JHR 87: 575-611 (2021) go Re, JOURNAL OF Seewtertensentan
doi: 10.3897/hr87.59794 MONOGRAPH () Hymenoptera
https://jhr.pensoft.net The Inarasional Society of Hymenopeeriss, RESEARCH

Revision of Phoenoteleia Kieffer
(Hymenoptera, Scelionidae, Scelioninae)

Zachary Lahey', Luciana Musetti? , Lubomir Masner’ , Norman FE. Johnson!”

| Department of Evolution, Ecology, and Organismal Biology, The Ohio State University, 1315 Kinnear Road,
Columbus, Ohio 43212, USA 2 Department of Entomology, The Ohio State University, 1315 Kinnear Road,
Columbus, Ohio 43212, USA 3 Agriculture and Agri-Food Canada, K.W. Neatby Building, Ottawa, Ontario
KIA 0C6, Canada

Corresponding author: Zachary Lahey (lahey. 18@osu.edu)

Academic editor: G. Broad | Received 23 October 2020 | Accepted 1 December 2020 | Published 23 December 2021
http://zoobank. org/F 16C4490-086F-4D88-AOBA-FDF13E995C4D

Citation: Lahey Z, Musetti L, Masner L, Johnson NF (2021) Revision of Phoenoteleia Kieffer (Hymenoptera,
Scelionidae, Scelioninae). In: Lahey Z, Talamas E (Eds) Advances in the Systematics of Platygastroidea III. Journal of
Hymenoptera Research 87: 575-611. https://doi.org/10.3897/jhr.87.59794

Abstract

The genus Phoenoteleia Kieffer is revised. Phoenoteleia canalis Dodd, P. rufa Kieffer, and P. rufescens (Ki-
effer) are redescribed, P. fusca (Kieffer) is transferred to Dicroscelio Kieffer as Dicroscelio fuscus (Kieffer),
comb. noy., and six species are described as new: P. buka Lahey, sp. nov. (Fiji), P. gunnelsi Lahey, sp. nov.
(Indonesia), P. halua Lahey, sp. nov. (Indonesia), P. kaca Lahey, sp. nov. (Indonesia), P. kuboa Lahey,
sp. nov. (Fiji), and P. sanma Lahey, sp. nov. (Vanuatu). The difficulty in associating male and female
specimens is discussed. A key is provided to differentiate between the species of Phoenoteleia and the puta-
tive closely related genera Mallateleia Dodd and Oxyteleia Kieffer.

Keywords

Australia, Dicroscelio, Indonesia, Malaysia, parasitoid, Platygastroidea, taxonomy

Introduction

The genus Phoenoteleia was described by J. J. Kieffer (1916) for a single female of the
type species P rufa Kieffer collected on the island of Mindanao, Philippines. In the
same publication, Kieffer (1916) erected the genus Plagioscelio Kieffer for two species

Copyright Zachary Lahey 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.

576 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

known only from males collected on the same island. Masner (1976) synonymized
Plagioscelio with Phoenoteleia based on shared similarities between the propodeum of
males and the first gastral tergite of females. He also suggested that P rufescens (Kieffer)
is the male of P rufa, although he did not synonymize the two species. Dodd (1929)
described an additional species, P canalis Dodd, from Queensland, Australia, as part of
his revision of the genus; however, this species was not mentioned by Masner (1976)
in his revisionary work on the family Scelionidae.

Masner (1976) included Phoenoteleia in the tribe Calliscelionini Masner based on
the following combination of characters: frons without a depression or central keel; an-
tenna 12-merous; skaphion absent; hind wing with tubular submarginal vein reaching
hamuli; tibial spur formula 1-1-1; T3 typically the longest of the metasomal tergites;
6 tergites and sternites visible externally in females; 8 tergites and 7 sternites visible
externally in males; T7 of female internal and exserted with the ovipositor (i-e., Scelio-
type) (Austin and Field 1997). The results of a phylogenomic analysis of the superfam-
ily places Phoenoteleia outside of Calliscelionini (Z. Lahey et al., unpublished data).
Instead, Phoenoteleia forms a clade with Mallateleia Dodd and Oxyteleia Kieffer, both
of which are members of Psilanteridini Kozlov, which is itself polyphyletic.

Phoenoteleia is a rare genus that is uncommon in collections despite being widely
distributed. Nearly all the specimens examined as a part of this study are female, mak-
ing the species descriptions biased towards characters present only in that sex. With the
few males we had available (n = 7), we were unable to reliably associate them with their
conspecific females, except for the single male of P. buka sp. nov., which we are conf-
dent belongs to that species due shared sculptural patterns. Images of male Phoenoteleia
not identified to the level of species are presented in Figures 63-74.

We are now able to build upon the work of Kieffer (1916) and Dodd (1929) based
on the accumulation of material collected over the last 50 years. Most of this new ma-
terial comes from Malaysia and the islands comprising the Indonesian Archipelago in
Southeast Asia, with additional specimens from as far west as Sri Lanka (South Asia),
as far north as the Amami Islands, Japan (East Asia), and as far east as Vanua Levu,
Fiji (Oceania). Photographs of the primary type of each species were examined, except
for P fusca (Kieffer), which is not in the Muséum National d’Histoire Naturelle, Paris,
France, and is feared lost.

Materials and methods

The numbers prefixed with “FBA”, “MNHN”, “OSUC”, “QM TYPE Hy/”and “US-
NMENT” are unique identifiers for the individual specimens (note the blank space
after some acronyms). Details of the data associated with these specimens may be ac-
cessed at the following link: https://mbd-s.asc.ohio-state.edu and entering the identi-
fier in the form.

Abbreviations and morphological terms used in the text: sensillar formula of cla-
vomeres: distribution of the large papillary sensilla (PS) on the ventral clavomeres of

Revision of Phoenoteleia 577

the female (Yang et al. 2016), with the segment interval followed by the number of
PS per segment (e.g., Al2—A8/1-2-2-2-1) (Bin 1981); EH: eye height, length of com-
pound eye measured parallel to dorsoventral midline of head; IOS: interocular space,
minimal distance on frons between compound eyes; LOL: lateral ocellar line, shortest
distance between the outer margins of the lateral and median ocelli (Masner 1980);
OD: ocellar diameter, greatest width of each ocellus; OOL: ocular ocellar line, short-
est distance between the inner orbit and outer margin of the lateral ocellus (Masner
1980); POL: posterior ocellar line, shortest distance between the inner margins of the
lateral ocelli (Masner 1980); T1, T2, ... [6: metasomal tergite 1, 2, ... 6; S1, S2, ... S6:
metasomal sternite 1, 2, ... 6. Morphological terminology generally follows Miké et al.
(2007). Morphological terms were matched to concepts in the Hymenoptera Anatomy
Ontology (Yoder et al. 2010) using the text analyzer function. A table of morphologi-
cal terms and URI links is provided in Suppl. material 1.

Most of the images were captured with a Leica MC170 HD digital camera at-
tached to a Leica Z16 APOA microscope using Leica Application Suite (version
4.12.0). The same software was also used to process image stacks into single montage
images. Montage images were postprocessed with Adobe Photoshop CC for brightness
and contrast. The images in Figures 1, 2, and 5 were produced following the methods
of ‘Talamas and Bufhngton (2015). The scanning electron micrographs of Oxyteleia
(Figures 3, 4) were taken following the methods of Talamas et al. (2016). The equip-
ment and software used to create Figure 6 is outlined in Lahey et al. (Lahey et al., this
volume). The distribution map (Figure 10) of the Phoenoteleia species treated in this
revision was created with SimpleMappr (Shorthouse 2010).

Author contributions

Z. Lahey: character definition and coding, generic concept development, imaging,
key development, manuscript preparation, species concept development; L. Musetti:
species concept development, key development, loan management; L. Masner: species
concept development, key development, provision of specimens; N. F. Johnson: spe-
cies concept development, key development, project coordination.

Collections

This work is based on specimens deposited in the following repositories:

AEIC = American Entomological Institute, Utah State University, Logan, UT, USA
BPBM Bernice Pauahi Bishop Museum, Honolulu, HI, USA

CNCI = Canadian National Collection of Insects, Ottawa, Ontario, Canada

MCZC Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA
MNHN Muséum National d'Histoire Naturelle, Paris, France

MZB Museum Zoologi Bogor, Bogor, Indonesia

OPPC Ovidiu Popovici Personal Collection, Iasi, Romania

578 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

OSUC CA. Triplehorn Collection, The Ohio State University, Columbus, OH, USA
QM Queensland Museum, Brisbane, Queensland, Australia

ROME Royal Ontario Museum, Toronto, Ontario, Canada

USNM_ Smithsonian National Museum of Natural History, Washington, D.C., USA

Abbreviations and characters annotated in the figures

arm armilla (Figure 9) mT1 membranous venter of horn

etc —_epitorular carina (Figure 32) (Figures 8, 9)

ctk central keel (Figure 4) occ occipital carina (Figure 7)

gen gena (Figure 7) prsl __ parapsidal line (Figure 7)

ffS2_— felt field of S2 (Figures 8, 9) sbc submedian carina (Figure 4)

horn horn of T1 (Figures 2, 9) scu mesoscutellum (Figures 3, 5, 7)

malp median area of the lateral propodeal shms_ mesoscutal suprahumeral sulcus
area (Figure 7) (Figure 7)

mch median channel of horn (Figure 7) sk skaphion (Figure 3)

msc mesoscutum (Figure 7) sss scutoscutellar sulcus (Figure 7)

msct metascutellum (Figure 7) vrx vertex (Figure 7)

mshs_ mesoscutal humeral sulcus (Figure 7)

Character discussion

Armilla

Armillae (sing. armilla) were ancient Roman military decorations (typically armbands
or bracelets) awarded to valorous soldiers. We introduce this term to refer to the carina
that marks the boundary between the horn (anterior) and basal (posterior) portions
of T1. The armilla is present in all species of Phoenoteleia, but it is not unique to the
genus. Certain species of Oxyteleia also have an armilla, as do members of several un-
described genera from the Neotropics and the West African species Stenotelea palustris
Huggert & Masner.

