JHR 86: 47-61 (202 1) or JOURNAL OF = *20ereevee openacces journal

doi: |0.3897/jhr.86.72690 RESEARCH ARTICLE () I Tymenopter a
e

https://jhr.pensoft.net The international Society of Hymenopteriss. REGEARCH

Revision of the genus Schoenlandella (Hymenoptera,
Braconidae, Cardiochilinae) in the New World, with
a potential biological control agent for a lepidopteran
pest of bitter gourd (Momordica charantia L.)

Ilgoo Kang', Michael J. Sharkey”, Rodrigo Diaz'

| Department of Entomology, Louisiana State University Agricultural Center, 404 Life Sciences Building,
Baton Rouge, LA, 70803, USA 2. The Hymenoptera Institute, 116 Franklin Ave., Redlands, CA, 92373, USA

Corresponding author: Ilgoo Kang (ikang1@lsu.edu)

Academiceditor: Jose Fernandez- Triana | Received6 August 2021 | Accepted 21 September 2021 | Published29 October 2021
http://zoobank.org/5617D391-3973-4570-9722-E67A3EDF477E

Citation: Kang I, Sharkey MJ, Diaz R (2021) Revision of the genus Schoenlandella (Hymenoptera, Braconidae,
Cardiochilinae) in the New World, with a potential biological control agent for a lepidopteran pest of bitter gourd
(Momordica charantia L.). Journal of Hymenoptera Research 86: 47-61. https://doi.org/10.3897/jhr.86.72690

Abstract

Schoenlandella Cameron, 1905 is the second largest genus of Cardiochilinae. Most members are recorded
from the Old World, with a small number of species in the New World. Herein, the New World species of
Schoenlandella are revised based on morphological data. This work entails a description of a new species:
S. montserratensis Kang, sp. nov. and potential lepidopteran host information of the new species associated
with bitter gourds on the Caribbean Island of Montserrat. Schoenlandella diaphaniae (Marsh, 1986) and
S. gloriosa Mercado & Wharton, 2003 are re-described, and a key to species of New World Schoenlandella

is provided. The taxonomic status of Schoenlandella is discussed.

Keywords

Caribbean Islands, Crambidae, melonworm moth, Neotropical region, parasitoid wasp

Introduction

Schoenlandella Cameron, 1905 is the second largest genus of the subfamily Cardi-
ochilinae (Yu et al. 2016). All the members of the genus, which host data are avail-
able, are solitary endoparasitoids of exposed-feeding lepidopteran larvae such as

Copyright Ilgoo Kang 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.

48 Ilgoo Kang et al. / Journal of Hymenoptera Research 86: 47-61 (2021)

Chrysodeixis includens (Walker, 1858) (Soybean looper; Noctuidae) and Chloridea vire-
scens (Fabricius, 1777) (Tobacco budworm; Noctuidae). Fifty-four species have been
included in the genus worldwide, and most members are recorded from the Old World,
especially from the Afrotropical and Australian regions (Dangerfield et al. 1999; Mer-
cado and Wharton 2003; Yu et al. 2016). Three species have been recorded from the
New World: S. diaphaniae (Marsh, 1986), S. gloriosa Mercado & Wharton, 2003, and
S. longimala (Mao, 1945).

Herein, the New World species of Schoenlandella are revised. This includes a new
species description, potential host information of S. montserratensis Kang sp. nov., and
a key to species of New World Schoenlandella. Previously described species of New
World Schoenlandella are re-described.

Material and methods

Specimen information

During 2019, braconid wasps collected from a bitter gourd field in Montserrat by Dr
Chris Malumphy’s team (Fera Science Ltd.) were shipped to the first author (IK). Two
specimens were identified as a new species of Schoenlandella based on morphological
data, and potential lepidopteran hosts were identified.

Specimens were borrowed from the following institutions:

¢ Fera Sciences Ltd (London, UK);

e Hungarian Natural History Museum (HNHM: Budapest, Hungary);

e Illinois Natural History Survey (INHS: Champaign, Illinois, USA);

¢ Museo de Insectos, University of Costa Rica (MICR: San José, Costa Rica);

e Texas AXM University Insect Collection (TAMU: College Station, Texas, USA).

