Corresponding author: Elijah J. Talamas (
Academic editor: M. Yoder
Talamas EJ, Johnson NF, Buffington M (2015) Key to Nearctic species of
A decade after its introduction into the United States in 1999, the economically destructive brown marmorated stink bug (BMSB),
BMSB is difficult to manage with pesticides because it feeds on interior plant tissues via its proboscis, bypassing ingestion of pesticides that are deposited on the surfaces of plant tissues. As a result, increased pesticide applications to combat BMSB disrupt ecosystem services, resulting in secondary pest outbreaks (
This paper is presented as an updated synthesis of the works of Norman
Although it may be impossible to fully predict which species will become introduced pests, educated decisions can be made about which species have the greatest potential, and similarly, which species have potential as biological control agents. While this paper was in review, a wild population of
The identification keys of
The contributions of the authors are as follows: E.J. Talamas: character definition and coding, imaging, manuscript preparation; N.F. Johnson: character definition and coding, manuscript preparation; M. Buffington: manuscript preparation, project coordination.
The locality data reported for primary types are not literal transcriptions of the labels: some abbreviations are expanded; additional data from the collectors are also included. The numbers prefixed with “USNMENT” or “OSUC ” are unique identifiers for the individual specimens (note the blank space after some acronyms). Details on the data associated with these specimens may be accessed at the following link,
Persistent URIs for each taxonomic concept were minted by xBio:D in accordance with best practices recommend by
Morphological terms were matched to concepts in the
Photographs were captured with a Z16 Leica®™ lens with a JVC KY-F75U digital camera using Cartograph®™ software, or a Leica®™ DMRB compound microscope with a GT-Vision®™ Lw11057C-SCI digital camera attached. In both systems, lighting was achieved using techniques summarized in
High quality optics and bright, diffuse lighting are critical for observing the characters in this key. The authors recommend fluorescent desk lamps, or fiber optic lamps with mylar sleeves affixed to the tips of the light pipes, or a mylar ‘shield’ between the tip of the light pipes and the specimen. Direct illumination of the specimen should be avoided. Additionally, some characters are better observed with appendages moved (especially the legs in couplet 5 and the wings in couplet 6 of the
This work is based on specimens deposited in the following repositories with abbreviations used in the text:
We coin this term to refer to the structure formed by the transaxillar, axillar, and axillular carinae located posterodorsal to the wing base (see Figs
In the Nearctic fauna, species in the
The episternal foveae of the
The presence of striae on the frons is typically weakly indicated or entirely absent, with a few exceptions. In some species the striae are present as shallowly incised short lines arising from the anterior articulation of the mandible (eg.
The mesopleural carina was used more extensively for species identification in the key of
Among the published descriptions and diagnoses, we have not encountered previous use of this character for species-level delimitation in
A1–12 antennomeres 1–12 (Fig.
ac acetabular carina (Figs
aem anteroventral extension of metapleuron (Figs
anfo antennal foramen (Fig.
aoc anterior ocellus (Fig.
as antennal scrobe (Figs
atcs antecostal sulcus (Figs
ats postacetabular sulcus (Figs
axcr axillar crescent (Figs
bs basiconic sensilla (Figs
cs clypeal setae (Fig.
ctk central keel (Figs
cx1 procoxa (Fig.
cx2 mesocoxa (Fig.
cx3 metacoxa (Fig.
eps episternal foveae (Figs
fs facial striae (Figs
gc genal carina (Figs
gen gena (Fig.
hoc hyperoccipital carina (Figs
iap interantennal process (Fig.
loc lateral ocellus (Figs
lt(s) laterotergite(s) (Figs
mc mesopleural carina (Fig.
mdb mandible (Fig.
mmc median mesoscutal carina (Fig.
mms median mesoscutal sulcus (Fig.
mpp mesopleural pit (Figs
ms malar sulcus (Figs
mscm mesoscutum (Figs
msct metascutellum (Figs
mshs mesoscutal humeral sulcus (Figs
mspl mesopleuron (Fig.
mtnm metanotum (Figs
mtpl metapleuron (Fig.
mtpm metapostnotum (Figs
nes netrion sulcus (Figs
not notaulus (Figs
of orbital furrow (Figs
pcxs paracoxal sulcus (Figs
ppm propodeum (Figs
prnm pronotum (Figs
prpl propleuron (Fig.