Median area of the lateral propodeal area

On either side of the horn of T1, directly posterior to the metanotum, lies an in-
verted triangular region we refer to as the median area of the lateral propodeal area
(Mik6 et al. 2021). This region is present in every species treated here, except for P
halua sp. nov.

Horn of T1

Complete bisection of the propodeum, metanotum, mesoscutellum, and at least the
osterior margin of the mesoscutum by the horn of T1 is one of the defining features
p g y. g

Revision of Phoenoteleia 579

of the genus. We observed considerable intra- and interspecific variation in the length
of the horn. Horn length did not appear to be correlated with specimen size, with some
of the smallest species possessing some of the longest horns (e.g., P. gunnelsi sp. nov.
and P. halua sp. nov.). In P. canalis, this character is particularly volatile; specimens
with the shortest and some of the longest horns belong to this species.

Key to Mallateleia, Oxyteleia, and Phoenoteleia

1 T3 longer than T2 (Figure 1); fore wing without tubular R vein (Figure 1);
‘TL Moftemale without hornt (Figures) crass serie Mallateleia Dodd

Figure |. Mallateleia sp., female (USNMEN1 42) | head, mesosoma, metasoma, dorsal view.
Scale bar in millimeters.

= T3 as long or shorter than T2 (Figures 2, 5); fore wing with tubular R vein
(rictiress2-y5) lel cotfemalesvithshor(Figuresi2:, 5) ieases. uli leteaceliaacats 2

treme Ba we oe J
ae fe: ae

Figures 2-4. Oxyteleia sp., female ( ) 2 head, mesosoma, metasoma, dorsolateral
view 3 mesosoma, dorsal view 4 head, anterolateral view. Scale bars in millimeters.

Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

Figures 5, 6. Phoenoteleia canalis Dodd, female (USNMENT01197920) 5 head, mesosoma, metasoma,

dorsal view 6 fore and hind wings, dorsal view. Scale bars in millimeters.

Skaphion present on mesoscutum (Figure 3) or absent; mesoscutellum trans-
verse, not present as two distinct lobes (Figure 3); tubular R vein of fore
wing straight (Figure 2); submedian carina and central keel present on frons
QE Cree hem ure. eens eum otic oes murs cele cone JO Oxyteleia Kieffer
Skaphion absent on mesoscutum (Figure 5); mesoscutellum bisected by horn
of Tl, present as two distinct lobes (Figure 5); tubular R vein of fore wing
bowed away from costal margin apically (Figure 5); submedian carina and
Central! KeeltabsertyO nM LOMSw ised eset Aims eioca retaliate. Phoenoteleia Kieffer

Key to species of Phoenoteleia (females)

1

Sculpture of vertex and dorsal frons sparsely punctate, with smooth areas
between’pulnceurest(Giemres: 42,4 455) SA) ath. sth n ded weet asta tedcea den dobea teas 2
Sculpture of vertex and frons rugose, transversely striate, or a combination of
boths(Figures Vel, 13 2G: 29-3216 2) srosoutcotseeaemoneishanott ee sea tet eadttss 3
Notaulus present (Figure 42); body black (Figures 42-44) oo... eee eeeeees
ha eee ae Phoenoteleia kuboa Lahey, sp. nov.
Notaulus absent (Figures 46, 49, 52); body reddish-brown (Figures 46-52) ...
Pete. irre Rae ST nn ty Decree ae REmenan reds Phoenoteleia rufa Kiefter
Sculpture of horn areolate throughout entire length or at least to transscutal
articttilationtthic tres I 5 3, SOR OU) wai aiaw dee cendencumrad esuceisaumce teats 4
Sculpture of horn areolate basally, otherwise transversely aciculate through-
OUP IC UIRGSe 277 350) Ra ee Nel st gigs RNIN glee SUNT MNO «a Senn GAME ARE BM 7

Revision of Phoenoteleia 581

Figures 7-9. Phoenoteleia canalis Dodd, female (OSUC 332073) 7 head, mesosoma, dorsal view 8 Horn
of T1, $1, $2, ventral view 9 Horn of T1, T2, S1, S2, lateral view. Scale bars in millimeters.

4 Posteromedial surface of vertex declivous, surface sculpture effaced (Figure 36);
lateral propodeal area without median area (Figure 36); clava 6-merous..........
ie 53.51 OUOnt galled 5 reel oa Biadlen 2.6 Eom. Phoenoteleia halua Lahey, sp. nov.

Posteromedial surface of vertex evenly rounded or weakly declivous, surface
sculpture continuous with vertex (Figures 11, 33, 60); lateral propodeal area
Withwmedianvarea(Picures, 1 l,/3.37 GO) Clava Sse OUS inns saute raitticst 5

5} Tergites 2 and 3 with weak longitudinal striae (Figure 33); body brown (Fig-
WTGSE 9D 9 Se ee ole. Satna 93 oie een Phoenoteleia gunnelsi Lahey, sp. nov.

582

Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

Tergites 2 and 3 with strong longitudinal striae (Figures 11, 63); body or-
ane-vellowgt Bicurres IU AGO) ea lus Akasa alana lawton esata eel ab anaes 6
Sculpture of frons transversely rugose (Figure 13); apical half of horn trans-
versely-aciculate (Figure Wy ii, tse tasebocvns. Phoenoteleia buka Lahey, sp. nov.
Sculpture of frons rugose (Figure 62); horn areolate throughout (Figure 60)...
5 ee: es ol eee > SO ee Phoenoteleia sanma Lahey, sp. nov.
Posteromedial surface of T5 nearly smooth (Figure 39); 16 distinctly lighter
in. color than T5 (Pigure:3 9)... sass Phoenoteleia kaca Lahey, sp. nov.
Posteromedial surface of T5 punctate-granulate (Figures 21-24); T6 con-
colorous with or darker than T5 (Figures 20-24) ooo. ceeeseeeeeeeeeeeeeeeeees
ih see reereri ps TOCA pets 74-0017 Se. Ao BE OO Phoenoteleia canalis Dodd

Key to species of Phoenoteleia (males)

1 Notauli-absente(Bigures:56; G0)... ae, ce asteveap cones tesa aeune ened one intakes 2.
— Notaulijpresent (Figure 66) «ciececvccessevcccsvvent Phoenoteleia sp. A (Sri Lanka)
2 Triangular portion of propodeum longitudinally striate (Figure 14); mesoscu-
telliunttncise (Ere ure MAN es Asst eek bes Phoenoteleia buka Lahey, sp. nov.
— Triangular portion of propodeum transversely rugose, sometimes with a sin-
gle central carina (Figure 56); mesoscutellum granulate (Figure 56)..............
aera an, Mallat enh ine tet An, Wet ar, a Phoenoteleia rufescens (Kieffer)
Taxonomy

Phoenoteleia Kieffer
http://zoobank.org/E1987CA4-0 1 FF-47E3-8A75-6606A 1099249

Phoenoteleia Kieffer, 1916: 62 (original description. Type: Phoenoteleia rufa Kieffer, by

monotypy and original designation); Kieffer, 1926: 265, 550 (description, keyed);
Dodd, 1929: 35 (description); Muesebeck and Walkley, 1956: 384 (citation of
type species); Baltazar, 1966: 185 (cataloged, catalog of species of the Philippines);
Masner, 1976: 31, 32 (description, key to males of Anteromorpha Dodd and Phoe-
noteleia Kieffer); Galloway and Austin, 1984: 7, 9, 20 (diagnosis, list of species
described from Australia, keyed); Johnson, 1992: 460 (cataloged, catalog of world
species); Austin and Field, 1997: 22, 68 (structure of ovipositor system, discussion
of phylogenetic relationships).

Plagioscelio Kieffer, 1916: 185 (original description. Type: Plagioscelio rufescens Kieffer,

by original designation. Synonymized by Masner (1976)); Kieffer, 1926: 266, 356
(description, keyed, key to species); Muesebeck and Walkley, 1956: 384 (citation
of type species); Baltazar, 1966: 176 (cataloged, catalog of species of the Philip-
pines); Masner, 1976: 32 (junior synonym of Phoenoteleia Kieffer).