The holotype and sole paratype of S. montserratensis Kang sp. nov. will be deposited
in the Natural History Museum in London (NHML: London, UK).

Morphological analysis

A Leica MZ75 stereomicroscope was used to examine specimens. Morphological
terminology used in this review are mostly based on Dangerfield et al. (1999) and
Sharkey and Wharton (1997), supplemented by terms on the Hymenoptera Anat-
omy Consortium website (http://portal.hymao.org/projects/32/public/ontology/,
accessed in September 2020). Terms for surface sculpture follow Harris (1979).
The following acronyms are used for morphological terms: T1: first metasomal
tergite; [2: second metasomal tergite; T'3: third metasomal tergite; T'4: forth meta-
somal tergite; [6: sixth metasomal tergite. Color images were taken with a Vision-

ary Digital BK Plus imaging system (Dun, Inc.) equipped with a Canon EOS

Revision of the genus Schoenlandella in the New World 49

5DS DSLR camera. Multiple images were stacked using Zerene Stacker v.1.04
(Zerene Systems LLC.) and edited in Adobe Photoshop CS 6 and Photoshop CC
(Adobe Systems, Inc). Body parts were measured also via Adobe Photoshop CS 6
and Photoshop CC. Each number in parentheses in species descriptions indicate
0.01x the actual size of each body part. The unit of length is mm. For example,
12 and 123 in parentheses (12:123) indicate 0.12 mm and 1.23 mm, respectively.
The distribution map of the new species was generated using QGIS 3.10.0 (QGIS
Development Team 2019). The Google terrain map was downloaded using Quick-
MapServices plugin.

Results

Taxonomy
Schoenlandella Cameron, 1905

Schoenlandella Cameron, 1905 (Cameron 1905a). Type Species: Schoenlandella ni-
gromaculata Cameron, 1905 (Cameron 1905a) by subsequent designation by
Viereck, 1914 (synonymized with Cardiochiles Nees, 1819 by Szépligeti, 1911).
Removed from synonymy by Whitfield & Dangerfield, 1997.

Ernestiella Cameron, 1905 (Cameron 1905b) synonymized with Schoenlandella Vi-
ereck, 1914. Type species: Ernestiella nigromaculata Cameron 1905 (Cameron

1905b).

Diagnosis (based on Dangerfield et al. (1999) with additions and modifications).
Diagnostic characters of Schoenlandella were described in Dangerfield et al. (1999),
based mostly on Old World species. The following are diagnostic features based on
both Old World and New World members.

Members of Schoenlandella can be distinguished from species of other cardiochiline
genera by the following characters: Head: possessing 32—44-segmented antenna; densely
setose eye (Figs 2C, 4C); two clypeal tubercles with sharp or smooth apical margin (Fig.
1C); short to elongate malar space; bidentate mandible; six-segmented maxillary palpus;
four-segmented labial palpus; absence of occipital carina; elongate galea, at least as long
as malar space in lateral view (Note: narrower than galeae of members of Cardiochiles
Nees, 1819) (Figs 1A, 2A, 4A); glossa elongate and deeply bilobed if entirely spread (Fig.
4A). Mesosoma: notaulus finely crenulate; scutellar sulcus crenulate; posterior scutel-
lum without cup-like pit; pronotum mostly smooth; mesopleuron mostly smooth; ep-
icnemial carina absent; metapleuron rugulose; mesosternal sulcus crenulate; propodeum
rugulose; propodeum with completely developed areola. Wings: stigma of forewing
moderate to broad; (RS+M)a of forewing present; lr of forewing absent; spectral 3r of
forewing reaching at basal fifth to half, if absent, examine characters of mouthparts and

hypopygium; RS vein of forewing angled or curved (Fig. 4F); 1a of forewing absent

50 Ilgoo Kang et al. / Journal of Hymenoptera Research 86: 47-61 (2021)

Figure |. Schoenlandella diaphaniae, non-type A lateral habitus B dorsal habitus C anterior head

D ventro-lateral metasoma E dorsal propodeum and mesonotum.

