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JHR 49: I-23 (2016) JOURNAL OF *0eerriewed opevaccersjoural
doi: 10.3897/JHR.49.7714 ME Hymenopter a

http://jhr.pensoft.net The inserational Society of Hymenopterists. RESEARCH

Revision of Dvivarnus (Scelionidae, Teleasinae)

Elijah J. Talamas', Istvan Mik6?, Robert S. Copeland?

I Systematic Entomology Laboratory, USDA/ARS clo NMNH, Smithsonian Institution, Washington DC, USA
2 Frost Entomological Museum, Pennsylvania State University, State College, PA, USA 3 International Centre
of Insect Physiology and Ecology, Nairobi, Kenya, and National Museums of Kenya, Nairobi, Kenya

Corresponding author: Elijah J. Talamas (elijah.talamas@ars.usda.gov)

Academic editor: /. Yoder | Received 6 January 2016 | Accepted 21 March 2016 | Published 28 April 2016
http://zoobank.org/3028A70D-A5F5-48C2-85C4-7466F 16E144F

Citation: Talamas EJ, Miké I, Copeland RS (2016) Revision of Dvivarnus (Scelionidae, Teleasinae). Journal of
Hymenoptera Research 49: 1-23. doi: 10.3897/JHR.49.7714

Abstract

Two new species, Dvivarnus elektrolythron Talamas & Miko, sp. n. and D. mikuki Talamas & Mik, sp.
n. are described. The genus is redescribed and a key is provided to separate Dvivarnus from other groups
in Teleasinae with mesoscutellar spines.

Keywords

Teleasinae, Dvivarnus, Trimorus, Gryonoides, mesoscutellum

Introduction

The subfamily Teleasinae is well defined morphologically by wing venation (elongate
marginal vein, short stigmal and postmarginal veins), the anterior pronotal process
and, in most cases, a compact ocellar triangle. Generic classification within the sub-
family is another matter and a thorough phylogenetic analysis is needed. ‘The vast ma-
jority of species are found in Trimorus Forster, a genus whose limits are poorly defined
with respect to many of the smaller genera. Dvivarnus Rajmohana & Veenakumari is
a well defined teleasine genus that morphologically falls well outside of Trimorus and
until now was monotypic.

Copyright Elijah J. Talamas et al. 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 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

We here expand knowledge about the species-level diversity in Dvivarnus with the
addition of two new species. We also provide additional characters to those of Veena-
kumari et al (2011) for its diagnosis relative to two lineages in Teleasinae that also have
mesoscutellar spines, Gryonoides Dodd and the Trimorus carus Nixon species group.
The analysis of Gryonoides follows the examination of 12 species conducted as part of
an active revision of this genus by the second author. Our treatment of the Trimorus
carus species group is based on examination of the holotype of 7: carus Nixon and two
undescribed species from the Central African Republic that share the presence of a
distally bifurcating metascutellar spine.

Materials and methods

The numbers prefixed with “USNMENT” or “OSUC ” are unique identifiers for the
individual specimens (note the blank space after some acronyms). Details on the data as-
sociated with these specimens may be accessed at the following link: purl.oclc.org/NET/
hymenoptera/hol, and entering the identifier in the form. Persistent URIs for each taxo-
nomic concept were minted by xBio:D in accordance with best practices recommended
by Hagedorn et al (2013). Morphological terms were matched to concepts in the Hyme-
noptera Anatomy Ontology (Yoder et al 2010) using the text analyzer function. A table
of morphological terms and URI links is provided in Suppl. material 1.

We represent natural language phenotypes in an Entity:Quality (EQ) format: Entity
attribute: value. Semantic statements of natural language phenotypes (Suppl. material
2.) were composed in Protégé 5.0 (http://protege.stanford.edu/) using the OWL Man-
chester syntax (http://www.w3.org/TR/owl2-manchester-syntax/) following Balhoff et
al (2013) and Miké et al (2014). The full data set, represented in OWL (Web Ontology
Language; http://www.w3.org/TR/owl2-overview/ last accessed February 4, 2014), was
deposited as a Resource Description Framework (RDF)-XML file (http://www.w3.org/
TR/REC-rdf-syntax/ in Figshare (https://dx.doi.org/10.6084/m9.figshare.2008203).

Taxonomic synopses and matrix-based descriptions were generated from the Hy-
menoptera Online Database (hol.osu.edu) and the online program vSysLab (vsyslab.
osu.edu) (matrix title: Revision of Dvivarnus) in the format of character: state. Mul-
tiple states for a character are separated by a semicolon. Characters shared among the
three species of Dvivarnus were exported as the generic description (OTU for generic
characters: Dvivarnus), those that were not shared among all species were exported as
species descriptions.

Photographs were captured with a Z16 Leica lens with a JVC KY-F75U digital
camera using Cartograph software. Single montage images were produced from image
stacks with the program CombineZP. In some cases, multiple montage images were
stitched together in Photoshop to produce larger images at high resolution and mag-
nification. Full resolution images are archived at the image database at The Ohio State

University (http://purl.oclc.org/NET/hymenoptera/specimage).

Revision of Dvivarnus (Scelionidae, Teleasinae) 3

Scanning electron micrographs were produced with a Hitachi TM300 Tabletop
Microscope. The specimen was disarticulated with a minuten probe and forceps and
mounted to 12 mm slotted aluminum mounting stub (EMS Cat. #75220) using car-
bon adhesive tabs (EMS Cat. #77825-12) by means of a fine paint brush and sputter
coated with approximately 70 nm of gold/palladium.