Revision of Phoenoteleia 583

Diagnosis. Phoenoteleia can be distinguished from other scelionines by the following
combination of characters: epitorular carinae present on frons; T1 of female always
produced into an elongate horn which fits into a deep channel bisecting the metas-
cutellum, mesoscutellum, and at least the posterior portion of the mesoscutum; venter
of horn membranous; T1 with armilla; T2 clearly longer than T3; central portion of
propodeum triangular, in same plane as mesoscutum, mesoscutellum, and metascutel-
lum (male only); submarginal vein strongly curving towards costal margin of fore wing
apically; marginal vein distinctly shorter than stigmal vein; postmarginal vein at least
twice as long as stigmal vein; basitarsus on hind leg at least twice as long as combined
length of remaining tarsomeres, distinctly incrassate in males.

Description. Length 2.21—3.95 mm; body elongate, gracile to robust.

Head. Head shape in dorsal view: transverse. Hyperoccipital carina: absent. Oc-
cipital carina: present, complete. Length of OOL: lateral ocellus < 1 OD from inner
margin of compound eye. Shape of upper frons: convex. Antennal scrobe: undifferen-
tiated from surrounding surface sculpture. Submedian carina: absent. Orbital carina:
absent. Course of inner orbits: diverging ventrally. IOS/EH: IOS shorter than EH.
Central keel: absent. Antennal foramen: oriented laterally on interantennal process.
Facial striae: present. Malar striae: present. Malar sulcus: present. Setation of com-
pound eye: present, short. Gena: convex, distinctly produced behind compound eye.
Shape of clypeus: convex, trapezoidal, lateral corners not produced. Ventral margin of
clypeus: rounded. Anteclypeus: absent. Postclypeus: absent. Labrum: transverse, visible
anteriorly. Number of mandibular teeth: 3. Arrangement of mandibular teeth: trans-
verse. Number of maxillary palpomeres: 3. Shape of maxillary palpomeres: cylindrical.
Number of labial palpomeres: 2. Sculpture of occiput: granulate.

Antenna. Number of antennomeres in female: 12. Number of antennomeres in
male: 12. Insertion of radicle into A1: parallel to longitudinal axis of Al. Shape of A1:
more or less cylindrical, not flattened. Length of A3 of female: longer than A2, approxi-
mately as long or slightly longer than radicle. Claval formula: 1-2-2-2-1; 1-2-2-2-2; 1-2-
2-2-2-1. Number of clavomeres: 5; 6. Arrangement of papillary sensilla: longitudinal.
Antennomeres bearing tyloids in male antenna: A5. Shape of male flagellum: filiform.

Mesosoma. Transverse pronotal carina: present. Posterior apex of pronotum in
dorsal view: articulate with tegula. Epomial carina: present. Anterior face of pronotum:
visible dorsally, short. Netrion: present. Shape of netrion: wide, closed dorsally,
open ventrally. Netrion sulcus: present. Anterior portion of mesoscutum: vertical,
flexed ventrally to meet pronotum. Shape of mesoscutum: pentagonal, excavate at
base of wings. Skaphion: absent. Parapsidial lines: absent; present. Antero-admedian
lines: absent. Transscutal articulation: developed, narrow. Shape of mesoscutellum:
transverse, present lateral to horn of T1 in females, complete in males. Transaxillar
carina: present. Axillular carina: present. Lateral mesoscutellar spine: absent. Median
mesoscutellar spine: absent. Axillular spine: absent. Surface of mesoscutellum: in
same plane as mesoscutum. Median longitudinal furrow on mesoscutellum: absent.
Metascutellum: transverse, lateral to horn of T1 in females, complete in males. Setation

584 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

of metascutellum: absent. Lateral propodeal projection: absent. Medial portion of
metascutellum in males: plate-like triangular, elevated relative to lateral portions.
Median propodeal projection: absent. Subacropleural sulcus: present, indicated by a
dorsoventral line of setae. Mesopleural carina: present. Mesepimeral sulcus: present.
Posterior mesepimeral area: present. Sculpture of posterior mesepimeral area: smooth.
Mesal course of acetabular carina: not separating fore coxae. Mesopleural pit: present.
Metapleural sulcus: present. Paracoxal sulcus: present.

Legs. Number of mesotibial spurs: 1. Number of metatibial spurs: 1. Relative
length of metabasitarsus: at least twice the length of remaining tarsomeres. Dorsal
surface of metacoxa: smooth. Shape of metacoxa: cylindrical, ecarinate. Trochantellus:
indicated by transverse sulcus on femur.

Wings. Wing development of female: macropterous. Wing development of male:
macropterous. Tubular veins in fore wing: present. Setation of R vein: present, sparse,
long. Bulla of fore wing R vein: absent. Length of marginal vein of fore wing: at least 2
times shorter than stigmal vein. Origin of stigmal vein (r-rs) in fore wing: arising from
marginal vein along costal margin. Development of R vein in hind wing: complete,
reaching frenal hooks.

Metasoma. Number of external metasomal tergites in female: 6. Number of exter-
nal metasomal sternites in female: 6. Number of external metasomal tergites in male:
8. Number of external metasomal sternites in male: 7. Shape of metasoma: fusiform,
narrowly constricted between Tl and T2 in females. Laterotergites: present. Lateroster-
nites: present. T'1 of female: produced into an elongate horn that bisects the metascutel-
lum, mesoscutellum, and at least the posterior margin of the mesoscutum. Armilla:
present. Relative size of metasomal terga: T2 longest (excluding horn of T1 in females),
T3 longer than T4. Transverse sulcus on T2: present. Metasomal terga with basal crenu-
lae: [2 in females, T1 and T2 in males. Sublateral carinae on metasomal terga: present.
Median longitudinal carina on metasomal terga: absent. Shape of T6 in female: widest
anteriorly. Anterior margin of S1: straight, not produced anteriorly. Felt fields on S2:
present. Felt fields on S3: absent. Ovipositor: Scelio-type (Austin and Field 1997).

Generic transfer of Phoenoteleia fusca (Kieffer)

Dicroscelio fuscus (Kieffer), comb. nov.

http://zoobank.org/7A825 1 D8-5870-4F3E-8932-A945B158733D

Plagioscelio fuscus Kieffer, 1916: 187 (original description); Kieffer, 1926: 356, 357
(description, figured, keyed); Baltazar, 1966: 176 (cataloged, distribution).
Phoenoteleia fusca (Kieffer): Johnson, 1992: 461 (cataloged).

Comments. Kieffer (1916) described Plagioscelio fuscus (=Phoenoteleia fusca) for a single
male collected in the Philippines. As far as we know, the type was never examined by
anyone other than Kieffer and the specimen was never deposited in MNHN, the primary
repository of his Hymenopteran type material. The single line drawing of the metasoma

Revision of Phoenoteleia 585

(Kieffer 1926; Suppl. material 2) and certain morphological characters mentioned in the
original description shed doubt on the placement of this species within Phoenoteleia. In-
stead, we propose that this species is more appropriately placed within Dicroscelio Kieffer,
a cosmopolitan genus well represented in Southeast Asia (Kieffer 1913; Masner 1976;
Yoder et al. 2009). Kieffer's drawing of the metasoma (incorrectly labeled as female!) con-
forms with that of a Dicroscelio species from New Caledonia (OSUC 185966; Suppl. ma-
terial 2). Similarly, his description of the wing venation, structure of the metanotum, and
length of the metabasitarsus is indicative of certain members of Dicroscelio and excludes
Phoenoteleia as a potential option for the placement of this species. It is important to note
that we are not identifying this Dicroscelio specimen as P. fusca. We are simply making a
comparison between this specimen and the description of P. fusca given by Kieffer (1916).

Species description

Phoenoteleia buka Lahey, sp. nov.
http://zoobank.org/ 1 DIOBEIE-1AA2-4C32-952F-3F9622D7B621
Figures 11-16

Description. Claval formula: 1-2-2-2-1. Number of clavomeres: 5. Color of antenna
in female: radicle and Al—AG light brown, A7—A12 brown. Color of A7 of female:
concolorous with clavomeres. Color of antenna in male: radicle and Al—A8 orange,
A9 light brown, A10—A12 brown. Color of head: light orange. Color of mesosoma:
mostly orange, darker on mesoscutum lateral to horn. Color of metasoma: horn orange
apically, becoming progressively darker posteriorly; T1 posterior to armilla burnt or-
ange; [2—T3 orange; [4 burnt orange; T5—T6 nearly brown. Color of legs: becoming
progressively darker distally. Setation of frons: short, sparse. Claval formula: 1-2-2-2-1.
Number of clavomeres: 5. Sculpture of frons: transversely rugose. Sculpture of ver-
tex: sharply areolate-rugose. Excavation on posteromedial vertex: absent. Sculpture of
posteromedial vertex: same as remainder of vertex. Sculpture of gena: rugose. Length
of LOL: < 1 OD. Length of POL: three times as long as LOL. Sculpture of dorsal
pronotal area: areolate-rugose. Sculpture of lateral pronotal area: transversely striate.
Sculpture of netrion: transversely striate. Notaulus: absent. Mesoscutal humeral sul-
cus: not clearly differentiated from surrounding surface sculpture. Mesoscutal supra-
humeral sulcus: not clearly differentiated from surrounding surface sculpture. Sculp-
ture of mesoscutum: longitudinally areolate-rugose. Parapsidial line: absent. Setation
of mesoscutellum: present. Sculpture of mesoscutellum: granulate throughout. Form
of metascutellum: two times wider than length, unsculptured, lateral corner acute.
Sculpture of propodeum: rugose. Setation of plical area: present. Median area of the
lateral propodeal area: present. Sculpture of mesofemoral depression: smooth. Setation
of mesepisternum ventral to mesofemoral depression: sparse. Sculpture of mesepister-
num ventral to mesofemoral depression: punctate. Setation of dorsal metapleural area:
absent. Sculpture of dorsal metapleural area: smooth throughout. Setation of ventral
metapleural area: absent. Sculpture of ventral metapleural area: smooth. Length of

586 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

© Phoenoteleia buka
() Phoenoteleia canalis

(©) Phoenoteleia gunnelsi
() Phoenoteleia halua

() Phoenoteleia kaca

@ Phoenoteleia kuboa
()Phoenoteleia rufa

() Phoenoteleia sanma

Figure 10. Geographic distribution of the Phoenoteleia species treated in this revision. Stars denote holo-
type localities for P canalis and P rufa.

