally, with 2 short setae along anterior margin of
relatively wide lateral panel. Propodeum smooth, with 2 propodeal setae close together.
Wings. Fore wing length (n=1) 723, width 259, length/width 2.98, longest marginal
setae 72. Marginal vein with about 11 microchaetae along its length; cubital line with
few setae extending to level of distal apex of retinaculum. Hind wing length 685, width
57, longest marginal setae 68. Legs. Femora and tibiae of all legs with faint longitudinal
reticulation. Protibia at mid length with 2 short pegs. Metasoma. Petiole 54 wide, 18
long. Gaster 212 wide, 597 long (to apex of apical tergum), with a few short setae on
most terga, the terga difficult to distinguish from one another (anterior and posterior
margins not distinct) and apparently with fine longitudinal wrinkles at least laterally; gt,
shorter than remaining terga, apparently with 1 lateral setae, gt, and gt, apparently with
1 submedian seta; gt, — gt, with about 4 submedian to sublateral setae in an irregular
transverse row; gt, acutely triangular, with a small apical seta. Cercus with the first two
cerci subequal in length, the third the longest and the fourth the shortest (Fig. 25). Hy-
popygium weakly sclerotized (almost transparent) dorsally and extending beyond gastral
apex by ~180, with a submedian and lateral row of about 8 setae. Ovipositor length
1074, its exserted part (posterior to hypopygium apex) 360.

Male. Unknown.

Material examined. Iwo females, collected in Malaise traps in combination with
yellow pan traps underneath. AUSTRALIA. Western Australia: Yanchep National
Park, 20—21.xii.1986, J.S. Noyes (12 on slide, CNC); Stirling Range National Park,
11-15.i.1987, J.S. Noyes (19 on point, BMNH).

14 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Neotriadomerus Huber, gen. n.
http://zoobank.org/5488307C-D58F-4E00-8 DC9-C2ACB542A103
Figs 32-152

Eustochomorpha: Lin et al., 2007: 33 (diagnosis in part, figs 132-134).

Type species. Neotriadomerus longiovipositor Huber, by present designation.

Diagnosis. Female. Antenna with funicle 8-segmented and clava 3-segmented
(Figs 39, 69). Venation about 0.85—0.90x fore wing length. Postmarginal vein distinct,
uniformly thick and about 1.3—1.6x marginal vein length (Figs 70, 122). Hypochaeta
present or, apparently, sometimes absent. Ovipositor strongly extending anteriorly un-
der mesosoma (Figs 59, 74) at least to level of hind wing base but not or only slightly
extending posteriorly beyond apex of gaster (Figs 75, 77). Male. Similar to female, but
flagellum with 11 relatively wide segments (Figs 40-43, 82, 95), each with several mps;
the segments progressively becoming narrower towards flagellar apex, fl,, sometimes
only a little over half as wide as fl,. Genitalia encapsulated, with thick walls (Figs 124,
143) and, in lateral view, paramere thick (Figs 65, 66).

Description. Female. Body 1380-5500 in length, excluding basal sac of gaster
(enclosing anterior extension of ovipositor). Colour. Generally brown (Figs 81, 128,
144, 146, 147) to dark brown, scape and pedicel, legs in part and sometimes basal sac
of gaster lighter brown to yellowish. Wings hyaline, the venation light brown. Head.
Head about 2.0—2.9x as wide as long, about 1.2—1.6x as wide as high and about 1.3-—
2.3x as high as long; in lateral view with anterior surface almost flat, at most barely re-
ceding ventral to eye, slightly depressed medially compared to laterally at preorbital
groove; posterior surface in lateral view slightly convex except slightly indented at pos-
torbital groove (Figs 33, 45). Face about 1.2—1.3x as wide as high (Fig. 32), in lateral
view almost flat, barely receding from ventral margin of eye to mouth opening; suban-
tennal grooves absent; preorbital groove ventral to level of torulus straight all the way to
lateral margin of mouth opening. Torulus in somewhat circular depression about 1.2x
as high as torulus width and separated by less than 0.4x torulus width from transverse
trabecula (Fig. 33). Vertex in lateral view usually almost in same plane as face, but
sometimes almost horizontal and forming a right angle with face, and vertex posteri-
orly forming an almost right angle with occiput and separated from it laterally by a
short transverse vertexal suture extending to or just medial to posterior ocellus but ab-
sent between posterior ocelli (Fig. 33). Ocellar triangle almost flat or raised slightly
above level of rest of vertex, the mid ocellus oblique to almost vertical, lateral ocelli
oblique, facing laterally or posterolaterally; ocelli with POL about 2.0x LOL and about
1.7-1.8x OOL (varying slightly with species); ocellar area (in cleared slide mounts)
with frontofacial suture (seen as white lines) between mid and lateral ocellus, frontal
suture (seen as white lines) from mid ocellus to corner of transverse trabecula, and
sometimes also a median longitudinal line from mid ocellus to middle of transverse
trabecula (Fig. 116). Transverse trabecula apparently separated (Fig. 67) or not (Figs 36,
37) from supraorbital trabecula; preorbital trabecula short, extending to about level of

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 15

ee! _—" - SSS
Figures 32-37. Neotriadomerus spp. 32 N. gloriosus female head, anterior 33 N. darlingi male head +
pronotum, dorsal 34 WN. gloriosus lower face + mouthparts, anterior 35 NV. gloriosus maxillae + labium,
anterior 36 NV. longissimus head, scape and mandibles, dorsoanterolateral 37 NV. longissimus radicle, ante-

rolateral (and showing 3-way junction of transverse, supraorbital and preorbital trabeculae). Scale bar for

32, 33, 36 = 100 um; 34 = 50 um; 35, 37 = 20 um.

dorsal margin of torulus; supraorbital trabecula in 2 equal or unequal sections, the an-
terior sections diverging posteriorly, the posterior, sometimes longer, sections slightly
converging, and the sutures outside posterior sections continuing onto occiput as short,
almost parallel lines, the occipital grooves, ventrally to level of foramen, thus separating
occiput medially from temple laterally (Fig. 68). Eye large with numerous small facets,

16 John T; Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

in lateral view at most about 1.4x as high as wide and clearly but narrowly separated
dorsally from back of head (temple in lateral view at most about 0.25x eye width).
Ocular apodeme short, straight, and thin (Fig. 119, inset). Malar sulcus absent. Gena
at level of ventral margin of eye at most as wide as length of malar space. Head posteri-
orly with curved postorbital groove extending across head above dorsal margin of oc-
cipital foramen to posterior margin of eye (Fig. 68) and separating occiput/temple
dorsally from gena/postgena ventrally. Mouthparts. Labrum with 4 or 6 setae; mandi-
ble with 4 uneven teeth, the ventral one the longest (Fig. 34). Antenna. Scape about
3.0—3.9x as long as wide, with radicle distinct from rest of scape and about 0.24—0.29x
total scape length; pedicel about 1.1—1.6x as long as wide, 0.72—1.09x as wide and
about 0.36—0.42x as long as entire scape; funicle 8-segmented (Fig. 69); clava 3-seg-
mented (Fig. 39), 1.07—1.22x as wide as apical funicle segment and 0.18—0.23x as long
as entire funicle. Mesosoma. About 1.8—2.4x as long as wide, 2.4—2.8x as long as high
and 1.0—1.3x as wide as high (card- or point-mounted specimens, air- or critical-point
dried). Pronotum entire, in dorsal view clearly visible, medially about 0.3—0.6x as long
as mesoscutum; collar almost horizontal, medially flat to slightly convex and without
evident shoulders (Fig. 44); neck not or barely differentiated from collar; in lateral view,
pronotum with lateral surface merging smoothly into dorsal surface, with a shallow,
oblique ventroanterior impression margined by an oblique groove posteriorly for recep-
tion of femur (when fore leg folded against body). Spiracle (Fig. 46) very slightly stalked,
at posterolateral angle of pronotum in a slight depression and facing posterodorsally.
Propleura near anterior apex not quite abutting then widening more anteriorly towards
the cervical sclerites (Fig. 141). Prosternum rhomboidal, almost completely divided
medially by longitudinal groove (Fig. 141). Mesoscutum about 1.1—1.6x as long as
scutellum, in dorsal view with narrow, distinctly diverging notauli (Fig. 44) appearing
in slide mounts wider and shallower near transscutal articulation (Fig. 72), in lateral
view almost flat except anteriorly (Figs 45, 81). Scutellum slightly longer than wide (at
widest point on frenum), the anterior scutellum narrower and slightly shorter to slight-
ly longer than transversely biconvex frenum and separated from it by an evenly curved
frenal line; campaniform sensilla as far apart from each other as to lateral margin of
anterior scutellum and slightly farther apart than their distance to transcutal articula-
tion, with apex of cone-shaped fenestra not extending to level of campaniform sensilla.
Axilla slightly advanced, the transscutal articulation laterally almost in line with median
section, and about as wide anteriorly as width of anterior scutellum at transcutal articu-
lation; axillar pit separated from anterior scutellum by curved axillular groove; meso-
phragma convex posteriorly, extending to posterior apex of propodeum (Fig. 72, faint-
ly visible under propodeum). Prepectus rather rectangular, at widest point about 0.8x
as wide as long. Mesopleuron somewhat spindle shaped, with shallow oblique depres-
sion separating mesepisternum from mesepimeron and almost straight groove extend-
ing from anterior margin to mesocoxal insertion (Fig. 92). Metanotum with distinct
biconvex (slide mount, Fig. 87) or somewhat triangular (SEM, Figs 44, 48) dorsellum
and lateral panel length at hind wing articulation about one-third to half length of
dorsellum. Metapleuron quadrangular, with almost straight sides (Figs 56, 123). Propo-

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 17

basal sac
/= of gaster

sppsilr im ae

Figures 38-43. Neotriadomerus spp. 38 N. longissimus female scape—base of fl, lateral 39 NV. sp. female
fl, + clava, lateral 40 NV. darlingi male apex of pedicel—base of fl,, lateral 41 N. darlingi male fl fh 3
lateral 42 N. darlingi male fia; lateral 43 NV. darlingi male apex of fl. lateral. Scale bar for 38 = 100 um;
39, 41 = 50 um; 40, 42, 43 = 20 um.

deum in lateral view weakly sloping, almost in same plane as dorsellum and in dorsal
view flat medially, without median or submedian grooves or other ornamentation (Figs
44, 46, 56, 72). Propodeal spiracle facing dorsolaterally, separated from anterior margin
of propodeum by much less than half its own diameter. Wings. Fore wing (Fig. 113)
about 2.6—3.7x as long as wide, evenly and densely covered with microtrichia from
wing apex proximally to at most about level of distal macrochaeta but almost without

18 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

humeral
plate

gu =

Figures 44-49. Neotriadomerus spp. 44 N. darlingi mesosoma, dorsal 45 NV. sp. mesosoma, dorsolateral
(arrows indicate pores) 46 J. sp. base of left wings (ventral) and surrounding mesosoma, dorsolateral
47 N. darlingi fore wing base, dorsal 48 N. longissimus fore wing base, ventral 49 NV. sp., hind wing at-
tachment to fore wing, ventral. Scale bar for 44, 45, 48 = 100 um; 46, 47, 49 = 50 um.

microtrichia behind submarginal vein (Figs 48, 113) and parastigma (Figs 50, 51) and
just behind much or all of marginal vein (Fig. 70); an oblique and usually distinct setal
line extending more or less horizontally from apical margin of wing to about halfway
towards base of marginal vein; cubital line ending proximally either at about same level
as remaining mictrotrichiae or extending to base of parastigma; marginal setae short,
the longest not more than about 0.13x wing width (Fig. 70). Venation complete (Fig.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 19

distal

proximal macrochaeta
macrochaeta y

peg-like
setae

ee COME” C™ tial
ee — ee comb
Yair / silejrer

\
basitarsal _ tibial
comb spur 53)

fi
Uf |

i HN / Ut
Figures 50-55. Neotriadomerus spp. 50 N. darlingi male parastigma, dorsal 51 N. longissimus female
parastigma, ventral 52 NV. darlingi male stigmal vein, dorsal 53 N. longissimus female calcar, lateral 54 N.

sp. female calcar, dorsolateral 55 UV. sp. female pretarsus, dorsolateral. Scale bar for 50, 53, 54 = 50 um;
51 = 100 um; 52,55 = 20 um.