Character annotations

cly clypeus (Figs 1—2)

ctk central keel (Figs 15-16, 18)

epc epomial carina (Fig. 25)

IpT3-T6 lateral patch on T3—T6 (Figs 21-24)
Ipc lateral propodeal carina (Figs 7, 9-10)
mc mesopleural carina (Fig. 43)

mcsp mesoscutellar spine (Figs 7—8)

mees mesepimeral sulcus (Fig. 34)

mmsp median mesoscutellar spine (Fig. 43)
ms marginal setae (Fig. 9)

mns metanotal trough (Figs 5—6)

msct metascutellar spine (Figs 5-8)

nc nuchal carina (Fig. 36)

nes netrion sulcus (Fig. 4)

net netrion (Figs 3-4, 34)

not notaulus (Fig. 46)

ple plica (Figs 6, 9-10)

ppp posterior propodeal projection (Figs 5—6)
psu posterior scutellar sulcus (Fig. 37)
pssu prespecular sulcus (Fig. 34)

r radicle (Figs 1-2, 17)

trt torular triangle (Figs 15—16, 18)
vmc ventral mesopleural carina (Fig. 43)
Specimens

This study is based on specimens from the following collections:

CNCI Canadian National Collection of Insects, Ottawa, Canada

BPBM Bernice P. Bishop Museum, Honolulu, HI, USA

ICIPE International Centre of Insect Physiology and Ecology, Nairobi, Kenya
NMKE _ National Museum of Kenya, Nairobi, Kenya

4 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

i ee

Pal

XN
\

Figure 1-4. | 7vimorus sp., female (OSUC 186090), head, anterior view 2 Gryonoides pulchellus Dodd,
female (USNMENT00872146), head, anterior view 3 Trimorus sp., female (OSUC 192417), pronotum,
anterolateral view 4 Gryonoides glabriceps Dodd, female (USNMENT00872142), pronotum, anterolat-

eral view. Scale bars in millimeters.

OSUC = CA. Triplehorn Collection, The Ohio State University, Columbus, OH, USA
USNM — Smithsonian National Museum of Natural History, Washington, DC, USA

Taxonomy

Dvivarnus Rajmohana & Veenakumari

http://bioguid.osu.edu/xbiod_concepts/305672

Dvivarnus Rajmohana & Veenakumari, 2011: 40 (original description. Type: Dvivar-
nus punctatus Rajmohana & Veenakumari, by monotypy. Diagnosis, keyed).

Description. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla
on A8: 2. Color of radicle: yellow. Length of radicle: shorter than apical width of
clypeus. Length of A3: as long as pedicel or longer. Number of basiconic sensilla on
A12: 1. Number of mandibular teeth: 3. Mandibular teeth: ventral tooth the longest.
Facial striae: present. Dorsal limit of facial striae: facial striae exceeding horizontal

Revision of Dvivarnus (Scelionidae, Teleasinae) 5

plane at margin of anterior ocellus. Torular triangle: present. Height of torular trian-
gle: less than height of clypeus. Central keel: present. Surface of dorsal frons in dorsal
view: convex. Orbital carina: present. Genal patch: absent. Vertex patch: absent. Hy-
peroccipital carina: absent. Anterior margin of occipital carina dorsally: crenulate.
Pronotal cervical sulcus: present. Sculpture of pronotal cervical sulcus: shallowly fo-
veolate. Pronotal suprahumeral sulcus: present. Sculpture of pronotal suprahumeral
sulcus: foveolate. Proximity of suprahumeral and pronotal cervical sulci: pronotal
suprahumeral sulcus terminating before reaching pronotal cervical sulcus. Posterior
pronotal sulcus: absent. Sculpture of propleural epicoxal sulcus: foveolate. Sculpture
of posterior scutellar sulcus: foveolate. Sculpture of mesoscutal suprahumeral sulcus:
foveolate. Length of mesoscutal suprahumeral sulcus: less than one half length the
distance from the tegula to the anterior apex of mesoscutum. Sculpture of mes-
oscutal humeral sulcus: foveolate. Lateral scutoscutellar sulcus: reaching transaxil-
lar carina. Transaxillar carina: present. Setae on lateral margin of mesoscutellum:
present. Posterior scutellar sulcus: present. Acropleural sulcus: present. Length of
acropleural sulcus: elongate. Subalar pit: present. Course of prespecular sulcus and
mesepimeral sulcus: not continuous dorsally. Mesopleural pit: present. Sculpture of
femoral depression: transversely rugose. Mesopleural carina: present. Proximity of
ventral apex of mesopleural carina and ventral mesopleural carina: carinae adjacent.
Sculpture of mesopleural epicoxal sulcus: foveolate. Sculpture of postacetabular sul-
cus: foveolate. Sculpture of mesopleuron below femoral depression: areolate rugose.
Episternal foveae: indistinguishable from surface sculpture. Sculpture of mesepime-
ral sulcus: foveolate. Metascutellar spine: present. Shape of metascutellar spine in
dorsal view: pointed. Length of metascutellar spine: longer than proximal striated
region of metascutellum. Apical semitransparent lamella on metascutellar spine: ab-
sent. Sculpture of metascutellum: longitudinally striate throughout. Proximal stria-
tion of metascutellum: extending onto surface of metanotal spine.Setation of central
propodeal area: present. Posterior propodeal projection: present. Metapleural sulcus:
present. Setation of metapleuron: Area delimited posteriorly by paracoxal and verti-
cal part of metapleural sulcus is covered with dense setae, remainder of metapleuron
glabrous. Sculpture of paracoxal sulcus: foveolate. Dorsal margin of T1 in lateral
view: concave. Length of pits on anterior T1: almost reaching posterior margin of
tergite. [Transverse line of pits on anterior T'1: present. Transverse line of pits on ante-
rior 2: present. Lateral patch on T2: present. Transverse line of pits on anterior T3:
absent. Width of T3: as wide or slightly wider than mesoscutum. Sculpture of T3:
punctate. Length of apical setae on T3: not longer than non-apical setae. Posterolat-
eral patch on T3: present. Lateral patch on T4: present. Lateral patch on T6: present.
Transverse line of pits on anterior S2: present. Felt field on S2: present. Transverse
line of pits on anterior S3: absent.

