Review of parasitoid wasps and flies (Hymenoptera, Diptera) associated with Limacodidae (Lepidoptera) in North America, with a key to genera.

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Review of Parasitoid Wasps and Flies (Hymenoptera, Diptera)
Associated with Limacodidae (Lepidoptera) in North America,
with a Key to Genera
Author(s) :Michael W. Gates, John T. Lill, Robert R. Kula, James E. O'Hara,
David B. Wahl, David R. Smith, James B. Whitfield, Shannon M. Murphy and
Teresa M. Stoepler
Source: Proceedings of the Entomological Society of Washington, 114(1):24-110.
2012.
Published By: Entomological Society of Washington
DOI: http://dx.doi.org/10.4289/0013-8797.114.1.24
URL: http://www.bioone.org/doi/full/10.4289/0013-8797.114.1.24
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PROC. ENTOMOL. SOC. WASH.
114(1), 2012, pp. 24–110
REVIEW OF PARASITOID WASPS AND FLIES (HYMENOPTERA,
DIPTERA) ASSOCIATED WITH LIMACODIDAE (LEPIDOPTERA)
IN NORTH AMERICA, WITH A KEY TO GENERA
M
ICHAEL
W. G
ATES
,J
OHN
T. L
ILL
,R
OBERT
R. K
ULA
,J
AMES
E. O’H
ARA
,D
AVID
B. W
AHL
,
D
AVID
R. S
MITH
,J
AMES
B. W
HITFIELD
,S
HANNON
M. M
URPHY
,
AND
T
ERESA
M. S
TOEPLER
(MWG, RRK, DRS) Systematic Entomology Laboratory, USDA, ARS, PSI, c/o
National Museum of Natural History, Washington, DC 20013-7012, U.S.A. (e-mail:
MWG michael.gates@ars.usda.gov, RRK robert.kula@ars.usda.gov, DRS dave.
smith@ars.usda.gov); (JTL, TMS) The George Washington University, Department
of Biological Sciences, 2023 G Street, NW, Suite 340, Washington, DC 20052, U.S.A.
(e-mail: JTL lillj@gwu.edu, TMS teresa_stoepler@gmail.com); (JEO) Canadian
National Collection of Insects, Agriculture and Agri-Food Canada, 960 Carling
Avenue, Ottawa, Ontario, Canada K1A 0C6 (e-mail: james.ohara@agr.gc.ca);
(DBW) American Entomological Institute, 3005 SW 56
th
Ave., Gainesville, Florida
32608 U.S.A. (e-mail: aei@aei.cfcoxmail.com); (JBW) Department of Entomology,
University of Illinois, Urbana-Champaign, Illinois 61801, U.S.A. (e-mail: jwhitfie@
life.uiuc.edu); (SMM) Department of Biological Sciences, University of Denver,
F. W. Olin Hall, 2190 E. Iliff Ave., Denver, Colorado 80208, U.S.A. (e-mail: Shannon.
M.Murphy@du.edu)
Abstract.—Hymenopteran and dipteran parasitoids of slug moth caterpillars
(Lepidoptera: Limacodidae) from North America are reviewed, and an illustrated
key to 23 genera is presented. Limacodid surveys and rearing were conducted during
the summer months of 2004–2009 as part of research on the ecology and natural
history of Limacodidae in the mid-Atlantic region of the U.S.A. Parasitoid rearing
involved a combination of collecting naturally occurring larvae in the field (at least
14 host species) and placing out large numbers of “sentinel” larvae derived from
laboratory colonies of three host species. Species in the following families are
documented from limacodids in North America as primary or secondary parasitoids
(number of genera for each family in parentheses; number of genera included in key
but not reared through this research in brackets): Chalcididae ([1]; Hymenoptera:
Chalcidoidea), Eulophidae (3; Chalcidoidea), Pteromalidae ([1]; Chalcidoidea),
Trichogrammatidae (1; Chalcidoidea), Braconidae (3 [1]; Hymenoptera: Ichneumo-
noidea), Ichneumonidae (7 [3]; Ichneumonoidea), Ceraphronidae (1; Hymenoptera:
Ceraphronoidea), Trigonalidae (2; Hymenoptera: Trigonaloidea), Bombyliidae ([1];
Diptera: Asilioidea), and Tachinidae (3; Oestroidea). We recovered 20 of 28 genera
known to attack limacodids in North America. Records discerned through rearing in
the mid-Atlantic region are augmented with previously published host-parasitoid
relationships for Limacodidae in North America north of Mexico. New records are
reported for the following parasitoids (total new records in parentheses): Uramya li-
macodis (Walker) (1), U. pristis (Townsend) (5), Austrophorocera spp. (6), Ceraphron
sp. (1), Alveoplectrus lilli Gates (1), Playplectrus americana (Girault) (10), Pediobi u s

crassicornis (Thomson) (1), Trichogramma (1), Mesochorus discitergus (Say) (1),
Hyposoter fugitivus (Say) (1), and Isdromas lycaenae (Howard) (5). The male of
Platyplectrus americana (Hymenoptera: Eulophidae) is redescribed, and the fe-
male is described for the first time. Incidental and miscellaneous host-parasitoid
associations are discussed, and it is concluded that most of these records are likely
parasitoids of contaminants accidentally introduced during the limacodid rearing
process. Triraphis eupoeyiae (Ashmead), new combination, is transferred from
Rogas (Hymenoptera: Braconidae).
Key Words: hyperparasitoid, parasitic, slug moth caterpillar, Acharia,Acrolyta,
Alveoplectrus,Ascogaster,Austrophorocera,Baryceros,Casinaria,
Ceraphron,Compsilura,Conura,Cotesia,Euclea,Hyposoter,Isa,
Isdromas,Isochaetes,Lithacodes,Lysibia,Mesochorus,Natada,
Orthogonalys,Packardia,Par a s a,Pediobius,Phobetron,Platyplectrus,
Prolimacodes,Psychophagus,Systropus,Taeniogonalos,Tortricidea,
Trichogramma,Triraphis,Uramya
Slug moth caterpillars (Lepidoptera:
Limacodidae) are mostly polyphagous
external foliage feeders on a broad array
of deciduous trees and shrubs in eastern
North America (Wagner 2005, Lill 2008).
Their broad diets, rather distinctive larval
morphologies, and often bright coloration
make them interesting subjects for eco-
logical and behavioral studies (e.g., Lill
et al. 2006, Murphy et al. 2010). Unlike
many more mobile caterpillars, limaco-
did larvae tend to remain on an individual
host plant for the duration of their de-
velopment: typically about two months,
going through numerous instars (7–12)
depending on the species, with some
species requiring more than 100 days
to complete development (J. Lill and
S. Murphy pers. obs.). Larvae tend to
occur in low densities (<1 caterpillar/
m
2
foliage; Lill et al. 2006, Lill 2008)
and can be highly cryptic, especially in
early instars, but they produce a rather
distinctive pattern of feeding damage
that facilitates their capture. Outbreaks
of North American limacodids are re-
ported rarely, but many tropical species
are important agricultural pests, partic-
ularly in banana and palm plantations
in South America and Southeast Asia
(Ostmark 1974, Cock et al. 1987, Godfray
and Chan 1990). Because many species
of Limacodidae possess stinging spines
during all or a portion of their larval de-
velopment(Epstein1988),thelarvaecan
be a nuisance when they occur in planta-
tions (Ostmark 1974) or on ornamental
palms near tourist attractions (e.g., Conant
et al. 2002). Their protracted larval de-
velopment period exposes the caterpillars
to attack from a diverse assemblage of
larval and larval-pupal parasitoids. How-
ever, research to-date on host-parasitoid
interactions in Limacodidae has focused
almost entirely on biocontrol of pest li-
macodids in tropical agricultural systems
(e.g., Cock et al. 1978) with the notable
exception of the massive caterpillar rear-
ing database compiled for tropical forests
of Area de Conservacio
´n Guanacaste in
Costa Rica (Janzen and Hallwachs 2009).
*Accepted by John W. Brown
VOLUME 114, NUMBER 1
25
DOI: 10.4289/0013-8797.114.1.24

By contrast, the hymenopteran/dipteran
parasitoid community that attacks North
American limacodids has been described
only anecdotally, and concerted rearing
efforts are mostly lacking (but see Le
Corff and Marquis 1999 and Stireman
and Singer 2003a, b for examples where
limacodids were reared as part of larger
community sampling).
In general, dipteran (i.e., Tachinidae)
and hymenopteran parasitoids are essen-
tial in regulating native forest macrolep-
idoptera and are reasonably well studied
(LaSalle 1993, McCullough et al. 1999).
Conversely, there is little information
regarding bombyliid parasitoids (i.e., in
Systropus) and their roles in regulating
native forest macrolepidoptera, likely
due to their relative rarity compared to
parasitic hymenopterans and tachinids.
Most hymenopteran parasitoids of mac-
rolepidoptera belong to the superfamilies
Ichneumonoidea and Chalcidoidea, and
species of Tachinidae are the most com-
mon dipteran parasitoids of microlepi-
doptera (Askew 1971, Quicke 1997). Two
recent studies in North America have
focused on Tachinidae and Hymenoptera
specifically and their host relationships in
eastern North American forests (Strazanac
et al. 2001, Petrice et al. 2004). Most
research on parasitoids of native forest
macrolepidoptera has focused on out-
break species such as fall webworm
(Hyphantria cunea Drury; Arctiidae),
eastern tent caterpillar (Malacosoma
americanum (F.); Lasiocampidae) (Kulman
1965, Morris 1976, Witter and Kulman
1979), and gypsy moth (Lymantria
dispar (L.); Lymantriidae) (Barbosa et al.
1975, Elkinton and Liebhold 1990 and
references therein). Few studies have
documented parasitoids in less abundant/
non-outbreak forest macrolepidoptera (in-
cluding limacodids) (Schaffner and Griswold
1934, northeasternU.S.A.; Raizenne 1952,
Ontario, Canada), and fewer still have
focused on Limacodidae (as mentioned
above).
Tachinidae and parasitoid Hymenoptera
are diverse in forest canopy communi-
ties in North America with new host-
parasitoid associations being recorded with
some frequency (Butler 1993, Strazanac
et al. 2001, Petrice et al. 2004). Bio-
logically, tachinids are recorded from
15 arthropod orders (Wood 1987), whereas
parasitoid Hymenoptera have been docu-
mented from 16 (LaSalle 1993). As a
group, parasitoid Diptera have been
viewed as secondary to hymenopterans
as effective parasitoids (Askew 1971,
Quicke 1997); however, both groups
are abundant in North American forests
where limacodids occur most commonly.
North American hymenopteran and dip-
teran parasitoid-host associations have
been cataloged by Krombein et al. (1979)
and Arnaud (1978), respectively. A world
catalog to Tachinidae is in preparation
(JEO), and databases are available for
Ichneumonoidea (Yu et al. 2005) and
Chalcidoidea (Noyes 2003). Recent large-
scale rearings of forest macrolepidoptera
by Butler (1993) documented 115 new
hymenopteran host associations, and
Strazanac et al. (2001) reported 60 new
tachinid host associations.
Many of the tachinid and hymenop-
teran parasitoids reported herein are
known to attack lepidopterans (or their
primary parasitoids) more generally, but
several specialize on Limacodidae, and
others are facultative (see Pediobius;
Peck 1985) or obligatory hyperparasitoids
(see Conura Spinola; Delvare 1992).
Given their specialization on Limacodidae,
many of these taxa, particularly certain
Chalcidoidea such as Platyplectrus Ferrie
`re
and Alveoplectrus Wijesekara and Schauff,
were encountered infrequently in North
America (Wijesekara and Schauff 1997).
This can be ascribed to the lack of major
limacodid pests in North America, the
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
26

difficulty of locating/handling their larvae,
and the lack of financial resources avail-
able to study non-outbreak pests. Most
of the parasitoids documented as a re-
sult of the rearing portion of this study
parasitize early limacodid instars.
The purpose of this study is to review
the parasitoids associated with limacodids
in America north of Mexico, including
introduced limacodids (i.e., Monema
flavescens Walker) and exotic parasitoids
recorded from limacodids elsewhere that
could potentially attack limacodids in
North America. We assemble these vari-
ous records herein and add host-parasitoid
associations generated from six years of
intensive rearing of larval limacodids in
the greater Washington, DC metropolitan
area (Table 1).
M
ATERIALS AND
M
ETHODS
Collection and Rearing
Beginning in mid- to late-June of each
summer of 2004–2009, we searched
manually for limacodid larvae on the
undersides of leaves of common woody
trees at the following field sites near
Washington, DC: Little Bennett Regional
Park, Plummers Island (Montgomery
County, MD), Patuxent National Wildlife
Refuge (Prince George’s County, MD),
Rock Creek Park (Washington, DC), and
the U.S. National Arboretum (Washington,
DC). More than a dozen host plants have
yielded limacodids, but most of our efforts
were focused on searching six common
host plants used by most species of
Limacodidae: American beech (Fagus
grandifolia Ehrh.), white oak (Quercus
alba L.), northern red oak (Quercus rubra
L.), black gum (Nyssa sylvatica Marsh.),
black cherry (Prunus serotina Ehrh.), and
pignut hickory (Carya glabra Miller).
Additional host plant species were sam-
pled less intensively (Table 1). For each
wild-caught larva, we recorded the species,
collection date, and host plant. In addition
to these wild-caught larvae, we also con-
ducted a series of field experiments as part
of a different project examining tri-trophic
interactions in Limacodidae that involved
placing out “sentinel” larvae on each of the
six common host plants described above
at the Little Bennett Regional Park site.
These larvae came from laboratory colo-
nies established through a combination of
larval and adult (ex ovo) collections; thus,
larvae were unparasitized when placed in
the field. These experimental larvae were
left exposed to parasitoid attack in the field
for one to several weeks depending on the
experiment and then brought back to the
lab for rearing.
Collected larvae were reared individu-
ally in 16 oz. clear plastic deli containers
containing a disk of moistened filter paper
to prevent host foliage from drying out.
Fresh leaves from the various host plants
were replaced as needed (typically twice
per week). Larvae showing signs of para-
sitism were checked routinely for parasit-
oid larvae/pupae, and emerging adults were
either placed in 95% ethanol (Hymenop-
tera) or frozen (Diptera) for later mounting/
pinning and identification. One genus of
tachinid flies (Austrophorocera spp.) exclu-
sively contains larval-pupal parasitoids, so
adults eclosed from overwintering cocoons
the following year (pupae were kept in an
environmental chamber in moist peat moss
during the winter months and then exposed
to spring conditions to induce fly pupation
and emergence). In addition, several of
the hymenopteran parasitoids reared from
limacodid hosts late in the season dia-
paused as pupae and emerged the follow-
ing summer.
Hymenopteran Parasitoid
Preparation and Imaging
Specimens in ethanol were dehydrated
through increasing concentrations of
VOLUME 114, NUMBER 1
27