122); submarginal vein with 1 proximal seta; parastigma (from distal macrochaeta to
base) about 0.5—0.7x submarginal vein length; marginal vein length (from distal mac-
rochaeta to junction of stigmal and postmarginal veins) about 1.2—1.5x parastigma
length; stigmal vein short but distinct, about 0.15—0.17x marginal vein length, curving
away from wing margin then closely paralleling it (Fig. 122), with 4 apical campani-
form sensilla (Fig. 52); postmarginal vein about 1.3—1.6x as long and as thick as mar-

20 John T; Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

ginal vein, with a few microchaetae along anterior margin; hypochaeta usually present,
occasionally apparently absent (broken off?), much closer to proximal than distal mac-
rochaeta; proximal campaniform sensillum near posterior margin of marginal vein just
apical to distal macrochaeta. Hind wing with membrane not extending to base of wing,
relatively wide medially, with a rounded apex, uniformly covered with microtrichia al-
most to base of membrane (Figs 49, 70). Legs. Profemur and mesofemur narrow;
metafemur widest, about 1.4x as wide as mesofemur (Figs 77, 92). Tarsi 5-segmented.
Calcar (moveable protibial spur) with setae along outer margin, and with inner tine
about 0.4x as long as outer tine (Figs 53, 54). Pretarsus normal (Fig. 55). Mid and hind
legs with tarsomere 1 at least as long as tarsomeres 2 + 3. Metasoma. Petiole usually
ring-like, about 0.4—0.5x as long as wide (Fig. 72) but in one (the largest) species 0.7—
1.0x as long as wide. Gaster 1.18—3.25x as long as mesosoma. Gaster with gt, and gt,
usually slightly the longest terga (Figs 56, 73, 89, 92). Cercus distinctly raied as a thin
flap above surface of gt, (Fig. 62) with 4 setae about equal in length (Fig. 65). Hy-
popygium distinct (Fig 56), extending about 0.3—0.6x length of gaster, sometimes as
far as level of spiracle. Ovipositor sheath barely projecting posteriorly beyond apex of
gaster but strongly projecting anteriorly (inside basal sac of gaster) from at least middle
of mesosoma (Fig. 74) to well in front of head (Fig. 126), apparently with 1 subapical
seta. Body sculpture and setation. Sculpture generally uniform and so faint that body
often appearing almost smooth and shiny (Figs 81, 82, 126-129, 144-147). Head with
engraved, slightly transverse (longitudinal on gena and postgena) or almost isodiamet-
ric reticulations becoming raised and slightly imbricate ventrally on face and on occiput
(Figs 32-34). Mesosoma with faint isodiametric to elongate reticulations either trans-
verse, mainly on pronotum, or longitudinal, mainly on scutellum. Metasoma appar-
ently without sculpture (Figs 56-62) except anterior apex of basal sac of gaster in at
least one species. Head with few to numerous setae on face ventral to toruli, and on
gena (Figs 32-34) but one (Fig. 36) or two (Fig. 32) just medial to torulus. Vertex with
2 setae, 1 between mid and lateral ocelli and 1 more laterally. Eye orbit with 3 setae
dorsoanteriorly and 3 dorsally and posterodorsally. Occiput with 1 submedial seta dor-
sally and, more ventrally, 1 seta sublaterally and 1 seta submedially. Gena/postgena with
several setae. Pronotum with about 3 setae along posterior margin and about 5 shorter
setae anteriorly. Propleuron with a few small scattered setae (Figs 44, 116). Prosternum
(Fig. 141) with 1-3 setae mainly in anterior half. Mesosoma with a few short setae (not
treated here) around insertions of wings and some longer setae as follows. Mesoscutum
with 1 adnotaular seta in posterior half of midlobe, 1 anteromedial and 1 posterolateral
seta on lateral lobe and a few small pits (Figs 44-46). Anterior scutellum with 1 seta at
lateral margin just anterior to axillular groove (represented by a curved medial edge at
junction with frenum). Axilla with 1 anteromedial seta and 1 lateral seta. Propodeum
with 2 or, occasionally, 3 propodeal setae laterally behind spiracle. Petiole apparently
with 1 minute seta ventrally on anterior margin. Gaster with setae on terga increasing
in number from gt, to gt, but apparently none on sterna except | or 2 short ones later-
ally on basal sac of gaster and longer ones on hypopygium (Fig. 56). Gt, with about 3
lateral and 1 dorsal in 1 row; gt, and gt, with a few more lateral and dorsal in 1 row;

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... ZA

A

nes

ovipositor
sheaths

hypopygium “"™

basal sac
of gaster

———_

Figures 56-60. Neotriadomerus spp. 56 N. sp. metasoma, lateral 57 NV. sp. metasoma, ventral 58 UN. sp.
apex of metasoma, ventral 59 NV. sp. mesosoma and ovipositor sac, ventral 60 JV. sp. apex of ovipositor

sac, lateral. Scale bar for 56, 57, 59 = 200 um; 58, 60 = 50 um.

gt,-gt, with even more in 2 or more irregular rows, those around spiracle often shorter;
gt, with about 10 setae in one row. Hypopygium with about 4 ventral setae. Cerci with
4 fairly long curved setae.

2D John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

anterior portion of
basal sac of gaster

Ss a

Figures 61-66. Neotriadomerus spp. 61 N. sp. apex of gaster, lateral 62 NV. sp. gt, + ovipositor apex,
lateral 63 NV. longissimus ovipositor sac, lateral 64 NV. sp. male ovipositor apex, lateral 65 N. darling apex
of gaster, posterolateral 66 N. powerae male genitalia, lateral. Scale bar for 61 = 100 um; 62 = 50 um;
63 = 200 um; 64-66 = 20 um.

Male. Similar to female. Body 1280-2560 in length (males still unknown for the
largest species). Flagellum with 11 relatively wide segments (Figs 40-43, 95). Gaster
(card- or point mounted specimens) narrower than mesosoma, laterally compressed,
with posterior apex in dorsal view wider and in lateral view usually higher than anterior
apex (at petiole). Sculpture and setation of male similar to female but with fewer setae

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 23

on gt. Genitalia encapsulated, with thick walls; aedeagal apodemes evenly curved an-
teriorly towards each other, and shorter than aedeagus; paramere shorter than aedeagus
and high in lateral view, with 3 or 4 short apical and ventral setae.

Etymology. The name is masculine. The prefix Neo is Greek for new or recent,
young, + Triadomerus, apparently the closest related genus.

Distribution. Neotriadomerus species occur only in Australia where seven are de-
scribed (below) and as many as four others are illustrated (Figs 144-152) but not
named. No fossil species are known.

Hosts and habitat. Hosts are unknown. ‘The estimated ovipositor length, when
fully extended, is about 12 mm for the largest species of Neotriadomerus. This suggests
that its host might be eggs of Orthoptera laid quite deeply inside plant tissue or in soil.
Perhaps only Orthoptera, but possibly Coleoptera or Cicadidae, would have eggs long
enough to host a developing female parasitoid (5.9 mm long) of this species. Specimens
of Neotriadomerus have been collected in a variety of habitats in all Australian states
except Victoria and Tasmania. Some specimens have been collected at light, suggesting
they may be active at night. Noyes and Valentine (1989: 28, and figs 49, 50) discussed
Australomymar Girault, illustrated two of the largest species, and suggested that eggs of
Orthoptera or Cicadidae might be their hosts. Several specimens (NZAC, UCRC) of
the species shown in Noyes and Valentine fig. 50, are ~55 mm long excluding the pos-
teriorly exserted section of the ovipositor and the longest one is 6.9 mm (NZAC) (S.
Triapitsyn, D. Ward, personal communication). A much smaller, undescribed speci-
men (CNC) of Australomymar, was reared from an unidentified taxon of Tettigoniidae
from Pinus radiata D. Don (Pinaceae) in Chile. I suggest here that Orthoptera might
also be the hosts of Neotriadomerus.

Key to species of Neotriadomerus. Females.

1 Body 5000 long; ovipositor at least 5900 long, extending anteriorly un-
der mesosoma to well beyond level of head (Fig. 126); mandible massive
CEE DON P ee ion Roars hanes etteitehs eaRepadctnatin pale Aaa son Sabaneta N. longissimus sp. n.

= Body at most about 2600 long; ovipositor at most 2240 long, extending
under mesosoma at most to level of head (Figs 114, 115); mandible much
Yon eeUlllesgud By toni aes So ANCOR Be a oe ORM Ae cee ece Soe Oy ert AUR re Ee CRI Ty ORR AO 2
2(1) Fore wing with cubital line of setae extending proximally to base of parastig-
ma, i.e., clearly proximal to other microtrichia on wing surface (Figs 70, 79,
jf Sz le 2) FA Red). ne et ea. came PY ORO AUR We eg Aen PY 3
- Fore wing with cubital line of setae extending proximally only to apex of par-
astigma, about level with other microtrichia on wing surface (Figs 86, 113)....9

3(2) Fl, with 2 mps; fl, with about 4 (5?) mps (Fig 69)........0+ N. burwelli sp. n.
a Fl, with at least 4 mps; fl, with at least 6 mps (Figs 78, 102, 131)... 4
(By) FL, with 2 barely overlapping whorls of mps (Figs 144, 147) .......ccesseeeseees 5
— El ponithplewhorl olonaps. (hie 02)5 neacectse gence aac thse tie an 6

John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Fly sleterthai wri chchie2, Gate PAA) EP tot, scastee, Bee aes cece Wars pateeng he Rese N. sp. 1
Fl length / width) 2,2 (Figs 14 A)... tecseseactsneneete ain deetansensseacetstacens N. sp. 3
Fl about 0.7x as long as fl, (Fig. 102); remaining funicle segments longer and
narrower, with fl, and ft, respectively, 2.5x and 2.0x as long as wide............
Segue gato tea 4a eaces aut ecah ane sea ccs pene oOR gach Meats e adores pe N. gloriosus sp. n.
Fl at least 0.9x as long as fl, (Figs 78, 131); remaining funicle segments
shorter and wider, with fl, and fl,, respectively, at most 2.1x and 1.9x as long

FSAAELCIG atte RR ee sera ore ene et See, oe tel eee, cee eee ees ce EL ree ane 7
Flt. each slightly more than 2.0 x as long as wide; clava about 3.2x as long
AS BUS ds snc cust aaah nssagouesetven bdiedee dubs seed cvitea datas tadyetbendas <n, N. powerae sp. n.
Fl —fL, each at most 1.6x as long as wide; clava about 2.6x as long as wide
IEF EO) Sesscroviens mnie tives lanes dommes ngeeotisrnecnesesindernenr cay wndasebins takyeabindicraese 8
Fifi eachtabout 16x48 loing as: wides.. ete tec toes cet cneeee N. crassus sp. n.
Fl flecach almost. Gita tate’ <, cuz. g2%cesewesacntdeteas—nsenhacusthadeatgteratledete N. sp. 2

Ovipositor extending anteriorly under mesosoma to level of head (Figs 114,
115), at least. about 3.6x as long as metatibia; funicle with at least 9 mps on
EACH ScOTMEMT OLD) 6 cshcsuedenapumseaabadeawedetnes ect. N. longiovipositor sp. n.
Ovipositor extending anteriorly under mesosoma to level of apex of procoxa
(Fig. 92), at most about 2.8x as long as metatibia; funicle with 6 mps on each

SCONICHE PIO Om aneee at seein unseat. ele nd eteaia anete eaten eee N. darlingi sp. n.

Key to species of Neotriadomerus. Males.

Males of darlingi, longiovipositor and powerae are known and almost certainly correctly

associated with the corresponding females; males are unknown for N. burwelli, N.

crassus, IN. gloriosus and N. longissimus. Two unnamed males are also keyed; one (sp. 4)

is not definitely associated with a female and the other (sp. 1) is definitely associated

with a female.

Fore wing with cubital line of setae extending at least to base of parastigma,
clearly proximal to other microtrichia on wing surface (Fig. 139)... 2
Fore wing with cubital line of setae extending at most to apex of parastigma
(level with distal macrochaeta), about level with other microtrichia on wing

ih EevoreiN G EIT Aas 45. ka dem Wey Uy pet Be PR Ov Den Pe hei Se Re RT Pe 4
Fl. wider, its length/width about 1.8 (Fig. 150).......... Neotriadomerus sp. 4
FlMnatroweb itglenechfwidth at least 2-0) (Rigel 38 )is,. co.cc .ncutedeseseeenghea eine 3
Bicabou tees. 2csic.cswasgne peeve steer geri reewaae ere N. powerae sp. n.
HEL SAD OU tS 59 sg ees Ba eco Seta he SPIE a cll ts rth N. sp. 1
Flagellomeres each with 2 barely overlapping whorls of shorter mps
(CE lerta 2 Ne) a east Ace Pata te Masha tone ats Mati eo Sarees N. longiovipositor sp. n.

Flagellomeres each with 1 whorl of longer mps (Fig. 95).....M. :darlingi sp. n.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 25

Neotriadomerus burwelli Huber, sp. n.
http://zoobank.org/B09A3418-3623-42B6-B88F-62C9729617E3
Figs 67—75

Type material. Holotype female (QMBA) on slide (Fig. 71) labelled: 1.“Mahogany
forest, 1200m Mt Moffatt Nat. Pk, C. Qld 24°56'S, 148°04'E 24-26 Feb 1996 CJ.
Burwell mv lamp”. 2. “Holotype Neotriadomerus burwelli Huber 2”.

Diagnosis. Neotriadomerus burwelli differs from the other small (body length less
than 2600) species of Neotriadomerus, as follows: fore wing with cubital line extending
proximally to base of parastigma (Fig. 70); fl, with 2 mps, fl, with 4 mps and almost
2x as long as fl, and fl —fl. each at least 2.0x as long as wide and with at most 6 long
mps in 1 whorl (Fig.69); ovipositor 2.7x as long as metatibia and extending anteriorly
under mesosoma as far as apex of procoxa (Fig. 74).