Diagnosis. Per the characters presented by Veenakumari et al (2011), Dvivarnus
can be differentiated from other teleasines by the combination of the dense punctation
found throughout T3 and S3, the presence of paired mesoscutellar spines, the absence
of lateral propodeal carina and the presence of an inverted U-shaped carina dorsally

6 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

surrounding the metasomal depression. Punctation on T3 can be found in some spe-
cies of Trimorus (Fig. 20), but the punctation is surrounded by rugulae of varying in-
tensity. In Dvivarnus, the punctation is uniform throughout most of the tergite and is
not accompanied by additional sculptural elements. Specimen USNMENT01109195
(Fig. 20) also has spines derived from the metapleural carina, which are not present in
Dvivarnus. Additional characters for the identification of Dvivarnus are presented in
the key to teleasines with mesoscutellar spines.

Comments. The species of Dvivarnus are extremely similar in most pleural char-
acters and differ primarily by features of the head, pronotum, and metasoma. Sexual
dimorphism is exhibited mostly in the pattern of setation and striation of the frons. In
males, the glabrous area above the interantennal process is less distinct and the density
of setation throughout the frons varies greatly. The facial striae in males extend dorsally
throughout the frons whereas in females the striation is absent from the center portion
of the frons.

We examined two morphospecies of males that we were unable to unambigu-
ously associate with the female of D. elektrolythron. One morphospecies (USN-
MENT01109164, Figs 46-50) shares with D. elektrolythron the pattern of striation on
the lateral pronotum (Fig. 50) and the longitudinal furrow on the metanotal trough
(Fig. 48). However, it has distinct notauli (Fig. 46) and D. elektrolythron has none, and
the posterior margin of the mesoscutellum between the mesoscutellar spines is con-
cave in USNMENT01109164, and medially pointed in D. elektrolythron. The other
morphospecies (USNMENT01109212, Figs 40-45) has the opposite arrangement of
characters: it shares with D. elektrolythron the absence of notauli and the presence of
a pointed posterior margin of the mesoscutellum (Figs 40, 43) but it has a foveolate
metanotal trough (Fig. 43) and the lateral pronotum is predominantly smooth (Fig.
45). In the absence of additional specimens that would allow us to thoroughly assess
intraspecific variability in males, or molecular or biological data, we consider it best
to document the morphology of these males and present them as undetermined at the
species-level.

Key to teleasines with mesoscutellar spines

i Radicle shorter than distal width of clypeus (Figs 15-19); lateral propodeal
carina absent (Figs 36, 43, 48); metasomal depression surrounded dorsally by
an inverted U-shaped carina (Figs 48, 43, 36); T3 and S3 densely punctate
(Figs 21-23, 25, 29); metascutellar spine longitudinally striate throughout
(Figs 5—6, 31, 36-37; T3 with lateral patch (Figs 21—23) .......0... Dvivarnus

- Radicle as long as distal width of clypeus (Figs 1-2); lateral propodeal carina
present (Figs 9-10); metasomal depression not surrounded dorsally by con-
tinuous carina (Figs 9-10); [3 and S3 not densely punctate (Figs 12, 14, 24),
or punctation with interstitial rugulae (Fig. 20); metascutellar spine smooth

in distal half (Figs 7-8); T3 without lateral patch (Figs 12, 14, 24)... 2

Revision of Dvivarnus (Scelionidae, Teleasinae) es

2 Metasomal depression without marginal setae (Figs 8, 10); netrion sulcus
distinct (Fig. 4); apex of metascutellar spine unbranched (Figs 8, 14) ...........
se tae ate veh te Uae Bip a ee Wed ae che Oa cau Ron cevetic cused Raveeeesta tb gedtdere Gryonoides

= Metasomal depression with marginal setae (Figs 7, 9); netrion sulcus not vis-
ible in lateral view (Fig. 3); metascutellar spine with bifurcate apex...............
Bo catty sviste uesletoacioaistcnrherteree crete teonrtreeesreermanctsmey Trimorus carus species group

Key to species Dvivarnus (females)

1 Frons with central keel separate from carinae of torular triangle (Fig. 15);
mesoscutellum with posterior margin between mesoscutellar spines medially
convex in dorsal view (Fig. 28, 31); pronotum with posterior portion trans-
versely striate (Fig. 30); lateral patch on T4 present as a dense tuft of setae
Pig. -22)5. ts wwith-lateral patch preserve Pig. 22). wens thacth.sesercasiaisceadeniocsnbeadcse
e Falt dh dict omni eater Sema diet Dvivarnus elektrolythron Talamas & Miko, sp. n.