Table 1. Parasitoids reared from host plant/limacodid host pairs in greater Washington, D. C., 2004–2009.
Limacodid Species Plant Species Parasitoid Species Notes
Acharia stimulea Acer negundo Uramya pristis
A. saccharinum U. pristis
P. americana*
Cotesia empretiae
Asimina triloba U. pristis
Pl. americana*
P.s crassicornis Hyperparasitoid of Pl. americana
Carya glabra U. pristis
A. cocciphila*
Pl. americana*
Pe. crassicornis Hyperparasitoid of Pl. americana
Fagus grandifolia A. cocciphila*
U. pristis
Pl. americana*
Pe. crassicornis Hyperparasitoid of Pl. americana
T. discoideus
Lindera benzoin Pe. crassicornis
Ly. mandibularis Pseudohyperparasitoid through
microgastrine.
Nyssa sylvatica U. pristis
Austrophorocera sp.*
Pe. crassicornis Hyperparasitoid of Pl. americana
Pl. americana*
Prunus serotina U. pristis
A. cocciphila*
Pl. americana*
Pe. crassicornis Hyperparasitoid of Pl. americana
Co. empretiae
Quercus alba U. pristis
A. cocciphila*
Pe. crassicornis Hyperparasitoid of Pl. americana
Pl. americana*
Co. empretiae
Quercus rubra Austrophorocera sp. Only egg observed
Ceraphron sp.* Hyperparasitoid, likely via
Pl. americana or Al. lilli
Pl. americana*
Acharia stimulea Quercus rubra T. discoideus
Pe. crassicornis Hyperparasitoid of Pl. americana
Acharia stimulea Quercus rubra Is. lycaenae*
Robinia pseudoacacia Pl. americana*
Co. empretiae
Adoneta spinuloides Carya glabra Pl. americana*
Fagus grandifolia U. pristis*
Nyssa sylvatica Pl. americana
Pe. crassicornis* Hyperparasitoid of Pl. americana
Quercus rubra U. pristis*
Al. lilli*
Pl. americana*
T. discoideus
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
28

Table 1. Continued.
Limacodid Species Plant Species Parasitoid Species Notes
Euclea delphinii Carpinus caroliniana Co. empretiae
Carya glabra U. pristis
A.einaris*
Pl. americana*
T. discoideus
Fagus grandifolia U. pristis
A.einaris*
Pl. americana*
T. discoideus
Nyssa sylvatica A.einaris*
Com. concinnata
T. discoideus
Taeniogonalos gundlachii Hyperparasitoid of Com. concinnata
Pl. americana*
Prunus serotina A.einaris*
U. pristis
Com. concinnata*
Pl. americana*
Quercus alba A.einaris*
Pl. americana*
Pe. crassicornis*
Trichogramma sp.* Egg parasitoid
T. discoideus
Quercus rubra A.einaris*
Euclea delphinii Quercus rubra Com. concinnata*
U. pristis
Al. lilli
Pl. americana*
Euclea delphinii Quercus rubra Pe. crassicornis* Hyperparasitoid of Pl. americana
T. discoideus
Diospyros virginiana Co. empretiae
Isa textula Acer saccharum U. pristis
Carya glabra U. pristis
Fagus grandifolia U. pristis
A. lilli
T. discoideus
Quercus alba U. pristis
Austrophorocera sp.*
Pl. americana*
Pe. crassicornis* Hyperparasitoid of Pl. americana
Is. lycaenae
T. discoideus
Quercus prinus U. pristis
Orthogonalys pulchella Hyperparasitoid of U.pristis
Quercus rubra U. pristis
Ceraphron sp. Hyperparasitoid of Pl.
americana*or Al. lilli*
Al. lilli
Pl. americana*
T. discoideus
Is. lycaenae* Likely hyperparasitoid of Triraphis
O. pulchella Hyperparasitoid of U.pristis
VOLUME 114, NUMBER 1
29

Table 1. Continued.
Limacodid Species Plant Species Parasitoid Species Notes
Isa/Natada Nyssa sylvatica Al. lilli
Isochaetes
beutenmuelleri
Fagus grandifolia U. pristis*
O. pulchella Hyperparasitoid of U.pristis
Quercus rubra U. pristis*
Lithacodes fasciola Acer negundo U. pristis*
Carya glabra U. pristis*
Pl. americana*
Fagus grandifolia Austrophorocera n. sp.*
Pl. americana*
Lithacodes fasciola Nyssa sylvatica Pl. americana*
Prunus serotina Austrophorocera n. sp.*
Pe. crassicornis* Hyperparasitoid of Pl. americana
T. discoideus
Quercus alba Austrophorocera sp.* Only egg observed
Pl. americana*
Quercus rubra Pl. americana*
T. discoideus
Lithacodes fasciola Quercus velutina Al. lilli
Lithacodes/Packardia Nyssa sylvatica Pl. americana
Natada nasoni Carya glabra Triraphis discoideus
Fagus grandifolia T. discoideus
Is. lycaenae*
Nyssa sylvatica T. discoideus
Prunus serotina Is. lycaenae*
Quercus alba T. discoideus
Quercus prinus Pe. crassicornis* Possible primary parasitoid
Quercus rubra Pl. americana*
T. discoideus
Is. lycaenae
Packardia geminata Fagus grandifolia U. pristis*
Pl. americana
Parasa chloris Quercus rubra Austrophorocera sp.* Only egg observed
T. discoideus
I. lycaenae*
Mesochorus discitergus Hyperparasitoid of Triraphis;
unconfirmed
Phobetron pithecium Fagus grandifolia Pl. americana*
Prolimacodes badia Carya glabra H. fugitivus*
Diospyros virginiana A.imitator*
Fagus grandifolia A.imitator*
Pl. americana*
Is. lycaenae*
Me. discitergus Hyperparasitoid of Triraphis
(unconfirmed)
Nyssa sylvatica Austrophorocera sp.* Only egg observed
Prunus serotina A.imitator*
T. discoideus
Quercus alba Pl. americana*
H. fugitivus*
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
30

ethanol to hexamethyldisilazane (HMDS)
(Heraty and Hawks 1998) before point- or
card-mounting. Images of specimens were
produced by scanning electron micros-
copy (SEM) and an EntoVision Imaging
Suite. Card- and point-mounted specimens
were examined using stereomicroscopes
with 10X or 25X oculars and fiber optic
light sources. Mylar film was used to dif-
fuse glare from fiber optic light sources to
reduce glare from the specimens. Scan-
ning electron microscope (SEM) images
weretakenwithanAmray1810(LaB
6
source).Specimenswereaffixedto12.7X
3.2 mm Leica/Cambridge aluminum SEM
stubs with carbon adhesive tabs (Electron
Microscopy Sciences, #77825-12). Stub-
mounted specimens were sputter coated
using a Cressington Scientific 108 Auto
with gold from at least three differ-
ent angles to ensure complete coverage
(;20–30 nm coating). Wings were re-
moved and slide-mounted in polyvinyl
alcohol prior to imaging. Wing and
habitus images were captured using an
EntoVision Imaging Suite, which includes
a firewire JVC KY-75 3CCD digital cam-
era mounted on a Leica M16 zoom lens
via a Leica z-step microscope stand. Ad-
ditionally, a GT-Vision Lw11057C-SCI
digital camera attached to a Leica DMRB
compound scope was used to feed image
data to a desktop computer. The program
Cartograph 5.6.0 (Microvision Instru-
ments, France) was used to merge an
image series (typically representing 15–
30 focal planes) into a single in-focus,
composite image. Lighting was achieved
using techniques summarized in Buffington
et al. (2005), Kerr et al. (2008), and
Buffington and Gates (2008).
Several images (see below) were ob-
tained with a Visionary Digital imaging
station. The station consists of an Infin-
ity Optics K2 long distance microscope
affixed to a Canon EOS 40D digital SLR
camera. Lighting was provided by a Dyna-
lite M2000er power pack and Microptics
ML1000 light box. Image capture soft-
ware is Visionary Digital proprietary ap-
plication with images saved as TIF with
the RAW conversion occurring in Adobe
Photoshop Lightroom 1.4. Image stacks
were montaged with Helicon Focus
4.2.1 for images of Conura nortonii
(Cresson), C. immaculata (Cresson),
Orthogonalys pulchella (Cresson), and
Taeniogonalos gundlachii (Cresson),
as well as habitus shots of Psychophagus
omnivorus (Walker), Platyplectrus ameri-
cana (Girault), and Pediobius crassicornis
(Thomson).
Final image plates for hymenopteran
and dipteran figures were prepared using
Table 1. Continued.
Limacodid Species Plant Species Parasitoid Species Notes
Quercus rubra A.imitator*
Pl. americana*
T. discoideus
Tortricidia sp. Quercus alba Pl. americana*
Fagus grandifolia T. discoideus
Prunus serotina U. pristis*
Quercus rubra U. pristis*
Unknown limacodid Carya glabra Pl. americana
(too small to ID) Pe. crassicornis
Unknown limacodid Fagus grandifolia Pl. americana
Unknown limacodid Quercus rubra T. discoideus
VOLUME 114, NUMBER 1
31

InDesign CS4. Figures 14–17, previously
unpublished, were used with the permis-
sion of D. M. Wood (Agriculture and
Agri-Food Canada, Ottawa). Figures 38
and 43–44 were reproduced from Townes
(1970) with permission.
Diptera Imaging
The image of Systropus macer Loew
was captured with a Nikon Coolpix 8800
and adapters through the ocular of a
Leica MZ9.5 stereoscope. Tachinid images
were taken with a Canon EOS 40D digital
SLR camera mounted on a Kaiser RS1
copy stand. A Canon EF 100 mm f/2.8
macro lens was used for full body images,
and a Canon MP-E 65 mm 1–5X macro
lens was used for images of body struc-
tures. A ring light consisting of 80 LEDs
and covered with a reflective dome
provided the lighting. Image stacks were
montaged using Syncroscopy’s Auto-
Montage, and the resultant images received
further treatment in Adobe Photoshop
CS4.
Terminology
Hymenopteran terminology for sur-
face sculpture follows Harris (1979) and
for morphology follows Wahl (1993a),
Gibson (1997), Sharkey and Wharton
(1997), and Deans et al. (2010). Several
measurements for chalcidoids were taken,
including the following: body length, in
lateral view from the anterior projection
of the face to the tip of the metasoma;
head width through an imaginary line
connecting the farthest lateral projection
of the eyes; head height through an
imaginary line from the vertex to the
clypeal margin bisecting both the median
ocellus and the distance between the tor-
uli; malar space, in lateral view between
the ventral margin of the eye and lateral
margin of the oral fossa; posterior ocellar
line (POL), the shortest distance between
the posterior ocelli; ocular ocellar line
(OOL), the shortest distance between the
lateral margin of the posterior ocellus
and the eye orbit; marginal vein, the
length coincident with the leading fore
wing edge to the base of the stigmal
vein; stigmal vein, the length between its
base on the marginal vein and its apex;
postmarginal vein, the length from the
base of the stigmal vein to its apex on
the leading fore wing edge. Mesosomal
and metasomal sclerites were measured
dorsally along the midline. The use of
“[ ]” in descriptions denotes structures
that are not visible in the specimens upon
which the description is based (observed
via SEM), whereas their use in the ma-
terial examined section refer to author
notes. For braconids metasomal terga
1, 2, 3, etc. are abbreviated as T1, T2, T3;
antennal flagellomeres are abbreviated
as F1, F2, F3, etc. The ovipositor of
ichneumonids was measured from the
structure’s base (observed or inferred)
to its apex. The ovipositor sheaths must
sometimes be separated by a fine needle
to expose the ovipositor valves. The junc-
ture of the occipital and hypostomal carina
above the mandibular base is measured
in posterolateral view; it is sometimes
necessary to remove the head to properly
measure. Dipteran terminology follows
McAlpine (1981).
The portion of the key pertaining to
Chalcidoidea is based upon the keys in
Gibson et al. (1997). That for Braconidae is
based on the keys of Sharkey (1997) and
Whitfield (1997). That for Ichneumonidae
is based on the key of Wahl (1993b). The
diagnosis for Cotesia Cameron is based on
the key in Whitfield (1997); the diagnosis
for Ascogaster Wesmael is based on the
key in Shaw (1997a). The diagnosis for
Triraphis Ruthe is based on the diagnoses for
Triraphis and Rogas Nees in van Achterberg
(1991), as well as the keys in van Achterberg
(1991) and Shaw (1997b).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
32