Description. Female. Body length ~ 1590 (holotype). Colour. Holotype body
almost uniformly dark brown; legs brown, with trochantelli, base and apex of femora
and entire protibia, base and apex of meso- and metatibiae, and tarsi except tar-
somere 5 paler, almost white; tarsomere 5 brown. Head. Head width 305 (Figs 67,
68). Antenna. FI about 0.5 as long as fl,, with 2 mps; fl, the longest funicle segment,
with 4 mps; fl,fl, with 4-6 mps; fl, with 7? mps. Clava with about 12 mps, 5 on
first, 2 on second and 5 on third segment (Fig. 69). Antennal measurements (n=1),
length/width (ratio of flagellar segments): scape 119/20 (3.91), pedicel 50/31 (1.62),
fl, 63/27 (2.32), fl, 82/29 (2.82/3.10 [left and right antenna different]), fl, 77/28
(2.80), fl, 73/30 (2.46), fl, 79/28 (2.79/2.36 [left and right antenna different], fl,
70/30 (2.38), fl, 70/31 (2.27), fl, 69/34 (2.00); entire clava 118/41 (2.89), with
segments 1—3 length [measured along dorsal margin] 44, 21, and 54. Mesosoma.
Width 250, length 514. Wings. Fore wing length 1074, width 386, length/width
2.78, longest marginal setae 60; cubital line extending to just proximal to base of
parastigma (Fig. 70). Hind wing length 862, width 118, longest marginal setae ~
66. Legs. Protibia with 5 short pegs along its length and a transverse row of 4 abut-
ting pegs apically (as in Fig. 53). Metasoma. Metasoma in dorsal view 300 at widest
point, gradually widening from petiole to about 0.6 of gaster length towards apex,
then more abruptly narrowing to cerci (Fig. 73). Gaster length 745, about 1.45x
as long as mesosoma; gt —gt., lengths about 220: 105: 100: 100; 130: 98 (Fig. 73);
hypopygium (apex difficult to see) extending about 0.7x length of gaster. Ovipositor
sheath length 1041, 2.64x metatibia length (394) and extending anteriorly to just
anterior apex of procoxae (Fig. 74) and posteriorly slightly beyond apex of gaster
Wigs: 73.75)

Male. Unknown.

Etymology. The species is named after Chris Burwell, curator of insects at the
Queensland Museum, who collected the only known specimen of this species.

26 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

frontofacial , frontal

suture a x suture

occipital
groove

Figures 67, 68. Neotriadomerus burwelli, holotype. 67 head, anterior 68 head, posterior. Scale bar =
100 pm.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... ZF

Q ote Babine vn
burwe \\y

Nubev &

QMAP

lMahogan MOR NE 1200m

Mi oatatt Nat. Pk, C.Qid
\24°56'S 148°04'E. «
24-26 Feb 1996
[C.J.Burwell_ my lamp,

Figures 69-71. Neotriadomerus burwelli, holotype. 69a right antenna, medial 69b same antenna, lateral
(as seen through antenna) 70 wings 71 holotype slide. Scale bar = 300 um.

28 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Figures 72, 73. Neotriadomerus burwelli, holotype. 72 mesosoma, dorsal 73 metasoma, dorsal. Scale
bar = 300 pm.

Neotriadomerus crassus Huber, sp. n.
http://zoobank.org/ DC4DB59A-207D-4DBD-9234-C944B9FD7 1AA
Figs 76-80

Type material. Holotype female (ANIC) on slide (Fig. 80) labelled: 1. “Black Mountain,
ACT 28-29 Mar. 1968 light trap”. 2. “Neotriadomerus crassus @ lateral holotype”.

Diagnosis. Neotriadomerus crassus differs from the other small (body length less
than 2600) species of Neotriadomerus as follows: fore wing with cubital line extending
proximally to about base of parastigma (Fig. 79); ffl, relatively short, each at most
1.65x as long as wide and with at least 8 mps (Fig. 78).

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 29

Figures 74, 75. Neotriadomerus burwelli, holotype. 74 mesosoma, ventral 75 metasoma, ventral (seen
through body). Scale bar = 300 pm.

Description. Female. Body length ~ 1685 (holotype). Colour. Body almost uni-
formly dark brown; legs brown, with trochantelli, base and apex of femora and entire
protibia, base and apex of meso- and metatibiae, and tarsi except tarsomere 5 of all
legs and metatarsomere | paler, almost white; tarsomere 5 brown and metatarsomere 1

30 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

——

¥,

Figures 76, 77. Neotriadomerus crassus, holotype. 76 head + anterior of mesosoma, dorsolateral 77 meso-

soma + metasoma (ovipositor broken off near base of hypopygium), dorsolateral. Scale bar for 76 = 200 um;
77 = 500 um.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 31

tiger AM rags
y [fi / f/f] ie Jp f ff >

— Black Mtn ACT,
™, 28-29 Har. 1960
- light trap

Figures 78-80. Neotriadomerus crassus, holotype. 78 antenna 79 wings 80 type slide. Scale bar = 500 um.

light brown. Head. Width not measurable (Fig. 76). Antenna. FI, slightly longer than
fl,, apparently with 8 mps; fl,—fl, with 8 mps (possibly 9 or 10 on fl, and fl,); clava
with about 22 mps, 8 on first, 6 on second and 8 on third segment (Fig. 78). Antennal
measurements (n=1), length/width (ratio of flagellar segments): scape 145/47 (3.10),
pedicel 58/42 (1.37), fl, 86/56 (1.53), fl, 84/52 (1.60), fl, 82/50 (1.63), fl, 81/50
(1.63), fl, 86/53 (1.62), fl, 82/51 (1.59), fl, 78/50 (1.57), fl, 76/48 (1.58); entire clava
143/55 (2.59), with segments 1—3 length [measured along dorsal margin], 46, 35, and
64. Mesosoma. Width not measurable, length 580. Wings. Fore wing length 1154,
width 397, length/width 2.91, longest marginal setae 46; cubital line extending to just
proximal to base of parastigma (Fig. 79). Hind wing length 942, width 130, longest
marginal setae 66. Legs. Protibia with 6 short, thick pegs along its length and trans-
verse row of 4 abutting pegs apically (as in Fig. 53). Metasoma. Metasoma (Fig. 77)
in lateral view 224 at highest point, gradually widening from petiole to about 0.6 of

32 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

gaster length towards apex, then more abruptly narrowing to cerci. Gaster length 765,
about 1.63x as long as mesosoma; gt —gt, lengths about ~ 156, ~ 94, 87, 112, 118,
152; hypopygium (Fig. 77, positioned at right angle to metasoma) extending about
0.7x length of gaster, extending to about halfway towards apex of tergum 5. Ovipositor
broken and partly missing, with estimated length (from basal loop to posterior apex
of sheath) ~ 864, about 2.11x metatibia length (410), extending anteriorly to about
level of apex of mesocoxae and extending posteriorly slightly beyond apex of gaster
(Fig. 77).

Male. Unknown.

Etymology. The species name, crassus, is Latin for thick or stout, referring to the
fairly short, thick funicle segments in females.

Neotriadomerus darlingi Huber, sp. n.
http://zoobank.org/12B05D86-1539-4CA9-B5F8-6895EA2E6952
Figs 33, 40-44, 47, 50, 52, 65, 81-99

Type material. Holotype female (ANIC) on slide (Fig. 84), labelled: 1. “Australia: Sth
Aust. 32 km N. Renmark, 263m 33°53'S, 140°43'E, 15.ii-15.iv.2000, DC Darling”.
2. “Bookmark Biosphere Rsv. Malaise trps (4) Amalia Dam xeric mallee scrub ROM
2000040”. 3. “Neotriadomerus darlingi Huber Holotype 9 dorsal”.

Paratypes. 3 females, 1 male. AUSTRALIA. South Australia: Same locality data
as holotype (1 Q and 14, ROM, CNC); 35 km N Renmark, 263m, 33°52'48"S;
140°43'30"E, 11-14.ii.2000, D.C. Darling, Bookmark Biosphere Reserve, Malaise
trap, Amalia Dam, mallee vegetation, ROM 2000030 (12, ANIC). Western Aus-
tralia: 85 km E. Southern Cross, Boorabbin National Park, 28.xii.1986, J.S. Noyes
(12, BMNH)

Two males with the following data questionably belong to this species so are not
labelled as paratypes. They were collected from South Australia, Brookfield Conservation
Park, 34.21°S, 139.29°E, 17 & 18.ii.1992, J. Cardale, A. Roach, light trap (24, ANIC)
and one is illustrated (Figs 93-99). Both sexes need to be collected together from Brook-
field Conservation Area to be more certain of their conspecificity with the holotype.

Diagnosis. Neotriadomerus darlingi differs from the other small (body length less
than 2600) species of Neotriadomerus as follows: fore wing with cubital line extending
proximally to about level of other microtrichia (Fig. 86); funicle segments with at most
6 mps (Fig. 85); ovipositor 2.57—2.74x as long as metatibia (Fig. 92).

Description. Female. Body length 1420-1560 (n=2, card-mounted paratypes).
Colour. Body (Fig. 81) almost uniformly dark brown; fore leg yellow except procoxa
brownish basally, ventral surface of profemur and tarsomere 5 brown; middle and hind
legs lighter brown with trochantelli, base and apex of femora and entire protibia, base
and apex of meso- and metatibiae, and tarsi except tarsomere 5 paler, almost white;
metafemur medially and tarsomere 5 brown; ovipositor sheath with lighter band sub-

apically (Figs 81, 88, 89). Head. Width ~ 320-346 (card-mounted) and 358 (slide

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 33

TES Se he a |
Figure 81. Neotriadomerus darlingi, paratype female, habitus lateral (35 km N. Renmark, 263 m, 11-14.
ii.2000). Scale bar = 1000 um.

mounted specimen from WA, Figs 90, 91). Antenna. Fl slightly shorter than to almost
equal to fl,, with 5 mps (Fig. 85); fl,—fl, with 6 mps. Clava with 10 or 11 mps, 5 or

6 on first, 2 on second and 4 on third segment. Antennal measurements (n=2, first

34 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Figure 82. Neotriadomerus darlingi, paratype male, habitus dorsal (35 km N. Renmark, 263 m, 15.ii-15.
iv.2000). Scale bar = 1000 um.

number is for the holotype, a slightly smaller specimen), length/width (ratio of flagel-
lar segments): scape 124—126/37—40 (3.37-2.51), pedicel 50-56/39—41 (1.29-1.36),
fl, 70-74/38-45 (1.85-1.65), fl, 80-75/34-40 (2.36-1.88), fl, 77-76/34-38 (2.26-
1.97), A, 78-75/35-38 (2.24-1.96), fl, 79-76/33-38 (2.39-2.01), fl, 76-74/33-37
(2.28-2.20), fl, 77-78/34-39 (2.23-2.13), fl, 76/3440 (2.25—-1.89); entire clava
141-142/39-42 (3.63—3.40), with segments 1—3 length [measured along dorsal mar-
gin], 48-50, 25, and 67-68. Mesosoma. Width 225 (holotype), length 628-656, with
2 propodeal setae (Fig. 87). Wings. Fore wing length 1045-1118, width 383-412,
length/width 2.71—2.73, longest marginal setae ~ 51-61; cubital line extending to just
proximal to base of parastigma (Fig. 86). Hind wing length ~ 870-890, width 121-
140, longest marginal setae ~ 58-62. Legs. Protibia with 4 or 5 short, thick pegs along
its length and a transverse row of 2 or 3 abutting pegs apically. Metasoma. Metasoma
in lateral view 255-360 at highest point, gradually widening from petiole to about
0.4—0.7 of gaster length towards apex, then more abruptly narrowing to cerci (Fig. 89).
Gaster length (holotype, second measurement) 776-858, 1.18—1.36x as long as meso-
soma; gt,—gt, lengths about ~ 162-170, ~ 136-144, 98-126, 113-177, 132-186,

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 35

aah ey
oe ae |
eres) Bai
$839 Ose
Zenk) S39
a ose ae
aoe 82

Figures 83-86. Neotriadomerus darlingi, holotype. 83 head, anterolateral 84 type slide 85 antenna

86 wings. Scale bar = 200 um.

36 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

basal sac
of gaster

“<—— ovipositor

f

ovipositor
sheath

y

‘a petiole... 2

Figures 87-89. Neotriadomerus darlingi, holotype. 87 mesosoma, dorsal 88 metasoma, lateral, with
ovipositor unsheathed 89 metasoma enlarged, lateral. Scale bar for 87 = 200 pm; 88, 89 = 500 um. Note:
88 and 89 are flipped horizontally (gaster apex pointing left in holotype slide).

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 37

——— EE | ' F ; ;

Figures 90-92. Neotriadomerus darlingi, paratype female (Boorabbin National Park). 90 head, anterior
91 head, posterior 92 mesosoma, legs and metasoma, lateral. Scale bar for 90,91 = 200 pum; 92 = 500 pm.

38 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

er eee es

Figures 93-96. Neotriadomerus ?darlingi, paratype male (Brookfield Conservation Area). 93 head, anterior
94 head, posterior 95 antenna 96 wings. Scale bar for 93-95 = 200 um; 96 = 500 um.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 39

aedeagal
apodeme

ae Ae
Figures 97-99. Neotriadomerus ?darlingi, paratype male (Brookfield Conservation Area). 97 mesosoma,
dorsal 98 metasoma, dorsal (gt, broken off) 99 genitalia, dorsal. Scale bar for 97, 98 = 300 um; 99 = 100 pm.