= Frons with central keel extending to torular triangle (Figs 16-18); mesoscu-
tellum with posterior margin concave between mesoscutellar spines in dor-
sal view (Figs 26, 33, 36); pronotum with posterior portion predominantly
smooth (Figs 25, 32, 34-35); lateral patch on T4 broad (Figs 39, 21, 23); T5
Withoutedlateral patch sti 24.525) seer ere sete snteree eateresnendlab vate onene se case Z

2 Torular triangle setose (Figs 16-17); sulcus in metanotal trough foveolate
(Fig. 5); lateral face of pronotum with epomial carina (Fig. 25) wo... eee
ah LNT: eee A Aten» ET Dn Sy. Dvivarnus agamades (Kozlov & Lé)

= Torular triangle without setation (Fig. 18); sulcus in metanotal trough pre-
sent as an elongate furrow (Figs 6, 36-37); lateral face of pronotum without
epomial carina (Fig. 34)... Dvivarnus mikuki Talamas & Mik6, sp. n.

Dvivarnus agamades (Kozlov & Lé)
http://bioguid.osu.edu/xbiod_concepts/343746
Figures 5, 16-17, 23, 25-26

Gryonoides agamades Kozlov & Lé, 1986: 100 (original description); Lé, 2000: 218
(description, type information).

Dvivarnus punctatus Rajmohana & Veenakumari, 2011: 44 (original description);
Talamas & Buffington, 2014: 104 (junior synonym of Dvivarnus agamades
(Kozlov & Lé)).

Dvivarnus agamades (Kozlov & Lé): Talamas & Buffington, 2014: 104 (description,

synonymy).

Description. Whorl of setae on flagellomeres in male: absent. Shape of A3—A11 in
male: cylindrical.

8 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

=~ 2 AMS
: oN ; f ~ P A
~ ) nnd | 1j if ,
——~ | y > 4“ ppp ih
= ‘ \
> as [oe

Figure 5-10.5 D. agamades, female (USNMENT01109190), mesosoma, posterolateral view 6 D. mikuki,
female (USNMENT01109213), mesosoma, posterolateral view 7 Trimorus sp., female (OSUC 186090),
posterior mesosoma, dorsolateral view 8 Gryonoides glabriceps Dodd, female (USNMENT00872142),
posterior mesosoma, dorsolateral view, anterolateral view 9 Trimorus sp., female (OSUC 1924417),
mesosoma, posterior view 10 Gryonoides glabriceps Dodd, female (USNMENT00872142), mesosoma,

posterior view. Scale bars in millimeters.

Color of antennae in female: Al—A2 orange, otherwise brown. Color of meso-
soma: dorsal mesoscutellum black, otherwise orange. Color of head: black. Number of
labial palpomeres: 1. Number of maxillary palpomeres: 4. Setation of torular triangle:
present. Continuity of torular triangle and central keel: torular triangle closed dorsally,
continuous complete central keel. Color of interantennal process: yellow. Setation of

Revision of Dvivarnus (Scelionidae, Teleasinae) 9

phe BE

Y= = aoe — ys SS ren Ss S&S é = 3
Figure 11-12. Il Zrimorus sp., female (OSUC 186090) head, mesosoma, metasoma, lateral view

12 Trimorus sp., male (OSUC 345677), head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

frons: transverse strip directly above interantennal process glabrous, otherwise setose.
Sculpture of dorsal frons: dorsoventrally strigose. Sculpture of vertex: rugulose; very
finely punctate. Epomial carina: present. Netrion in lateral view: present only at ventral
limit of pronotum. Sculpture of vertical face of pronotum: smooth. Ventral propleural
area: smooth. Sculpture of propleural cervical sulcus: foveolate. Sculpture of mesoscu-
tum: punctate to finely areolate, coarser in posterior half. Notaulus: weakly indicated
posteriorly amid sculpture. Sculpture of scutoscutellar sulcus: smooth; smooth along
posterior margin of mesoscutum, anterior margin of mesoscutellum with ridges later-
ally. Orientation of transaxillar carina: parallel to longitudinal axis of body. Sculpture
of mesoscutellum: punctate to areolate; rugose. Density of setae on lateral margin of
mesoscutellum: dense. Posterior margin of mesoscutellum: concave between mesoscu-
tellar spines. Median mesoscutellar spine: present. Lateral extreme of posterior scutos-
cutellar sulcus: foveae extending to axillula. Sculpture of metanotal trough: foveolate.
Sculpture of lateral propodeal area: irregularly rugose. Lateral propodeal carina: pre-
sent. Plica: present. Forewing pattern in female: wing uniform in color. Forewing color

10 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

es ; S a | \
Hest. x Sing _ o : : ; \ : o ’
a = ee “ a —— — * ~~ SS ae ata ions a. fs
fen scat a, Z Sa ee ar ee ee
14 se - ae Sng ie a
P \ a wa hy A “ 4 . * ray

Figure 13-14. Gryonoides glabriceps, female (USNMENT00872142), 13 head, mesosoma, metasoma,

lateral view 14 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

in male: uniform throughout. Sculpture of metapleural sulcus: smooth. Sculpture of
dorsal metapleural area: smooth; transversely rugose. Sculpture of ventral metapleural
area: transversely rugose. Number of setae on lateral T1: 2; 3; 4; many. Sculpture of
T4: smooth. Lateral patch on T5: present.

Diagnosis. Dvivarnus agamades can be separated from D. elektrolythron and D.
mikuki by the foveolate metanotal trough (Fig. 5). Females may be separated on the
basis of the non-patterned forewing.