(Note: if present, the vein is nebulous); 2—1A of hind wing usually absent. Legs: tarsal
claw pectinate with sharp or obtuse apical tooth; hind basitarsus cylindrical (Fig. 1A)
or slightly expanded (Fig. 4A) (Note: not nearly as expanded as in Hartemita Cameron,
1910); Metasoma: lateral suture of T1 absent posteriorly; T2 and T3 entirely smooth;
hypopygium apically acute in lateral view (Fig. 1D); hypopygium uniformly sclerotized
(Fig. 1D); median longitudinal fold of hypopygium absent (Fig. 1D) (Note: if present,
the fold is entirely sclerotized or only slightly desclerotized (Figs 2D, 4D)); ovipositor
slightly downcurved (Figs 1A, 2D); ovipositor sheath < ~0.6 x length of hind tibia.

Key to species of the genus Schoenlandella of the Neotropical region

1 A. Body mostly imelanicts2,.t5, sss: anet mae S. longimala (Mao, 1945) Note:
Based on images of the holotype on the NMNH website (https://collections.
nmnh.si.edu/search/ento/?ark=ark:/65665/3a9805bdd5 1964c16838984ab0
e197ed9, accessed September 2021)

Br Bodyimostliorangerotyellow pales. Bea. acca ee ene lta Pes onsale dy ne 4
2 A. Malar space < ~0.33 x longer than eye height in anterior view, AA. fore-
Wineventirely iniliscate-wer. mes Metco S. montserratensis Kang, sp. nov.

- B. Malar space about ~0.50 x longer than eye height in anterior view, BB.
forewing. apical lyaintiuscaterssiy.5. 0. niis decane emeee tae s RO ceat oy nce al 3

Revision of the genus Schoenlandella in the New World 51

3 A. Forewing basally hyaline, and stigma entirely melanic, AA. apex of clypeal
tubercle sharply aneledh de sescseaschs. tung tootegee + S. diaphaniae (Marsh, 1986)
- B. Forewing basally pale yellow, and stigma entirely pale, BB. apex of clypeal
tubercle relatively smooth ............... S. gloriosa Mercado & Wharton, 2003

Schoenlandella diaphaniae (Marsh, 1986)
Fig. [A-E

Cardiochiles diaphaniae Marsh, 1986 (Marsh 1986)
Schoenlandella diaphaniae (Marsh, 1986) (Dangerfield et al. 1999)

Material examined. Non-type specimen:TRINIDAD AND Tosaco ® 19; Curepe, Trini-
dad and Tobago; 21 Jul. 1978. Malaise Trap. Deposited in INHS.

Diagnosis. Members of S. diaphaniae are distinguished from members of S. glo-
riosa by having shorter lower face and malar space (Fig. 1C); basally hyaline forewing
(Fig. 1A); stigma entirely melanic (Fig. 1A).

Description. Marsh (1986) described color of the species and some morphological
characters in his species description. Here, the species is re-described based on a speci-
men collected in Trinidad and Tobago.

Body length 4.5—5.8 mm (Marsh 1986). Antenna length: ~-4.2 mm. Forewing
length: ~5.5 mm. Head: Antenna 34-segmented. Eye length ~0.45 x longer than its
height (40:89). Dorsal width of lower face as long as its height (81:81); Malar space
~0.40 x longer than height of eye in anterior view (32:80), ~2.13 x longer than basal
width of mandible (32:15) (Fig. 1C). Clypeus ~1.53 x longer than its width (49:32);
clypeal tubercles with sharp margins (Fig. 1C). Galea as long as malar length in lateral
view (32:32), with curved apical margin (Fig. 1A). Mesosoma: Scutellar sulcus with five
to six crenulae. Postscutellar depression present. Propodeum rugulose; median areola
of propodeum diamond-shaped, median length as long as its width; propodeum with
median transverse carina reaching lateral margin. Pronotum mostly smooth with in-
complete posteroventral carina reaching posterior margin. Mesopleuron mostly smooth
and polished; precoxal sulcus medially present with five crenulae. Metapleuron rugu-
lose. Mesosternal sulcus crenulate. Legs: Basal spur on mid tibia ~0.83 x mid-basitarsus
(49:59). Hind femur medially ~0.33 x broader than its length (45:137). Basal spur on
hind tibia ~0.66 x longer than hind basitarsus (60:90). Hind basitarsus cylindrical.
Hind tarsal claw pectinate with five teeth; apical tooth basally rounded and apically
angled; basal four teeth sharp. Wings: Forewing second submarginal cell its maximum
width ~2.87 x longer than its maximum length (89:31); 3r absent; Rs broken basally
and angled at a basal sixth; stigma about ~3.44 x longer than width medially (93:27).