All binominals for hosts and parasitoids
are reported in their current nomenclatural
combinations. Although this is not
necessarily straightforward in any single
publication, we cite the most recent/
comprehensive nomenclatural authorities
where required. For Limacodidae, nomen-
clatural references include: Fletcher and
Nye (1982), Davis (1983), and Becker and
Epstein (1995). Host records from the lit-
erature for Tachinidae list the current
combination, as well as the combinations
under which the hosts and tachinids were
originally cited. New host records result-
ingfromtherearingeffortsofJTL,SMM
and TMS for a particular parasitoid are
denoted with *inthetextandTable1.
Those newly reported herein from other
sources are denoted with
‡
.Wereportnew
host records at the parasitoid species level
only rather than higher taxonomic levels.
Thus, if a parasitoid species is newly re-
corded for a particular host genus or spe-
cies, we report it as new even if congeneric
parasitoids were previously recorded from
that host genus or species.
Abbreviations for collections are as
follows: AEI (American Entomological
Institute, Gainesville, Florida, U.S.A.),
ANSP (Academy of Natural Sciences,
Philadelphia, Pennsylvania, U.S.A.), BMNH
(The Natural History Museum, London,
United Kingdom), CNC (Canadian
National Collection of Insects, Ottawa,
Ontario, Canada), IRSNB (Institut Royal
des Sciences Naturelles de Belgique,
Brussels, Belgium), MZLU (Zoological
Museum, Lund University, Lund, Sweden),
MCZ (Museum of Comparative Zoology,
Harvard University, Cambridge, Massa-
chusetts, U.S.A.), MSUC (Michigan State
University, East Lansing, Michigan,
U.S.A.), NHMW (Naturhistorisches
Museum Wien, Vienna, Austria), SEMK
(Snow Entomological Museum, Univer-
sity of Kansas, Lawrence, Kansas, U.S.
A.), ULQC (Universite
´Laval, Quebec,
Canada), USNM (National Museum of
Natural History, Smithsonian Institution,
Washington, DC, U.S.A.), and ZMUC
(Zoologisk Museum, Copenhagen,
Denmark).
R
ESULTS AND
D
ISCUSSION
Key to Primary and Secondary
Parasitoids Known From Limacodids
in North America
1. Hind wing modified as haltere (Fig. 1),
hamuli absent, mouthparts sponging (Fig. 1)
........................2(Diptera)
1’ Hind wing not modified as haltere, hamuli
present (Figs. 92–93), mouthparts mandib-
ulate (e.g., Figs. 39, 82) . .............
................... 5(Hymenoptera)
2. (1) Abdomen narrow and elongate, >2.0X
as long as broad and swollen apically
(Fig. 3). Wings shorter than abdomen.
Antennae and proboscis much longer than
head ..........................
..Systropus (macer Loew; Bombyliidae)
2’ Abdomen stout, less than 2.0X as long as
broad (Figs. 1, 11). Wings longer than ab-
domen. Antennae and proboscis at most as
long as head . . .........3(Tachinidae)
3. (2’) Facial ridge bare except for a few small
and decumbent setae on lower third or less
(Fig. 2). Metathoracic spiracle fringed with
plumose hairs of about equal length along
both anterior and posterior edges, leaving a
V-shaped middorsal opening (Fig. 4). Pros-
ternumbare(Fig.6)...................
Uramya (pristis (Walker), limacodis (Townsend))
3’ Facial ridge with row of stout setae on lower
one-half or more (Fig. 9). Metathoracic
spiracle with posterior lappet much larger
than anterior one (Fig. 5). Prosternum haired
laterally(Fig.7)....................4
4. (3’) Ocellar seta vestigial or absent (Fig. 9).
Abdominal terga 3 and 4 each with 1 pair of
median discal setae. Bend of vein M obtuse
(Fig. 8). Female with sickle-shaped, pierc-
ing ovipositor (Fig. 8) . . .............
......Compsilura (concinnata (Meigen))
4’ Ocellar seta well developed, similar in size
to outer vertical seta (Fig. 10). Abdominal
terga 3 and 4 without median discal setae
(Fig. 11). Bend of vein Malmost a right angle
(Fig. 11). Female with short, non-piercing
ovipositor(Fig.11)..................
..Austrophorocera (cocciphila (Aldrich and
VOLUME 114, NUMBER 1
33

Webber), coccyx (Aldrich and Webber), A.
einaris (Smith), A. imitator (Aldrich and
Webber), A. n. sp.)
5 (1’) Fore wing venation complete, with at
least 2 closed cells (Figs. 18, 28, 34) . . . . .
. . . . . . 6 (Trigonaloidea, Ichneumonoidea)
5’ Fore wing venation reduced, with fewer
than 2 closed cells (Figs. 77–78, 96, 113)
.....17(Chalcidoidea,Ceraphronoidea)
6. (5) Fore wing with veins C and SC+R
touching/fused, costal cell absent (Figs.
63, 65). Metasomal sterna less strongly
sclerotized than terga (Figs. 19, 23, 27,
29) .............7(Ichneumonoidea)
6’ Fore wing with veins C and SC+R separate,
costal cell present (Fig. 18). Metasomal sterna
and terga equally sclerotized (Figs. 70–73) . .
...................16(Trigonaloidea)
7. (6) Fore wing with vein 2m-cu present
(Fig. 28, 35) ........8(Ichneumonidae)
7’ Fore wing with vein 2m-cu absent (Fig.
56) ............... 14(Braconidae)
8. (7) Areolet of fore wing closed, large and
rhombic (Figs. 20, 26). Ovipositor delicate,
needlelike, sheaths thick and rigid (Figs. 19,
22). Female hypopygium prominent and tri-
angular in lateral view (Fig. 22). Male gon-
oforceps produced into elongate process.
Spiracle of metasomal segment 1 near or just
behind middle, glymma large and deep
(Fig. 21) . . . ......................
.........Mesochorus (discitergus (Say))
8’ Areolet of fore wing open (vein 3rs-m ab-
sent) or closed, if closed then cell obliquely
quadrate and petiolate (Figs. 26, 34).
Ovipositor always stouter than above, sheaths
thin, often curved if ovipositor ;2.0X as
long as metasomal apical depth. Female
hypopygium small and quadrate in lateral
view (Fig. 23). Male gonoforceps not
produced. Spiracle of metasomal segment
1 beyond middle, glymma small or absent
(Fig.27) .......................9
9. (8’) Ventral posterior corner of pro-
pleuron with strongly produced, more or
less angulate lobe touching or overlapping
pronotum (Fig. 24). Sternaulus about 0.3X
as long as mesopleuron (Fig. 30). Clypeus
not separated from supraclypeal area by
distinct groove (Fig. 32). Areolet of fore
wing closed, obliquely quadrate and peti-
olate (Fig. 26) . . . ................10
9’ Ventral posterior corner of propleuron not
developed as distinct lobe, not angulate, at
most with weak groove delimiting it from
main area of propleuron (Fig. 25). Sternaulus
of mesopleuron present and reaching middle
coxa (Fig. 31). Clypeus separated from su-
praclypeal area by groove (Fig. 33). Areolet
of fore wing open and pentagonal (Fig. 34).
...............................11
10. (9) Petiole of fir st met asomal segment
long and cylindrical in cross-section;
midpoint of petiole with tergo-sternal
suture at midline; T1 without trace of glymma
(Fig. 28) . . . .Casinaria (grandis Walley)
10’ Petiole of first metasomal segment shorter
and quadrate in cross-section; midpoint of
petiole with tergo-sternal suture close to
ventral margin; T1 with glymma present as
pitlike impression (Fig. 27) . . . .......
...........Hyposoter (fugitivus (Say))
11. (9’) Body color (excluding legs) black with
white markings (Fig. 29). Mesosoma with
coarse punctures (Fig. 37). Dorsal margin
of pronotum with strong swelling at dorsal
end of epomia (Fig. 37). Fore wing 5.0–8.5
mm long . . ......................
........Baryceros (texanus (Ashmead))
11’ Body color (excluding legs) ranging from
uniformly black/dark brown to having
brownish red areas on T2–T3 (Figs. 34–36).
Mesosoma with punctures ranging from
fine to absent. Dorsal margin of pronotum
without strong swelling at dorsal end of
epomia.Forewing1.9–3.8mmlong.....
..............................12
12. (11’) Apical 0.3 of clypeus turned inward
at 90°(Fig. 43). Vein 2-Cu of hind wing
basally incomplete . . ...............
....Lysibia (mandibularis (Provancher))
12’ Apical 0.3 of clypeus flat, not turned in-
ward (Fig. 44). Vein 2-Cu of hind wing
complete . . ....................13
13. (12’) Occipital and hypostomal carinae
meeting at mandibular base (Fig. 41) . . .
Acrolyta (nigricapitata (Cook and Davis))
13’ Occipital carina meeting hypostomal ca-
rina above mandibular base, juncture sep-
arated from base by about 0.2X basal
mandibular width (Fig. 42) ...........
........ Isdromas (lycaenae (Howard))
14. (7’) Labrum visible through gap between
ventral margin of clypeus and mandibles
and concave (as in Fig. 39) (cyclostome
Braconidae). Epicnemial carina present
(Fig. 47); median carina present on T1 and
usually extending posteriorly from dorsal
carinae; T2 striate to striate-rugose (Figs.
52, 57); fore wing vein 1m-cu basad or in
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
34

line with fore wing vein 2RS (Fig. 56) . .
.......... Trira ph is Ruthe (eupoeyiae
(Ashmead), discoideus (Cresson), harri-
sinae (Ashmead))
14’ Labrum concealed by clypeus or if vis-
ible not concave (Figs. 40 (as in), 60)
(non- cyclostome Braconidae). Without
combination of characters found in
Triraphis .......................
........Cheloninae, Microgastrinae, 15
15. (14’) Occipital carina present, fore wing
vein RS reaching wing margin as tubular
vein, T1–T3 fused into carapace cover-
ing all other terga (Figs. 48, 63). Other
characters not in combination found in
Cotesia (seebelow) ..............
..Ascogaster (quadridentata Wesm a el )
15’ Occipital carina absent, fore wing vein RS
not reaching wing margin as tubular vein
(Fig. 65), T1–T3 variable (Microgastrinae).
Fore wing vein r-m absent, areolet ab-
sent (Fig. 65). Ovipositor and sheaths
short, weakly extending beyond tip of
hypopygium (Fig. 67). Hypopygium evenly
sclerotized medially. Propodeum with-
out areola, often with distinct medial
longitudinal carina and usually rugose
(Fig.66)........................
.. Cotesia (empretiae (Viereck), phobetri
(Rohwer), schaffneri (Rohwer))
16. (6’) Antenna black with white or light
yellow band at center. Head and mesosoma
with black and white pattern. Metasoma
and legs mostly orange (Figs. 70–71).
Metasoma thin, smooth, impunctate . . .
.....Orthogonalys (pulchella Cresson)
16’ Antenna yellowish brown without lighter
band at center. Head and body black with
yellow markings (Figs. 72–73). Meta-
soma black and yellow, legs yellow with
dark brown to black on femur, tibia
dusky in apical third. Metasoma stout,
punctate........................
..Taeniogonalos (gundlachii (Cresson))
17. (5’) Fore wing with tubular submarginal
vein basally on anterior margin, stigmal
vein curving distally (Fig. 77). Female an-
tenna with 7–8 flagellomeres (Fig. 76), male
with 8–9 (Fig. 74). Dorsum flat in lateral
view ............................
......Ceraphron Jurine (Ceraphronidae)
17’ Fore wing lacking tubular vein basally on
anterior margin (Fig. 78). Other features
not as above . ..................18
18. (17’) Hind femur enlarged (<3.0X as long
as broad) (Figs. 79–80). Prepectus reduced
to a small sclerite along dorsal margin of
mesopleuron (Fig. 83). Gaster petiolate
(Figs. 79–80) . ....................
....Conura (camescens (Cameron), nig-
ricornis (F.), nortonii (Cresson))
18’ Hind femur not enlarged (>3.0X as long as
broad). Prepectus larger, triangular (Figs.
87, 109). Gaster indistinctly or not petiolate
...... Eulophidae,Trichogrammatidae,
Pteromalidae, 19
19. (18’) Legs with 5 tarsomeres. Clypeus
shallowly bilobed apically (Fig. 88).
Pedicel ;2.0X as long as broad, antennal
formula 11263 (Fig. 89). Propodeum re-
ticulate-rugose, with median carina in-
complete (Fig. 90). Gaster subcircular to
short-ovate(Fig.91).................
.....Psychophagus (omnivorus (Walker))
19’ Legs with 3 or 4 tarsomeres. Other features
not as above . ....................
. . . . Eulophidae, Trichogrammatidae, 20
20. (19’) Legs with 4 tarsomeres. Fore wing
lacking setal tracks (Fig. 113) .........
....................Eulophidae, 21
20’ Legs with 3 tarsomeres (Fig. 94). Fore
wing with setal tracks, broad with sig-
moid venation and distinctive Rs1 setal
track(Fig.96)...................
...... Trichogramma (minutum Riley)
21. (20’) Scutellum with one pair of setae (Fig.
95). Submarginal vein with two setae on
dorsal surface. Propodeum with paired
submedian carinae that diverge posteriorly
(Fig.95) ................. ........
......Ped i obi u s (crassicornis (Thomson))
21’ Scutellum with two pairs of setae (Figs.
104–105). Submarginal vein with three or
more setae on dorsal surface (Fig. 113).
Propodeum with single carina, or if with
paired submedian carinae then carinae not
arcuately divergent (Figs. 104–105) . . . . . .
..................... Eulophinae,22
22. (21’) Propodeum with two submedian
carinae (Fig. 104). Scutellum with scu-
tellar grooves broad, contiguous with
posterior margin (Fig. 104). Hind tibia
withonespur...................
..........Alveoplectrus (lilli Gates)
22’ Propodeum with single median carina
(Fig. 105). Scutellum with scutellar
grooves narrow (Fig. 105). Hind tibia with
twospurs.......................
....Platyplectrus (americana (Girault))
VOLUME 114, NUMBER 1
35

Diptera
Systropus Weidemann (Bombyliidae:
Toxophorinae: Systropodini)
(Fig. 3)
Diagnosis.—
Systropus can be identi-
fied in North America using the key by
Hall (1981) or Kits et al. (2008; Mel-
anderella Cockerell not extant) and can be
distinguished from other North American
genera (Dolichomyia Weidemann) of Sy-
stropodini by the following characters:
pedicel at least 2–3 times longer than wide
(aswideaslonginDolichomyia); first
flagellomere flattened (cylindrical in
Dolichomyia); and anal lobe of wing
not reduced (extremely reduced in Do-
lichomyia). However, Dolichomyia is
not known from eastern North America.
A third genus, Zaclava Hull, is repre-
sented by four species known only from
Australia. Typically, Systropus spp. mimic
sphecid wasps and thus are often elongate,
largely orange and black with the abdo-
men long and swollen apically (Fig. 3).
The wings are conspicuously shorter than
the abdomen, and both the antennae and
proboscis are much longer than the head
(Fig. 3).
Fauna.—
Systropus is a large genus
containing 154 species most commonly
encountered in tropical areas of all con-
tinents (Evenhuis and Greathead 1999).
The majority of the species are known
from the African and Oriental regions
(Evenhuis and Greathead 1999). Species
from other biogeographic regions have
not been treated via comprehensive tax-
onomic publications. In North and Cen-
tral America, the 23 described species
range throughout tropical and subtropical
areas (Painter and Painter 1963, Adams
and Yanega 1991), with three found north
of Mexico.
Biology.—
Specific biology related to
host location and oviposition behavior
has not been reported for Systropus spp.,
although they are documented to oviposit
directly onto their hosts by adhering eggs
to the host’s integument (Genty 1972). In
at least one instance, oviposition of black
eggs is accompanied by rapid movement
of the abdomen of the fly on the host
(Genty 1972). Systropus spp. are specialists
on limacodid larvae and are placed in
Toxophorinae, one of only two bomb-
yliid subfamilies containing true endo-
parasitoids (the other is Anthracinae)
(Greathead 1987, Yeates and Greathead
1997). Hosts are known only for S. macer
in North America (see below). Extra-
limitally, host associations have been re-
ported from Africa (Hull 1973), Central/
South America (Adams and Yanega 1991,
Aiello 1980, Dyar 1900), and Asia
(Greathead 1987). Adults routinely take
nectar from flowers (Robertson 1928,
Painter and Painter 1963) and are com-
monly reported from woodland glades
and various habitats in the wet tropics
(Painter and Painter 1963, Greathead
1987). Two species are known from
Australia (Evenhuis and Greathead 1999).
Literature.—
Painter and Painter (1963)
revised the Systropus of North and
Central America treating 23 species,
including three found north of Mexico.
Extralimitally, Enderlein (1926) keyed
13 species from Southeast Asia, but 48
species are known from the region today
(Evenhuis and Greathead 2003). The group
has been cataloged along with a full bibli-
ography as part of larger works on Bomb-
yliidae (Evenhuis and Greathead 1999),
and these data are now available on the
Internet (Evenhuis and Greathead 2003).
North American Records:
Systropus macer Loew (Fig. 3): Systropus
macer Loew, 1863: 375. Lectotype /,
MCZ, designated by Evenhuis (1982) (not
examined).
Systropus macer is diagnosed in the key
of Painter and Painter (1963) by:
swollen part of propleura bright yellow,
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36