Link to distribution map. http://hol.osu.edu/map-large.html?id=343746

Material examined. Paratype: VIETNAM: Gia Lai Prov., K’Bang Dist., rice /
sweet potato, Budén Ludi, 26.X1.1978, X. H. Lé (1 male, OSUC 184258 (ZIN)).
Other material: (19 females, 8 males) INDIA: Delhi Union Terr., Indian Agricul-
tural Research Institute (IARI) area, New Delhi, 26.X.1979, Boucek (1 male, USN-
MENT01109192 (CNCI)). Karnataka St., Bangalore, 21.VI-30.VI.1987, K. Ghor-
pade (1 female, USNMENT01109190 (CNCI)). Karnataka St., Indian Council of
Agricultural Research (ICAR), Bangalore, XII-2003, Malaise trap, J. Poorani (1 male,

Revision of Dvivarnus (Scelionidae, Teleasinae) 1a

USNMENT01109189 (CNCI)). Karnataka St., grass / roadside, Malur, 28.1V.1988
(1 male, OSUC 230647 (OSUC)). Tamil Nadu St., Nilgiris Dist., Gudalur,
19.V1.1987 (1 male, OSUC 59262 (OSUC)). LAOS: Vientiane Prov., Gi Sion, Ban
Na (Ban Tha Ngon Na), 7.II-21.11.1965 (1 female, USNMENT00877588 (BPBM)).
NEPAL: Central Develop. Reg., nr. Birganj, MT #25, Lothar, 450ft, 29.VII-5.
[X.1967, Malaise trap (1 male, USNMENT01109175 (CNCI)). TAIWAN: Taiwan
Proyv., Nantou Co., Wushe, 1150m, 13.IV.1983, flight intercept trap, H. Townes (1
female, PSUC_000096246 (CNCI)). Taiwan Prov., Nantou Co., Wushe, 1150m,
22.V.1983, flight intercept trap, H. Townes (1 female, PSUC_000096141 (CNCI)).
Taiwan Prov., Nantou Co., Wushe, 1150m, no date, Malaise trap, H. Townes &
M. Townes (1 female, USNMENT01109188 (CNCI)). Taiwan Prov., Nantou
Co., Wushe, 1150m, no date, H. Townes (4 females, USNMENT01109183, USN-
MENT01109184, USNMENT01109185, USNMENT01109187 (CNCI)). THAI-
LAND: Chaiyaphum Prov., Taad Fah Waterfall, water supply station, T862, 245m,
15°56.468'N, 102°05.855'E, Tad Ton (Tat Tone) National Park, 8.IX-9.IX.2006,
yellow pan trap, T. Jaruphan & O. Budsawong (1 female, OSUC 342789 (OSUC)).
Chaiyaphum Prov., dry dipterocarp forest, T16, 250m, 15°59.037'N, 102°02.103'E,
Tad Ton (Tat Tone) National Park, 28.VI.2006, Malaise trap, C. Nichumnan (1 fe-
male, OSUC 374197 (OSUC)). Kanchanaburi Prov., Khong Kraborg, # 4781, 210m,
14°29.972'N, 98°53.035'E, Khuean Srinagarindra National Park, no date, Malaise
trap, Boonnam & Phumarin (1 male, UJNMENT01109177 (CNCI)). Kanchanaburi
Prov., Mae Kamint River, headquarters, # 3466, 14°38.123'N, 98°59.657'E, Khuean
Srinagarindra National Park, no date, Malaise trap, Somboon & Daorueng (1 female,
USNMENT01109173 (CNCI)). Kanchanaburi Prov., Mae Kamint River, tourist
center, 14422, 210m, 14°38.312'N, 98°59.643'E, Khuean Srinagarindra National
Park, no date, Malaise trap, Somboon & Daorueng (1 male, USNMENT01109169
(CNCI)). Khon Kaen Prov., Disturb (Moob Cave), T2, 296m, 16°44.837'N,
102°00.160'E, Phu Pha Man National Park, 13.VI-20.VI.2006, Malaise trap, R.
Phatai (1 female, OSUC 374198 (OSUC)). Nakhon Si Thammarat Prov., TV aerial,
T3108, 966m, 08°14.262'N, 99°48.289'E, Namtok Yong National Park, no date,
Malaise trap, Yai && Amnad (1 female, USNMENT01109170 (CNCI)). Phetchabun
Prov., Kaeng Krachan Nat. Park 12932.141'N, 99°27.914'E T4540, no date, Ma-
laise trap, Thongbai (1 female, USNMENT01109172 (CNCI)). Phetchabun Prov.,
helicopter landing ground, T266, 890m, 16°43.156'N, 101°35.118'E, Nam Nao Na-
tional Park, 8.VH-9.VII.2006, pan trap, N. Hongyothi (1 female, OSUC 284994
(OSUC)). Phetchaburi Prov., Huai Palao Forest Unit 3, Pa La-U Waterfall, T4564,
12°32.149'N, 99°28.265'E, Kaeng Krachan National Park, no date, pan trap, Thong-
bai (1 female, USNMENT01109180 (CNCD). Phetchaburi Prov., Huai Palao For-
est Unit 3, Pa La-U Waterfall, T4566, 12°32.149'N, 99°28.265'E, Kaeng Krachan
National Park, no date, Malaise trap, Thongbai (2 females, USNMENT01109174,
USNMENT01109178 (CNCD). Phetchaburi Prov., km33 / helipad, T4693, 735m,
12°50.177'N, 99°20.688'E, Kaeng Krachan National Park, no date, Malaise trap, Siri-
chai (1 male, USNMENT01109179 (CNC)J).

12 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

Dvivarnus elektrolythron Talamas & Mik6, sp. n.
http://zoobank.org/OCBCE486-CEBD-4924-9AF2-74EEA6A222B0
http://bioguid.osu.edu/xbiod_concepts/403212

Figures 15, 22, 27-31

Description. Color of antennae in female: brown throughout with dense white setae
on A2—A4. Color of mesosoma: mesoscutellum and propodeum black, metascutellar
spine brown, otherwise red. Color of head: except interantennal process, black. Seta-
tion of torular triangle: present. Continuity of torular triangle and central keel: torular
triangle opened dorsally, not continuous reduced central keel. Color of interantennal
process: yellowish brown. Setation of frons: transverse strip directly above interanten-
nal process glabrous, otherwise setose. Sculpture of dorsal frons: strigose. Sculpture of
vertex: strigose.