Figures 11-13. Phoenoteleia buka Lahey, female holotype (FBA136312) I 1 head, mesosoma, meta-

soma, dorsal view 12 mesosoma, lateral view 13 head, anterior view. Scale bars in millimeters.

Revision of Phoenoteleia 587

Figures 14-16. Phoenoteleia buka Lahey, male paratype (FBA136318) 14 head, mesosoma, metasoma,

dorsal view 15 mesosoma, lateral view 16 head, anterior view. Scale bars in millimeters.

horn on T1: reaching apex of mesoscutum. Sculpture of horn on T1: areolate-rugose
basally and along margin of armilla, otherwise transversely aciculate. Sculpture of T1
posterior to armilla: longitudinally striate throughout, interstices rugose. Sculpture of
T2: longitudinally striate, interstices rugose. Sculpture of T3: longitudinally striate
throughout, interstices reticulate medially, granulate laterally. Sculpture of T4: longitu-
dinally striate laterally, granulate medially. Sculpture of T5: smooth. Sculpture of T6:
smooth. Length of T6 in female: 1.5 times maximum width. Relative length of hind
basitarsus in female: 2.5 times as long as remaining tarsomeres. Relative length of hind
basitarsus in male: 2.5 times as long as remaining tarsomeres.

Diagnosis. Phoenoteleia buka sp. nov. comes closest to P. sanma sp. nov. from
which it differs by having the frons transversely rugose and the apical surface of the T1
horn with aciculate sculpture. Other characters that allow for its recognition include
the darkened T5 and T6, and the margins of the median channel that are distinctly
elevated relative to the lateral mesoscutum. The male of P. buka sp. nov. shares the
transversely rugose sculpture on the frons and is easily separated other Phoenoteleia
males by the rugose mesoscutellum.

Etymology. Buka is the Fijian word for fire and is used to refer to the coloration of
this species. The epithet is treated as a noun in apposition.

588 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

Material examined. Holotype, female: Fy: Central Div., Rewa Prov., Viti Levu
Isl., 3.5km N Veisari Settlement, MT3, 300m, 18.068°S, 178.367°E, logging road to
Waivudawa, 14.[I—8.III.2003, Malaise trap, E. Schlinger & M. Tokota’a, FBA136312
(deposited in BPBM). Paratypes: Fiyt: 3 females, 1 male, FBA136316, FBA136318
(BPBM); FBA136327 (CNCI); FBA136330 (OSUC).

Comments. The vertex is slightly declivous in females, most likely due to the
length of the horn which surpasses the anterior margin of the mesoscutum.

Phoenoteleia canalis Dodd
http://zoobank.org/4B75D 182-A436-41B1-9E79-7CD9ED997C9 1
Figures 5-9, 17-32

Phoenoteleia canalis Dodd, 1929: 35 (original description); Galloway, 1976: 101 (type
information); Johnson, 1992: 461 (cataloged, type information).

Description. Claval formula: 1-2-2-2-1; 1-2-2-2-2. Number of clavomeres: 5. Color of
antenna in female: radicle and Al—A7 light brown, A8—A12 brown. Color of A7 of fe-
male: distinctly lighter than clavomeres. Color of antenna in male: unknown. Color of
head: mostly orange, interocellar space and most of vertex brown. Color of mesosoma:
reddish-brown. Color of metasoma: reddish-brown. Color of legs: light yellow-brown,
fore legs darkest. Setation of frons: short, sparse. Sculpture of frons: punctate-rugose
dorsally, transversely striate ventrally. Sculpture of vertex: punctate-rugose. Excavation
on posteromedial vertex: absent. Sculpture of posteromedial vertex: same as remainder
of vertex. Sculpture of gena: longitudinally striate. Length of LOL: < 1 OD. Length of
POL: < two times as long as LOL. Sculpture of dorsal pronotal area: areolate-rugose.
Sculpture of lateral pronotal area: rugose. Sculpture of netrion: transversely striate. No-
taulus: absent. Mesoscutal humeral sulcus: not clearly differentiated from surround-
ing surface sculpture. Mesoscutal suprahumeral sulcus: not clearly differentiated from
surrounding surface sculpture. Sculpture of mesoscutum: areolate-rugose. Parapsidial
line: absent; present. Setation of mesoscutellum: present. Sculpture of mesoscutellum:
granulate throughout. Form of metascutellum: twice as wide as long, unsculptured,
lateral corner acute. Sculpture of propodeum: rugose. Setation of plical area: present.
Median area of the lateral propodeal area: present. Sculpture of mesofemoral depres-
sion: transversely rugose. Setation of mesepisternum ventral to mesofemoral depres-
sion: sparse. Sculpture of mesepisternum ventral to mesofemoral depression: smooth
dorsomedially, otherwise punctate-rugose. Setation of dorsal metapleural area: absent.
Sculpture of dorsal metapleural area: smooth anteriorly, rugose posteriorly. Setation
of ventral metapleural area: present posteriorly. Sculpture of ventral metapleural area:
smooth. Length of horn on T1: reaching apex of mesoscutum; reaching middle of
mesoscutum. Sculpture of horn on T1: rugose basally, otherwise faintly transversely
aciculate. Sculpture of T1 posterior to armilla: rugose medially, longitudinally striate
laterally, interstices rugose. Sculpture of T2: longitudinally striate, interstices rugose.

Revision of Phoenoteleia 589

Figures | 7-20. Phoenoteleia canalis Dodd, female holotype (QMTYPE Hy3293) 17 mesosoma, dorsal view

18 mesosoma, dorsal view 19 mesosoma, lateral view 20 mesosoma, lateral view. Scale bars in millimeters.

Sculpture of T3: longitudinally striate throughout most of length, interstices rugose,
posteromedially granulate. Sculpture of T4: longitudinally striate laterally, faintly
granulate medially; longitudinally striate laterally, granulate medially. Sculpture of T5:
weakly longitudinally striate laterally, faintly granulate medially; weakly longitudinally
striate laterally, punctate medially. Sculpture of T6: faintly granulate; punctate. Length
of T6 in female: 1.5 times maximum width; 1.25 times maximum width. Relative
length of hind basitarsus in female: 2.5 times as long as remaining tarsomeres; 2 times
as long as remaining tarsomeres. Relative length of hind basitarsus in male: unknown.

Diagnosis. Phoenoteleia canalis is separated from its congeners by the rugose frons,
concolorous T'5 and T6, and the horn which is areolate-rugose basally and aciculate
throughout most of its length.

Material examined. Holotype, female: Austratia: QLD, Dunk Island, VIII-
1927, H. Hacker, QM TYPE Hy/3293 (deposited in QM). Other material: BRu-
NEI: OSUC 332072 (CNCI); OSUC 332088 (OSUC); INDongsia: OSUC 181592,
332115-332117, 491273 (ROME); OSUC 331984, 331985, 332067, 332069,
332081—332083, 332096, 332118 (BMNH); OSUC 332068, 332073-332078,
332080, 332084, 332085, 332115 (CNCI); OSUC 332070, 332079, 332086; Ma-
Laysta: OSUC 149608, 202439 (AEIC); OSUC 332087, 332089-332092, 332094,
332095, 332119-332122, 491275 (CNCI); OSUC 332093, 491274 (OSUQC);

Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

Figures 21-24. Phenotypic variation within female Phoenoteleia canalis Dodd, habitus, dorsal view
21 Indonesia, West Java (OSUC 332116) 22 Brunei, Belait District (OSUC 332072) 23 Papua New
Guinea, New Britain Province (OSUC 331975) 24 Papua New Guinea, Milne Bay Province (OSUC
331983). Scale bars in millimeters.