metasternum solid brown/black, meso-
notum with small yellow spots, legs light
brown/black, and the caudal edge of the
8
th
sternum notched/undulating.
Distribution: Northeastern North
America (Painter and Painter 1963).
Limacodidae: Adoneta spinuloides,
Apoda sp., Euclea delphinii (Boisduval),
Lithacodes fasciola (H. –S.), Proli-
macodes badia (Adams and Yanega
1991), Parasa indetermina (Boisduval)
(Epstein 1997).
Other hosts: No records.
Uramya Robineau-Desvoidy
(Tachinidae: Dexiinae: Uramyini)
(Figs. 1–2, 4, 6)
Diagnosis.—
Uramya can be identi-
fied in North America using the key by
Wood (1987) and can be distinguished
from Compsilura and Austrophorocera
by the following combination of characters:
ocellar seta weak or absent; facial ridge
bare except for a few small, decumbent
setae on lower third or less (Fig. 2); pros-
ternum bare (Fig. 6); 3 large postsutural
dorsocentral setae; metathoracic spiracle
fringed with plumose hairs of about equal
size along anterior and posterior edges,
leaving V-shaped middorsal opening
(Fig. 4); scutellum with 3 pairs of strong
marginal setae, including pair of strong,
crossed apicals; vein M sharply angled
at bend (Fig. 1); abdomen with 1–2
pairs of median discal setae on each of
tergites 3 and 4 (Fig. 1); female ovi-
positor short and pointed.
Fauna.—
Uramya isaNewWorldgenus
of about 20 described species ranging from
Canada to northern Argentina. Twelve
species were recognized from South
America by Guimara
˜es (1980), and seven
species were recorded from North America
by O’Hara and Wood (2004); however,
there is no recent revision or key available
to these species. Janzen and Hallwachs
(2009) recorded several undescribed mor-
phospecies of Uramya from Guanacaste
(Costa Rica); all were reared from
limacodids.
Biology.—
Females of Uramya are
ovolarviparous on lepidopteran larvae.
Hosts are known for four of the seven
species in North America. Two of the
four, U. limacodis and U. pristis, attack
limacodids (Arnaud 1978, see below).
Uramya halisidotae (Townsend) is most
commonly reared from Lophocampa
argentata (Packard) (as Halisidota argen-
tata; Arctiidae), but there is a single record
from Malacosoma sp. (Lasiocampidae)
(Aldrich 1921, Arnaud 1978). Stireman
and Singer (2003a, b) reared U.indita
(Walker) from Norape tenera Druce
(Megalopygidae) in Arizona.
Two South American species have
been reared in Venezuela: U. longa
(Walker) from Ammalo helops (Cramer)
(Arctiidae) and U. sibinivora Guimara
˜es
from Acharia sp. (as Sibine; Limacodidae)
(Guimara
˜es 1980). Cruttwell (1969)
documented the biology of U.brevicauda
Curran, reared from Pareuchaetes pseu-
doinsulata Rego Barros (Arctiidae) in
Trinidad. Eggs are laid directly on a host
and hatch immediately; the first instar
penetrates the host’s integument within
a few seconds. Cruttwell (1969) found
that larvae could develop gregariously in
lab-reared hosts, but wild hosts were
generally parasitized by single maggots
continuously throughout the year in Tri-
nidad. Schaffner (1959) reported that the
North American species U. limacodis
and U. pristis develop as solitary parasit-
oids in their recorded hosts (listed below),
have one generation per year, and
overwinter in the pupal stage. All of the
U. pristis reared as part of this study
followed this pattern.
About 10 morphospecies of Uramya
were reared from five genera of lima-
codids (Acharia Hu
¨bner, Euclea Hu
¨bner,
VOLUME 114, NUMBER 1
37

Isochaetes Dyar, Natada Walker, a n d
Parasa Moore) by Janzen and Hallwachs
(2009).
Literature.—
Aldrich (1921) revised the
Uramya of the Americas under the ge-
neric names Uramya and Pseudeuantha
Figs. 1–3. Tachinidae and Bombyliidae. 1, Uramya pristis, male, habitus (white arrows: (r), haltere;
(l), mouthparts; black arrow, discal setae); 2, U.pristis, male, head, lateral (arrow, setae absent on facial
ridge); 3, Systropus macer, female, habitus (arrows: (u), antenna; (l), mouthparts).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
38

Townsend. Twelve species were treated,
including only four of the seven species
currently recognized from North America.
Curran (1930) followed the taxonomy of
Aldrich (1921) and provided a key to the
three species of Pseudeuantha then known
from the U.S.A. but excluded without
comment P. j o h n s o n i . Reinhard (1935)
described two additional species, one in
Pseudeuantha and the other in Uramya.
Sabrosky and Arnaud (1965) reorganized
the species under Anaporia,Pseudeuantha
and Uramya,alsoaddingindita Walker to
Pseudeuantha and synonymizing john-
soni with pristis under Anaporia. Wood
(1987) synonymized Anaporia and Pseu-
deuantha with Uramya and keyed the
genus among other Nearctic genera. This
classification of seven species of Uramya
inNorthAmericawasfollowedinthe
catalog by O’Hara and Wood (2004).
O’Hara and Wood (2004; catalog) pro-
vided type and distributional information.
North American Records:
Uramya limacodis (Townsend) (Fig. 13):
Aporia limacodis Townsend, 1892: 275.
Holotype ?, SEMK (O’Hara and Wood
2004) (not examined).
ThereisnokeytotheUramya of North
America, but the species can be readily
separated into three groups. The first com-
prises U. halisidotae and U. aldrichi,
which are characterized in the male by
an unusual taillike projection of ab-
dominal tergite 5 that extends dorsally
beyond the male terminalia by about the
preceding length of tergite 5. The second
group comprises U. indita and U. um-
bratilis. These species are recognized
easily by their bi-colored wings in both
sexes: yellow basally and blackish on
apical half (particularly close to veins). The
final three species, U. limacodis,
U. pristis,andU. rubripes, are more or-
dinary in appearance although the last
(known only from Florida and Texas) is
the only species in which the legs and
portions of the abdomen are typically
reddish yellow. Uramya limacodis and U.
pristis are the only two species of the genus
in North America that have been reared
from limacodids. They are typically 7–12
mm long, and only the males are easily
distinguished. In males the long slender
hairs on the anepisternum, from the central
portion of the sclerite to behind the ane-
pisternal setae, are black in U. limacodis
(Fig. 13) and pale in U. pristis (Fig. 12).
Distribution: Saskatchewan to Nova
Scoti a, south to Wisconsin and Georgia
(O’Hara and Wood 2004).
Limacodidae: E.delphinii
‡
(CNC [North
Burgess Township, Lanark County,
Ontario, 1973]), Isa textula (H.-S.)
(Schaffner 1959, as Anaporia limacodis
ex Sisyrosea textula), Limacodes sp.
(Townsend 1892, as Aporia limaco-
dis), Li.fasciola (Schaffner 1959, as
Anaporia limacodis;CNC[NorthBur-
gess Township, Lanark County, Ontario,
1971]), Tortricidia flexuosa (Grote)
(Aldrich 1921, as Pseudeuantha co-
quilletti Aldrich; Johnson 1925, as
Pseudeuantha coquilletti; Schaffner
1959, as Anaporia limacodis).
Uramya pristis (walker) (Figs. 1–2, 4,
6, 12): Dexia pristis Walker, 1849: 841.
Holotype ?, BMNH (O’Hara and Wood
2004) (not examined).
See U. limacodis for differential diagnostics.
Distribution: Michigan to Quebec and
New Hampshire, south to Florida; also
Arizona (O’Hara and Wood 2004).
Limacodidae: Acharia stimulea
(Clemens) (Aldrich 1921, as Pseudeuan-
tha pristis ex. Empretia stimulea; Johnson
1925, as Pseudeuantha pristis ex. Em-
pretia stimulea), E.delphinii (Coquillett
1897, as Exorista blanda (Osten Sacken)
ex Euclea cippus (Cramer); Johnson,
1925, as Pseudeuantha pristis ex Eu-
clea “ceppus”[=cippus] (Cramer)), I.
textula (Coquillett 1897, as Exorista
isae Coquillett ex Isa inornata (Grote
VOLUME 114, NUMBER 1
39

and Robinson)), Phobetron pithecium (J.
E. Smith) (Schaffner 1959, as Anaporia
pristis). Five additional new limacodid
hosts are reported herein (Table 1): Ad.
spinuloides*,Isochaetes beutenmuelleri*,
Li.fasciola*,Pack a rdia geminata
(Packard)*,Tortricidia sp*.
Other hosts: Megalopyge crispata
(Packard) (Megalopygidae).
Misidentification: The record by
Coquillett (1897) and cited by Arnaud
(1978) of U. pristis (as Macquartia
pristis) reared from the arctiid Lopho-
campa argentata (as Halisidota ar-
gentata) is in error. The tachinid reared
was U.halisidotae (Aldrich 1921), a
well known parasitoid of L. argentata.
Compsilura Bouche
´(Tachinidae:
Exoristinae: Blondeliini)
(Figs. 5, 7–9)
Diagnosis.—
Compsilura concinnata
(Meigen) is the single species of the genus
in North America. It is typically 7–8 mm
in length but can be as short as 4 mm. It
can be identified in North America using
the key by Wood (1987) and can be dis-
tinguished from Uramya and Austro-
phorocera by the following combination of
characters: ocellar seta usually absent; fa-
cial ridge with row of stout setae on lower
one-half or more (Fig. 9); prosternum haired
laterally (Fig. 7); 4 postsutural dorsocentral
setae; metathoracic spiracle with posterior
lappet much larger than anterior one (Fig.
5); scutellum with 4 pairs of setae, apicals
crossed and shorter than laterals, subapicals
usually divergent; vein M obtusely-angled
at bend (Fig. 8); abdomen with 1 pair of
median discal setae on each of tergites 3
and 4 (Fig. 8); female with short spines
ventromedially on tergites 3 and 4, and
sicklelike piercing ovipositor (Fig. 8).
Fauna.—
Compsilura contains four spe-
cies, mostly of Old World distribution.
Compsilura concinnata was introduced
from Europe into North America on
multiple occasions beginning in the early
1900s to control a variety of lepidopteran
pests, most notably gypsy moth (L. dispar)
and browntail moth (Euproctis chrysor-
rhoea (L.)) (Boettner et al. 2000).
Biology.—
Only Com. concinnata has
reported host records. This species is
extremely polyphagous, attacking many
species of Lepidoptera in a variety of
families and some species of Hymenoptera
(Symphyta) (Herting 1960, Europe; Arnaud
1978, North America; Shima 2006, Japan).
Over 180 hosts have been recorded in North
America, including a single limacodid, M.
flavescens (Schaffner and Griswold 1934,
Schaffner 1959, repeated in Arnaud 1978;
all as Cnidocampa flavescens).
Compsilura species are ovolarvipar-
ous, injecting eggs with fully developed
and ready-to-hatch first instars. In Com.
concinnata, there is a peculiar adaptation
known in few other tachinids: the first
instar occupies the narrow space between
the peritrophic membrane and the wall of
the midgut and remains there throughout
most of its three larval instars (Ichiki and
Shima 2003). Bourchier (1991) suggested
that living in the midgut allows the de-
veloping maggot to evade host immune
response.
The life history of Com. concinnata
is varied according to host species and
geographic area. There are two to four
generations per year, often involving al-
ternate hosts throughout the season. In
larger hosts, multiple maggots can develop
gregariously. The species overwinters in
the larval stage within a host (Culver 1919,
Schaffner 1959) or possibly in the pupal
stage away from a host (Tothill 1922).
Literature.—
Shima (1984) diagnosed
Compsilura and the single Japanese
species Com. concinnata in a paper on
Japanese blondeliine genera Blondelia
Robineau-Desvoidy and Compsilura.
Wood (1985) revised and keyed the
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40

Figs. 4–8. Species of Uramya and Compsilura.4,Uramya pristis, metathoracic spiracle (arrow:
posterior lappet); 5, Compsilura concinnata, metathoracic spiracle (arrow: posterior lappet); 6, U. pristis,
prosternum, (arrow: no setae); 7, Com. concinnata, female, prosternum (arrow: setae); 8, Com. concinnata,
female, habitus (arrows: (u), bend of vein M; (m), median discal setae present; (l), piercing ovipositor).
VOLUME 114, NUMBER 1
41

Figs. 9–11. Males. 9, Compsilura concinnata, head, lateral (arrows: (u), no ocellar setae; (l), setae
on facial ridge); 10, Austrophorcera n. sp., head, lateral (arrows: (u), ocellar seta; (l), setae on facial
ridge); 11, A. n. sp., female, habitus (arrows: (u), bend of vein M; (m), median discal setae absent;
(l), non-piercing ovipositor).
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42