40 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

209-146, 53-58; hypopygium extending about 0.6x length of gaster, about to apex of
tergum 4. Ovipositor sheath length ~ 946-1050, about 2.57—2.74x metatibia length
(368-383), extending anteriorly to apex of procoxa (or anterior margin of frenum) and
extending posteriorly to a little beyond apex of gaster (Figs 88, 89, 92).

Male. Body length 1330 (point-mounted paratype) and 1230 (slide-mounted
specimen, Brookfield Conservation Area). Colour as for female but with legs almost
entirely brown (Fig. 82). Head. If correctly associated, as for female (Figs 93, 94). An-
tenna. Fl, with about 13 mps and about 1.5x as wide as fl,, with about 7 mps (Fig. 95);
total length of flagellum 475. Antennal measurements length/width (slide-mounted
specimen): scape 120/41 (2.90), pedicel 47/43 (1.09), fl, 92/53 (1.75), fl, 105/48
(2.24), fl, 108/48 (2.24), Al, 104/44 (2.38), A, 109/43 (2.54), fA, 100/42 (2.41), fl
98/41 (2.39), fl, 97/43 (2.24), fl, 96/4142.35), fl, 86/395(2:22), fl 92/35: (2.64); f1.
with about 11 mps. Mesosoma. Length 620, width 290 (Fig. 97). Wings. If correctly
associated, as for female (Fig. 96). Metasoma. Pedicel length/width 30/75 (Fig. 98).
Gaster length ~ 785. Genitalia with capsule thick-walled, aedeagal apodeme thick and
shorter than aedeagus; paramere thick and apically curved medially (Fig. 99).

Etymology. ‘The species is named after Chris Darling, curator of Entomology at
the Royal Ontario Museum, Toronto, who collected the type series.

Neotriadomerus gloriosus Huber, sp. n.
http://zoobank.org/33067489-E0B9-488E-9AAD-591B903E3F00
Figs 32, 34, 35, 100-107

Type material. Holotype female (QMBA) on slide (Fig. 107), labelled: 1. “Mt. Glori-
ous, S.E. Qld. Hiller March 1982 Malaise trap”. 2. “Neotriadomerus gloriosus Huber
Q dorsal holotype”.

Diagnosis. Neotriadomerus gloriosus differs from the other small (body length less
than 2600) species of Neotriadomerus, as follows: fore wing with cubital line extending
to just proximal to base of parastigma (Fig. 103); ffl, each at least 2.0x as long as
wide and with at most 6 long mps in 1 whorl (Fig. 102).

Description. Female. Body length ~ 1840 (holotype). Colour. Holotype body
almost uniformly dark brown; legs brown, with trochantelli, base and apex of femora
and entire protibia, and base and apex of meso- and metatibiae, and tarsi except tar-
somere 5 paler, almost white; tarsomere 5 brown. Head. Head width 374 (Figs 100,
101). Antenna. Fl, about 0.7x as long as fl,, with 4 mps; fl, slightly the longest funicle
segment, with 8 mps; ffl, with 8 (92 on fl.) mps; clava with about 18 mps, 8 on
first, 4 on second and 6 (8?) on third segment (Fig. 102). Antennal measurements
(n=1), length/width (ratios of flagellar segments, different for each antenna so left an-

tenna, except scape of right antenna, measured—the left antenna has narrower funicle
segments): scape 152/49 (3.10), pedicel 57/43 (1.32), fl, 91/50 (1.84), fl, 118/48
(227), she VIA IAGs( AS), He I/AG (2.45) tO 5146 42.29), 99145: @., 24) oie
99/47 (2.09), fl, 92/46 (1.98); entire clava 148/52 (2.82), with segments 1-3 length

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 4]

——
SSS =

— SSS
——S -

\
(ee

se

Figures 100-103. Neotriadomerus gloriosus, holotype. 100 head, anterior 101 head, posterior
102a right antenna 102b left antenna 103 wings. Scale bar for 100, 101 = 200 um; 102, 103 = 500 um.

42 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

: —» r :

basal sac.
of gaster

| tie
anterior extension
of ovipositor
|

er)

(Hi |
March

Mt. ero tos FE OIE |
1
982

Figures 104-107. Neotriadomerus gloriosus, holotype. 104 mesosoma, dorsal 105 metasoma, lateral
106 metasoma, median plane 107 type slide. Scale bar = 200 um.

[measured along dorsal margin] 56, 28, and 64. Mesosoma. Width 324, length 711,
with 2 propodeal setae (Fig. 104). Wings. Fore wing length 1367, width 486, length/

width 2.81, longest marginal setae 77; cubital line extending to just proximal to base of

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 43

parastigma (Fig. 103). Hind wing length 1109, width 157, longest marginal setae 78.
Legs. Protibia with 6 or 7 short, thick pegs along its length and a transverse row of 4
abutting pegs apically (as in Fig. 53). Metasoma. Metasoma height in lateral view 225
(excluding lowered hypopygium) (Figs 105, 106). Gaster length 1002, about 1.41x as
long as mesosoma; gt gt, lengths about 170: 166: 151: 118: 130: 209 [measurement
of a tergum excludes the telescoped portion inside another (more anterior) tergum];
hypopygium extending posteriorly to about level of spiracle. Ovipositor sheath length
1274, ~ 2.54x metatibia length (= 502), extending anteriorly to about level of apex of
procoxa and extending posteriorly only slightly beyond apex of gaster (Fig. 105, 106)
[apparently extending more than normal because ovipositor not at its normal resting
position, i.e., filling entire length of basal sac of gaster, whose almost membranous apex
is distinctly folded over (Fig. 105, arrow)].

Male. Unknown.

Etymology. The species is named after the type locality, Mt. Glorious National Park.

Neotriadomerus longiovipositor Huber, sp. n.
http://zoobank.org/1 CDB865F-9F 10-446D-913D-C76B7E6501C2
Figs 108-125

Type material. Holotype female (ANIC) on slide (Fig. 111) labelled: 1. “15.178
145.10E 5Km WbyN Rounded Hill nr Hope Vale Mission Q 7 Oct. 1980 J.C. Card-
ale ex ethanol”. 2. “Neotriadomerus longiovipositor Huber Holotype 9 dorsal”.

Paratypes. 4 females, 4 males, 1 deformed male (gynandromorph?). AUS-
TRALIA. Queensland: Batavia Downs, 12.40°S; 142.39°E, 22.vi-23.viii.1992, P.
Zborowski & J. Cardale, flight interception trap (1d, ANIC); NW of Chillagoe near
historical mine site, 17°08'40"S; 144°30'16"E, 30.iii.1992, E.C. Dahms & G. Sarnes
(13, QMBA); 18.5 km W. Gordonvale, 13.xi.1979, E.C. Dahms, J.B. Woolley &
J. LaSalle, open forest (12, QDPC); Heathlands, 11.45°S; 142.35°E, 15—26.i1.1992,
I. Naumann, T. Weir, at light (12, ANIC); Millstream Falls National Park, 17.41°S;
145.26°E, 24—25.v.1980, I.D. Naumann, J.C. Cardale (1 deformed 3, ANIC); 5 km
W. by N. Rounded Hill near Hope Vale Mission, 15.17°S, 145.10°E; 7-10.v.1981,
I.D. Naumann (19, ANIC); 15.16°S; 144.59°E 14 km W. by N. Hope Vale Mis-
sion. 8-18.x.1980, J.C. Cardale, collected at light (13, ANIC); Ross River resevoir, 10
km S. Townsville, 19° 27'S; 146° 44'E, 27.xi.1991, C.J. Burwell (14, QMBA); near
Swamp Mitchell Plateau airfield, 14.47°S; 125.49°E, 18.v.1983, I.D. Naumann, J.C.
Cardale (12, ANIC).

Diagnosis. Neotriadomerus longiovipositor differs from the other small (body
length less than 2600) species of Neotriadomerus, as follows: fore wing with cubital
line extending proximally to about level of other microtrichia (Fig. 113); ovipositor
extending anteriorly under mesosoma at least to level of head (Figs 114, 115); funicle
segments with mps in two widely overlapping whorls (Fig. 112). Male flagellomeres
with mps in two scarcely overlapping whorls (Fig 121).

44 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

tentorium

4

Figures 108-111. Neotriadomerus longiovipositor, holotype. 108 head, anterior 109 mouthparts and
tentorium I 10 head, posterior 111 holotype slide. Scale bar = 200 um.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 45

ee a ee ey a ee eee
eee a ee 1] a
ae eee Ss fe see es ee t |
=, 2: = —_ — a ie Se - Ae
= aa Fie a! roghil > spol ae 423 4 F
Sie s ee Oe = i a s Hi zs —, /
nee = ~“ ey oe? a - - a! iis. i}
Spyg ee pe ee ghd co /
/ A

>
TI 2G +

MMV YY 13)

Ze” ” 15)
| R | A

Figures 112-115. Neotriadomerus longiovipositor, holotype. 112 antenna 113 wings 114 mesosoma +

metasoma, dorsal (inset shows fenestra of scutellum) 115 mesosoma + metasoma, ventral as seen dorsally

through body. Scale bar = 500 pm.

46 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

» Cercus

Figures | 16-118. Neotriadomerus longiovipositor, female paratypes. 116 head + anterior of mesoso-
ma, dorsal (near Swamp Mitchell Plateau Airfield) 116 metasoma, lateral (18.5 km W. Gordonvale)
1 17 metasoma, median plane I 18 metasoma, lateral. Scale bar for 116 = 200 um; 117, 118 = 1000 um.

Description. Female. Body length ~ 1700-2250 (n=6). Colour. Body (before
slide mounting one specimen) uniformly shiny black except mouthparts brown; an-
tenna dark brown but scape and pedicel ventrally slightly lighter; pro- and mesocoxa
dark brown except extreme apices yellowish, metacoxa dorsally almost black; the rest
of each leg yellowish except for femur ventrally of fore- and mid leg, entire femur ex-

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 47

A
| Ocular
- apodeme

Figures 119, 120. Neotriadomerus longiovipositor, male paratype (14 km W. by N. Hope Vale Mission).
119 head, anterior (inset shows ocular apophysis or apodeme) 120 head, posterior. Scale bar = 200 um.

48 John T; Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

postmarginal vein

MEELIS IDG Tp Dot
er -
a eee ts = Se
— 5, eae < = = = 5
LL pip ed ee 5. . F ,
MA ii ypee Co ypae?
VY) if ii | fe
sm |

Figures 121, 122. Neotriadomerus longiovipositor. \2\ male antenna, paratype (14 km W. by N. Hope
Vale Mission) 122 wings. Scale bar = 500 pm.

cept extremities of hind leg, and apical tarsomere of all legs brown. Head. Head width
344-412 (Figs 108, 110). Antenna. FI, slightly shorter than fl,, with 8 (9?) mps; ff,
slightly the longest funicle segment, with 9-12 mps; fll, with 9-12 (13?) mps; clava
with 22 mps, 8 on first, 6 on second and 8 on third segment (Fig. 112). Antennal
measurements (n=4), length/width (ratio of flagellar segments): scape 174—194/47-65
(2.99-3.78), pedicel 57-66/40-46 (1.30-1.67), fl, 85-104/44-57 (1.83-1.95), fl,
88-108/45—52 (1.88-2.19), fl, 80-109/43-52 (1.72-2.11), fl, 7284/4447 (1.65—
1.97), fl, 70-102/42—47 (1.66-2.19), fl, 68-97/43—-50 (1.56—-1.94), fl, 65-96/41—44
(1.54-2.17), fl, 63-64/42-46 (1.41-1.49); entire clava 126—128/49/53 (2.43-2.59),

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n....

49

—— ¢ \ F A,

Figures 123, 124. Neotriadomerus longiovipositor, male paratype (14 km W. by N. Hope Vale Mission).

123 mesosoma, legs + metasoma, lateral 124 genitalia, lateral. Scale bar for 123 = 1000 um; 124 = 200 um.

with segments 1—3 length [measured along dorsal margin] 40-46, 29-31, and 50-60
(the paratype from 18.5 km W. Gordonvale was the largest specimen but had fl, and

clava missing from both antennae so the maximum lengths in the ranges given above

50 John T; Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

paramere”

aedeagus“

Figure 125. Neotriadomerus longiovipositor, male paratype (near historical mine site, NW of Chillagoe).

Gaster, median plane showing genitalia. Scale bar = 300 um.

for those segments are smaller than they should be). Mesosoma. Mesosoma length
617-678, width 317-327 (n=3, slide mounts), height ~ 280 (critical point dried
paratype). Wings. Fore wing length (n=3) 1118-1208, width 407-432, length/width
2.71—2.80, longest marginal setae ~ 54-62; cubital line extending to just proximal
to base of parastigma (Fig. 113). Hind wing length ~ 914-1052, width 120-136,
longest marginal setae ~ 50-67. Legs. Protibia with 5 or 6 (on specimen with only
2 on one leg) short, thick pegs along its length and a transverse row of 3 or 4 abut-
ting pegs apically (as in Fig. 53). Metasoma. Metasoma height in lateral view ~ 480
(including slightly lowered hypopygium, critical point dried paratype) or 320-400
(three slide mounted paratypes). Gaster length 918-1453, about 1.49—2.12x as long as
mesosoma; gt,—gt, lengths about 204—244, 190-236, 136-266, 130-207, 114-209,
218-320, ~ 24-59 [measurement of a tergum excludes the telescoped portion inside
another (more anterior) tergum]; hypopygium extending posteriorly to about apex of
gt, to halfway between anterior and posterior margin of gt,. Ovipositor length 1445-
2238, = 3.58—4.82x metatibia length (~ 380-472) and extending anteriorly to level of
head or beyond (Figs 114, 115) and not extending posteriorly beyond apex of gaster
(Figs 114, 115), except in one paratype (Figs 117, 118).