52 Ilgoo Kang et al. / Journal of Hymenoptera Research 86: 47-61 (2021)

Figure 2. Schoenlandella gloriosa, non-type Honduras A lateral habitus B dorsal habitus C anterior head

D ventral metasoma; arrow: median invagination on hypopygium E dorsal propodeum and mesonotum

F wings.

Hind wing 2-1A present at basal third (Fig. 1A). Metasoma: Medial length of T1
~2.59 x longer than medial length of T2 (75:29). Medial length of T2 ~0.22 x longer
than its apical width (29:129). T3 about ~1.66 x longer than T2 medially (48:29).
Hypopygium entirely sclerotized, medially with shallow area, without median longitu-
dinal fold (Fig. 1D). Ovipositor moderately downcurved. Protruded ovipositor sheath
moderately downcurved, ~0.57 x longer than hind tibia (101:176), slightly broadened
apically, anterior 2/5 depilous and posterior 3/5 pilose apically with long setae.

Color. Body mostly bright yellow; the following areas melanic: flagellomeres, pedi-
cel mostly, outer scape; ocellary field and frons dorsally, labrum, apical mandible, galea
mostly, mid tibia apically, hind coxa apically, hind trochanter and trochantellus, hind

Revision of the genus Schoenlandella in the New World 53

Map data ©2015 Google

ore
ian.
. ¥ 4
ee ‘| y
e * ! i d 2

3 Schoenlandella montserratensis

Figure 3. Distribution map of S. montserratensis sp. nov. in Montserrat.

femur basally and apically, hind tibia apically, hind tarsomeres apically, entire oviposi-
tor sheath. T4—T6 medially (Fig. 1E). Wings basally hyaline and apically infuscate,
C+SC+R vein in forewing mostly melanic, stigma mostly melanic.

Male. See Marsh (1986).

Hosts. Diaphania nitidalis (Stall) and D. hyalinata (L.) (Marsh 1986).

Distribution. Colombia, Venezuela, and Trinidad and Tobago.

Schoenlandella gloriosa Mercado & Wharton, 2003
Fig. 2A—F

Material examined. Paratypes: Mexico ® 19; 3 mi E Papantla, Veracruz, Mexico; 7
Jun. 1965; leg. Burke, Meyer, Schaffner ¢ 19; 2 mi SE Tecpan de Galeana, Guerrero,
Mexico; 14 Jul. 1966; leg. RM and PK Wagner ¢ 24; Hotel Covandonga, 12 km S
Valles, Ruta 85, San Luis Potosi, Mexico; 27—29 Jun. 1981; leg. C. Porter, L. Stange.
Deposited in TAMU. Non-type material: Costa Rica ¢ 19; 10 km NW Cafas, Moji-
ca, Guanacaste, Costa Rica; 26 Sep.—10 Oct. 2011; leg. P. Hanson. Deposited in MICR.
Honduras ¢ 19; Tela, Lancetilla, Atlantida, Honduras; 15°43'N, 87°27'W; 30 Apr.
1995; leg. R. Cave. 12; same as previous except for the collecting date, 15 May. 1995.
14; same as previous except the collecting date, 31 Aug. 1995. Deposited in HNHM.