Revision of Phoenoteleia 59.1

Figures 25-28. Phenotypic variation within female Phoenoteleia canalis Dodd, mesosoma, lateral view
25 Indonesia, West Java ( ) 26 Brunei, Belait District ( ) 27 Papua New

Guinea, New Britain Province ( ) 28 Papua New Guinea, Milne Bay Province (

). Scale bars in millimeters.

Papua NEw GUINEA: ; ; — ;
(CNCI; : ; (OSUC); THAILAND:
(OSUQ); (CNCI).

Comments. Dodd (1929) described P. canalis for a single female specimen collected
on Dunk Island in Queensland, Australia. The holotype is in relatively good condition,
despite the mesosoma and metasoma having become separated. ‘The head, however, has
been missing for over 40 years (Galloway 1976), precluding our ability to examine what is
perhaps the most important tagma for species level identification of Phoenoteleia. Dodd's
description of the cephalic characters of P. canalis provides enough detail to reliably
separate it from P. halua sp. nov., P. kuboa sp. nov., and P. rufa, and its constellation of
somal characters is incongruent with the remainder of the species treated in this revision.

Our concept of P. canalis is that it is a highly variable, widespread species. We docu-
mented morphological variation between P. canalis populations from Brunei, Indonesia,
Malaysia, Papua New Guinea, and Thailand (Figures 21-32). The most glaring morpho-
logical difference is the length of the horn, a character we have found to be highly variable
in the type species of the genus (P. rufa). Specimens of P canalis ranged in color from

reddish-purple to light orange-brown. Dodd (1929) described the color of this species as

592 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

1: >
te
,

- —
ete
ie

Se oe

Xe

may
Wk
‘

Figures 29-32. Phenotypic variation within female Phoenoteleia canalis Dodd, head, anterior view
29 Indonesia, West Java (OSUC 332116) 30 Brunei, Belait District (OSUC 332072) 31 Papua New
Guinea, New Britain Province (OSUC 331975) 32 Papua New Guinea, Milne Bay Province (OSUC
331983). Scale bars in millimeters.

bright red-brown, but images of the holotype suggest that its color has faded considerably
over the past century. Such fading could explain the variation in color between specimens.

While this article was in press, a collaborator (Dr. Ovidiu Popovici, Alexandru Ioan
Cuza University, Iasi, Romania) provided the first author with images of two female
Phoenoteleia specimens that match our concept of P. canalis. One of the specimens is
part of a long series from Sulawesi collected by Dr. John Noyes, and the other is from
Australia. The Australian specimen is the second Phoenoteleia known to us from that
continent and is virtually identical to a female we examined from Milne Bay Province
in Papua New Guinea (OSUC 331983; Figures 24, 28, 32).

Transcribed data labels for these specimens are included below. Both specimens are
deposited in the OPPC.

AUSTRALIA: Queensland; Daintree; James Cook University, rainforest site;
1LGrO64 53.25, 145227 13082Es altel 9m 1 Owiii-9 ix. 2014;tlec. D> Rentz26c 2.
Tripotin (MT).

Sutawes!: Utara; Dumoga-Bone N.P. Toraut; 220m; 9-16.v.1985, leg. J.S. Noy-
es [voucher specimen used by Popovici et al. 2017 to illustrate the mouthparts of
Phoenoteleia].

Revision of Phoenoteleia 593

Phoenoteleia gunnelsi Lahey, sp. nov.
http://zoobank.org/F4587 DAC-5 FFE-4D66-9EAA-0701285E86C6
Figures 33-35

Description. Claval formula: 1-2-2-2-1. Number of clavomeres: 5. Color of antenna
in female: radicle and Al—A7 light brown, A8—A12 brown. Color of A7 of female:
distinctly lighter than clavomeres. Color of antenna in male: unknown. Color of
head: reddish-brown. Color of mesosoma: reddish-brown. Color of metasoma: red-
dish-brown. Color of legs: becoming progressively darker distally. Setation of frons:
long, sparse. Sculpture of frons: punctate-rugose dorsally, transversely striate ventral-
ly. Sculpture of vertex: punctate-rugose. Excavation on posteromedial vertex: absent.
Sculpture of posteromedial vertex: same as remainder of vertex. Sculpture of gena:
longitudinally striate. Length of LOL: 1 OD. Length of POL: two times as long as
LOL. Sculpture of dorsal pronotal area: areolate-rugose. Sculpture of lateral pronotal
area: transversely striate. Sculpture of netrion: transversely rugose. Notaulus: absent.
Mesoscutal humeral sulcus: not clearly differentiated from surrounding surface sculp-
ture. Mesoscutal suprahumeral sulcus: not clearly differentiated from surrounding
surface sculpture. Sculpture of mesoscutum: areolate-rugose. Parapsidial line: absent.
Setation of mesoscutellum: present. Sculpture of mesoscutellum: granulate posteri-
orly. Form of metascutellum: twice as wide as long, foveolate. Sculpture of propo-
deum: rugose. Setation of plical area: present. Median area of the lateral propodeal
area: present. Sculpture of mesofemoral depression: smooth. Setation of mesepister-
num ventral to mesofemoral depression: sparse. Sculpture of mesepisternum ventral
to mesofemoral depression: smooth dorsomedially, otherwise punctate-rugose. Seta-
tion of dorsal metapleural area: absent. Sculpture of dorsal metapleural area: smooth
anteriorly, rugose posteriorly. Setation of ventral metapleural area: absent. Sculpture
of ventral metapleural area: smooth. Length of horn on T1: reaching apex of mesos-
cutum. Sculpture of horn on T1: unsculptured apically, otherwise areolate. Sculp-
ture of T1 posterior to armilla: longitudinally striate throughout, interstices rugose.
Sculpture of T2: longitudinally weak medially, replaced by areolate surface sculpture,
interstices areolate-rugose. Sculpture of T3: longitudinally striate, interstices areolate.
Sculpture of T4: finely longitudinally striate, otherwise areolate. Sculpture of T5: lon-
gitudinally striate throughout, interstices punctate. Sculpture of T6: punctate. Length
of T6 in female: 1.5 times maximum width. Relative length of hind basitarsus in
female: 2 times as long as remaining tarsomeres. Relative length of hind basitarsus in
male: unknown.

Diagnosis. Phoenoteleia gunnelsi sp. nov. most closely resembles P. halua sp. nov.
due to similarities in their coloration, size, and length of the horn on T1. Noticeable
differences include the sculpture of the vertex, number of clavomeres, and presence of
the median area on the lateral propodeal area.

Etymology. Named for Dr. Charles W. Gunnels IV (Florida Gulf Coast Univer-
sity, Fort Myers, Florida) in recognition of his generosity, zeal for teaching, and love for
the natural world. The epithet is treated as a noun in the genitive case.

594 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

° - / AJ ‘ * |
+ sp on , . ies : asi
. ‘ eS ee ey — in ha i We pares |
a wa i % a ' s , ; ated
. ns r fs. F 7 4 Ps r BS is
aa ay + ~ 4 =
We 0.1 a. = a
iss al ow 2 f a“ oe

Figures 33-35. Phoenoteleia gunnelsi Lahey, female holotype (OSUC 359935) 33 head, mesosoma,

metasoma, dorsal view 34 mesosoma, lateral view 35 head, anterior view. Scale bars in millimeters.

Material examined. Holotype, female: INDONEsIA: Seram, Maluku, 10 km NW
Waisarisa, 3°S, 128°E, 300m, 17—25 Jan 1995, DC Darling IIS 950006 | Universitas
Pattimura Forest Area, 2° forest selectively logged, 1983, Malaise trap — no pans, Light
gap, clearing, OSUC 359935 (deposited in MZB).

Comments. We erect this species for a single female specimen. Our first consid-
eration when treating this species was that is represented variation within P. halua
sp. nov.; however, the differences between this specimen and P. halua sp. nov. exceed
the range of morphological variation observed within other species of the genus. It is
for this reason that we feel justified in describing P. gunnelsi sp. nov.