Male. Colour. As for female. Head. Head (Figs 119, 120) width 380-436 (n=4).
Antenna. Measurements, length/width (n=3): scape 170-—214/56-67, pedicel 57—
68/46—52, flagellar segments: fl, 117122155263; fl, 109-134/52-58, fl, 103-136/50-—
62, fl, 105-130/47-56, fl, 102-125/48-55, fl, 98-127/50, fl, 96-120/48-52, fl, 89-
120/44—53, fl, 84-112/45-48, fl, 76-104/42-48, fl, 90-118/36—43; total flagellar
length 1068-1349; fl length/width ratio 1.98—2.54, with about 17-19 mps in two
slightly overlapping whorls (Fig. 121). Mesosoma. Mesosoma length 780-860, about
0.80—1.22x as long as metasoma length (680-1020); propodeum with 2 propodeal se-
tae, rarely 3 on one side. Wings. Fore wing (Fig. 122) length (n=3) 1191-1320, width

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 51

446-495, length/width, 2.56—2.67, longest marginal setae 58—68; cubital line extend-
ing to about level of other microtrichia. Hind wing length 921-1024, width 132-158,
longest marginal setae 52-65. Metasoma. Petiole length 35-36, width 70-76 (n=2).
Gaster length 780-860; gt,—gt. each with a row of about 5 lateral and dorsal setae on
each side (apparently without dorsal setae on gt,), and gt, with about 6 long dorsal setae
on each side and about 20 short lateral setae (Fig. 123). Genitalia (Fig 124, 125) with
paramere in lateral view higher than wide and blunt apically, with 2 short apicoventral
setae close together and 2 or 3 short subapical setae more widely spaced.

Etymology. The name is a noun in apposition, referring to the long ovipositor (the
second longest in the genus) that extends anteriorly to the head.

Neotriadomerus longissimus Huber, sp. n.
http://zoobank.org/B79EB 1 F3-8092-4382-96C7-A966E9549C50
Figs 36-38, 48, 51, 53, 63, 126, 127

Type material. Holotype female (ANIC) on point labelled: 1. “Blundells ACT 27 Feb
1951 H M Cage”. 2. “Holotype Neotriadomerus longissimus Huber 2”.

Paratype. | female. AUSTRALIA. Queensland: Brisbane Forest Park, 27°25'04"S;
152°49'48"E, 23-29.x.1998, N. Power, dry sclerophyll, MT (12, CNC).

Diagnosis. Neotriadomerus longissimus differs from other species of Neotriadomerus
as follows: body 5000—almost twice the length of any other species (Fig. 126); head
cuboidal, with vertex horizontal, almost at right angle to the face (Fig. 126); mid ocel-
lus almost in contact with the transverse trabecula; mandible massive (Fig. 36); gaster
with basal sac extending anteriorly well anterior to head (Figs 38, 63, 126, 127).

Description. Female. Body length 5000-5450 (n=2, card and point-mounted
holotype and paratype), excluding basal sac of gaster (5900-6300 if this included).
Colour. Holotype body black except for brown mandibles, scutellum, and most of
middle segments of metasoma; legs and apex of anterior extension of ovipositor brown-
ish yellow; paratype body (Figs 126, 127) uniformly brown, except for almost black
head and scape brown, legs mostly yellow except for partly or almost entirely brown
coxae and metafemur, anterior extension of ovipositor from level of head almost white
(Figs 126, 127). Head. Head width 640-690. Vertex horizontal, forming almost right
angle with face, with ocelli in anterior half and mid ocellus in contact with transverse
trabecula. Eye almost circular, slightly narrower ventrally than dorsally. Gena at dorsal
and ventral margins of eye almost equally wide. Mandible massive, extending ventral
to head by at least half face height. Antenna. Each funicle segment in lateral view at
least twice its width as in dorsal view, with about 25-30 short mps in 3 more or less
overlapping whorls (Fig. 38). Clava with about 10 mps on each segment. Antennal
measurements (n=2), length/width (ratio of flagellar segments): scape 297-317/109—
119 (2.58-2.91), pedicel 54-69/59-69 (1.08-1.07), fl, 228/109-119 (1.92-2.09),
fl, 262—267/89-109 (2.45-2.94), fl, 248-257/89-99 (2.60-2.78), fl, 238-248/79
(3.00-3.13), fl, 228/69—74 (3.07-3.29), H, 198-218/69 (2.86-3.14), fl, 178-198/69

52 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

\

e

K-

Figures 126, 127. Neotriadomerus longissimus, paratype, lateral (Brisbane Forest Park). 126 habitus,
lateral 127 anterior body, enlarged. Scale bar for 126 = 2000 um; 127 = 1000 pm.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 53

(2.57-2.86), fl, 168—-178/69 (2.23-2.57), entire clava 267/64—69 (3.86-4.05), with
segments 1—3, respectively, 109, 69 and 89-99 long. Mesosoma. Mesosoma length
110-120. Wings. Fore wing length (n=2) 2460-2790, width 670-845, length/width
3.30-3.72, longest marginal setae 100; cubital line of setae extending halfway between
apex and base of parastigma (about midway between level of proximal and distal mac-
rochaetae). Hind wing length 1970-1790, width 205-230, longest marginal setae
100-130. Metasoma. Metasoma (paratype only) in dorsal view thin, 260 at its widest
(near apex) and 80 at its narrowest, its length (near middle) 345 and 390 (holotype
and paratype), 3.18—3.25x as long as mesosoma; in lateral view 435 high (near apex)
and 205 (near middle); gt —gt, lengths (paratype only) 715: 690: 155: 715: 740: 435:
80; hypopygium extending about 0.3x length of gaster. Ovipositor sheath length (n=2)
5900-6300, 6.23—7.9x metatibia length (795-950) and extending anteriorly to well
in front of head but barely extending posterior to apex of gaster (Fig. 126).

Male. Unknown.

Etymology. ‘The species name, /ongissimus, is Latin for longest, referring the ex-
tremely long gaster.

Neotriadomerus powerae Huber, sp. n.
http://zoobank.org/AEDF 1 141-A5A7-48EF-935A-A47373FE4329
Figs 128-143

Type material. Holotype female (ANIC) on slide (Fig. 130) labelled: 1. “Australia:
Qld. Brisbane Forest Park, 27°25'04"S 152°49'48"E 29.xi-5.xii.1997, dry sclerophyll,
N. Power, Mt”. 2. “Neotriadomerus powerae Huber @ dorsal holotype”.

Paratypes. 1 female, 3 males. AUSTRALIA. Queensland: Same data as holotype
but 14-20.iii.1998 (12 and 14, CNC), 28.xi.1998 (13, ANIC), 26.ix-2.x.1999 (18,
ANIC). The collector stated that the specimens were collected in a Malaise trap set
across a creek bed running through the base of an occasionally flooded gully between
two sections of dry sclerophyll.

Diagnosis. Neotriadomerus powerae differs from other small (body length less than
2600) species of Neotriadomerus as follows: fore wing with cubital line extending to
just proximal to base of parastigma (Fig. 132); fl—fl, each just over twice as long as
wide, with at least 8 mps and clava with 6 mps on each segment (Fig. 131).

Description. Female. Body length ~ 1560 (holotype). Colour. Body (Fig. 128)
almost uniformly dark brown; legs brown, with trochantelli, base and apex of femora
and entire protibia, and base and apex of meso- and metatibiae, and tarsi except tar-
somere 5 paler, almost white; tarsomere 5 brown. Head. Head width 363 (holotype).
Antenna. EF almost as long as iN with 8 mps; fl, about equal to fl, or following fu-
nicle segments except f1, or fl,, with 8 mps; fl,—fl with 8 mps, fl, and fl, apparently
with 9 or 10 mps. Clava with 18 mps, 6 on first, 6 on second and 6 on third segment
(Fig. 131). Antennal measurements (n=1), length/width (ratio of flagellar segments):

54 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

|
Figure 128. Neotriadomerus powerae, paratype female habitus, dorsal (Brisbane Forest Park). Scale bar
= 1000 pm.

scape ~ 138/44 (= 3.15), pedicel 54/43 (1.27), fl, 84/48 (1.77), fl, 88/43 (2.07), A,
90/44 (2.06), fl, 88/42 (2.10), fl. 88/40 (2.19), fl. 89/40 (2.23), fl 86/42 (2.05), fl,
82/43 (1.91); entire clava 151/47 (3.23), with segments 1—3 length [measured along
dorsal margin] 50, 37, and 64. Mesosoma. Width 284, length 578, with 3 propodeal
setae (Fig. 133). Wings. Fore wing length 1173, width 417, length/width 2.81, longest
marginal setae 52. Hind wing length 940, width 130, longest marginal setae 62. Legs.
Protibia with 5 short, thick pegs along its length and a transverse row of 4 abutting
pegs apically (as in Fig. 53). Metasoma. Metasoma height in lateral view 270. Gaster
length 865, about 1.5x as long as mesosoma; gt gt, lengths about 169: 106: 99: 113:
162: 170 (Figs 134, 135) [measurement of a tergum excludes the telescoped portion
inside another (more anterior) tergum]; hypopygium (difficult to see) extending pos-
teriorly to level of apex of tergum 5. Ovipositor sheath length 1113, = 2.73x metatibia

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 55

eee se

Figure 129. Neotriadomerus powerae, paratype male habitus, lateral (Brisbane Forest Park). Scale bar =
1000 pm.

length (~ 51), extending anteriorly to about level of apex of procoxa and extending
posteriorly slightly beyond apex of gaster (Figs 134, 135).

Male. Colour. As for female (Fig. 129). Head. Head (Figs 136, 137) width 346
(n=1). Antenna. Measurements, length/width: scape 124/45, pedicel 49/43, flagellar
segments: fl, 97/62, fl, 108/54, fl, 104/56, fl, 102/50, fl. 108/50, fl, 102/50, fl 98/53,
fe 99/51, fl, 94/50, fi; 86/47, fl, 90/45; total flagellar length 1089; fl. length/width

56 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

2 dor sa\
HOLOTYPE

| Hie Pe p< ae 3 me Be 1 rt
| Nnry i rrr /

Figures 130-132. Neotriadomerus powerae, holotype. 130 type slide 131 head + antenna, anterior
132 wings. Scale bar = 500 pm.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 57

Figures 133-135. Neotriadomerus powerae, holotype. 133 mesosoma, dorsal 134 mesosoma, lateral
135 mesosoma, median plane. Scale bar for 133 = 200 pm; 134, 135 = 500 um.

58 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Cin

Figures 136-139. Neotriadomerus powerae, male paratype (Brisbane Forest Park). 136 head, anterior
137 head, posterior 138 antenna 139 wings. Scale bar for 136, 137 = 200 um; 138, 139 = 500 um.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... a9

Figures 140-143. Neotriadomerus powerae, male paratype (Brisbane Forest Park). 140 mesosoma, dorsal

141 mesosoma, ventral 142 metasoma, dorsal 143 genitalia, dorsal seen through gaster. Scale bar = 200 pm.

60 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

2.04, with 12 mps (Fig. 138). Mesosoma. Mesosoma length 610, about 1.24x as long
as metasoma length (494); propodeum with 3 propodeal setae (Fig. 140). Wings. Fore
wing length (n=1) 1043, width 386, length/width 2.70, longest marginal setae 76; cu-
bital line extending to just proximal to base of parastigma (Fig. 129). Hind wing length
834, width 132, longest marginal setae 62. Metasoma. Petiole length 13, width 29.
Gaster length 468; gt —gt, with 1 long dorsal setae and 4 long lateral setae on each side
and gt. with about 5 long dorsal setae, 3 long lateral setae, and about 20 short lateral
setae on each side (Fig. 142). Genitalia as in Fig. 143.

Etymology. The species is named after Narelle Power, who ran a Malaise trap for
many months in Brisbane Forest Park.

Neotriadomerus sp. 1.

Figs 144, 145

Material examined. AUSTRALIA. Australian Capital Territory: 3 km E. Piccadilly
Circus, Blundells Creek, 35.22°S, 148.50°E, 850m, xii.1984, Weir, Lawrence, Johnson
(19 and 13, ANIC).

Comment. The female (Fig. 144) and male (Fig. 45) almost certainly are conspe-
cific, with body lengths 2460 and 2480, respectively. Female antenna with 2 widely
overlapping whorls of mps (each segment with about 10? mps); male antenna with 2
narrowly overlapping whorls of mps (each segment with about 20? mps). This species
most resembles NV. longiovipositor but differs in that the cubital line of setae extends
proximal to base of parastigma, unlike in NV. longiovipositor.

Neotriadomerus sp. 2
Fig. 146

Material examined. AUSTRALIA. Western Australia: 29 km SE by E of Coolgardie,
31.07°S, 121.24°E, 5.v.1983, E.S. Nielsen, E.D. Edwards (12, ANIC).