Epomial carina: indistinguishable from dorsoventral striation. Netrion in lat-
eral view: extending dorsally to proximity of mesothoracic spiracle. Netrion sulcus:
complete, extending dorsally to posterior margin of pronotum. Sculpture of vertical
face of pronotum: dorsoventrally strigose anteriorly, longitudinal striate posteriorly.
Sculpture of mesoscutum: finely punctate. Density of setation on medial mesoscu-
tum: dense. Notaulus: absent. Sculpture of scutoscutellar sulcus: smooth. Orientation
of transaxillar carina: projecting posterolaterally. Sculpture of mesoscutellum: finely
areolate. Density of setae on lateral margin of mesoscutellum: sparse. Posterior margin
of mesoscutellum: convex between mesoscutellar spines. Median mesoscutellar spine:
present. Lateral extreme of posterior scutoscutellar sulcus: foveae terminating below
mesoscutellar spine. Sculpture of metanotal trough: smooth with elongate furrow in
ventral half. Lateral propodeal carina: absent. Forewing pattern in female: wing mem-
brane and setae brown posterior to marginal vein and in distal third, separated by a
band of hyaline membrane and white setae. Sculpture of metapleural sulcus: smooth.
Sculpture of dorsal metapleural area: transversely rugose. Sculpture of ventral meta-
pleural area: transversely rugose. Number of setae on lateral T1: 3. Sculpture of T4:
punctate. Lateral patch on T5: present. Number of apical setae on T7: 2.

Diagnosis. Dvivarnus elektrolythron may be separated from females of D. mikuki
and D. agamades by the incomplete central keel on the frons (Fig. 15), the form of the
lateral patch on T4 (Figs 22, 29), and by the medially convex posterior margin of the
mesoscutellum.

Etymology. The epithet for this species refers to the bright red color on the meso-
soma of this species. It is derived from the words elektron which in Classical Greek
means “amber” and, by extension in modern times, “electricity”, and /ythron, meaning
“gore”. The name is treated as a noun in apposition.

Link to distribution map. http://hol.osu.edu/map-large.html?id=403212

Material examined. Holotype, female: WORY COAST: Savanes Rég., Korhogo
Dept., Konborodougou, 18.HI—21.H1.1984, M. Matthews, USNMENT01109168
(deposited in CNCI).

Revision of Dvivarnus (Scelionidae, Teleasinae) 13

419 q aa )
Figure 15-19. 15 Dvivarnus elektrolythron, female holotype (USNMENT01109168), head and an-
tennae, anterior view 16 D. agamades, female (USNMENT01109190), head, anterior view I7 D.
agamades, male (USNMENT01109177), head, anterior view 18 D. mikuki, female paratype (USN-
MENT01109214), head and antenna, anterior view 19 D. mikuki, male (USNMENT01109158), head,

anterior view. Scale bars in millimeters.

14 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

IpT6

SVR ia K

* Ree NY S a A be aS
Figure 20-24. 20 7rimorus sp., female (USNMENT01109195), head, mesosoma, metasoma, dorsal
view 21 Dvivarnus mikuki, female (USNMENT01109213), metasoma, dorsolateral view 22 D. elektro-
lythron, female (USNMENT01109168), metasoma, dorsolateral view 23 D. agamades, female (USN-
MENT01109174), metasoma, dorsolateral view 24 Trimorus sp., female (OSUC 186090), metasoma,

posterodorsal view. Scale bars in millimeters.

(i
F a

=

Dvivarnus mikuki Talamas & Miké, sp. n.
http://zoobank.org/1EC45732-63E1-40D5-94DE-453A560A9EEF
http://bioguid.osu.edu/xbiod_concepts/40321 1

Figures 6, 18-19, 21, 32-39

Description. Whorl of setae on flagellomeres in male: absent. Shape of A3—A11 in
male: cylindrical. Color of antenna in male: brown. Color of antennae in female:

Revision of Dvivarnus (Scelionidae, Teleasinae) ul)

= ee 5
is ~ cee = “San -
OD

Figure 25-26. Dvivarnus agamades. 25 female (USNMENT01109183), head, mesosoma, metasoma,
lateral view 26 female (USNMENT01109174), head, mesosoma, metasoma, dorsal view. Scale bars in
millimeters.

Al brown, A2 brown to yellow, A3—A5 yellow with white setae, AG6B—A16 brown.
Color of mesosoma: black, with yellow metascutellar spine. Color of head: black.
Setation of torular triangle: absent. Continuity of torular triangle and central keel:
torular triangle closed dorsally, continuous complete central keel. Setation of frons:
area above interantennal process glabrous, otherwise setose. Sculpture of dorsal
frons: dorsoventrally strigose. Sculpture of vertex: smooth; concentrically strigose.
Epomial carina: absent. Netrion in lateral view: present only at ventral limit of
pronotum. Ventral propleural area: smooth. Sculpture of propleural cervical sulcus:
smooth. Sculpture of mesoscutum: punctate to finely areolate, coarser in posterior
half. Density of setation on medial mesoscutum: dense. Notaulus: absent; weakly

16 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

Figure 27. Dvivarnus elektrolythron, female holotype (USNMENT0109168), head, mesosoma, meta-

soma, lateral view. Scale bar in millimeters.