pvm posteroventral portion of metapleuron (Figs
r radicle (Figs
scu mesoscutellum (Figs
ss sublateral seta (Figs
T1–6 mediotergite (Figs
tga tegula (Figs
The following key includes platygastroids with host records indicating emergence from pentatomoid eggs. More associations are certain to exist, particularly in
1 | Metasoma with laterotergites tightly appressed to sternites, forming a sharply angled lateral margin (Fig. |
|
– | Metasoma with laterotergites wide and loosely attached to sternites, metasoma without sharp lateral margin (Fig. |
|
2 | Frons with central keel extending from interantennal process to anterior ocellus (Figs |
|
– | Frons without central keel or keel short, not extending to anterior ocellus (Figs |
|
3 | Mesoscutum with notauli (Fig. |
|
– | Mesoscutum without notauli (Fig. |
|
4 | T2 longer than wide (Fig. |
|
– | T2 wider than long (Fig. |
|
1 | Metapleuron with posteroventral portion glabrous (Figs |
|
– | Metapleuron with posteroventral portion setose (Figs |
( |
2 | Vertex with hyperoccipital carina (Figs |
( |
– | Vertex without hyperoccipital carina (Fig. |
( |
3 | Frons between antennal scrobe and anterior ocellus with parallel, arched rugae (Figs |
|
– | Frons between antennal scrobe and anterior ocellus smooth or with rugae that are not parallel and arched (Figs |
|
4 | Inner margin of eye with orbital furrow constricted ventrally (Fig. |
|
– | Inner margin of eye with orbital furrow expanded near intersection with malar sulcus (Figs |
|
5 | Clypeus with 4 setae (Fig. |
|
– | Clypeus with 2 setae (Fig. |
|
6 | Female with antennal flagellum distinctly bicolored: A3–A6 yellow, A7–A11 dark brown (Fig. |
|
– | Female with antennal flagellum (A3–A11) infuscate throughout (Fig. |
|
7 | Mesopleuron with anteroventral portion rugulose (Figs |
|
– | Mesopleuron with anteroventral portion smooth or with shallowly impressed microsculpture (Figs |
|
8 | Mesoscutellum coarsely rugose (Figs |
|
– | Mesoscutellum smooth or with coriaceous microsculpture (Figs |
|
9 | Radicle yellow (Fig. |
|
– | Radicle dark brown to black (Fig. |
|
10 | Metapleuron with paracoxal sulcus visible in ventral half (Fig. |
|
– | Metapleuron with paracoxal sulcus absent or obscured by coarse rugae in ventral half (Figs |
|
11 | T2 smooth or with faintly impressed striation posterior to antecostal sulcus (Figs |
|
– | T2 with pronounced striae posterior to antecostal sulcus (Figs |
|
12 | Metapostnotum invaginated near propodeal spiracle, not separating propodeum from metanotum near metascutellum (Figs |
|
– | Metapostnotum invaginated near metascutellum, separating propodeum from metanotum near metascutellum (as in Fig. |
|
13 | Mesoscutellum with distinct coriaceous microsculpture and setal bases usually pustulate (Fig. |
|
– | Mesoscutellum entirely smooth and setal bases not strongly raised (Fig. |
|
14 | Gena in lateral view bulging (Fig. |
|
– | Gena in lateral view narrow, often with genal carina extending dorsally from base of mandible (Fig. |
|
15 | Mesoscutellum covered with shallowly impressed coriaceous microsculpture (Fig. |
|
– | Mesoscutellum entirely smooth, without microsculpture (Fig. |
|
16 | Frons outside of antennal scrobes with raised, irregular rugulae (Fig. |
|
– | Frons outside of antennal scrobes coriaceous, without raised rugulae but with more or less well-defined setigerous punctures, (Fig. |
|
17 | Gena in lateral view bulging (Fig. |
|
– | Gena in lateral view narrow (Figs |
|
18 | Mesopleural carina absent ventrally (Fig. |
|
– | Mesopleural carina complete (Fig. |
|
Lucid Key Server edition (only web browser required):
Applet edition (requires installation of Java Runtime Environment):
Within the New World species of the
[
Emerged from egg of
Lectotype, female,
The last known examination of the lectotype of
[
emerged from
Neotype, female,
[
Emerged from egg of
Holotype, female,
[
Emerged from egg of
Lectotype, female:
[
Emerged from egg of
Holotype, female:
[
collected on
The separation of
[
Collected on
Lectotype, female,
[
Emerged from
As previous authors have stated (
[
Emerged from egg of
Holotype, female,
This species may be distinguished from other species in the
[
Collected on
Holotype, female:
[
Collected on
Holotype, female:
[
Emerged from egg of
[
Holotype, female:
[
Emerged from egg of
[
Emerged from egg of / host egg of
Holotype, female:
[
Emerged from egg of
Lectotype, female:
[
Emerged from
Lectotype, female,
Female body length: 0.97–1.11 mm (n=6). Color of radicle: yellow; brown; pale brown. Number of mandibular teeth: three. Number of clypeal setae: 6. Facial striae: absent. Shape of gena in lateral view: receding posteriorly. Genal carina: extending dorsally from base of mandible. Macrosculpture of frons outside of antennal scrobe: absent; irregularly rugose. Orbital furrow: narrow to absent near malar sulcus. Hyperoccipital carina: absent. Preocellar pit: present.