Diagnosis. Members of the S. gloriosa are nearly identical to S. diaphaniae. The
following combination of characters differentiate S. gloriosa from S. diaphaniae: face

54 Ilgoo Kang et al. / Journal of Hymenoptera Research 86: 47-61 (2021)

concave; malar space relatively elongate (~0.50 x longer than height of eye in anterior
view) (Fig. 2C); two clypeal tubercles with smooth margins (Fig. 2C); forewing basally
yellow and apically infuscate; stigma entirely pale; forewing with junction of angled Rs
not swollen (Fig. 2F).

Description. Modified from Mercado and Wharton (2003), including additional
characters.

Body 5.5—8.0 mm. Head: Antenna 32—34-segmented. Eye length ~0.41 x longer
than its height (35:85) in lateral view. Dorsal width of lower face ~1.05 x longer
than height of lower face (79:75). Clypeus 1.20—1.53 x longer than its width, two
clypeal tubercles with smooth margin (Fig. 2C). Malar space 0.47—0.50 x longer than
height of eye in anterior view (35:75—40:80), 2.75—3.42 x longer than basal width
of mandible (33:12—41:12). Galea 1.30-1.40 x longer than malar length as viewed
laterally (43:33-56:40) (Fig. 2A). Mesosoma: Scutellar sulcus with 5—7 crenulae.
Propodeum with median transverse carina reaching lateral margin. Pronotum weakly
carinate medially. Mesopleuron mostly smooth; precoxal sulcus smooth, not reaching
posterior margin. Legs: Basal spur on mid tibia 0.83—0.90 x mid-basitarsus length.
Hind femur medially 0.31-0.32 x broader than its length (52:164—48:155). Wings:
Forewing: maximum width of second submarginal cell ~2.26x longer than maximum
length (113:50); 3r absent; Rs vein angled at basal fourth (Fig. 2F). Hind wing 2-1A
present as basal stump (Fig. 2F). Metasoma: Medial length of T1 2.00—2.47 x longer
than medial length of T2 (78:39-75:30). Medial length of T2 0.21-0.26 x long-
er than its apical width (30:140-39:149). T3 about 1.46—-1.60 x longer than T2
medially (57:39-48:30). Hypopygium evenly sclerotized, median longitudinal fold
absent (Note: A weakly depressed medial longitudinal area is present in females col-
lected in Honduras, but never membranous and folded) (Fig. 2D). Ovipositor mod-
erately downcurved. Protruded ovipositor sheaths moderately downcurved, 0.45—
0.50 x longer than hind tibia, broadened apically, anteriorly depilous and moderately
pilose apically with long setae.

Color. See Mercado and Wharton (2003). Melanic areas of the Costa Rican spec-
imen (female) and Mexican specimens are slightly darker than specimens collected
in Honduras.

Male. See Mercado and Wharton (2003).

Host. Unknown.

Distribution. Costa Rica, Honduras, and Mexico.

Schoenlandella longimala (Mao, 1945)

Cardiochiles longimala Mao, 1945 (Mao 1945)
Schoenlandella longimala (Mao, 1945) (Dangerfield et al. 1999)

Type material. Holotype: Mexico * 3, Guadalajara, Mexico; 2 Aug. 1914; Deposited
in NMNH.

Revision of the genus Schoenlandella in the New World 3)

Diagnosis (based on images of the holotype on the NMNH website). Body
mostly black except for legs. Malar space shorter than basal width of mandible. Mouth-
parts moderately elongated. Scutellar sulcus with six crenulae. Lateral side of scutellum
mostly rugulose. Pronotum medially rugose, with median areola anteriorly angled.
Forewing entirely infuscate; stigma entirely melanic; 3r vein absent; la present as a
nebulous vein.

Description. See Mao (1945).

Host. Unknown.

Distribution. Mexico.

Schoenlandella montserratensis Kang, sp. nov.
http://zoobank.org/D22783D0-15B6-422E-9989-34760DA85579
Fig. 4A—F

Material examined. Holotype: MONTSERRAT ® 2; 16°45'34.19"N, 62°13'1.58"W;
leg. Elvis Gerald (Ref. CM-Mt-2019-41) woodlands, private farm. Single adult on
Momordica charantia L. (bitter melon), hand caught in a plastic pot. Paratype same
data as for holotype.