Blondeliini of North and Central America
and the West Indies. The genus was keyed
among other Nearctic Tachinidae by
Wood (1987) and among other Palearctic
Tachinidae by Tschorsnig and Richter
(1998). O’Hara and Wood (2004) pro-
vided type and distributional information.
North American Records:
Compsilura concinnata (Meigen) (Figs. 5,
7–9): Tachina concinnata Meigen, 1824:
412. Holotype /, NHMW (Herting 1972,
O’Hara et al. 2009) (not examined).
Compsilura concinnata is the most po-
lyphagous species of Tachinidae. It has
recently been implicated in the decline
of native species of saturniid moths in
New England (Boettner et al. 2000, El-
kinton and Boettner 2004). Historical
evidence suggests it may have success-
fully controlled browntail moth, one of
the invasive pest species it was originally
introduced against (Elkinton et al. 2006).
Distribution: Widespread throughout
all biogeographic regions of the Old
World (Shima 1984). Introduced into
North America and currently known
from most of southern Canada and
western and northeastern U.S.A.
(O’Hara and Wood 2004). The species
may still be expanding its range in
North America.
Limacodidae: M.flavescens (Webber
and Schaffner 1926, Schaffner and
Griswold 1934, Schaffner 1959; all
as Cnidocampa flavescens)andE.
delphinii*.
Other hosts: Over 180 species of Lepi-
doptera and Symphyta in North America
(Arnaud 1978, Boettner et al. 2000).
Austrophorocera Townsend
(Tachinidae: Exoristinae: Exoristini)
(Figs. 10–11, 14–17)
Diagnosis.—
The genus can be identi-
fied in North America using the key by
Wood (1987) and can be distinguished
from Uramya and Compsilura by the fol-
lowing combination of characters: ocellar
seta well developed (Fig. 10); facial ridge
with row of stout setae on more than lower
one-half (Fig. 10); prosternum haired lat-
erally (as in Fig. 7); 4 postsutural dorso-
central setae; metathoracic spiracle with
posterior lappet much larger than anterior
one (as in Fig. 5); scutellum with 4 pairs of
setae, apicals crossed and subequal in length
to laterals; vein M right-angled at bend (Fig.
11); abdomen without median discal setae
ontergites3and4(Fig.11);femalewith
non-piercing ovipositor (Fig. 11).
Fauna.—
There are 10 described spe-
cies of Austrophorocera in North America
(O’Hara and Wood 2004), and at least 15
species are described from the Neo-
tropical Region (Wood and Zumbado
2010), although most of the latter are
currently assigned to other genera. Janzen
and Hallwachs (2009) recorded about 20
morphospecies of Austrophorocera in
Guanacaste, Costa Rica, of which only
two are referable to named species. There
is no modern key to the New World spe-
cies of Austrophorocera. There are five
species in the Oriental and Australasian/
Oceanic regions (Crosskey 1976, Cantrell
and Crosskey 1989), with two of those
species reaching the Palearctic part of
China (O’Hara et al. 2009).
Biology.—
Austrophorocera species are
oviparous. Eggs are laid directly on a host
and hatch within a few days; the first in-
star then uses its mouth hooks to burrow
into the host. Two or three species have
been reared in North America (one nom-
inal), and except for one record from
a notodontid, the hosts have all belonged
to Limacodidae (see below). Janzen and
Hallwachs (2009) documented 307
Austrophorocera specimens (20 morpho-
species, two nominal) reared from 14 gen-
era and >20 species of Limacodidae in
Guanacaste. A widespread species in the
Oriental and Australasian regions, A.
VOLUME 114, NUMBER 1
43

Figs. 12–18. Species of Uramya,Austrophorocera, and Taeniogonalos. Figs. 12–13. Anepisternum.
12, Uramya pristis; 13, U. limacodis. Figs. 14–17. Male genitalia, lateral and posterior views, Austro-
phorocera spp. 14, A. cocciphila (arrows: (l), surstylus; (r), syncercus); 15, A. coccyx; 16, A. einaris;
17, A. n. sp. Fig. 18. Taeniogonalos gundlachii, fore wing.
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44

grandis (Macquart) has been reared from
several species of Thosea Walker and Dor-
atifera Duncan (Limacodidae) (Crosskey
1976, Chadwick and Nikitin1985), as well
as Heliothis sp. (Noctuidae) (Cantrell
1986).
Schaffner (1959) briefly reviewed the
life history of A. coccyx parasitizing two
limacodid species (see below). He re-
ported one parasitoid per host, one gen-
eration per year, and larva overwintering
in the prepupal stage of the host. Murphy
et al. (2009) reported on the undescribed
A. sp. recorded in the present study, stat-
ing that it is a larval-pupal parasitoid that
parasitizes the larva and emerges from
the host’s pupa the following spring. Host
pupation appears to be a trigger for de-
velopment of A. sp., because flies have
repeatedly been observed to eclose from
non-diapausing host cocoons in the one
North American limacodid species with
a partial second generation, E.delphinii.
Literature.—
Aldrich and Webber
(1924) treated seven of the 10 species
of Austrophorocera recognized currently
from North America as Phorocera
Robineau-Desvoidy (subgenus Paraseti-
gena Brauer and Bergenstamm) in part.
Sabrosky and Arnaud (1965) dispersed
the species of P.(Parasetigena) among
several genera but kept together six
of the seven Austrophorocera species
in the genus Palpexorista Townsend.
These authors added three additional
species to Palpexorista that had been
described in Phorocera after Aldrich
and Webber (1924). Wood (1987) synon-
ymized Palpexorista under Austrophoro-
cera, a genus formerly known only from
the Old World. Thus, the nine species
assigned to Palpexorista by Sabrosky and
Arnaud (1965) were treated under Austro-
phorocera by O’Hara and Wood (2004).
A tenth species was added as a new North
American record by O’Hara and Wood
(2004).
North American Records:
Austrophorcera cocciphila (Aldrich and
Webber): Phorocera (Parasetigena)
cocciphila Aldrich and Webber, 1924:
53. Holotype ?, USNM (O’Hara and
Wood 2004) (not examined).
Two of the North American species
of Austrophorocera,A. cocciphila and
A. einaris, are easily distinguished from
the other species by having two katepis-
ternal setae instead of three. The male
syncercus of these two species is densely
setose posteriorly (Figs. 14, 16). The se-
tae are black, and their outer portions
have a crinkled appearance. The setae
form a thicker and tighter mat in A. ei-
naris (Fig. 16) than in A. cocciphila (Fig.
14), and the mat in A. einaris is positioned
more centrally on the more elongate syn-
cercus of that species. The syncercus of A.
einaris in lateral view is distinctly bent
subapically, compared to slightly curved
subapically in A. cocciphila.
Distribution: District of Columbia,
Maryland, Ohio, Tennessee (O’Hara
and Wood 2004).
Limacodidae: Ac. stimulea.
Other hosts: None known.
Austrophorocera coccyx (Aldrich and
Webber) (Fig. 15): Phorocera (Para -
setigena)coccyx Aldrich and Webber,
1924: 64. Holotype ?, USNM (O’Hara
and Wood 2004) (not examined).
Austrophorocera coccyx and A. n. sp. be-
long to a group of seven North American
Austrophorocera species that have three
katepisternal setae and a patch of short, fine,
wavy yellowish hairs within a depressed
area on the basal portion of the posterior
surface of the male syncercus. Two of the
seven species, A. pellecta (Reinhard)
(Arizona) and A. tuxedo (Curran), are
smaller than the rest at about 6–9 mm
long; the former has an unusually broad
parafacial, and the latter is yellowish on
the upper portion of the head (fronto-
orbital plate) and the thoracic dorsum.
VOLUME 114, NUMBER 1
45

The remaining five species comprise
A. alba (Townsend), A. coccyx,A. stol-
ida (Reinhard), A. virilis (Aldrich and
Webber), and A. n. sp. These species are
moderately large tachinids (about 10–13
mm long except for A. n. sp., which is
8.5–10.0 mm long) and are grayish black
in coloration. They are most easily
separated by distinct differences in
their male syncercus: A. alba (and other
forms that may represent undescribed
species) has the posterior surface of the
syncercus expanded laterally at mid-
length to form a lobe on either side, and
the apical portion of the syncercus
is short and narrow; in A. coccyx (Fig. 15)
the syncercus in posterior view is
abruptly narrowed at midlength and ta-
pered to a pointed tip; in A. stolida the
syncercus is gently narrowed at mid-
length, tapered to a rounded tip, and
bears a tuft of dense black setae basally
above the area with the short yellow
hairs; in A. virilis the syncercus is smoothly
tapered from base to tip; in A.n.sp.(Fig.
17) the syncercus is broadened on lower
third and abruptly narrowed subapically
to a pointed tip.
Distribution: Ontario to New York,
south to Florida, also Arizona and
Texas (O’Hara and Wood 2004).
Limacodidae: Ac.stimulea (Aldrich and
Webber 1924, as Pho.coccyx; Schaffner
1959, as Pho.longiuscula (Walker)) and
Pho. pithecium (Schaffner and Griswold
1934, as Pho. longiuscula; Schaffner
1959, as Pho. longiuscula).
Other hosts: Datana ministra (Drury)
(Notodontidae).
Austrophorcera einaris (Smith): Phor-
ocera einaris Smith, 1912: 119. Holotype
?, USNM (O’Hara and Wood 2004) (not
examined).
See A. cocciphila for differential diagnostics.
Distribution: “Michigan to Quebec and
Massachusetts, south to Texas and
Florida” (O’Hara and Wood 2004).
Limacodidae: E. delphinii.
Other hosts: None known.
Austrophorcera imitator (Aldrich and
Webber): Phorocera (Parasetigena)im-
itator Aldrich and Webber, 1924: 63.
Holotype ?, USNM (O’Hara and Wood
2004) (not examined).
Austrophorocera imitator and another
North American species, A. sulcata
(Aldrich and Webber), have three kate-
pisternal setae and only black setae
posteriorly on the male syncercus. The
setae on the syncercus are shorter and
more regular in appearance than in A.
einaris and A. cocciphila. The syncercus
in both species is slender and evenly
tapered in posterior view; in lateral view
the apex is slightly clubbed and bent
posteriorly in A. imitator and tapered to
a point and slightly curved anteriorly in
A. sulcata. The surstylus of A. imitator is
distinctive among the North American
species of the genus in being tightly ap-
pressed against, and possibly fused with,
the syncercus along its length, giving the
appearance on first inspection of being
absent.
Distribution: Missouri to Massachusetts,
south to Texas and Florida (O’Hara
and Wood 2004).
Limacodidae: Pr. badia.
Other hosts: None known.
Austrophorocera n. sp. (Fig. 17) Murphy
et al. 2009 (Austrophorocera n. sp.).
See A. coccyx for differential diagnostics.
Distribution: Eastern U.S.A.
Limacodidae: Reported from “Lima-
codidae” in Murphy et al. (2009) and
Li. fasciola*(Table 1).
Austrophorocera “sp. 1”: Stireman and
Singer 2003a, b.
Not examined and characters not known.
Distribution: Studied in Arizona.
Limacodidae: Pa.chloris (H.-S.)
(Stireman and Singer 2003b), Pro.
trigona (Edwards) (Stireman and Singer
2003a, b).
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46

Hymenoptera
Ceraphron Jurine
(Ceraphronoidea: Ceraphronidae)
(Figs. 74–77)
Diagnosis.—
Vertex rounded (Fig. 74),
ocellar triangle small, distant from the
eyes; median furrow present on the mes-
oscutum (Fig. 75) (Dessart and Cancemi
1986), and dorsum of the latter flat in lat-
eral view (Austin 1987b). Adult speci-
mens of Ceraphron, particularly males,
are difficult to identify to species due to
their simplified genitalia (I. Miko
´pers.
comm.) and the dearth and age of pub-
lished information for the Nearctic Re-
gion (Ashmead 1904, Brues 1906).
Fauna.—
Thereislittleknownabout
the diversity of Ceraphron; no compre-
hensive treatments have been published
regionally or globally. Twenty-four spe-
cies were recorded from North America
by Brues (1906).
Biology.—
Species of Ceraphron are
commonly considered hyperparasitic on
primary hymenopteran and dipteran par-
asitoids based largely on records from the
related genus Aphanogmus (recorded from
Ichneumonidae and Braconidae by Austin
(1987b)). Other species of Ceraphron have
been reared as hyperparasitoids (through
Hymenoptera) from both Aphididae (Evans
and Stange 1997) and Cecidomyiidae
(Franklin 1919), among other hosts.
Literature.—
Austin (1987b) recorded
asingleCeraphron sp. hyperparasitoid
from one limacodid rearing from Southeast
Asia. Brues (1906) published a key to
Nearctic Ceraphron and followed this
with a key to genera and species of Cera-
phronidae (Ceraphronoidea) in Connecticut
(Brues 1916). Ashmead (1893) reported on
the genera and species of Ceraphronidae
known in North America, and Dessart
(1981) proposed species groups of Cera-
phron. The most recent key to the world
genera of Ceraphronoidea (incl. subgenera
of Ceraphron) was published by Dessart
and Cancemi (1986).
North American Records:
Ceraphron sp. (Figs. 74–77): Diagnostics
unavailable for North American species.
Distribution: This cosmopolitan genus
is widespread in North America.
Limacodidae: Hyperparasitoid that
emerged from a primary parasitoid
(possibly Pl.americana*or Al.lilli*)
of Ac.stimulea on Q.rubra.
Other hosts: Unknown for this species.
Conura Spinola (Chalcidoidea:
Chalcididae: Chalcidinae)
(Figs. 78–86)
Diagnosis.—
Conura mayberecognized
by postmarginal vein longer than stigmal
vein (Fig. 78); gaster distinctly petiolate
(Figs. 79–80); and propodeal spiracle
subvertical to nearly longitudinal.
Fauna.—
Conura is a large genus of
;296 species restricted to the New
World, primarily the Neotropical Region
(283 species), with only 12 species in the
Nearctic (Noyes 2003). Species have been
recovered as far north as Alaska and as far
south as Chile and Argentina (Noyes
2003). For a discussion of the classification
of Conura and related genera, see Delvare
(1992). Of the known species from the
Nearctic, two (C. nigricornis,C. nortonii)
are specialists on Limacodidae and belong
to the nigricornis group, whose members
exclusively attack limacodids. The third
(C. camescens) is apparently polyphagous,
also known as a secondary parasitoid of
Hymenoptera through Lepidoptera (see
below), and belongs to the immaculata
group. Delvare (1992) recorded the biology
of this group as hyperparasitoids of ich-
neumonoids through lepidopterans. The
members of this group are recognized by
the superficial malar sulcus (Fig. 82),
interantennal projection a weak carina,
shallow scrobal depression (Fig. 81), and
VOLUME 114, NUMBER 1
47

Figs. 19–22. Mesochorus discitergus, female. 19, Habitus, scale =1.0 mm (arrows: (u), areolet; (l),
ovipositor sheaths); 20, Aerolet; 21, Metasoma, anterior lateral, scale =0.5 mm (arrows: (l), glymma; (r),
spiracle); 22, Ovipositor and hypopygium (arrow).
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48