Comment. Body length 1380. Female antenna with lwhorl of mps on each seg-
ment and the shortest and widest funicle segments (Fig. 146) of any Neotriadomerus
species; fore wing with cubital line of setae extending to base of parastigma.

Neotriadomerus sp. 3
Fig. 147

Material examined. AUSTRALIA. New South Wales: Kosciusko National Park,
Leather Barrel Creek, 0.8 km SW. Picnic Area, 36°32'S, 148°11'E, 1080m, 7-21.
ii.1993, A. Newton, M. Thayer, open Eucalyptus forest (gum + delegatensis) with
shrubby understory, window trap (192, ANIC).

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 61

Figures 144, 145. Neotriadomerus sp. 1 (Australia, ACT, 3 km E. Piccadilly Circus, Blundells Creek,

35.22°S, 148.50°E, 850m, xii.1984. 144 female habitus, dorsolateral 145 male habitus, dorsal. Scale bar
= 1000 pm.

62 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Figure 146. Neotriadomerus sp. 2, female habitus, lateral (Australia, WA, 29 km SE by E of Coolgardie,
31.07°S, 121.24°E, 5.v.1983). Scale bar = 1000 um.

Comment. Body length 1920; fore wing with cubital line of setae extending to
base of parastigma. This specimen is similar to the female from Blundells Creek but fl,
is even shorter than fl, so it is provisionally treated as distinct.

Neotriadomerus sp. 4
Figs 148-152

Material examined. AUSTRALIA. Northern Territory: 12 km WNW Ross River,
Tourist: Camp, 23°32°5,7134.23" 5, 13.v1978,).€2 Cardale (14, ANIC).
Description. Male. Body length ~ 1330. Colour. Body almost uniformly dark
brown; legs brown, with trochantelli, base and apex of femora and tibiae, and tarsi
except tarsomere 5 paler; tarsomere 5 brown. Antenna. Measurements, length/width:
scape 139/46, pedicel length/width 52/46, flagellar segment length: fl, 100/68, 41,
94/675 197161, fh, TOOTS G98. 100/56, 98/54, HO 96/54, i. 96/53; T9927 50h.
87/46, fl, 94/37; total flagellar length 1053; fl, length/width 1.81, with 11 (12?) mps

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 63

eles Aa IES Tt j

Figure 147. Neotriadomerus sp. 3, female habitus, lateral (Australia, NSW, Kosciusko National Park,
Leather Barrel Creek, 0.8 km SW. Picnic Area, 36°32'S, 148°11'E, 1080m, 7—21.ii.1993). Scale bar =
1000 pm.

(Fig. 150). Mesosoma. Mesosoma (Figs 148, 149) length 640, 1.16x as long as meta-
soma length; propodeum with 3 propodeal setae (Fig. 149). Wings. Fore wing length
1077, width 398, length/width 2.71, longest marginal setae 60; cubital line extending
to just proximal to base of parastigma (Fig. 151). Hind wing length 900, width 138,

64 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

a

»
2

Figures 148-150. Neotriadomerus sp. 4, male (Australia, NT, 12 km WNW Ross River, Tourist Camp).
148 head and anterior mesosoma, lateral 149 mesosoma, lateral 150 antenna. Scale bar for 148, 149 =

200 um; 150 = 400 um.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 65

— ames r WI

Pe Ss ae" ==
peace FS ee

rn
Iie Oe ee eae yy |
Yap iri G)
er re aes

Figures 151-152. Neotriadomerus sp. 4, male (same data as 148-150). 151 wings 152 metasoma,
lateral. Scale bar = 300 um.

longest marginal setae 64. Metasoma. Gaster length 517, with setae present on sterna
but fewer than on terga (Fig. 152). Genitalia length 220 (Fig. 152).

Comment. The flagellar segments are wider than those of NV. darlingi (Figs 82,
952), NV. longiovipositor (Fig. 121), and N. powerae (Figs 129, 138) and WN. sp. 1; the
number and distribution of mps are most similar to NV. darlingi or N. powerae and the
propodeum has 3 setae as in NV. powerae. The condensed description given above may
help associate the male with a conspecific female when they are collected, preferably
together with more males.

66 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Proarescon Huber, gen. n.
http://zoobank.org/9D52630E-F7D4-4DE0-A3D 1-0469F28531E2
Figs 153-176

Type species. Borneomymar primitivum Huber, by present designation.

Diagnosis. Female. Antenna with funicle 8-segmented (in Avescon 5-segmented)
and clava 1-segmented, gradually narrowing apically to a point (Figs 156, 166). Both
sexes. Fore wing with microtrichia more densely spaced except for oval area along
posterior margin (in Avescon with microtrichia usualy more sparsely spaced, as shown
in Triapitsyn [2016]).

Description. Female. Body 635—720 in length (critical point dried). Colour. Body
generally light brown with some areas yellow to creamy white; darker brown are mouth
margin, trabeculae, ocellar triangle, clava except apex, dorsellum, meso- and metapleuron,
propodeum, and gt,-gt. (Figs 165, 168, 170). Wings hyaline except for light brown be-
hind venation (Fig. 169 and Huber 2002, fig. 5). Head. Head about 1.50—1.59x as wide
as long, about 1.29—1.35x as wide as high, and 1.17—1.18x as high as long; in lateral view
with anterior surface slightly convex, flat at level of toruli, then evenly curved to mouth
margin; posterior surfaces convex and evenly curved from vertex to mouth margin. Face
about 0.9x as wide as high; subantennal groove absent; preorbital groove ventral to level of
torulus straight then more ventrally curving slightly medially to lateral margin of mouth
opening (Figs 153, 171—male). Torulus in slight triangular depression about 1.7x as high
as torulus width and separated by about 2.0x its width from transverse trabecula (Fig.
171—male). Vertex in lateral view horizontal, forming a right angle with face, posteriorly
almost at right angle with occiput and separated from it by medially divided tranverse
vertexal suture extending behind posterior ocelli almost from eye to eye; occiput separated
from gena by a short, oblique posterior extension of supraorbital suture extending from
lateral apex of vertexal suture and curving ventrally to dorsolateral corner of occipital fo-
ramen. Ocellar triangle small, slightly raised, with mid ocellus almost vertical and lateral
ocelli oblique and facing posteriorly; ocelli with POL about 1.0x LOL and about 0.67x
LOL,; ocelli on stemmaticum (Fig. 154)—-seen as white lines in cleared slide mounts (Fig.
165)—these are, respectively, a short, transverse groove in front of mid ocellus, continuing
anterolaterally as the frontal suture to midpoint of supraorbital trabecula (apparently di-
vided medially by an unscletotized area), a groove between the lateral margins of mid and
lateral ocelli—the frontofacial suture, and a medially divided transverse groove behind the
lateral ocelli, the vertexal suture, extending almost from eye to eye. Transverse trabecula
darkly sclerotized medially and at each apex apparently not separated from supraorbital
trabecula; preorbital trabecula extending ventrally about halfway between dorsal and ven-
tral margins of torulus to where torulus nearest to eye; supraorbital trabecula in 2 almost
equal sections, the anterior sections diverging posteriorly, the posterior sections parallel
(Fig. 165). Eye large with numerous small facets, in lateral view about as high as wide and
clearly separated dorsally from back of head (temple about 0.3x eye width). Ocular ap-
odeme long and straight, needle-like. Malar sulcus present. Gena at level of ventral margin
of eye slightly wider than malar space. Occiput separated from temple by occipital groove

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 67

(Fig. 165) but otherwise not separated from gena/postgena. Mouthparts. Labrum with 1?
seta; mandible with 4 uneven teeth (Fig. 153). Antenna. Scape 3.44.7 as long as wide,
with radicle distinct from rest of scape and about 0.36—0.37x total scape length; pedicel
about 2.0x as long as wide, almost as wide as and 0.26—0.27x as long as entire scape; fu-
nicle 8-segmented; clava 1-segmented, 0.98—1.10x as wide as apical funicle segment and
0.41—0.63x as long as entire funicle (Figs 156, 166, and Huber 2002, fig. 6). Mesosoma.
About 1.7x as long as wide, 1.8x as long as high and 1.2x as wide as high. Pronotum en-
tire (Fig. 157), in dorsal view clearly visible, medially about 0.5x as long as mesoscutum,
with collar bell-shaped in lateral view pronotum sloping down towards junction with
head and neck almost absent (not separable), and lateral panel somewhat rectangular and
overlapping anterior margin of mesoscutum, with lateral surface merging smoothly into
dorsal surface, with a shallow, oblique groove for femur. Spiracle (Fig. 157) flat with sur-
face of pronotum, facing posterodorsally, and apparently slightly closer to anterior apex
of notaulus than to posterolateral angle of pronotum. Propleura near anterior apex not
quite abutting, then gap widening slightly more anteriorly. Prosternum rhomboidal and
completely divided medially by faint longitudinal groove. Mesoscutum about 1.8x as long
as scutellum, in dorsal view with shallow, thin, slightly diverging notauli a little wider and
shallower posteriorly (Figs 157, 158, 168), in lateral view almost flat. Scutellum slightly
wider than long, the anterior scutellum about 0.9x as long as frenum and separated from
it by a shallow, medially straight frenal depression (Fig. 157); campaniform sensilla about
as far from each other as to lateral margin of anterior scutellum; fenestra small, almost
circular, and posterior to campaniform sensilla (Fig. 168, inset). Axilla distinctly advanced,
the transscutal articulation laterally forming a distinct angle with median section (Fig.
168); axillula short, separated from anterior scutellum by concave axillular groove; meso-
phragma fairly narrowly convex posteriorly, extending to posterior apex of propodeum.
Prepectus apparently narrowly triangular; mesopleuron somewhat rectangular, with shal-
low depression separating mesepisternum from mesepimeron. Metanotum with distinct
triangular (Fig. 159) or lens-shaped (Fig. 168, in cleared slide mounts) dorsellum and
lateral panel length toward hind wing articulation about one third length of dorsellum.
Metapleuron triangular, the margin at junction with mesopleuron almost straight and
posterior margin straight and vertical. Propodeum without carinae, with 1 propodeal seta
(Fig. 159). Wings. Fore wing wide, with microtrichia on most of membrane beyond and
partly behind venation to level of second macrochaeta except for a bare area medially
along posterior margin (Fig. 169). Venation complete; submarginal vein with 1 proximal
macrochaeta but no distal seta; parastigma 0.73x submarginal vein length; marginal vein
present, its length about 1.42x parastigma length, with a second macrochaeta about mid-
way between first distal macrochaeta and stigmal vein; stigmal vein distinct, about 0.28x
length of marginal vein, with anterior margin of stigma parallel or converging with wing
margin and with 4? apical campaniform sensilla in a line; postmarginal vein present, ap-
parently about 1.1x as long and almost as thick as marginal vein; hypochaeta fairly close
(about 0.3x length of parastigma) to proximal macrochaeta; proximal campaniform sen-
sillum near posterior margin of parastigma just next to first distal macrochata. Hind wing
normal (Fig. 169). Legs. Profemur slightly wider than meso-and metafemora; metafemur

68 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

about 1.2x mesofemur width. Tarsi 5-segmented. Calcar (moveable protibial spur) with
about 2 setae along outer margin, and with inner tine about 0.45x outer tine length. Mid-
dle and hind legs with tarsomere 1 as long as tarsomere 2. Metasoma. 1.95x as long as
wide, 2.18x as long as high and 1.12x as wide as high; its length, excluding exserted part of
ovipositor, about 1.37x that of mesosoma. Petiole ring-like, about 0.36x as long as wide.
Gastral terga about equal in length except gt, slightly longer (Figs 162, 170). Cercus flat,
with 4 setae, the second-most dorsal one longest (Figs 162, 170). Hypopygium short, ex-
tending about one-third ovipositor length (Fig. 170). Ovipositor sheath exserted beyond
gastral apex by about 0.2x total sheath length, with 1 subapical seta.

Male. Body length ~ 585 (slide mounted paratype). Colour. Similar to female
but with slightly more extensive brown on mesosoma (Fig. 173), and metasoma with
brown apically instead of medially (Fig. 175). Head. As for female, mandible with 4
teeth (Fig. 171). Antenna. Scape (n=1) about 3.00x as long as wide, with radicle about
0.35x scape; pedicel 1.28x as long as wide; flagellum 11-segemented, with fl, shorter
and wider than other segments, fl —fl,, subequal, each flagellomere with 4 mps (Fig.
172). Mesosoma. As for female (Figs 159, 160, 173). Wings. Fore wing (Fig. 174)
with proximal campaniform sensillum near posterior margin of marginal vein about
midway between first and second distal macrochata. Metasoma. Petiole length/width
0.38; gaster about 0.82x as long as mesosoma (Fig. 162). Genitalia with long para-
meres and apparently no digiti (Figs 163, 164, 176).

Etymology. The genus is masculine. The prefix, pro- is Latin for in front of, earlier
or first, + Arescon, apparently the most closely related genus.

Key to species of Proarescon. Females.

1 Clava 4.7x times as long as wide, with ventral margin distinctly concave
(Figs 156, 166); fore wing with cubital line extending proximally to about
level of second distal macrochaeta (Fig. 169), not much further towards wing
base than remaining microtrichia on wing surface.... P. similis Huber, sp. n.