Diagnosis. Members of Schoenlandella montserratensis sp. nov. are distinguished
from other New World Schoenlandella species by having shorter malar space (Fig. 4C);
longer mouthparts (Fig. 4C); stigma basally pale and apically melanic (Fig. 4F); 3r vein
of forewing present at basal half (Fig. 4F).

Description. Body 4.21—4.40 mm. Forewing length: ~4.45 mm (holotype). An-
tenna length: ~3.25 mm (paratype). Head: Antenna 34-segmented (paratype). Eye
length ~0.52 x longer than its height (31:60). Malar space slightly shorter than basal
width of mandible. Clypeus ~2.08 x longer than its width (50:23); two clypeal tu-
bercles with smooth margins. Galea ~2.12 x longer than malar space in lateral view
(36:17), apically narrowed. Mesosoma: Scutellar sulcus with five crenulae. Propodeal
median transverse carina reaching lateral margin. Pronotum medially crenulate, ven-
trally costate. Mesopleuron mostly smooth; precoxal sulcus strongly crenulate with ~ 10
crenulae, not reaching posterior margin. Legs: Basal spur on mid tibia ~0.88 x mid-ba-
sitarsus (30:34). Basal spur on hind tibia ~0.68 x longer than hind basitarsus (36:53).
Hind basitarsus laterally broaden. Tarsal claw pectinate with five teeth; apical tooth
obtuse, other remaining teeth sharp. Wings: Forewing second submarginal cell width
~2.79 x longer than height (78:28); 3r apparently present at basal half and slightly
curved; Rs angled at basal two-fifths; stigma about ~3.36 x longer than wide medially
(74:22); 1a absent (Fig. 4F). Hind wing 2—1A absent (Fig. 4F). Metasoma: Medial
length of T1 ~2.68 x longer than medial length of T2 (59:22). Medial length of T2
~0.20 x longer than its apical width (22:110). 3 entirely smooth, ~1.27 x longer than
T2 medially (28:22). Hypopygium surface entirely sclerotized with a distinct median
longitudinal fold (Fig. 4D). Ovipositor slightly downcurved; protruded ovipositor
sheath ~0.40 x longer than hind tarsus, broadened apically.

56 Ilgoo Kang et al. / Journal of Hymenoptera Research 86: 47-61 (2021)

7 _|
ovate farm. 18908524 197,
DIOLS SSW, Sage sauten
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Figure 4. Schoenlandella montserratensis, sp. nov. A lateral habitus B dorsal habitus C anterior head
D ventral metasoma; arrow: median fold on hypopygium E dorsal propodeum and mesonotum F wings;

arrow: 3r vein on forewing.

Revision of the genus Schoenlandella in the New World 57

Color. Body mostly pale orange; the following areas melanic: apical scape, pedicel,
flagellomere, apical mandible, hind tarsus, external ovipositor sheaths. Wings entirely
lightly infuscate, stigma dark brown at apical half.

Etymology. This species is named after the collecting site, “Montserrat”, a volcanic
Caribbean Island.

Host. Unknown but see details in the following discussion section.

Distribution. Schoenlandella montserratensis sp. nov. is only known from Montser-

rat (Fig. 3).

Discussion

Taxonomic status of Schoenlandella and character discussion

The validity of the taxonomic status of Schoenlandella Cameron has fluctuated several
times before the current work. Dangerfield et al. (1999) indicated that species of Sch-
oenlandella could be easily distinguished from all other cardiochiline genera based on
the following five characters: conspicuously setose eyes; long and deeply bilobed glossa;
elongate blade-like galea; presence of spectral 3r of forewing (if absent, members of the
genus have relatively short mouthparts); evenly sclerotized hypopygium (if a median
longitudinal fold is present, the surface is not membranous). The genus was resolved
as a monophyletic group in the phylogeny in Dangerfield et al. (1999) based on mor-
phological data. However, Mercado and Wharton (2003) and Papp (2014) indicated
difficulty in distinguishing members of Schoenlandella from those of Cardiochiles Nees.
Dangerfield et al. (1999) and Mercado and Wharton (2003) retained Schoenlandella as
a valid genus, and the latter author subsumed the genus into Cardiochiles as a subge-
nus. Edmardash et al. (2018) treated Schoenlandella as a valid genus.