indicated posteriorly amid sculpture. Sculpture of scutoscutellar sulcus: smooth.
Orientation of transaxillar carina: parallel to longitudinal axis of body. Shape of
axillula: bent ventrolaterally distally. Sculpture of mesoscutellum: punctate to areo-
late. Density of setae on lateral margin of mesoscutellum: sparse. Posterior margin of
mesoscutellum: concave between mesoscutellar spines. Median mesoscutellar spine:
absent. Lateral extreme of posterior scutoscutellar sulcus: foveae extending to axil-
lula. Sculpture of metanotal trough: smooth with elongate furrow in ventral half.
Lateral propodeal carina: absent. Plica: present. Forewing pattern in female: wing
membrane and setae brown posterior to marginal vein and in distal third, separated
by a band of hyaline membrane and white setae. Forewing color in male: uniform
throughout. Sculpture of metapleural sulcus: transverse portion smooth, dorsoven-
tral portions foveolate. Sculpture of dorsal metapleural area: smooth. Sculpture of
ventral metapleural area: transversely rugose; smooth. Number of setae on lateral
T1: 3; 4. Sculpture of T4: smooth. Lateral patch on T5: absent. Number of apical
setae on I’7: 4. Transverse line of pits on anterior S1: present.

Diagnosis. Dvivarnus mikuki can be separated from D. agamades and D. elektro-
lythron by the glabrous torular triangle and by the color of the mesosoma, which is
entirely black except for the metascutellar spine. Additionally, D. mikuki can be sepa-
rated from D. agamades by the absence of an epomial carina, the form of the metanotal
trough, which is non-foveolate and is dorsoventrally divided by a transverse furrow,

Revision of Dvivarnus (Scelionidae, Teleasinae) 17

metasoma, dorsal view 29 metasoma, dorsal view 30 pronotum, anterolateral view 31 posterior meso-

soma, dorsal view. Scale bars in millimeters.

and by the banding pattern on the wings of females. From D. elektrolythron it can be
separated by the broad lateral patch on T4 and the convex posterior margin of the
mesoscutellum between the mesoscutellar spines.

Etymology. The word “mikuki” means “spears” in Swahili, the language of Kenya
where the holotype specimen originates, and refers to the many spines found on the
mesosoma. The name is treated as noun in apposition.

Link to distribution map. http://hol.osu.edu/map-large.html?id=403211

Material examined. Holotype, female: KENYA: Nairobi Co., International Cen-
tre of Insect Physiology and Ecology (ICIPE) campus, nr. stream / meadow / de-
graded shrub-grassland, 1600m, 01.22317°S 36.89653°E, Kasarani, 27.V-3.V1.2014,
Malaise trap, R. Copeland, USNMENT01059120 (deposited in NMKE). Paratypes:
(11 females, 8 males) BENIN: 25km N Cotonou, Abomey-Calavi, XH-1988, J. S.
Noyes (1 male, USNMENT01109165 (CNCI)). KENYA: Kilifi Co., 3.30958° S
39.96538°E, Arabuko-Sokoke Forest 80 m, 22.VIII-5.[X.2014, Malaise trap, R.
Copeland (1 male, USNMENT01109193 (USNM)). Kilifi Co., indigenous forest
/ secondary forest, 19m, 03.30946°S 40.01941°E, Gede Forest, 11.XJI-25.XII.2011,
Malaise trap, R. Copeland (2 females, USNMENT01059122-01059123 (USNM)).

18 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

soma, lateral view 33 female paratype (USNMENT01059121), head, mesosoma, metasoma, dorsal view.

Scale bars in millimeters.

Kwale Co., indigenous forest, 39.52462°E, Muhaka Forest, 41m 4.32664°S, 13.XII-
27.X1.2014, Malaise trap, R. Copeland (1 female, USNMENT01109214 (NMKE)).
Kwale Co., indigenous forest, Muhaka Forest, 41m, 4.32664°S, 39.52462°E, 27.XII-
10.1.2015, Malaise trap, R. Copeland (1 female, USNMENT01109213 (NMKE)).
Kwale Co., indigenous forest, 76m, 04.52814°S 39.24028°E, Marenje Forest, 11.VH-
25.VII.2014, Malaise trap, R. Copeland (1 male, USNMENT01109194 (NMKE)).
Kwale Co., indigenous forest, 76m, 04.52814°S 39.24028°E, Marenje Forest,
25.VII-8.VIII.2014, Malaise trap, R. Copeland (3 females, USNMENT01059121,
01059124, 01059135 (USNM)). NIGERIA: Oyo St., International Institute of Trop-
ical Agriculture (IITA), Ibadan, X-1987, screen sweeping, J. S. Noyes (1 male, USN-
MENT01109162 (CNCI)). Oyo St., International Institute of Tropical Agriculture
(IITA), Ibadan, no date, screen sweeping, J. S. Noyes (1 male, USNMENT01109163
(CNCI)). ZIMBABWE: Harare (Salisbury), no date, A. Watsham (1 female, 2 males,

Revision of Dvivarnus (Scelionidae, Teleasinae) 19

ee

oe
——

Ne
Se
=
SS

\, »
\
\ a;
f Sy
P
Ne
7 (SSS
L pe
a
>
1 ay
ee
On
\Noke
4 i\ hey
* We
SS
~ BRS
/
,
{ t

~
Se
Se
:

Figure 34-39. Dvivarnus mikuki, female paratype (USNMENT010591135) 34 mesosoma, antero-
lateral view 35 posterior pronotum, lateral view 36 mesosoma, posterior view 37 posterior mesosoma,
posterolateral view 38 metasoma, dorsolateral view 39 T3—1T5, dorsolateral view. Scale bars in millimeters.