Figs. 23–27. Species of Hyposoter and Lysibia. Figs. 23–24, 26–27. Hyposoter fugitivus, female. 23,
Habitus; 24, Propleuron, lateral (arrow); 26, Aerolet; 27, Metasoma, lateral (arrows: (l), glymma; (r),
spiracle). Fig. 25. Lysibia mandibularis, propleuron, lateral (arrow).
VOLUME 114, NUMBER 1
49

Figs. 28–29. Species of Casinaria and Baryceros. 28, Casinaria grandis, female, habitus, scale =
2 mm (arrow: 2m-cu crossvein); 29, Baryceros texanus, female, habitus, scale =2 mm.
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50

Figs. 30–34. Species of Hyposoter and Isdromas. Figs. 30, 32. Hyposoter fugitivus. 30, Meso-
pleuron, lateral (arrow: sternaulus); 32, Head, anterior (arrow: no groove). Figs. 31, 33–34. Isdromas
lycaenae, female. 31, Mesopleuron, lateral (arrows: sternaulus); 33, Head, anterior (arrows: (u), shallow
groove; (l), impressed clypeal margin); 34, Habitus, (arrow: aerolet), scale =1 mm.
VOLUME 114, NUMBER 1
51

reticulo-rugose to rugose mesoscutum.
Burks (1940) defined the nigricornis group
based on deep and coarse body sculpture,
the presence of conspicuous long pilosity
(Figs. 84–85), and a moderately deep
scrobal depression with slightly cari-
nate m arg ins . Delvare (1992) refined the
definition of this group by noting notauli
Figs. 35–38. Species of Lysibia,Acrolyta, and Baryceros. Fig. 35. Lysibia mandibularis, female,
habitus (arrow: vein 2m-cu), scale =1 mm. Fig. 36. Acrolyta nigricapitata, female, habitus, scale =1mm.
Figs. 37–38. Baryceros texanus, female. 37, Mesosoma, lateral (arrow: epomia); 38, Ovipositor tip.
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52

Figs. 39–45. Species of Braconidae, Acrolyta,Isdromas, and Lysibia. 39, Cyclostome braconid,
hypoclypeal depression (arrow: labrum); 40, Non-cyclostome braconid, head, anteroventral (arrow: la-
brum); 41, Acrolyta nigricapitata, head, posterolateral, scale =0.5 mm (arrow: hypostomal/occipital carina
junction); 42, Isdromas lycaenae, head, posterolateral, scale =0.5 mm (arrow: hypostomal/occipital carina
junction); 43, Lysibia mandibularis, head, anterior; 44, Is. lycaenae, head, anterior. 45, Braconidae, hind
wing (arrows: (l), M+CU; (r), 1M).
VOLUME 114, NUMBER 1
53

Figs. 46–49. Species of Trirhaphis and Ascogaster. Figs. 46–47, 49. Trirhaphis discoideus, female.
46, Mesopleuron (arrow: epicnemial carina); 47, Tarsal claw. Fig. 48. Ascogaster quadridentata, met-
asoma, dorsal. Fig. 49. Mesosoma, dorsal.
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54

Figs. 50–51. Species of Trirhaphis. 50, Trirhaphis eupoeyiae, female, habitus; 51, T. harrisinae,
female, habitus. Scales =1.0 mm.
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Figs. 52–55. Trirhaphis harrisinae, female. 52, Metasoma, anterodorsal (arrows, median carina);
53, Mesosoma, dosal; 54, Mesosoma, lateral, scale =0.25 mm; 55, Propodeum.
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56

deep, broad, and crenulate (Fig. 84); pro-
notum with recurved lamina anteriorly
(Fig. 85); and mesopleuron with ventral
carina (Fig. 83).
Biology.—
Generally, species of Conura
are documented as parasitoids of larvae
and pupae of different insects, mostly of
Lepidoptera. However, some species of
Figs. 56–57. Triraphis discoideus, female. 56, Habitus (large arrows: (u), vein 2m-cu absent; (l),
vein m-cu; small arrows: (u), 2CUa; (l), 1CUa); 57, Metasoma, anterodorsal (arrow: (l) groove between
T2 and T3; (r) median carina).
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57

Figs. 58–62. Species of Trirhaphis and Ascogaster. Figs. 58–59. Trirhaphis discoideus, female. 58,
Propodeum (arrow: divergent carinae); 59, Hind tibia, apex (arrow: flattened setae). Figs. 60–62 As-
cogaster quadridentata, female. 60, Head, anterior; 61, Head, dorsal (arrow: occipital carina); 62,
Mesosoma, dorsal (arrow: scutellar disc).
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Figs. 63–65. Ascogaster quadridentata and Cotesia empretiae.63,Ascogaster quadridentata, female
habitus, scale =1.0 mm (arrows: (l), vein C+SC+R; (m), vein (RS+M)a; (r), vein 3RS); 64, Cotesia em-
pretiae, female, habitus; 65, Co. empretiae, wings (arrows: (l), vein C+SC+R; (r), areolet absent).
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59

Figs. 66–69. Cotesia spp. 66, Co. empretiae, female, propodeum and mesosoma (arrows: propodeal/
metasomal sculpture); 67, Co. empretiae, lateral metasoma (arrow: ovipositor and sheaths); 68, Co.
phobetri, female, habitus, scale =0.5 mm; 69, Co.schaffneri, female, habitus, scale =0.5 mm.
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60

Conura are recorded from Coleoptera,
Hymenoptera, Diptera, and Neuroptera,
and others are known secondary para-
sitoids of ichneumonoids and tachinids
through lepidopterans (Delvare 1992,
Boucˇek and Halstead 1997). Three spe-
cies that occur in North America are
known to attack limacodids, two of
these exclusively (see below). Known
biologies indicate chalcidid host special-
ization as solitary parasitoids, but some
smaller species of Conura and Brachy-
meria are gregarious.
Literature.—
Boucˇek (1992; key to
genera) revised the New World genera,
Delvare treated the New World Chalcidini
(1992; key to species groups of Conura;
side group keyed to species), and Burks
(1940) offered an earlier treatment of
Chalcidini, with keys to genera, species
groups, and species.
North American Records:
Figs. 70–71. Orthogonalys pulchella. 70, Habitus, female; 71, Habitus, male. Scales =4.0 mm.
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61

Conura nigricornis (F.): Chalcis nigricornis
F., 1798: 243. Lectotype /, ZMUC, des-
ignated by Boucˇek and Delvare (1992)
(not examined).
This species belongs to the rarely col-
lected nigricornis group (Burks 1940),
which contains 14 nominal (at least 16
undescribed) species, all of which are
Figs. 72–73. Taeniogonalos gundlachii. 72, Habitus, female (arrow: armature); 73, Habitus, male.
Scales =2.0 mm.
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62

known solitary parasitoids of Limaco-
didae and Megalopygidae (Delvare
1992). This species is yellow with black
maculae on the body as reported by
Boucˇek and Delvare (1992: 29), the
scutellum has a median groove, and the
frenal carina is present and not fused
with the apical margin of the scutellum.
The propodeum has distinct lateral teeth
(fig. 42 in Boucˇek and Delvare 1992).
Burks (1940) separates C. nigricornis
from C. nortonii by the presence of five
Figs. 74–78. Species of Ceraphron and Conura. Figs. 74–77. Ceraphron sp. 74, Habitus, male,
scale =0.5 mm. 75, Mesosoma, female, dorsal; 76, Antenna, female, scale =0.5 mm. 77, Fore wing.
Fig. 78. Conura nortonii, male, fore wing, scale =1.0 mm.
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63

to six hamuli on the hind wing and the
presence of an inner tooth on the hind
femur as compared to C. nortonii (see
below).
Distribution: South America (Brazil,
Guyana) and eastern U.S.A.
Limacodidae: Adoneta sp. (Boucˇek and
Delvare 1992), Ad.spinuloides (Peck 1963,
Figs. 79–80. Species of Conura. 79, Conura nortonii, male, habitus. 80, Conura immaculata, fe-
male, habitus. Scales =2.0 mm.
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64

Herting 1976, Burks 1979), Apoda sp.
(Peck 1963, Burks 1979, Boucˇek and
Delvare 1992), Par a s a sp. (Boucˇek and
Delvare 1992), Pa.indetermina (Hert-
ing 1976, Burks 1979).
Conura nortonii (Cresson) (Figs. 78–
79, 83–86): Smicra nortonii Cresson,
1872: 45. Neotype ?,USNM,des-
ignated by Delvare (1992: 239)
(examined).
Figs. 81–86. Speceis of Conura. Figs. 81–82. Conura immaculata, female. 81, Head, anterior, scale =
0.5 mm; 82, Lower face, anterior, scale =0.5 mm (arrow: superficial malar sulcus). Figs. 83–86. Conura
nortonii, male. 83, Mesopleuron (arrows: (u), prepectus; (l), ventral carina); 84, Mesosoma, anterodorsal
(arrow: notaulus); 85, Mesosoma, anterodorsal (arrow: recurved pronotum); 86, Lower face, anterior (ar-
row: malar sulcus).
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65

Figs. 87–91. Psychophagus omnivorus, female. 87, Habitus (arrow: prepectus), scale =1.0 mm; 88,
Head, anterior; 89, Antenna; 90, Mesosoma, dorsal; 91, Metasoma, dorsal.
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66

Figs. 92–96. Species of Psychophagus,Trichogramma, and Pediobius. Figs. 92–93. Psychophagus
omnivorus, female. 92, Hind wing (arrow: hamuli); 93, Hamuli. Figs. 94, 96. Trichogramma sp., female.
94, Tarsus; 96, Fore wing (arrow: Rs1 setal track). Fig. 95. Pediobius crassicornis, female, propodeum
(arrows: (u), scutellar setal socket; (l), divergent carinae).
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67

This species also belongs to the nigricornis
group. Schaffner (1959) reported that two
specimens, one per host, were reared
from Ph.pithecium cocoons collected in
Bolton, Connecticut. The base color of
this species is yellow with black mac-
ulation, although the neotype in the
USNM is more orange brown (perhaps
killed with cyanide) than yellow. Com-
pared to C. nigricornis, the propodeum
lacks lateral teeth and the scutellum
lacks a median groove, although there is
minute lamina apically (Burks 1940).
Conura nortonii has three hamuli on the
Figs. 97–99. Pediobius crassicornis, female. 97, Habitus, scale =1.0 mm; 98, Pupal remnants (arrow)
within limacodid larval cadaver; 99, Pupal remnants (arrow) within pupa of Platyplectrus americana.
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hind wing and lacks the inner tooth on the
hind femur.
Distribution: Eastern U.S.A., west to
Texas (Burks 1940).
Limacodidae: Apoda sp. (Peck 1963,
Burks 1979), Ph. pithecium (Schaffner
1959, Peck 1963, Herting 1976, Burks
1979), Pr.badia (Herting 1976, Burks
1979).
Conura camescens Delvare: Ceratosmicra
flavescens Cameron, 1913: 114. Conura
camescens is a replacement name for
Conura flavescens (Cameron 1913) nec
(Andre
´1881). Lectotype ?, BMNH,
designated by Delvare (1992: 217) (not
examined).
This species belongs to the immaculata
subgroup of the immaculata group
(Delvare 1992). Delvare (1992) reported
this species from the U.S.A. without fur-
ther locality information. A specimen
from Venezuela in the USNM identified
by Delvare has label data indicating
Manduca sexta (Joh.) as a host [possibly
on primary parasitoid]. Two additional
specimens, also from Venezuela, indicate
Alabama argillacea (Hu
¨bner), “par. by
Rogas.” This species is reported as a pale
brownish yellow with areas of the face,
pronotum, mesoscutal sutures, and scu-
tellum in basal half more yellow in color
(Cameron 1913). The ;13 teeth of the
hind femur are black, the basal tooth
triangular with the apical teeth closely
appressed.
Distribution: New World. (Delvare 1992).
Limacodidae: Acharia sp. (as Sibine;
hyperparasitoid of Cotesia sp.) (Delvare
1992).
Other hosts: Prodenia eridania
(Cramer) (Noctuidae); Plutella xy-
lostella (L.) (Plutellidae); Apanteles
sp., Cotesia sp., Co. plutellae (Kurd-
jumov), and Rogas sp. (Braconidae)
(De Santis 1979, De Santis and Fidalgo
1984, Delvare 1992, Cha
´vez et al.
1993).
Alveoplectrus Wijesekara and Schauff
(Chalcidoidea: Eulophidae: Eulophinae)
(Fig. 104)
Diagnosis.—
Alveoplectrus may be re-
cognized by the presence of two sub-
median carina on the propodeum (Fig.
104), scutellum with lateral grooves broad
and continuous posteriorly (Fig. 104),
dorsellum produced medially as cuplike
carina (Fig. 104), and hind tibia with
single elongate spur. The related genus
Platyplectrus (see below) has a single
median carina on propodeum, scutellum
with narrow lateral grooves discontinuous
posteriorly, dorsellum not produced as
triangular point posteromedially, and hind
tibia with two elongate spurs (>0.5X
lengthofhindtarsus).
Fauna.—
Alveoplectrus is a small ge-
nus of four species restricted to the
New World, primarily the eastern Nearctic
and Neotropical regions (Wijesekara and
Schauff 1997, Gates and Stoepler 2010).
Species of Alveoplectrus were previously
placed in Euplectrus (Ashmead 1904)
or Euplectromorpha (see Schauff et al.
1997). The published distribution for Al-
veoplectrus includes Brazil, Costa Rica,
and the U.S.A. (Florida) (Wijesekara and
Schauff 1997). For a discussion of the
classification of Alveoplectrus and re-
lated genera, see Wijesekara and Schauff
(1997).
Biology.—
Known biologies indicate
host specialization as solitary ectopar-
asitoids of larvae of Limacodidae. One
species, Al.floridanus Wijesekara and
Schauff, is recorded from label data for
undetermined species in USNM indicat-
ing Neotropical Euprosterna elaea (as
elaesa) Dyar and Euprosterna sp. (Li-
macodidae) as hosts.
North American Records:
Alveoplectrus floridanus Wijesekara and
Schauff: Wijesekara and Schauff, 1997:
105. Holotype /, USNM (examined).
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69