Only one study partially indicated genus-level relationships of Cardiochilinae.
Murphy et al. (2008) presented three phylogenies based on seven genes. Even though
the phylogenies focused on subfamily-level relationships of microgastroid subfamilies,
genus-level relationships of five cardiochiline genera were included. Cardiochiles and
Schoenlandella were resolved as polyphyletic in a clade with a rare Australian genus,
Gwenia Dangerfield, Austin & Whitfield in their Maximum Parsimony phylogeny.
However, the other two phylogenies resulting from Maximum Likelihood and Bayes-
ian phylogenetic analyses using the same molecular data indicated that members of
each genus were well-clustered even though they were recovered as paraphyletic. It
should be noted that Murphy et al. (2008) treated C. minutus (Cresson) as a member
of Schoenlandella and indicated Cardiochiles and Schoenlandella were polyphyletic. We
treat S. minuta in Murphy et al. (2008) as C. minutus (Cresson) since Mercado and
Wharton (2003) transferred the species to Cardiochiles.

In the current work, we examined specimens of both New World and Old World
Schoenlandella species. Old World members mostly possess and share the five diagnos-
tic characters defined by Dangerfield et al. (1999). We confirmed that specimens of

58 Ilgoo Kang et al. / Journal of Hymenoptera Research 86: 47-61 (2021)

S. montserratensis sp. nov. collected in Montserrat are the first New World species hav-
ing the five diagnostic characters. Of the other three New World species, specimens of
S. diaphaniae and S. gloriosa were examined, and morphological characters of S. longi-
mala were confirmed based on descriptions of Mao (1945) and images of the holotype
on the NMNH website (https://collections.nmnh.si.edu/search/ento/, accessed Sep-
tember 2020). All New World species have dense eye setae consistent with Old World
members. Dangerfield et al. (1999) indicated that most Schoenlandella members pos-
sess elongate mouthparts. Tucker et al. (2012) discussed two hypotheses regarding the
elongation of mouthparts of the Afrotropical agathidine genus Camptothlipsis Ender-
lein and preferred the hypothesis that the members with elongate mouthparts are spe-
cialized nectar feeders, dependent on flowers that their host caterpillars feed on. ‘This
has been confirmed in the case of Agathis malvacearum Latreille (Juhala 1967). Most
other cardiochilines as well as members of Heteropteron Brullé, which was resolved as
the most ancestral cardiochiline genus (Dangerfield et al. 1999; Murphy et al. 2008)
possess short mouthparts. The elongate mouthparts most likely evolved independently
in Schoenlandella. Some Old World members and the three New World species in the
current work possess relatively shorter mouthparts than most Schoenlandella species
(but narrower and longer than the typical mouthparts of Cardiochiles). This seems to
be a secondary loss due to environmental changes like host flower traits. In addition,
IK confirmed that females of S. diaphaniae and S. gloriosa do not possess the median
membranous fold or area in the hypopygium. ‘The hypopygial character of S. longimala
could not be confirmed because the holotype is male. Additionally, most Schoenlan-
della members possess an angled RS vein in the forewing, which is not present in
Cardiochiles. They also have relatively longer second submarginal cells than members
of Cardiochiles. Despite the contradictory opinions of Papp (2014) and Murphy et al.
(2008), we treat Schoenlandella as a valid genus as in Dangerfield (1999) and Mercado
and Wharton (2003) and place the New World species in the genus. IK is conducting
molecular analysis to generate a phylogeny to determine genus-level relationships of
cardiochilines based on a larger taxon sample than previous research. We expect that
the phylogeny will help resolve the taxonomic issue in the near future and predict that
Schoenlandella will be confirmed as a valid genus with support of shared morphological
characters such as elongate mouthparts, angled Rs vein and elongated submarginal cell
of the forewing, and evenly sclerotized surface of the hypopygium.