= Clava less than 3.5x as long as wide, with ventral margin almost straight
(Huber 2002, fig. 6); fore wing with cubital line extending proximally almost
to level of proximal macrochaeta, distinctly further towards wing base than
remaining microtrichia on wing surface (Huber 2002, fig. 5)... cesses
Cee Fe em en ter, ea ae ene ee P. primitivus (Huber)

Proarescon primitivus (Huber), comb. n.

Figs 153-176

Borneomymar primitivum Huber, 2002: 49 (description, figs 5, 6).

Comment. In Huber (2002) the female and male symbols in the material examined
sections were inadvertently either deleted or were replaced by a question mark. All

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 69

the specimens of the three species mentioned in that chapter were females except
for 2 males on points and 2 males on slides of Borneomymar discus Huber from
Malaysia, Sarawak, Gunung Buda. ‘The brief description below complements the
original description, which did not include antennal segment widths or hind wing
measurements.

Description. Female. Body length 634 (paratype). Antenna. Length/width meas-
urements (holotype): scape 184/35, pedicel 50/26, fl, 16/13, fl, 16Al2; fl, 23/16, fl,
26/16, fl. 29/18, fl. 62/25, fl 59/26, fl, 63/31, clava 120/35.Wings. Fore wing length
583, width 154, length/width 3.78, longest marginal setae 122. Hind wing length
560, width 23, longest marginal setae 90.

Material examined. THAILAND: Chanthaburi, Khao Kitchakut Nat. Park,
Khao Prabaht Peak, 12°50.45'N 102°9.81'E, 875m, 27.ii-6.1i.2009, MT, Suthida and
Charoenchai, #4046 (12, CNC).

Proarescon similis Huber, s

http://zoobank. ore/7CEE8233- "EC3D- 4B9D-AB79-CCEFA137E27D
Figs 153-176

Type material. Holotype female (QSBG) on slide (Fig. 167) labelled: 1. “Proarescon si-
milis Huber 9 dorsal Holotype”. 2. “Thailand: Surat Thani, Khao Sok Nat. Park, Klong
Morg Unit, 87 m, 8°53.725'N 99°38.025'E, MT 10-17.ii.2009, Phongphan, #3937”.

Paratypes. 1 female and male. THAILAND. Nakhon Si Thammarat. Namtok
Yong Nat. Park, TV aerial, 966m 8°14.262'N; 99°48.289'E, 15-22.ix.2008, Malaise
trap, Paiboon, #3540 (14 on slide, QSBG); Namtok Yong Nat. Park, Klong Jang
waterfall, 8°16.283'N; 99°38.702'E, 154m, Malaise trap, 23.ii-2.iii.2009, S. Sam-
naokan, #4665 (13, CNC). Surat Thani. Khao Sok Nat. Park, 122m, 8°54.555'N;
98°30.522'E, Malaise trap, 13-20.i1.2009, Pongphan, #3914 (12, CNC).

Diagnosis. Female. Clava 4.7x times as long as wide, with ventral margin dis-
tinctly concave (in P primitivus, clava about 3.4x as long as wide, with ventral margin
almost straight); fore wing with cubital line not extending proximally beyond first api-
cal macrochaeta, barely proximal to remainder of microtrichia (in P primitivus, cubital
line extending proximally almost to level of proximal macrochaeta, distinctly proximal
to other microtrichia).

Description. Female. Body length 740 (holotype), 634 (paratype on card). Head.
Head width 182 (holotype). Face with weak elongate reticulate sculpture, vertical lat-
erally becoming horizontal medially, with thin setae distributed on each side as follows:
2 medial to torulus and 7 ventral to torulus, the 2 setae submedially above mouth
margin longer and thicker than the others (Fig. 153). Vertex with transverse elongate
reticulate sculpture, with 1 or 2 setae on orbit of eye lateral to transverse trabecula and
2 setae between eye and posterior section of supraorbital trabecula and 1 seta lateral
to posterior apex of vertexal suture; ocellar triangle (inside stemmaticum) with 1 seta
between mid and lateral ocelli and 1 setae lateral to mid ocellus but outside triangle

70 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Figures 153-158. Proarescon similis. 153 male head, ventroanterior 154 female head and pronotum,

posterodorsal 155 male lower face and prosternum 156 female antennae 157 female mesosoma, dorsal

158 male mesosoma, dorsolateral. Scale bar for 153, 154, 157, 158 = 50 um; 155= 20 um; 156 = 100 um.

(Fig. 165). Malar space with 2 setae. Occiput with engraved transverse reticulate sculp-
ture, with 1 short setae submedially and 1 sublaterally above occipital foramen. Gena
with longitudinally reticulate sculpture laterally, becoming isodiametric around fora-
men, and with about 5 setae sublaterally and laterally. Antenna. Fl —fl, without mps,
Fl —fl, each with 2 mps; clava with 6 mps. Length/width measurements: scape 166/-,
pedicel 39/-, fl, 11/12, A, 13/12, A, 12/13, fl, 15/14, A, 18/16, A, 50/16, A, 51/21, A
54/31, clava 143/30. Mesosoma. Width 156 and length 287. Pronotum with raised

transverse to isodiametric reticulate sculpture, with 3 long setae near posterior margin

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 71

Figures 159-164. Proarescon similis, male. 159 mesosoma, dorsal 160 base of wings + axilla, dorsal
161 fore wing, dorsal 162 mesosoma, laterodorsal 163 apex of gaster + genitalia, dorsolateral 164 apex of
gaster + genitalia, ventral. Scale bar for 159 = 50 um; 160, 163, 164 = 20 um; 161 = 200 um; 162 = 100 pm.

and 2 short setae towards anterior margin. Propleuron faintly, longitudinally reticulate,
with 1 seta medially and 1 seta laterally about midway between anterior and posterior
apices. Prosternum apparently smooth, with 1 seta submedially almost at anterior mar-
gin. Mesoscutum with sculpture isodiametric anteriorly and on lateral lobes to longi-
tudinally reticulate posteriorly on midlobe, with 1 setae along inner margin midway
between anterior and posterior apices of notaulus, and 1 setae at lateral angle of lateral
lobe. Anterior scutellum smooth, with 1 setae laterally at dorsal margin of axillula,
and with campaniform sensilla about 2x their diameter from transscutal articulation

72 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

TRAILANO? Surat
Thani, khao Sok
Nat. Pack, klong
Morg Unit, S%m
8° $3.226"

993. 2-025 EMT
Jo-)F.11.2 004
Phenarhan 313? F

Figures 165-167. Proarescon similis, holotype. 165 head, dorsal 166a antenna without scape, lateral
166b antenna, dorsal 167 type slide. Scale bar = 100 um.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... i)

Figures 168, 169. Proarescon similis, holotype. 168 metasoma, dorsal (inset: enlargment of central area

of scutellum showing fenestra) 169 wings. Scale bar = 100 pm.

74 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Figures 170-172. Proarescon similis. 170 holotype metasoma, lateral 171 male paratype head, anterior
172 male paratype antenna. Scale bar for 170, 172 =200 um; 171 =100 um.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... ia)

Figures 173, 174. Proarescon similis, male paratype. 173 mesosoma, dorsal 174 wings. Scale bar =
100 pm.

76 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Figures 175, 176. Proarescon similis, male paratype. 175 metasoma, dorsal 176 genitalia, dorsal (seen
through body). Scale bar for 175 = 100 um; 176 = 50 pm.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... Fal.

and about 7x their diameter from each other; fenestra small, transversely oval and pos-
terior to campaniform sensilla; frenum with faint, longitudinal wrinkles. Axilla with
isodiametric reticulations and with 1 seta dorsally on apparently smooth lateral panel;
axillula smooth. Metanotum smooth with 1 short seta on anterior margin closer to
lateral apex of lateral panel than to dorsellum and 2 minute seta on anterior margin
at apex of lateral panel. Propodeum apparently smooth medially, with faint engraved
longitudinal reticulations laterally, with 1 propodeal setae midway between spiracle
and posterior margin. Wings. Fore wing length (n=1) 563, width 146, length/width
3.86, longest marginal setae 107; cubital line of setae extending halfway towards base
of parastigma, about midway between level of proximal and distal macrochaetae. Hind
wing length 542, width 26, longest marginal setae 104. Legs. Femora and tibiae of all
legs at most with faint longitudinal reticulation. Protibia with about 14 short, thick
pegs along its length. Metasoma. Petiole width 26, length 11. Gaster (Fig. 170) height
155, length 352, apparently without sculpture (Figs 162, 163) and with 2 short setae
on gt, and 3-6 longer lateral and dorsal setae on remaining terga. Cercus with 4 setae,
the first seta about 2x as long as the remaining setae. Hypopygium about 0.35x gaster
length. Ovipositor 380 long, its exserted part (56) about 0.2x entire sheath length,
with 1 subapical seta.

Male. Colour. As in generic description. Head. Head width 192 (n=1). Wings.
Fore wing length (n=1) 563, width 146, length/width, 3.88, longest marginal setae
107. Hind wing length 542, width 26, longest marginal setae 114. Antenna. Measure-
ments (n=1): scape length/width 74/22, pedicel length/width 30/37, flagellar segment
length: fl, 44, fl, 59, fl, 62, fl, 61, fl, 60, fl, 60, fl, 64, A, 61, fl 61, fl,, 62, fl, 58; total
flagellar length 652; fl, length/width 2.73, with 4 mps (Fig. 172). Metasoma. Gaster
(Fig. 175) apparently without sculpture, with setae dorsolaterally and laterally and
a few minute spicules medially on gt, and gt.. Genitalia (Figs 163, 164, 176) with
curved aedeagal apodeme about 0.6x as long as aedeagus; paramere thin, with 3 sub-
apical and apical setae.

Etymology. ‘The species name, similis, is Latin for similar, referring to the similar-
ity of this species to the only other described species in Proarescon.

Borneomymar Huber

Figs 176-187

Type species. Borneomymar discus Huber 2002: 45, by original designation.
Diagnosis. Female. Head without stemmaticum (Fig. 178); occipital groove
almost transverse (Fig. 178). Antenna with funicle 8-segemented, only fl,-fl, with
mps, and clava 1-segmented (Fig. 179); mandible with 3 teeth; fore wing length
1006, width 235, length/width 4.28, longest marginal setae 203 (Fig. 180); venation
= 0.75x wing length (apex of postmarginal vein difficult to determine); submarginal
vein 2.03 x as long as parastigma, parastigma with hypochaeta much closer to prox-
imal than to distal macrochaeta; marginal vein 1.94x as long as parastigma; postmar-

78 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

,

unrolled tran

Figures 177-180. Borneomymar madagascar, female. 177 head, anterior (vertex detached anteriorly,
with trabeculae unrolled) 178 head, posterior 179 antenna 180 wings. Scale bar for 177, 178 = 100 um;
179, 180 = 500 um.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 79

Figures 181-183. Borneomymar madagascar, female. 181 body (without head), habitus, dorsal
182 mesosoma, dorsal 183 metasoma, dorsal (slightly twisted). Scale bar for 181 = 1000 pm; 182, 183 =
100 pm.

80 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Figures 184-187. Borneomymar madagascar, male. 184 head, dorsal 185 antenna, dorsal, twisted lat-
erally after fl, 186 metasoma (gt, missing) 187 genitalia, dorsal. Scale bar for 184, 187 = 100 um;
185, 186 = 200 um.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 81

ginal vein = 0.95x as long as marginal vein; hind wing length 648, width 32, longest
marginal setae 122; ovipositor extremely long, projecting posterior to apex of hypopy-
gium by ~ 2.2—2.4x body length (Fig. 181). Male. Similar to female except for gaster
(Fig. 186) and antenna. Head (Fig. 184). Fore wing length 1134, width 303, length/
width 3.74, longest marginal setae 220; hind wing length 797, width 32, longest
marginal setae 32. Antenna with flagellum 11-segmented, all segments with several
mps (Fig. 185); genitalia (Fig. 187), encapsulated, with long aedeagal apodeme and
digitus with 3 teeth.

Borneomymar madagascar Huber

Figs 176-187

Borneomymar madagascar: Huber, 2002: 48 (original description); Engel et al. 2013: 2
(comparison with a fossil species).

Diagnosis. This species differs from, B. discus Huber, the only other extant species now
included in the genus, as follows: radicle about as long as wide, scape 2.84x as long
as wide (in B. discus radicle much longer than wide, scape 5.86x as long as wide); fore
wing uniformly hyaline (Fig. 180) (in B. discus fore wing with distinct brown areas);
gaster in lateral view about as 0.5x as high as long (almost circular in B. discus).

Description. Female. Body length 922-998 (n = 4, critical point dried specimens,
measured to apex of hypopygium), ovipositor length ~ 2022-2330 (not completely
straight so probably slightly longer). Head (Fig. 184) width 266. Antenna measure-
ments (n = 1), length/width (ratio): scape 101/36 (2.84), pedicel 56/35 (1.59), f,
68/20 (3.35), fl, 8.17/30:(2-73), fl, 82729: (2813 fl 86/30 (2.85), fl. 79/32 (2.43), fl,
76/34 (2.25), fl, 72/32 (2.27), fl, 74/34 (2.20), clava 176/52 (3.40); ffl, each ap-
parently with 3 mps, clava with 12 mps (Fig. 179; hypopygium extending distinctly
beyond posterior apex of gaster. Male. Body length ~ 1315 (excluding gt, removed
to extract genitalia). Head (Fig. 184) width 262. Antenna measurements (n= 1, slide
mounted), length/width [scape—fl, are vertical so width cannot be measured]: scape
104/ - , pedicel 56/ -, fl, 106/ -, ,108/ -, fl, 106/ -, A, 108/- , , 111/~ 35, fl, 101/38,
fl 92/42, fl, 96/41, fl, 94/41, fl, 91/39, Be 88/35; total flagellar length 1102; fl.
length/width 2.66, with 9? mps.