This species is differentiated from Al.
lilli by the absence of a short, transverse
carina beneath the anterior ocellus and
the presence of a transepimeral sulcus
(compare with Gates and Stoepler 2010:
fig. 2).
Distribution: U.S.A. (Florida) (Wijesekara
and Schauff 1997).
Figs. 100–105. Species of Pediobius,Alveoplectrus, and Platyplectrus. Figs. 100–103. Pediobius
crassicornis, female. 100, Head, anterior; 101, Mesosoma, dorsal; 102, Female, antenna; 103, Male,
antenna. Fig. 104. Alveoplectrus lilli, female, mesosoma, posterodorsal (arrows: (l) dorsellum pro-
duction, (r) submedian carina). Fig. 105. Platyplectrus americana, female, mesosoma, dorsal (arrows:
(l) scutellar groove, (r) median carina).
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Limacodidae: Ala. slossoniae (Packard)
(Wijesekara and Schauff 1997).
Alveoplectrus lilli Gates (Fig. 104): Gates
and Stoepler 2010: 209. Holotype /,
USNM (examined).
This species is differentiated from Al. flor-
idanus by the presence of a short, trans-
verse carina beneath the anterior ocellus
and the absence of a transepimeral sul-
cus (figured in Gates and Stoepler 2010:
fig. 2).
Distribution: Eastern U.S.A. (Gates
and Stoepler 2010).
Limacodidae: I.textula,Li. fasciola,
E.delphinii (Gates and Stoepler 2010),
and Ad. spinuloides*.
Platyplectrus Ferrie
`re (Chalcidoidea:
Eulophidae: Eulophinae)
(Figs. 105–113)
Diagnosis.—
Platyplectrus is most
likely to be confused with Alveoplectrus
(among parasitoids of limacodids in North
America) and Euplectrus, although spe-
cies in the latter genus are not known from
limacodids in North America. Species of
Platyplectrus may be recognized by the
following combination of features: single
median carina on propodeum (Fig. 105),
scutellum with narrow lateral grooves dis-
continuous posteriorly (Fig. 105), dorsel-
lum not produced (Fig. 105), and hind tibia
with two elongate spurs. Alveoplectrus has
two submedian carinae on propodeum
(Fig. 104), scutellum with lateral grooves
broad and continuous posteriorly (Fig.
104), dorsellum produced medially (Fig.
104), and hind tibia with single elongate
spur. Euplectrus, aside from its host pref-
erences, is most similar to Platyplectrus in
overall coloration and habitus but lacks
scutellar grooves.
Fauna.—
One species is known from
North America of 46 described worldwide.
Highest species richness is in southeastern
Asia, but several species are known from
Australia and eastern Europe (Noyes
2003).
Biology.—
Known biologies indicate
host specialization as solitary or gre-
garious ectoparasitoids of larvae of
Limacodidae. Laboratory studies of the
extralimital Pl. natadae Ferrie
`re (Gadd
et al. 1946) revealed this species engaged
in host feeding once for every four eggs
laid, and a host used for host feeding was
never parasitized. Oviposition occurs on
the lateral surfaces below any tubercles of
the host after the host is paralyzed with a
sting. The egg, positioned externally, is held
in place by a pedicel within the host’s body.
In nature, this species usually lays one or
rarely lays two eggs per host with only one
larva typically reaching maturity. However,
in the present study, the number of eggs
per host appears to be size-dependent,
with larger hosts (or later instars) yielding
as many as 14 mature larvae. A newly
eclosed larva migrates to the venter of the
host’s body where feeding begins imme-
diately and results in the death of the host.
Pupation occurs under the host with the
host cadaver providing a protective cover-
ing (Gadd et al. 1946, M. Gates pers. obs.).
Literature.—
Only eight species have
associated host records, with five of those
records being from limacodids. The
widespread Pl. laeviscuta (Thomson)
ranges from Sweden and China south to
Australia and is cited from Stigmella rufi-
capitella (Haworth) (Nepticulidae) (Noyes
2003), but we find no record of it in the
original paper (Hansson 1987). Platyplec-
trus chlorocephalus (Nees) is recorded
from a zygaenid (Rhagades pruni (Denis
and Schiffermu
¨ller)), and this is perhaps
unsurprising given the relationship be-
tween Zygaenidae and Limacodidae
(Epstein 1996, Wahlberg et al. 2005,
Niehuis et al. 2006). The third non-limacodid
record published by Zhu and Huang
(2004) was based on label data: Cifuna
locuples Wlkr (Lymantriidae).
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71

Figs. 106–110. Platyplectrus americana, female. 106, Habitus, scale =1.0 mm; 107, Head, anterior;
108, Head, posterior; 109, Mesosoma, lateral (arrow: prepectus); 110, Mesosoma, ventral.
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Redescription of Platyplectrus americana
(Figs. 105–113):
Diagnosis.—
Platyplectrus americana is
the sole representative of the genus in
North America and thus will not be con-
fused with congeners. Platyplectrus can be
differentiated from related eulophid genera
by the generic characters outlined above.
Note.—
Girault (1916) described this
species from a single male. The original
description is short, and additional mor-
phological details of the male require
documentation, as given below. The fe-
male is newly described.
Description.—
Female. Body length:
2.4 mm (1.2–3.5 mm; n =20). Color:
Black except for the following golden—
scape, flagellum in ventral half, fore coxa
in apical third, legs, apical tarsomere, and
basal half gaster dorsomedially (Fig. 106);
brown—dorsal half flagellum, tegula,
metafemur in apical half except extreme
apex golden, pretarsus, gaster apically, lat-
erally, and ventrally, wing veins; white—
basal three tarsomeres.
Head: 1.7X as broad as high, clypeal
region transversely impressed along mar-
gin, smooth, straight ventrally; malar sul-
cus fine, slightly curved, malar space 0.53X
eye height; mandibles reduced, not touching
medially; toruli with upper margin posi-
tioned one-half torular diameter above
lower ocular line, separated by ;2torular
diameters (Fig. 107). Scrobal depression
smooth and shallow, shaped as an inverted
V. Antenna (Fig. 111) with scape reaching
three-quarters distance to anterior ocellus;
ratio of scape (minus radicle):pedicel:
anellus: F1:F2: F3:F4:club as 27:8:1:16:
14:12:12:18; funicular segments with 2
Figs. 111–113. Platyplectrus americana, female. 111, Antenna; 112, Fore wing; 113, Antenna, male.
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73

irregular rows of interdigitated longitu-
dinal sensilla and evenly setose; clava
apparently bisegmented with fusion com-
plete. Ratio of lateral ocellus:OOL:POL
as 7:7:14. [Head posteriorly lacking post-
occitpital carina, hypostomal carinae pres-
ent ventrally but faint (Fig. 108).]
Mesosoma (Fig. 105): 1.3X as long as
broad; midlobe of mesoscutum 0.9X as
long as broad; scutellum 1.0X as long
as broad; notauli fine, complete; midlobe
of mesoscutum shallowly bilobate post-
eromedially; scutellum shallowly bilobate
anteromedially, sublateral grooves mesad
scutellar setae, extending from scutoscu-
tellar suture to near posterior edge scu-
tellum; scutoscutellar suture obliterated
medially; axilla smooth with few setae
posteromedially; lateral surface of pre-
pectus triangular, smooth; [subventral ca-
rinae of prepectus steplike, prepectus with
ovate depression ventromedially] (Fig. 109).
Mesepimeron smooth, femoral depression
a sigmoidal sulcus, small linear depression
posteromedially. Mesepisternum smooth
(Figs. 109–110), convex, defined poste-
riorly by carina, discrimen medially be-
tween mesofurcal pit and anterior margin
mesopectus. Dorsellum smooth with post-
eromedial truncation, subequal in length
with propodeum medially, slightly pro-
jecting over propodeum medially and
excavated submedially (Fig. 105). Meta-
pleuron and lateral areas of propodeum
(Fig. 109) smooth; callus setose; propo-
deum smooth with elevated median carina
broadest anteriorly and posteriorly broadly
joined to nucha, with fine rugulosity;
flangelike carina extending laterally from
nucha toward metasomal foramen and spi-
racle; spiracle about one-third its greatest
diameter from dorsellum. All coxae im-
bricate, smoother laterally. Fore wing with
ratio of marginal vein:postmarginal vein:
stigmal vein as 64:45:23 (Fig. 113).
Metasoma: Smooth; petiole ;2.0X
as broad as long, finely sculptured dor-
sally; Gt1 depressed anterodorsally near
petiole occupying about one-third the
length of gaster; remaining terga and syn-
tergum smooth; [petiole transverse in dor-
sal view, smooth; lateral carinae on petiole
body meeting ventrally as slightly pointed
collar]; measurements of gastral terga
along midline as 30:9:9:7:5:7:8.
Male. Body length: 1.5 mm (1.2–2.8 mm;
n=5). Color: As described for female,
but with legs and venter of gaster paler,
whitish. Sculpture as described for fe-
male. Antenna with funicular segments
(Fig. 112) minutely pedicellate, each with
multiple rows of suberect setae and about
1.0X as long as width of segment; ratio of
scape (minus radicle):pedicel:anellus:
F1:F2:F3:F4:F5:club as 16:7:1:8:8:8:8:15;
scape with ventral plaque not apparent.
Gastral petiole in lateral view roughly
cylindrical, roughly trapezoidal in dorsal
view, length about 0.8X as long as
greatest width.
Variation.—
Most variation occurs in
coloration and size. Smaller male spec-
imens (<1.8 mm; range =1.1–1.8 mm)
tend toward a browner coloration rather
than black (especially the head and
mesosoma) with brownish maculation
on the femora only a faint suffusion
(especially fore and midfemora) and the
fore femur may be entirely golden/whitish
whereas the hind femur may approach
being entirely dark brown. The whitish
maculation dorsally on the gaster may be
confined only to that area but can extend
laterally and ventrally. Two males are
almost golden in body color. For females
(range =1.3–2.3 mm), the coloration
patterns seen for males based on size seems
to hold true. One specimen is medium
brown with white legs, except for brown
suffusion on the hind femur; the antenna
is whitish. For both sexes, there are ex-
ceptions to the size/color trend but typically
only at the smaller size range (dark, small
specimens).
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74

Type Material.—
20471 [red label,
handwritten]; Euplectromorpha americana
?, Type, Girault [handwritten] (USNM,
examined).
Material Examined.—
DISTRICT OF
COLUMBIA: Rock Creek Park, col-
lected 9.VIII.2005, J. Lill et al. 05-607,
ex Acharia stimulea on Robinia sp. (1/
USNM); same locality, collected 12.
VII.2006, emerged 23.VII.2006, J. Lill,
et al., 06-281, ex Limacodidae on Carya
glabra (?USNM); same locality, col-
lected 23.VII.2006, emerged 14.VIII.2006,
J. Lill, et al. 06-313, ex Lithacodes fasciola
on Nyssa sylvatica (/?USNM); same
locality, collected 14.VIII.2006, emerged
21.VIII.2006, J. Lill et al. 06-326, ex Pro-
limacodes badia on Quercus alba (/
USNM); same locality, collected 13.
VIII.2006, emerged 3.IX.2006, J. Lill et al.,
06-336, ex Acharia stimulea on Acer sac-
charinum (/USNM); same locality, col-
lected 20.VIII.2006, emerged 3.IX.2006, J.
Lill et al. 06-387, ex Euclea delphinii on
Nyssa sylvatica (10/USNM); same lo-
cality, collected 20.VIII.2006, J. Lill et al.,
06-389, ex Adoneta spinuloides on Nyssa
sylvatica (6/USNM); same locality, col-
lected 25.VIII.2006, J. Lill et al., 06-408,
ex Lithacodes fasciola on Nyssa sylvatica
(3/USNM); same locality, collected 26.
VI.2007, emerged 17.VII.2007, J. Lill et al.
07-120, ex Lithacodes fasciola on Quercus
rubra (/USNM); same data, but collected
30.VI.2007, emerged 17.VII.2007, 07-140,
ex Lithacodes fasciola on Nyssa sylvatica
(?USNM); same data, 07-143, ex
Limacodidae sp. on Fagus grandifolia (?
USNM); same locality, collected 30.
VI.2007, emerged 17.VII.2007, J. Lill
et al., 07-144, ex Limacodidae sp. on
Fagus grandifolia (?USNM); same
data, 07-148 (/USNM); same data, 07-
149 (?USNM); same data, 07-152 (?
USNM). MARYLAND: Montgomery
Co.: Little Bennett Regional Park, col-
lected 25.VII.2007, J. Lill et al. 07-320,
ex Acharia stimulea on Quercus alba
(1/USNM); same locality, collected
25.VII.2007, emerged 21.VIII.2007, J.
Lill et al. 07-319, ex Acharia stimulea on
Prunus serotina (2/?USNM); same
data, 07-316, (1/USNM); same data,
07-315, (2/USNM); same locality,
collected 25.VII.2007, J. Lill et al. 07-
340, ex Acharia stimulea on Quercus
rubra (1/USNM); same data, 07-339,
(1/USNM); same locality, collected
25.VII.2007, emerged 21.VIII.2007, J.
Lill et al. 07-338, ex Acharia stimulea on
Quercus rubra (1/USNM); same lo-
cality, collected 9.VII.2008, emerged 25.
VII.2008, J. Lill et al. 08-152, ex Lith-
acodes fasciola on Fagus grandifolia
(1/USNM); same locality, collected 9.
VII.2008, emerged 18.VII.2008, J. Lill
et al. 08-155, ex Euclea delphinii on
Quercus alba (1/USNM); same local-
ity, collected 10.VII.2008, emerged 25.
VII.2008, J. Lill et al. 08-175, ex Euclea
delphinii on Fagus grandifolia (1/
USNM); same data, 08-175 (1/USNM);
same data, 08-176, (1/USNM); same
data, 08-177, (1/USNM); same data, 08-
178 (1/USNM); same locality, collected
10.VII. 2008, emerged 28.VII.2008, J.
Lill et al. 08-186, ex Euclea delphinii on
Carya glabra (1/USNM); same locality,
collected 10.VII.2008, emerged 21.
VII.2008, J. Lill et al. 08-187, ex Euclea
delphinii on Carya glabra (1/USNM);
same locality, collected 11.VII.2008,
emerged 25.VII.2008, J. Lill et al. 08-
200, ex Euclea delphinii on Quercus ru-
bra (1/USNM); same locality, collected
14.VII.2008, emerged 28.VII.2008, J.
Lill et al. 08-217, ex Euclea delphinii on
Nyssa sylvatica (1/USNM); same lo-
cality, collected 19.VII.2007, emerged
21.VIII.2007, J. Lill et al., 07-260, ex
Lithacodes fasciola on Quercus rubra
(3/USNM); same locality, collected 19.
VII.2007, emerged 4.VIII.2007, J. Lill
et al., 07-292, ex Phobetron pithecium on
VOLUME 114, NUMBER 1
75