Potential host information of S. montserratensis sp. nov.

Female adults of S. montserratensis sp. nov. were collected in a bitter gourd field in
Montserrat in 2019. Four potential lepidopteran host species were collected in the
same location in the same period: Diaphania hyalinata (L., 1767) (Melonworm
moth; Crambidae); Plutella xylostella (L. 1758) (Diamondback moth; Plutellidae);
Calpodes ethlius (Larger canna leafroller; Hesperiidae), and unidentified bagworm
(Psychidae) (Dr Chris Malumphy pers. comm.). Among these species, the mel-
onworm moth caterpillars were the most serious pests in the field. Because other
Schoenlandella species chiefly attack caterpillars of Crambidae and Noctuidae (Ta-

Revision of the genus Schoenlandella in the New World 59

Table |. Host information of each Schoenlandella species prior to the current study.

Schoenlandella species Lepidopteran host Source(s)
S. diaphaniae (Marsh, 1986) Diaphania hyalinata (L., 1767) (Crambidae) (Marsh 1986)
S. fulva (Cameron, 1907) Omiodes indicata (F., 1775) (Crambidae), (Odak and Dhamdhere 1968)

Spoladea recurvalis (F., 1775) (Crambidae),
Pramadea lunalis (Guenée, 1854) (Crambidae)

S. goosei (Dangerfield & Austin, 1995) Crocidolomia pavonana (F., 1794) (Crambidae) (Dangerfield and Austin 1995)

S. hymeniae (Fischer & Parshad, 1968) Spoladea recurvalis (E., 1775) (Crambidae) (Fischer and Parshad 1968)

S. nigromaculata Cameron, 1905 Helicoverpa armigera (Hubner, 1808) (Noctuidae) (Nyiira 1970)

S. sahelensis (Huddleston & Walker, 1988) Heliocheilus albipunctella (de Joannis, 1925) (Bhatnagar 1988) (Huddleston

(Noctuidae) and Walker 1988)

S. trimaculata Cameron, 1905 Helicoverpa armigera (Hubner, 1808) (Noctuidae) (Coaker 1959) (Risbec 1960)

S. uniformis (Turner, 1918) Helicoverpa sp. (Noctuidae) (Chadwick and Nikitin 1976)
(Dangerfield and Austin 1995)

S. variegata (Szépligeti, 1913) Helicoverpa armigera (Hiibner, 1808) (Noctuidae) (Bhatnagar 1988) (Huddleston

and Walker 1988)

ble 1), the melonworm moth caterpillar may be the preferred host of S. montser-
ratensis sp. nov. If females of the new species are host specific and attack the melon-
worm moth caterpillars as expected, females of S. montserratensis sp. nov. should be
developed as effective biological control agents for the melonworm moth. Detailed
biology of S. montserratensis sp. nov. and its host range will need to be confirmed
in further investigations (i.e., rearing caterpillars and collecting parasitoids of the
host caterpillars). Future studies should investigate the effectiveness of S. montser-
ratensis sp. nov. to parasitize the melonworm moth and determine its potential as a
biological control agent against melonworm moth.

Acknowledgements

Thanks to Dr Chris Malumphy (Fera Science Ltd.) who led the collecting team in
Montserrat as well as Mr Rob Deady (Fera Science Ltd.) who sent the specimens of
S. montserratensis sp. nov. We are also grateful to LSU AgCenter for financial support
and Dr Karen Wright in TAMU, Dr Paul Hanson in the MICR, Dr Tommy McElrath
in the INHS, and Dr Zoltan Vas in the HNHS for the loans of specimens. We also
thank Dr James Whitfield who donated cardiochiline specimens to INHS, Dr Michael
Stout for his support, and Mr Josh Snook for reviewing the manuscript. In particular,
IK would like to acknowledge Dr Chris Carlton and Ms Victoria Bayless for their

encouragement and supports.

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