Phoenoteleia halua Lahey, sp. nov.

http://zoobank.org/B9F8B02B-AA05-4C23-A04A-3E0636BEBD44
Figures 36-38

Description. Claval formula: 1-2-2-2-2-1. Number of clavomeres: 6. Color of an-
tenna in female: radicle and Al—AG light yellow, A7—A12 brown. Color of A7 of
female: concolorous with clavomeres. Color of antenna in male: unknown. Color
of head: reddish-brown. Color of mesosoma: reddish-brown. Color of metasoma:

Revision of Phoenoteleia 595

Figures 36-38. Phoenoteleia halua Lahey, female holotype (OSUC 359932) 36 head, mesosoma, meta-

soma, dorsal view 37 mesosoma, lateral view 38 head, anterior view. Scale bars in millimeters.

reddish-brown. Color of legs: light yellow-brown, fore legs darkest. Setation of frons:
short, sparse. Sculpture of frons: punctate-rugose dorsally, transversely striate ven-
trally. Sculpture of vertex: punctate-rugose. Excavation on posteromedial vertex: pre-
sent. Sculpture of posteromedial vertex: smooth. Sculpture of gena: rugose. Length of
LOL: 1 OD. Length of POL: two times as long as LOL. Sculpture of dorsal pronotal
area: areolate-rugose. Sculpture of lateral pronotal area: transversely striate. Sculpture
of netrion: transversely rugose. Notaulus: absent. Mesoscutal humeral sulcus: not
clearly differentiated from surrounding surface sculpture. Mesoscutal suprahumeral
sulcus: not clearly differentiated from surrounding surface sculpture. Sculpture of
mesoscutum: areolate-rugose. Parapsidial line: absent. Setation of mesoscutellum: ab-
sent medially. Sculpture of mesoscutellum: granulate posteriorly. Form of metascutel-
lum: twice as long as wide, unsculptured. Sculpture of propodeum: rugose. Setation
of plical area: present. Median area of the lateral propodeal area: absent. Sculpture of
mesofemoral depression: transversely rugose. Setation of mesepisternum ventral to
mesofemoral depression: sparse. Sculpture of mesepisternum ventral to mesofemo-
ral depression: smooth dorsomedially, otherwise punctate-rugose. Setation of dorsal
metapleural area: absent. Sculpture of dorsal metapleural area: smooth anteriorly,
rugose posteriorly. Setation of ventral metapleural area: absent. Sculpture of ven-
tral metapleural area: smooth. Length of horn on T1: reaching apex of mesoscu-

596 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

tum. Sculpture of horn on T1: areolate-rugose basally and along margin of armilla,
otherwise transversely aciculate. Sculpture of T1 posterior to armilla: transversely
striate-rugose medially, longitudinally striate laterally, interstices rugose. Sculpture of
T2: longitudinally weak medially, replaced by areolate surface sculpture, interstices
areolate-rugose. Sculpture of T3: longitudinally striate, interstices areolate. Sculpture
of T4: transversely striate laterally, faintly areolate medially. Sculpture of T5: weakly
longitudinally striate laterally, punctate medially. Sculpture of T6: punctate. Length
of T6 in female: 1.25 times maximum width. Relative length of hind basitarsus in
female: 2 times as long as remaining tarsomeres. Relative length of hind basitarsus in
male: unknown.

Diagnosis. Three characters are unique to P. /alua sp. nov.: (1) the posterior ver-
tex is excavated and devoid of surface sculpture; (2) the clava is 6-merous; and (3) the
median area on the lateral propodeal area is absent. Otherwise, P. halua sp. nov. is most
likely to be confused with P. gunnelsi sp. nov. based on its size and habitus similarities.

Etymology. The species name is taken from the Indonesian word for smooth, in
reference to the sculpture of the declivity on the vertex. The epithet is treated as a noun
in apposition.

Material examined. Holotype, female: INDoNgsia: W. Kalimantan Gunung
Palung Nat. Pk. 15 JUN-15 AUG 1991 Darling, Rosichon, Sutrisno | Cabang Panti
Res. Sta. 1 rainforest.100—400m Alluvial light gap 1°15'S, 110°5'E Malaise trap head
IIS 910122, OSUC 359932 (deposited in MZB). Paratypes: INDoNEsIA: 2 females,
OSUC 359933 (CNCI); 359934 (ROME).

Comments. The three females that comprise the type series are virtually identical.

Phoenoteleia kaca Lahey, sp. nov.
http://zoobank.org/223 FD7FF-717C-41 DE-99F3-87A4F980ED8E
Figures 39-41

Description. Claval formula: 1-2-2-2-1. Number of clavomeres: 5. Color of antenna
in female: radicle and Al—A7 yellow, A8—A12 brown. Color of A7 of female: distinctly
lighter than clavomeres. Color of antenna in male: unknown. Color of head: light
yellow anteriorly, light yellow anterodorsally, burnt orange posterodorsally. Color of
mesosoma: light yellow-orange laterally, burnt orange-brown dorsally. Color of meta-
soma: T'1 including horn orange apically, becoming progressively darker posteriorly;
T2-T4 brown laterally, burnt orange-orange sublaterally, brown medially; T5 mostly
brown, yellow-orange anterolaterally; T6 orange. Color of legs: coxae, trochanters,
femurs light yellow; tibiae orange; tarsi burnt orange-brown. Claval formula: 1-2-2-
2-1. Number of clavomeres: 5. Setation of frons: long, moderate. Sculpture of frons:
areolate-rugose. Excavation on posteromedial vertex: absent. Sculpture of posterome-
dial vertex: same as remainder of vertex. Sculpture of vertex: areolate-rugose. Sculp-

ture of gena: rugose. Length of LOL: 1 OD. Length of POL: two times as long as

Revision of Phoenoteleia 597

Figures 39-41. Phoenoteleia kaca Lahey, female holotype (OSUC 359936) 39 head, mesosoma, meta-

soma, dorsal view 40 mesosoma, lateral view 41 head, anterior view. Scale bars in millimeters.

LOL. Sculpture of dorsal pronotal area: areolate-rugose. Sculpture of lateral pronotal
area: rugose. Sculpture of netrion: transversely rugose. Notaulus: absent. Mesoscutal
humeral sulcus: not clearly differentiated from surrounding surface sculpture. Mesos-
cutal suprahumeral sulcus: not clearly differentiated from surrounding surface sculp-
ture. Sculpture of mesoscutum: areolate-rugose. Parapsidial line: present. Setation of
mesoscutellum: present. Sculpture of mesoscutellum: granulate throughout. Form of
metascutellum: twice as long as wide, unsculptured. Sculpture of propodeum: rugose.
Setation of plical area: present. Median area of the lateral propodeal area: present.
Sculpture of mesofemoral depression: transversely rugose. Setation of mesepisternum
ventral to mesofemoral depression: dense. Sculpture of mesepisternum ventral to mes-
ofemoral depression: punctate. Setation of dorsal metapleural area: absent. Sculpture
of dorsal metapleural area: smooth anteriorly, rugose posteriorly. Setation of ventral
metapleural area: absent. Sculpture of ventral metapleural area: rugose. Length of horn
on T1: reaching middle of mesoscutum. Sculpture of horn on T1: areolate-rugose
basally and along margin of armilla, otherwise transversely aciculate. Sculpture of T1
posterior to armilla: rugose medially, longitudinally striate laterally, interstices rugose.
Sculpture of T2: longitudinally striate, interstices rugose. Sculpture of T3: longitudi-
nally striate throughout most of length, interstices rugose, posteromedially granulate.

598 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

Sculpture of T4: longitudinally striate laterally, granulate medially. Sculpture of T5:
weakly longitudinally striate laterally, smooth throughout remainder. Sculpture of T6:
smooth. Length of T6 in female: 1.25 times maximum width. Relative length of hind
basitarsus in female: 3 times as long as remaining tarsomeres. Relative length of hind
basitarsus in male: unknown.

Diagnosis. Phoenoteleia kaca sp. nov. can be distinguished from other species in
the genus by having T5 smooth and T6 lighter than T5.

Etymology. Kaca is the Indonesian word for glass and is used to reference the ap-
pearance of T5. The epithet is treated as a noun in apposition.

Material examined. Holotype, female: Mataysta: Sabah: Danum Valley, Silam
Rd. km 57, X1.86, MT, P. Eggleton, OSUC 359936 (deposited in CNCI). Paratypes:
InponesiA: 10 females, OSUC 256840 (ROME); OSUC 359937-359942, 359945
(CNCI); OSUC 359943, 359944 (OSUC).

Phoenoteleia kuboa Lahey, sp. nov.
http://zoobank.org/FOD01A7F-6B35-40E8-B9ED-6844187A4171
Figures 42-44

Description. Claval formula: 1-2-2-2-1. Number of clavomeres: 5. Color of antenna
in female: radicle and Al—AG light brown, A7—A12 brown. Color of A7 of female: con-
colorous with clavomeres. Color of antenna in male: unknown. Color of head: black.
Color of mesosoma: black. Color of metasoma: black. Color of legs: dark yellowish-
brown, coxae darkest. Claval formula: 1-2-2-2-1. Number of clavomeres: 5. Setation
of frons: short, sparse. Sculpture of frons: sparsely punctate ventrolaterally and dor-
sally, unsculptured medially. Sculpture of vertex: sparsely puncate anteriorly, mostly
smooth posteriorly except for short, longitudinal striae posteromedially. Excavation on
posteromedial vertex: absent. Sculpture of posteromedial vertex: longitudinally striate.
Sculpture of gena: sparsely punctate dorsally, longitudinally striate ventrally. Length
of LOL: 3 OD. Length of POL: two times as long as LOL. Sculpture of dorsal pro-
notal area: transversely punctate. Sculpture of lateral pronotal area: transversely stri-
ate. Sculpture of netrion: transversely striate. Notaulus: present. Mesoscutal humeral
sulcus: present. Mesoscutal suprahumeral sulcus: present. Sculpture of mesoscutum:
mostly smooth; coriaceous adjacent to notaulus between median channel. Parapsidial
line: absent. Setation of mesoscutellum: absent medially. Sculpture of mesoscutel-
lum: granulate posteriorly. Form of metascutellum: twice as long as wide, foveolate.
Sculpture of propodeum: lateral propodeal area smooth; plical area rugose. Setation
of plical area: present. Median area of the lateral propodeal area: present. Sculpture of
mesofemoral depression: smooth. Setation of mesepisternum ventral to mesofemoral
depression: sparse. Sculpture of mesepisternum ventral to mesofemoral depression:
canaliculate anteromedially, otherwise smooth. Setation of dorsal metapleural area:
absent. Sculpture of dorsal metapleural area: smooth throughout. Setation of ventral