Material examined. MADAGASCAR. Antananarivo. Botanic garden near en-
trance to Andasibe National Park, 1025 m, 18°55.58'S; 48°24.47'E, 1-5.xi.2001, R.
Harin Hala, tropical forest, Malaise trap, lot # 007164 (12 on slide, CAS). Diego-Suar-
ez. Montagne d’Ambre National Park, 975 m, 12°31'S; 49°11'E, 4-19.iii.2001, M.
Irwin, E. Schlinger, R. Harin’Hala, Malaise trap, lot # 007149 (13 on slide, CAS).
Fianarantsoa. Massif de Andringitra, 975m, 21°57'50"S; 46°55'59"E, 7-19. xii.1999,
M.E. Irwin et al., Malaise trap (49. on cards, CNC, UCRC).

82 John T. Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Tribe Triadomerini stat. n.

Notes. Yoshimoto (1975) treated the Cretaceous amber fossils from Canada and classi-
fied Triadomerus Yoshimoto in his new subfamily Triadomerinae, Carpenteriana Yoshi-
moto and Macalpinia Yoshimoto in Mymarinae, and Enneagmus Yoshimoto in Tricho-
grammatidae. He defined his Triadomerinae as “having the submarginal and marginal
veins distinctly separated at the junction of the fore wing and antennae 13-segmented
in the female”. Poinar and Huber (2011) keyed the Cretaceous genera of Mymaridae,
adding only Myanmymar Huber from Burmese amber to bring the number of genera
to five, including Enneagmus, which Huber (2005) had reclassified in Mymaridae.

I treat Yoshimoto's Triadomerinae as a tribe, defined as follows, based mainly on
extant genera and species: mandible with 3 (Borneomymar, Eustochomorpha) or 4 teeth;
pronotum entire; fore wing wide, with marginal setae much shorter than wing width;
venation more than 85% of wing length, with marginal vein present and longer than
submarginal vein, and with postmarginal vein present and longer than marginal vein;
hypochaeta, when present, closer to proximal than to distal macrochata; hind wing
wide with marginal setae shorter than wing width; tarsi 5-segmented, with tarsomere
1 distinctly longer than any of the others; petiole clearly shorter than wide, ring like.
Female. Antenna with flagellum at most 11 segmented (funicle 8-segmented and clava
1—3-segmented); ovipositor usually greatly exserted beyond either posterior (Borneo-
mymar, Eustochomorpha) or anterior (Neotriadomerus) apex of body but in the extinct
genera not projecting either anteriorly or posteriorly. Male. Antenna 11-segmented,
the flagellomeres each with several mps; genitalia encapsulated, with short, thick para-
meres, apparently without digiti (in Neotriadomerus) but thinner walled and with digiti
(in Borneomymar)

Triadomerini is treated here as the sister clade to the remainder of Mymaridae. The
only apomorphy that defines the tribe is reduction in number of flagellar segments
(at most 11) relative to Rotoitidae, whose species have a 12-segmented flagellum in
females of both included extant genera. An additional diagnostic feature of the extant
species of Triadomerini is the exserted cerci on a distinct prominence, similar to that of
Torymidae. The occurrence of elevated cerci, number of teeth in mandibles, and sev-
eral other features cannot definitely be determined from the fossil specimens studied.
The lack of a hypochaeta apparently occurs in Triadomerus and Eustochomorpha and
apparently also in at least one of the Neotriadomerus species.

Triadomerus is known only from Cretaceous amber from present day western Can-
ada (Yoshimoto 1975). Eustochomorpha and Neotriadomerus species are confined to
Australia, and extant Borneomymar species occur in the islands of Borneo and Mada-
gascar whereas the one extinct Borneomymar species is from Eocene amber from the
Baltic region (Engel et al. 2013). Macalpinia and especially Carpenteriana doubtfully
belong in Triadomerini but perhaps are better classified here than elsewhere. Even
though Carpenteriana has 7-segmented funicle and an entire clava its fore wing vena-

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 83

tion appears to be similar to Macalpinia. Macalpinia has an 8-segmented funicle and
3-segmented clava, as in Triadomerus and Neotriadomerus, so on the basis of this fea-
ture is classified fairly well in the tribe even though it appears to have no postmarginal
vein and apparently only 4-segmented tarsi (tarsi are difficult to see; they may, in fact,
be 5-segmented). Thus, I classify six genera in Triadomerini: four genera definitely—
Borneomymar, Eustochomorpha, Neotriadomerus, Triadomerus, and two genera tenta-
tively—Carpenteriana and Macalpinia.

Tribe Aresconini

Notes. Viggiani (1988) proposed Aresconini for Avescon Walker and defined the tribe
on the characteristic male genitalia, i.e., encapsulated, with relatively long, narrow par-
ameres but apparently no digiti. Based on mainly extant genera and species, Aresconini
is defined by the following features: mandible with 4 teeth; pronotum entire; venation
long but postmarginal vein usually short; funicle in females 4-, 5- or 8-segmented;
clava entire or 2-segmented; male genitalia usually with long parallel-sided parameres. |
classify members of the extant genera Kzkiki Huber and Beardsley (Huber and Beards-
ley 2000), Tinkerbella Huber & Noyes (Huber & Noyes 2013), Proarescon (this paper),
and the extinct genus Myanmymar (Poinar and Huber 2011) in the tribe based on simi-
larities in head features, fore wing shape or venation, and male genitalia where known.

Arescon is almost worldwide (Triapitsyn 2016) whereas Proarescon is only in
southeastern Asia. Lin et al. (2007) and Huber (2009) had first classified Kzkiki
in the Al/aptus group of genera then, with Tinkerbella described as a separate genus
(Huber and Noyes 2013), in the Azagrus group of genera. In both cases, similari-
ties in various features were used to justify their placement. Now, however, I believe
these two genera are best classified in Aresconini. The male genitalia of Kikiki have
fairly long and thick parameres (Huber and Noyes 2013, fig. 35), the vertex has a
distinct stemmaticum (Huber and Noyes 2013, figs 12, 41), the fore wing has a long
venation, and the mandible has 4 teeth, all features that occur also in Avescon and
Proarescon. Kikiki may be almost worldwide (Australia, India, northern South Amer-
ica, Hawaiian Islands) and Tinkerbella is so far known only from Costa Rica (Hu-
ber and Noyes 2013). Enneagmus may fit in Aresconini better than in Triadomerini
because of its apparently short venation, and 4-segmented funicle and 3-segmented
tarsi as in Kikiki. The 3-segmented clava may simply be an ancestral feature, reduced
to two segments in A7zkiki and Tinkerbella. However, the short venation, apparently
much less than half the wing length is unlike the other genera in Aresconini. Thus,
six genera are here classified in Aresconini: Avescon, Enneagmus, Kikiki, Myanmymar,
Proarescon, and Tinkerbella. lf Minutoma Kaddumi (Kaddumi 2005) is indeed a
mymarid, it may also belong here; its small body length is comparable to that of
Tinkerbella.

84 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

Key to extant genera of Aresconini and Triadomerini. Females.

See Poinar and Huber (2011) for key to the extinct Cretaceous genera.

1 Clava 2 or 3-segmented; propodeum with 2 or 3 setae......sceesesceeeseeeeeees 2
- Clava 1-segmented (i.e., entire); propodeum with 1 seta... eee 5
2(1) CEE WORE Teled st Tel vigcra Bey yee aie Ra In re ered ea ara s anita Rem send ny 3

— Clava 3-segmented [ovipositor extending anteriorly well under mesosoma
but not exserted posteriorly much beyond gastral apex (Figs 105, 126)] .......
Cee Bde RANG Oh se hoor ease Min son Meee mayen Ae. Neotriadomerus Huber, gen. n.
3(2) Ovipositor extending posteriorly well beyond posterior apex of gaster (Figs 2, 4);
tarsi 5-segmented; funicle 8-segmented ....... Eustochomorpha haeckeli Girault
= Ovipositor not or barely exserted beyond posterior apex of gaster; tarsi 3- or

A-Seoimeniteds fMiicleA—OF J=SECMIENCE sick sao consceosvasdnctonbdemenstlvemenenstenatleet 4
4(3) Tarsi 4-segmented; funicle 5-segmented........... Tinkerbella Huber & Noyes
- Tarsi 3-segmented; funicle 4-segmented............. kikiki Huber & Beardsley
5(1) Remicle’2sco mi ciated ssc ....00.cavies.pesssate cseoutampaababalhcah naniconns te Arescon Walker
- [Bur attal Lomo iicieree patch ¢\rcca by eB Me Re, Oe OD, Ne A as, PR OO ee, 6
6(5) Fl —fl, each much shorter than fl, —fl, (Fig. 166) .....Proarescon Huber, gen. n.
_ Fl —-fl, as long as fl fl, (Huber 2002, figs 2, 4)... Borneomymar Huber
Discussion

Huber (2002) treated Triadomerus (extinct) and Eustochomorpha (extant) as sister genera to
the remaining extant genera of Mymaridae based, in females (males are unknown), on an
8-segmented funicle and 3- or 2-segmented clava, respectively, i.e., 11 or 10 flagellar seg-
ments in total. In Neotriadomerus, males are known and have the same number of flagellar
segments as in females, i.e., 11, and this is considered to be as the most plesiomorphic
feature possible in Mymaridae. Macalpinia and Triadomerus also have an 8-segmented
funicle and 3-segmented clava. Species in all other genera (all extant only, except for one
fossil Borneomymar species) whose males are known have a different number of segments
between females and males, with at least one less segment in females compared to males.
But because males of the species of most extant genera have an 1 1-segmented flagellum,
it is probably safe to assume that males in the extinct genera also have an 11-segmented
flagellum, as in females. On this basis, 7iiadomerus, Macalpinia and Neotriadomerus are
treated as the sister clade to other genera within Triadomerini and to all remaining genera
of Mymaridae, both extinct and extant. At present the relationships among these three
genera cannot be determined but, among extant genera only, Neotriadomerus is clearly
the sister genus to the all remaining genera, including Eustochomorpha. Loss of one claval
segment in Eustochomorpha, to give a 10-segmented flagellum in its females, makes this
the sister genus to the remaining genera of extant Mymaridae other than Neotriadomerus.

Eustochomorpha Girault Neotriadomerus, gen. n., and Proarescon gen. n.... 85

Conclusions

The distribution of the extinct and extant genera that can definitely be placed in
Mymaridae encompasses a time span of almost 100 my. Kaddumi (2005) classified
Minutoma, found in 120-135 my Cretaceous amber from the Zarga River basin,
in Mymaridae but Huber in Heraty et al. (2013: 2) suggested it might be better
placed in Tetracampidae; its clava appears to be 4-segmented. But if it is a member
of Mymaridae then the family extends back into the early Cretaceous, well after the
beginning of the breakup of Pangea into Laurasia in the north and Gondwana in
the south, beginning 200 million years ago. The present centre of diversity of My-
maridae is in the Australasian region, particularly Australia and New Zealand, but
the fossil evidence shows that the family was already quite diverse in the Cretaceous
and that the origin of Mymaridae is not necessarily there. In Triadomerini, all the
Cretaceous genera (known only from Laurasia) eventually became extinct whereas
Eustochomorpha and Neotriadomerus are extant in Australia (part of Gondwana). Bor-
neomymar, in contrast, is represented by two extant species in the islands of Borneo
and Madagascar (both part of Gondwana) and one extinct species from Baltic amber,
ie., from Laurasia (Engel et al. 2013). In Aresconini, the only extinct genus is Myan-
mymar (Laurasia?); all the rest are extant and, except for Proarescon and Tinkerbella,
widespread.

The widespread distribution of Mymaridae already existed at least 80 my ago
(described taxa from Canadian and Burmese Cretaceous amber) and the family is
currently worldwide, except for Antarctica. Two scenarios may explain this: 1) The
family may have evolved before the breakup of Pangaea, 200-180 million years ago,
and was already widespread throughout the supercontinent wherever suitable hosts
occurred, which would suggest a Jurassic origin; 2) The family is more recent, hav-
ing originated in only part of Pangea, probably Gondwana, and some species then
spread to Laurasia after the two parts became well separated from each other. This
is quite possible because Mymaridae are small and easily carried long distances on
wind and some would have survived the trip. Mymaridae as a recognizable taxon
may therefore be a lot older than the present evidence shows, going back well into
the early Cretaceous.

Acknowledgements

The following curators and collectors are thanked for sending specimens for study: J.
Cardale (ANIC), C. Burwell and G. Daniels (QMBA), C. Darling (ROM), J.E Don-
aldson (QDPC), and S. Triapitsyn (UCRC). I thank particularly J. Read (CNC, now
retired) for proof reading, taking the scanning electron micrographs and digital images,
and arranging them into plates.

86 John T: Huber / Journal of Hymenoptera Research 57: 1-87 (2017)

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