USNMENT01109157, USNMENT01109158, USNMENT01109161 (CNCI)).
Harare (Salisbury), no date, pan trap, A. Watsham (1 female, USNMENT01109159
(CNCI). Harare (Salisbury), no date, yellow pan trap, A. Watsham (1 female, USN-
MENT01109156 (CNCD). Harare (Salisbury), Chishawasha, no date, A. Watsham
(1 female, USNMENT01109167 (CNCI)). Harare (Salisbury), Chishawasha, no
date, pan trap, A. Watsham (1 male, USNMENT01109160 (CNCD).

20 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

ANS /
ay *<
“ZZ

Mo eit

Figure 40-45. Dvivarnus sp., male (USNMENT01109212). 40 head and mesosoma, dorsal view
41 metasoma, dorsal view 42 head and mesosoma, lateral view 43 mesosoma, posterodorsal view 44 head,

anterior view 45 pronotum, anterolateral view Scale bars in millimeters.

Dvivarnus sp., male

Figures 40-45

Dvivarnus sp., male

Figures 46-50

Revision of Dvivarnus (Scelionidae, Teleasinae) P|

Figure 46-50. Dvivarnus sp., male (USNMENT01109164) 46 head, mesosoma metasoma, dorsal view

47 head and mesosoma, lateral view 48 mesosoma, posterolateral view 49 head, anterior view 50 prono-

tum, anterolateral view. Scale bars in millimeters.

Acknowledgements

We thank Lubomir Masner, Andy Bennet (CNCI), and Luciana Musetti (OSUC)
for the loans of material that made this revision possible. We thank Norman Johnson
(OSUC) for developing and maintaining the Hymenoptera Online Database, Specim-
age, and vSysLab, which underlie the cybertaxonomy conducted here and Joe Cora

(OSUC) for providing invaluable database support. Ashton Smith, Trenton Jung, and

22 Elijah J. Talamas et al. / Journal of Hymenoptera Research 49: 1-23 (2016)

Pei Luo (USNM) contributed many of the photographs used to illustrate these spe-
cies. Luke Kresslein (USNM) contributed scanning electron micrographs. We thank
Juliet Muriuki, Mzee Hamisi Muhamed Shee, Kazungu Katana, Beatrice Elesani and
Mathias Ngonyo for excellent field work. Dr. Miguel Alonso-Zarazaga (MNCN) pro-
vided valuable input on etymology. This work was made possible by funding from the
Systematic Entomology Lab, USDA-ARS. The USDA does not endorse any commer-
cial product mentioned in this research. USDA is an equal opportunity provider and
employer. Istvan Mik6 was supported by the U. S. National Science Foundation (grant
DBI-1356381).

References

Balhoff JP, Miké IM, Yoder MJ, Mullins P, Deans AR (2013) A semantic model for species
description applied to the ensign wasps (Hymenoptera: Evaniidae) of New Caledonia.
Systematic Biology 62: 639-659. doi: 10.1093/sysbio/syt028

Hagedorn G, Catapano T, Giintsch A, Mietchen D, Endresen D, Sierra S, Groom Q, Biserkov
J, Glockler F, Morris R (2013) Best practices for stable URIs. http://wiki.pro-ibiosphere.eu/
wiki/Best_practices_for_stable_URIs

Kozlov MA, Lé XH (1986) [New species of scelionids (Hymenoptera, Scelionidae) of the fauna
of Vietnam.] Trudy Zoologicheskogo Instituta Akademii Nauk SSSR 159: 89-102.

Lé XH (2000) Egg-parasites of family Scelionidae (Hymenoptera). Fauna of Vietnam (vol. 3).
Science and Technics Publishing House, Hanoi, 386 pp.

Miko I, Copeland RS, Balhoff JP, Yoder MJ, Deans AR (2014) Folding Wings like a Cock-
roach: A Review of Transverse Wing Folding Ensign Wasps (Hymenoptera: Evaniidae:
Afrevania and Trissevania). PLoS ONE 9: e94056. doi: 10.137 1/journal.pone.0094056

Nixon GEJ (1936) The African species of Teleasinae (Hym., Proctotrupoidea, Fam. Scelionidae).
Annals and Magazine of Natural History 17: 114-191.

Talamas EJ, Buffington ML (2014) Updates to the nomenclature of Platygastroidea in the Zo-
ological Institute of the Russian Academy of Sciences. Journal of Hymenoptera Research
39: 99-117. doi: 10.3897/JHR.39.7698

Veenakumari K, Rajmohana K, Manickavasagam S, Mohanraj P (2011) On a new genus of
Teleasinae (Hymenoptera: Platygastridae) from India. Biosystematica 5: 39-46

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Hymenoptera. PLoS ONE 5(12): e15991. doi: 10.137 1/journal.pone.0015991

Revision of Dvivarnus (Scelionidae, Teleasinae) 23

Supplementary material |

URI table of HAO morphological terms

Authors: Elijah J. Talamas, Istvan Mik6, Robert S. Copeland

Data type: Microsoft Excel Spreadsheet (-xls)

Explanation note: This table lists the morphological terms used in this publication and
their associated concepts in the Hymenoptera Anatomy Ontology.

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Supplementary material 2

Semantic representations of phenotypes in Manchester syntax format of Dvivarnus

Authors: Elijah J. Talamas, Istvan Mik6, Robert S. Copeland

Data type: Microsoft Rich Text Format (.rtf)

Explanation note: Semantic representations of phenotypes in Manchester syntax format
of the taxonomic treatment of World Dvivarnus (Hymenoptera: Platygastroidea:
Teleasinae).

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.