Revision of Phoenoteleia 599

Figures 42-44. Phoenoteleia kuboa Lahey, female holotype (FBA040453) 42 head, mesosoma, meta-

soma, dorsal view 43 mesosoma, lateral view 44 head, anterior view. Scale bars in millimeters.

metapleural area: absent. Sculpture of ventral metapleural area: smooth. Length of
horn on T1: reaching apex of mesoscutum. Sculpture of horn on T1: rugose basally,
otherwise faintly transversely aciculate. Sculpture of T1 posterior to armilla: trans-
versely striate medially, longitudinally striate laterally. Sculpture of T2: longitudinally
striate, interstices rugose. Sculpture of T3: longitudinally striate, interstices rugose.
Sculpture of T4: longitudinally striate laterally, faintly granulate medially. Sculpture of
T5: weakly longitudinally striate laterally, faintly granulate medially. Sculpture of T6:
faintly granulate. Length of T6 in female: 1.5 times maximum width. Relative length
of hind basitarsus in female: 3 times as long as remaining tarsomeres. Relative length
of hind basitarsus in male: unknown.

Diagnosis. The presence of notauli on the mesoscutum and/or the completely smooth
frons can be used to distinguish P kuboa sp. nov. from other members of the genus.

Etymology. The epithet was inspired by the Fijian word for smoke (kubou) and is
intended to be treated as a noun in apposition.

Material examined. Holotype, female: Fit: Northern Div., Bua Prov., Vanua Levu
Isl., FJ-58A, 146m, 16°48.927'S 178°59.110'E, Kilaka Village, 3. VI-10.V1.2004, Ma-
laise trap, M. Irwin, E. Schlinger & M. Tokota’a, FBA040453 (deposited in BPBM).
Paratypes: Fy: 1 female, FBA047851 (CNCI).

600 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

Figures 45-48. Phoenoteleia rufa Kieffer, female holotype (MNHN 0025) 45 mesosoma, metasoma,

dorsal view 46 mesosoma, dorsal view 47 mesosoma, metasoma, lateral view 48 mesosoma, lateral view.

Scale bars in millimeters.

Phoenoteleia rufa Kieffer
http://zoobank.org/098F839B-1E69-4170-A3FD-7E704FA492DF
Figures 45-54

Phoenoteleia rufa Kieffer, 1916: 63 (original description); Kieffer, 1926: 551 (descrip-
tion); Baltazar, 1966: 185 (cataloged, type information, distribution); Johnson,
1992: 461 (cataloged, type information).

Phoenateleia rufa Kieffer: Kelner-Pillault, 1958: 151 (type information, spelling error).

Description. Claval formula: 1-2-2-2-1. Number of clavomeres: 5. Color of antenna
in female: radicle and Al—A7 yellow, A8—A12 brown. Color of A7 of female: dis-
tinctly lighter than clavomeres. Color of antenna in male: unknown. Color of head:
reddish-brown. Color of mesosoma: reddish-brown. Color of metasoma: reddish-
brown. Color of legs: light yellow-brown, fore legs darkest. Setation of frons: short,
sparse. Sculpture of frons: transversely rugose ventrally, smooth dorsally, punctate
throughout. Sculpture of vertex: punctate throughout, mostly smooth, reticulate
along margin of inner eye and minute, transverse sulci between punctures posteriorly.
Excavation on posteromedial vertex: absent. Sculpture of posteromedial vertex: same

Revision of Phoenoteleia 601

Figures 49-51. Phoenoteleia rufa Kieffer, female (OSUC 376916) 49 head, mesosoma, metasoma, dor-

sal view 50 mesosoma, lateral view 51 head, anterior view. Scale bars in millimeters.

as remainder of vertex. Sculpture of gena: punctate throughout, longitudinally rugose
ventrally, granulate dorsally. Length of LOL: 1 OD. Length of POL: two times as
long as LOL. Sculpture of dorsal pronotal area: areolate-rugose. Sculpture of lateral
pronotal area: transversely striate medially, rugose ventrally, granulate dorsolaterally.
Sculpture of netrion: transversely striate. Notaulus: absent. Mesoscutal humeral sul-
cus: not clearly differentiated from surrounding surface sculpture. Mesoscutal supra-
humeral sulcus: not clearly differentiated from surrounding surface sculpture. Sculp-
ture of mesoscutum: punctate throughout, rugose posterolaterally and along mar-
gins, granulate anteromedially. Parapsidial line: present. Setation of mesoscutellum:
present. Sculpture of mesoscutellum: granulate throughout. Form of metascutellum:
twice as wide as long, unsculptured, lateral corner acute. Sculpture of propodeum:
rugose. Setation of plical area: present. Median area of the lateral propodeal area:
present. Sculpture of mesofemoral depression: transversely rugose. Setation of mese-
pisternum ventral to mesofemoral depression: dense. Sculpture of mesepisternum
ventral to mesofemoral depression: punctate. Setation of dorsal metapleural area:
absent. Sculpture of dorsal metapleural area: smooth anteriorly, rugose posteriorly.
Setation of ventral metapleural area: present posteriorly. Sculpture of ventral meta-
pleural area: punctate posteriorly, otherwise smooth. Length of horn on T1: reaching
middle of mesoscutum. Sculpture of horn on T1: areolate-rugose basally and along

602 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

Figures 52-54. Phoenoteleia rufa Kieffer, female ( ) 52 head, mesosoma, metasoma, dorsal

view 53 mesosoma, lateral view 54 head, anterior view. Scale bars in millimeters.

margin of armilla, reticulate apically. Sculpture of T1 posterior to armilla: transversely
striate-rugose medially, longitudinally striate laterally, interstices rugose. Sculpture of
T2: longitudinally striate, interstices rugose. Sculpture of T3: longitudinally striate
throughout most of length, interstices rugose, posteromedially granulate. Sculpture
of T4: longitudinally striate laterally, granulate medially. Sculpture of T5: weakly
longitudinally striate laterally, faintly granulate medially. Sculpture of T6: punctate.
Length of T6 in female: 1.25 times maximum width. Relative length of hind basitar-
sus in female: 2.5 times as long as remaining tarsomeres.

Diagnosis. Phoenoteleia rufa is most similar to P. canalis from which is can be
separated by the sculpture of the frons and vertex.

Material examined. Holotype, female: PHitippines: Mindanao Isl., Butuan Char-
tered City, Baker, MNHN 0025 (deposited in MNHN). Other material: PHILIPPINEs:
2 females, (ROME); (MCZC). Tarwan: 2 females,

5 (CNCI).

Comments. Kieffer (1916) proposed the genus Phoenoteleia for a single female
collected in the Philippines. The head of the holotype is missing (Baltazar 1966),
but the meso- and metasoma are in good condition. We examined four specimens
that we consider conspecific with the holotype; two are from the Philippines and
two are from Taiwan. Additional specimens of P. rufa have been collected from the

Revision of Phoenoteleia 603

Figures 55-58. Phoenoteleia rufescens (Kieffer), male holotype (MNHN 0026) 55 head, mesosoma,
metasoma, dorsal view 56 head, mesosoma, dorsal view 57 head, mesosoma, metasoma, lateral view

58 head, mesosoma, lateral view. Scale bars in millimeters.

Amami Islands, part of the Ryukyu Island chain, in Japan (Y. Komeda, personal
communication). The Japanese material represents the most northernly distribution
of the genus.

Phoenoteleia rufescens (Kieffer)
http://zoobank.org/156FFD2A-9558-4BAD-ABD4-D2ED3A1E4C2A
Figures 55-59

Plagioscelio rufescens Kieffer, 1916: 186 (original description); Kieffer, 1926: 356,
357 (description, keyed); Kelner-Pillault, 1958: 151 (type information); Baltazar,
1966: 177 (cataloged, type information, distribution).

Phoenoteleia rufescens (Kieffer): Masner, 1976: 33 (generic transfer); Johnson, 1992:
461 (cataloged, type information).

Description. Color of antenna in female: unknown. Color of antenna in male: radi-
cle and A1l—A6 lighter than A7—A12. Color of head: light orange. Color of mesosoma:

mostly reddish-brown, with lighter areas on mesoscutum, mesopleuron, netrion, and

604 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

Figure 59. Phoenoteleia rufescens (Kieffer), male holotype (MNHN 0026) head, anterior view. Scale bar

in millimeters.

pronotum. Color of metasoma: mostly reddish-brown, except for T3 and T4 which
are lighter throughout most of their length. Color of legs: light yellow-brown, fore legs
darkest. Setation of frons: long, sparse. Sculpture of frons: transversely rugose ventral-
ly, smooth dorsally, punctate throughout. Excavation on posteromedial vertex: absent.
Length of LOL: 1 OD. Notaulus: absent. Setation of mesoscutellum: present. Sculpture
of mesoscutellum: granulate throughout. Form of metascutellum: transverse. Sculpture
of propodeum: rugose. Setation of plical area: present. Sculpture of mesofemoral de-
pression: smooth. Setation of mesepisternum ventral to mesofemoral depression: dense.
Setation of dorsal metapleural area: present posteriorly. Setation of ventral metapleural
area: present posteriorly. Sculpture of ventral metapleural area: transversely rugose an-
terodorsally, otherwise smooth. Sculpture of T2: longitudinally striate, interstices rugose.
Relative length of hind basitarsus in male: 2.5 times as long as remaining tarsomeres.