Epomial carina: present. Netrion sulcus: complete. Mesoscutal suprahumeral sulcus: indicated by cells. Mesoscutal humeral sulcus: indicated by cells. Pattern of mesoscutal microsculpture: antero-posteriorly uniform. Macrosculpture of mesoscutum: coriacious. Area bounded by axillar crescent: smooth. Parapsidal line: absent. Notaulus: present. Median mesoscutal carina: absent; present. Median mesoscutal sulcus: absent.
Sculpture of mesoscutellum: smooth. Postacetabular sulcus: comprised of cells. Shape of episternal foveae: round; antero-posteriorly elongate. Number of episternal foveae: 3–5. Course of episternal foveae ventrally: abutting cells of postacetabular sulcus. Course of episternal foveae dorsally: extending dorsally to mesopleural pit. Sculpture of anterior mesepisternum: smooth or with shallowly impressed microsculpture. Mesopleural epicoxal sulcus: comprised of cells. Mesopleural carina: complete. Speculum: transversely striate. Paracoxal sulcus in ventral half of metapleuron: absent or indistinguishable from sculpture. Anteroventral extension of metapleuron: short, not reaching mesocoxa. Line of pits along metapleural carina: present. Setation of metapleuron: present. Metapostnotum: invaginated near edge of metascutellum and separating metanoum from propodeum. Color of legs beyond coxae: yellow; femora brown, otherwise variably yellow to brown. Metasomal depression: punctate or crenulate dorsally.
Sublateral setae on T1: absent. Setation of laterotergite 1: absent. Sculpture of T2 posterior to antecostal sulcus: smooth or with very faintly impressed striation; distinctly striate posterior to basal costae.
The epithet “valkyria” is Old Norse for “chooser of the slain” and refers to the female figures in Norse mythology that selected which soldiers would die in battle. The name is to be treated as a noun in apposition.
[
Holotype, female:
Female body length: 1.28–1.41 mm (n=20). Male body length: 1.18 mm (n=1). Color of radicle: brown. Number of mandibular teeth: three. Number of clypeal setae: 6. Facial striae: present as 3 or more rugulae extending onto lateral frons. Shape of gena in lateral view: receding posteriorly. Genal carina: extending dorsally from base of mandible. Macrosculpture of frons outside of antennal scrobe: irregularly rugose. Orbital furrow: narrow to absent near malar sulcus. Hyperoccipital carina: absent. Preocellar pit: present.
Epomial carina: present. Netrion sulcus: complete. Mesoscutal suprahumeral sulcus: indicated by cells. Mesoscutal humeral sulcus: indicated by cells. Pattern of mesoscutal microsculpture: antero-posteriorly uniform. Macrosculpture of mesoscutum: reticulate anteriorly, longitudinally rugulose posteriorly. Area bounded by axillar crescent: smooth. Parapsidal line: absent. Notaulus: absent. Median mesoscutal carina: absent. Median mesoscutal sulcus: absent. Sculpture of mesoscutellum: coriaceous. Postacetabular sulcus: comprised of cells. Shape of episternal foveae: irregular; round. Number of episternal foveae: 1–2. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: distinctly separated from mesopleural pit. Sculpture of anterior mesepisternum: faintly rugulose; finely reticulate. Mesopleural epicoxal sulcus: present as a smooth furrow; comprised of cells. Mesopleural carina: complete; well defined in anterior half, posterior half poorly defined to absent. Speculum: transversely striate. Paracoxal sulcus in ventral half of metapleuron: indicated by a line of distinct foveae. Anteroventral extension of metapleuron: long, extending to mesocoxa. Line of pits along metapleural carina: present. Setation of metapleuron: absent. Metapostnotum: invaginated near edge of metascutellum and separating metanoum from propodeum. Color of legs beyond coxae: femora and tibiae brown, otherwise variably yellow to brown. Metasomal depression: punctate or crenulate dorsally.
Sublateral setae on T1: absent; present. Setation of laterotergite 1: absent. Sculpture of T2 posterior to antecostal sulcus: smooth or with very faintly impressed striation.
[
Emerged from
Holotype, female:
We are grateful to: Luciana Musetti (OSUC), Sara Hemly (OSUC), Manuela Vizek (NHMW), Hege Vårdal (NHRS), Lubomir Masner (CNCI), Andy Bennett (CNCI), Tim Haye (CABI), Kim Hoelmer (BIIRU) and Christine Dieckhoff (BIIRU) for loans and specimens deposited in USNM; David Notton (BMNH) for specimen loans and commentary on nomenclature, Joe Cora (OSUC) for critical database support and making taxonomic literature available; Istvan Miko for commentary on morphological characters; and to Ian Realo and Samantha Fitzsimmons-Schoenberger for photography and transcribing label data. This work was made possible by funding from the Systematic Entomology Lab, USDA-ARS, and the Beneficial Insect Introduction Research Laboratory. Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the USDA; USDA is an equal opportunity provider and employer.
doi:
doi:
URI table of HAO morphological terms
Microsoft Excel Spreadsheet (.xls)
This table lists the morphological terms used in this publication and their associated concepts in the