Diagnosis. ‘The granulate mesoscutellum and sculpture of the triangular propode-
al plate distinguishes P. rufescens from the male of P. buka sp. nov. We did not observe
any characters that would reliably separate P. rufescens from the male of P canalis.

Material examined. Holotype, male: PHmipprnes: Mindanao Isl., Butuan Char-
tered City, Baker, MNHN 0026 (deposited in MNHN).

Comments. The holotypes of P. rufescens and P. rufa were collected on the Philippine
island of Mindanao in Butuan, probably as part of the same collecting event. This led Mas-

Revision of Phoenoteleia 605

= a y Dd A £ PPA Wa
Figures 60-62. Phoenoteleia sanma Lahey, female holotype (OSUC 192538) 60 head, mesosoma, meta-

soma, dorsal view 61 mesosoma, lateral view 62 head, anterior view. Scale bars in millimeters.

ner (1976) to suggest that P. rufescens is the male of P. rufa. This is probably true, but we
refrain from synonymizing P. rufescens with P. rufa because we did not physically examine
the holotype, and the male is indistinguishable from specimens of the same sex that were
collected from locations where P. rufa females were not examined (Figures 63-65, 69-74).

In the original description of P. rufescens, Kieffer (1916) stated that the rear ocelli
are separated from the inner margin of the compound eye by twice their diameter. Im-
ages of the holotype show this is not the case: the rear ocelli appear to be positioned less
than 1 OD from the inner margin of the compound eye (Figure 56). Kieffer (1916)
also mentioned that the postmarginal vein (postmarginalis) of the fore wing is absent.
This is incorrect. Like all Phoenoteleia, the postmarginal vein of P. rufescens is at least
twice as long as the stigmal vein.

Phoenoteleia sanma Lahey, sp. nov.
http://zoobank.org/5CEFC24D-EE0E-4A6B-B873-99 194DA74003
Figures 60-62

Description. Claval formula: 1-2-2-2-1. Number of clavomeres: 5. Color of antenna

in female: radicle and Al—A7 light brown, A8—A12 brown. Color of A7 of female:

606 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

soma, metasoma, dorsal view 64 mesosoma, lateral view 65 head, anterior view. Scale bars in millimeters.

distinctly lighter than clavomeres. Color of antenna in male: unknown. Color of
head: light orange. Color of mesosoma: mostly orange, darker on mesoscutum lateral
to horn. Color of metasoma: horn orange apically, becoming progressively darker
posteriorly; T1 posterior to armilla burnt orange; T2—T3 orange; T4 burnt orange;
T5-T6 nearly brown. Color of legs: becoming progressively darker distally. Setation
of frons: long, moderate. Sculpture of frons: punctate-rugose. Sculpture of vertex:
sharply areolate-rugose. Excavation on posteromedial vertex: absent. Sculpture of pos-
teromedial vertex: same as remainder of vertex. Sculpture of gena: punctate-rugose.
Length of LOL: 1 OD. Length of POL: two times as long as LOL. Sculpture of dorsal
pronotal area: areolate-rugose. Sculpture of lateral pronotal area: transversely stri-
ate. Sculpture of netrion: transversely striate. Notaulus: absent. Mesoscutal humeral
sulcus: not clearly differentiated from surrounding surface sculpture. Mesoscutal
suprahumeral sulcus: not clearly differentiated from surrounding surface sculpture.
Sculpture of mesoscutum: areolate-rugose. Parapsidial line: absent. Setation of mesos-
cutellum: absent medially. Sculpture of mesoscutellum: granulate throughout. Form
of metascutellum: twice as wide as long, unsculptured, lateral corner acute. Sculpture
of propodeum: rugose. Setation of plical area: present. Median area of the lateral pro-
podeal area: present. Sculpture of mesofemoral depression: smooth anteriorly, punc-
tate-foveolate posteroventrally. Setation of mesepisternum ventral to mesofemoral

Revision of Phoenoteleia 607

Figures 66-68. Phoenoteleia sp. A, male (OSUC 359927; Kandy, Sri Lanka) 66 head, mesosoma, meta-

soma, dorsal view 67 mesosoma, lateral view 68 head, anterior view. Scale bars in millimeters.

depression: sparse. Sculpture of mesepisternum ventral to mesofemoral depression:
punctate. Setation of dorsal metapleural area: absent. Sculpture of dorsal metapleural
area: smooth anteriorly, rugose posteriorly. Setation of ventral metapleural area: pre-
sent posteriorly. Sculpture of ventral metapleural area: foveolate-rugose. Length of
horn on T1: reaching apex of mesoscutum. Sculpture of horn on T1: unsculptured
apically, otherwise areolate. Sculpture of T1 posterior to armilla: longitudinally stri-
ate throughout, interstices rugose. Sculpture of T2: longitudinally striate, interstices
rugose. Sculpture of T3: longitudinally striate, interstices rugose. Sculpture of T4:
longitudinally striate, interstices rugose. Sculpture of T5: smooth. Sculpture of T6:
punctate. Length of T6 in female: 1.25 times maximum width. Relative length of
hind basitarsus in female: 2.5 times as long as remaining tarsomeres. Relative length
of hind basitarsus in male: unknown.

Diagnosis. The sculpture of the frons, horn of T1, and ventral metapleural area
distinguishes P. sanma sp. nov. from its closest relative, P. buka sp. nov.

Etymology. The epithet refers to the collection locality (Sanma Province, Vanuatu)
and is meant to be treated as a noun in apposition.

Material examined. Holotype, female: Vanuatu: Santo Island, Penaoru 900B ~
900m, 18-30.xi.06, moist lowland forest ground, C. Villemant, MT; MG09B2, DNA
Voucher N116, OSUC 192538 (deposited in MNHN).

608 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 575-611 (2021)

Figures 69-71. Phoenoteleia sp. nr. rufescens, male (OSUC 331982; Milne Bay, Papua New Guinea)
69 head, mesosoma, metasoma, dorsal view 70 mesosoma, lateral view 71 head, anterior view. Scale bars

in millimeters.

Figures 72-74. Phoenoteleia sp. nr. rufescens, male (OSUC 232280; East Kalimantan, Indonesia)
72 head, mesosoma, metasoma, dorsal view 73 mesosoma, lateral view 74 head, anterior view. Scale bars

in millimeters.

Revision of Phoenoteleia 609

Comments. We describe this new species for a single female and the only speci-
men collected on the island nation of Vanuatu. The holotype is in excellent condition
but A9-A12 are missing from the right antenna.

Acknowledgements

We thank Sara Hemly (OSUC) for critical assistance with specimen handling and
databasing. Dr. Elijah Talamas (Florida State Collection of Arthropods, Gainesville,
Florida) is acknowledged for providing the image used in Figure 6 and for constructive
comments on Dicroscelio. Dr. Yoto Komeda (Teiso Kasei Co. Ltd., Shizuoka, Japan)
is thanked for discussions on the distribution of the genus, providing images of Phoe-
noteleia rufa from Japan, and for his careful critique that improved the quality of the
manuscript. We thank Dr. Ovidiu Popovici (Alexandru Ioan Cuza University, Iasi,
Romania) for providing images of P. canalis from Australia and Sulawesi and for allow-
ing us to publish collection data associated with those specimens. Lastly, we thank the
collectors for their efforts in capturing these magnificent animals.

References

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Revision of Phoenoteleia 611

Supplementary material |

Supplementary material for Revision of Phoenoteleia (Hymenoptera, Scelionidae,

Scelioninae)

Authors: Zachary Lahey, Luciana Musetti , Lubomir Masner , Norman E Johnson

Data type: Morphological terminology

Explanation note: List and definitions of morphological terms used in Revision of
Phoenoteleia (Hymenoptera, Scelionidae, Scelioninae).

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.

Link: https://doi.org/10.3897/jhr.87.59794.suppl1

Supplementary material 2

Figures $1-S4

Authors: Zachary Lahey, Luciana Musetti , Lubomir Masner , Norman E Johnson

Data type: Word doc file

Explanation note: Supplementary Figures S1-S4. Dicroscelio Kieffer, male (OSUC
185966; New Caledonia) $1 head, mesosoma, dorsal view $2 metasoma, fore wing
venation, dorsal view (inset: dorsal metasoma of Phoenoteleia fusca (=Plagioscelio
fuscus), line drawing of Kieffer (1926)) $3 mesosoma, lateral view $4 head, anterior
view. Scale bars in millimeters.

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.

Link: https://doi.org/10.3897/jhr.87.59794.suppl2