Fagus g r a n d i f o l i a (3?USNM); same
data, 07-293 (?USNM); same data, 07-
295 (2?USNM); same locality, collected
25.VII.2007, emerged 21.VIII.2007, J. Lill
et al. 07-307, ex Euclea delphinii on Pru-
nus serotina (3/USNM); same locality,
collected 25.VII.2007, J. Lill et al. 07-351,
ex Acharia stimulea on Nyssa sylvatica
(1/USNM); same data, 07-346, ex
Acharia stimulea on Carya glabra (1/
USNM); same data, 07-343 (2/USNM);
same data, 07-345 (2/2?USNM);
same data, 07-349, ex Acharia stimulea
on Nyssa sylvatica (1?USNM); same
data, 07-354 (4/?USNM); same lo-
cality and date, emerged 21.VIII.2007, J.
Lill et al. 07-363, ex Lithacodes fasciola
on Fagus grandifolia (/?USNM);
same data, 07-367, ex Acharia stimulea
on Fagus grandifolia (/?USNM);
same data [no emergence date], 07-368
(1/USNM); same locality, collected 1.
VIII.2007, J. Lill et al. 07-466, ex Acha-
ria stimulea on Quercus alba (2/?
USNM); same data, 07-467 (2/USNM);
same data, emerged 21.VIII.2007, 07-470
(/?USNM); same data [no emergence
date], 07-471, ex Acharia stimulea on
Quercus rubra (1/USNM); same data,
07-489, ex Adoneta spinuloides on Carya
glabra (?USNM); same locality, col-
lected 27.VII.2007, J. Lill et al. 07-499,
ex Euclea delphinii on Quercus rubra
(1/USNM); same locality, collected 3.
VIII.2007, emerged 21.VIII.2007, J. Lill
et al. 07-558, ex Prolimacodes badia on
Fagus grandifolia (1/USNM); same
data, 07-561 (?USNM); same data, 07-
563 (?USNM); same data, 07-564 (1/
USNM); same locality, collected 14.
VIII.2007, emerged 1.VI.2008, J. Lill et al.
07-581, same host (1/USNM); same lo-
cality, collected 31.VII.2008, emerged 11.
VII.2008, J. Lill et al. 08-528, ex Acharia
stimulea on Nyssa sylvatica (3/USNM);
same data but emerged 26.VIII.2008, 08-
536, ex Acharia stimulea on Carya glabra
(2/USNM); same locality, emerged 2.
VIII.2007, 07-996, ex Euclea delphinii on
Prunus serotina,(3/?USNM); same
data, 07-995, (?USNM); same data, 07-
997, (1/USNM); same locality, [No
date], 07-993, ex Euclea delphinii on
Nyssa sylvatica (2/2?USNM); same
data, emerged 2.VIII.2007, 07-999, ex
Euclea delphinii on Quercus rubra, (1/
USNM); same data, 07-994, ex Euclea
delphinii on Fagus grandifolia (2/
USNM); same data, 07-991, ex Acharia
stimulea on Quercus alba (1/USNM);
same locality, collected 3.VIII.2007,
emerged 21.VIII.2007, J. Lill et al. 07-
526, ex Acharia stimulea on Fagus gran-
difolia (1/USNM); same data, 07-522,
ex Acharia stimulea on Prunus serotina
(1/USNM); same locality, collected 5.
VIII.2008, emerged 18.VIII.2008, J. Lill
et al. 08-632, ex Acharia stimulea on
Nyssa sylvatica (3/USNM); same lo-
cality, collected 31.VII.2008, emerged 13.
VIII.2008, J. Lill et al. 08-527, ex Acharia
stimulea on Nyssa sylvatica (1/USNM);
same locality, collected 18.VII.2007,
emerged 2.VIII.2007, J. Lill et al. 07-998,
ex Euclea delphinii on Quercus rubra (1/
USNM); same locality, 23.VIII. 2007,
emerged 1.VI.2008, J. Lill et al. 07-672, ex
Prolimacodes badia on Quercus rubra
(1/USNM); same locality, collected 3.
VIII. 2007, emerged 21.VIII.2007, J. Lill
et al. 07-524, ex Acharia stimulea on Fa -
gus grandifolia (1/USNM); same local-
ity, collected 14.VII.2008, emerged 1.
VIII.2008, J. Lill et al. 08-226, ex Euclea
delphinii on Nyssa sylvatica (/?
USNM); same data, 08-234 (2/USNM);
same locality, collected 28.VII.2008,
emerged 11.VIII. 2008, J. Lill et al. 08-
328, ex Acharia stimulea on Quercus
rubra (3/USNM); same locality and
collection date, emerged 19.VIII.2008, J.
Lill et al. 08-326, ex Acharia stimulea on
Nyssa sylvatica (4/USNM); same lo-
cality, collected 31.VII.2008, emerged 18.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
76

VIII.2008, J. Lill et al. 08-525, same host
(1/USNM); same locality, collected 31.
VII. 2008, emerged 11.VII.2008, J. Lill
et al. 08-510, ex Acharia stimulea on Pru-
nus serotina (3/?USNM). Plummers
Island, collected 18.VIII.2006, J. Lill et al.
06-ak, ex Acharia stimulea on Asimina
triloba (/USNM); same data, but
emerged 25.VIII.2006, J. Lill et al. 06-am
(6/USNM); same data, but emerged 28.
VIII.2006, 06-ab, ex Acharia stimulea on
Quercus alba (/USNM; 6 pupae beneath
host, 4 pupae on leaf); same locality, col-
lected 18.VIII.2006, J. Lill et al. 06-aq-V,
ex Acharia stimulea on Quercus rubra
(?USNM); same data, but emerged 3.
IX.2006, 06-ai, ex Acharia stimulea on
Asimina triloba (? USNM; specimen
damaged); same data [no emergence date],
06-ae-II, ex Acharia stimulea on Asimina
triloba (/?USNM); same data, 06-an-
II, ex. Acharia stimulea on Quercus
rubra (/USNM); collected 3.VII.2007,
emerged 17.VII.2007, J. Lill et al. 07-
221, ex Lithacodes fasciola on Carya
glabra (/USNM); same data, 07-222
(?USNM). Prince Georges County:
Patuxent NWR, collected 21.VII.2005,
emerged 9.VIII.2005, J. Lill et al. 05-
501, ex Lithacodes fasciola on Fagus
grandifolia,(2/?USNM); same lo-
cality, collected 17.VIII.2005, emerged 1.
IX.2005, J. Lill et al. 05-730, ex Pack-
ardia geminata on Fagus grandifolia,(/
USNM); same locality, collected 17.
VIII.2005, emerged 27.VIII.2005, J. Lill
et al. 05-742, ex Lithacodes fasciola on
Fagus grandifolia,(/USNM); same lo-
cality, collected 17.VIII.2005, emerged
30.VIII.2005, J. Lill et al. 05-761, ex
Lithacodes fasciola on Fagus grandifolia,
(2/USNM (5 pupae)); same locality,
collected 17.VIII.2005, emerged 10.
VI.2006, J. Lill et al. 05-779, ex Proli-
macodes badia on Fagus grandifolia
(/USNM); same locality, collected 1.
IX.2005, emerged 20.IX.2005, J. Lill
et al. 05-899, ex Lithacodes fasciola on
Fagus grandifolia (3/?USNM); same
locality, collected 6.IX.2005, emerged
10.VI.2006, J. Lill et al. 05-916, ex
Lithacodes fasciola on Fagus grandifolia
(2?USNM (1 pupa)); same data, but
emerged 11.VI.2006, 05-921 (2/USNM);
same locality, collected 11.VII.2006, J. Lill
et al. 06-252, ex Lithacodes fasciola on
Fagus grandifolia (?USNM); same data,
emerged 31.VII.2006, 06-260 (/USNM);
same data, 06-268 (/USNM); same data,
emerged 1.VIII.2006, 06-274, ex Limaco-
didae on Carya glabra (/USNM);
collected 30.VIII.2007, emerged 1.
IX.2007, J. Lill et al. 07-841, ex Proli-
macodes badia on Fagus grandifolia,
(1/USNM); same locality, collected 8.
VIII.2007, emerged 16. X .2007, J. Lill
et al. 07-611, ex Lithacodes fasciola on
Quercus alba,(
/?USNM); same
data, no emergence, 07-608, same host,
(4/?USNM); same locality, collected
14.IX. 2007, emerged 16.X.2007 J. Lill
et al. 07-957, ex Isa textula on Quercus
alba (2/?USNM).
North American Records:
Platyplectrus americana (Girault) (Figs.
105–113): Euplectromorpha americana
Girault, 1917: 114. Holotype ?,USNM
(examined).
Based on the significant new rearing
data (Table 1), Platyplectrus (and most
eulophid parasitoids) preferentially attack
early limacodid instars. Platyplectrus
americana is a solitary/gregarious ecto-
parasitoid with a single wasp emerging
from a host 41% of the time, two emerging
26% of the time, three (12%), four (10%),
five (4%), six (4%), and greater than six
(4%). The position for larval feeding is as
previously described for Pl. natadae (Gadd
et al. 1946). Pupation occurs beneath the
host cadaver (as in Fig. 98; cadaver re-
moved), and the pupa and cadaver are
adhered to the substrate with the cadaver
providing protection for the parasitoid
VOLUME 114, NUMBER 1
77

pupa. No in situ observations for ovipo-
sition are yet available for Pl. americana.
Distribution: Eastern U.S.A.
Limacodidae: Previously reported from
Packardia geminata (Gates and Burks
2003). Newly recorded hosts (Table 1)
include: Ac.stimulea*, Ad.spinuloides*,
E.delphinii*, I.textula*, Li.fasciola*,
N.nasoni*, Ph.pithecium*, Pr.badia*,
and Tortricidia sp.*
Pediobius Walker (Chalcidoidea:
Eulophidae: Entedoninae)
(Figs. 95, 97–103)
Diagnosis.—
The following combina-
tion of features is diagnostic: propodeum
medially with two subparallel carinae (Fig.
95), with distinct plicae; petiole ventrally
pointed; notauli incomplete (Fig. 101);
antenna with three funiculars and two
claval segments (Figs. 102–103).
Fauna.—
Pediobius is a large genus of
277 species worldwide with 36 known
from the Nearctic (Noyes 2003). New
World species of Pediobius range from
central Canada to Argentina. Extralimitally,
Cock (1987) summarized the Pediobius as-
sociated with limacodids in Southeast Asia:
Pe. imbreus (Walker) (as detrimentosus
(Gahan)), Pe. elasmi (Ashmead), and Pe.
ptychomyiae (Ferrie
`re). They are recorded
from a tachinid, Apanteles (Ap. sp. and Ap.
parasae; Braconidae), and Pa.philepida
Holloway (Limacodidae), respectively.
Noyes (2003) summarized additional
host records including primary lepi-
dopteran hosts and secondary dipteran/
hymenopteran hosts.
Biology.—
Most commonly associated
with species of Lepidoptera, Coleoptera,
Diptera, and Hymenoptera as primary or
secondary parasitoids (Peck 1985). Some
species are known from spider egg sacs
where they may act as secondary parasitoids
(Peck 1985). Limacodid host associations in
North America are newly reported herein.
Literature.—
Burks (1966) published a
key to 23 species known from North
America, 24 were treated in a catalog
(Burks 1979), and Peck (1985) provided
a key to 32 species from North America.
North American Records:
Pediobius crassicornis (Thomson) (Figs.
95, 97–103): Pleurotropis crassicornis
Thomson, 1878: 255. Holotype /,
MZLU (not examined).
This Holarctic species has a broad host
range, but surprisingly, it had not been
reported as associated with limacodids
previously (Noyes 2003). It has, however,
been reported previously from primary
parasitoids (Ichneumonidae: H. fugiti-
vus, Tachinidae: Com. concinnata)that
are reported from limacodids herein. It
is most commonly recovered as a pri-
mary (Lepidoptera) or secondary para-
sitoid of hymenopterans through lepi-
dopterans. From the rearing results
reported herein, Pe.crassicornis com-
monly parasitize Pl. americana (Fig. 98).
When attacking Pl. americana,adultPe .
crassicornis emerge from the anterior end
of the pupa and escape the limacodid
cadaver laterally or dorsally, depending
upon the degree of adherence of the li-
macodid cadaver to the substrate. In the
latter case, circular emergence holes are
chewed through the dorsum of the li-
macodid when the margins of the cadaver
are tightly appressed to the substrate. It is
possible that Pe. crassicornis also parasit-
izes Al. lilli. There are a few specimens
associated only with the remains of lima-
codid or other species of primary parasit-
oid (ichneumonoids), but these records
are suspect due to the nature of the
emergence hole or the lack of pupal
remnants reminiscent of Pe.crassi-
cornis. In the former instance, the re-
mains of Ac. stimulea are mummified
and a very large emergence hole is
chewed posterolaterally, roughly equal to
4–5 body widths of Pe. crassicornis,and
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
78

no pupal remnants are visible inside. In
the latter (same limacodid host), micro-
gastrine cocoons associated with Pe.
crassicornis areemptyandappeartohave
emerged normally, but no primary host
remains are associated with them. There
is a series of Co. schaffneri in the USNM
(no label data) bearing a handwritten
identification of, “On Tortricidia testacea
Dyar ‘97” that has several larvae with
cocoons of Cotesia adhered. A series of
five (formerly seven as two are missing)
co-mounted Pe.crassicornis appear to
correspond with 10 cocoons of Cotesia,
eight of which bear emergence holes lat-
erally and contain pupal remains. Ped-
iobius crassicornis can be separated from
other Pediobius by the following combi-
nation of characters: occipital carina pres-
ent, frons coarsely reticulate (Fig. 100),
male with three funiculars (Fig. 103),
scutellum smooth medially (Fig. 95),
basal gastral tergum alutaceous.
Distribution: Widespread across North
America from the west to east coasts
into Canada in the north. Also found in
northern Europe and Japan (Peck
1985, Noyes 2003).
Limacodidae: Associated with the fol-
lowing species but appears to be acting
as a hyperparasitoid in most instances:
Ac. stimulea,Ad. spinuloides,N.nasoni,
E.delphinii,I.textula, and Li.fasciola.
Natada nasoni may serve as a host in
one instance as the larva is mummified
and an emergence hole is visible post-
erolaterally, roughly equal to two body
widths of Pe. crassicornis, and pupal
remnants are visible inside.
Other hosts: The numerous hosts are
listed by Noyes (2003) and span 22
families across the following four insect
orders: Diptera (2), Coleoptera (2),
Lepidoptera (10), and Hymenoptera
(8). Herein it is newly recorded as a
parasitoid of Pl. americana*, al-
though it is possible that it is attacking
additional taxa (see species treatment
above).
Psychophagus Thomson (Chalcidoidea:
Pteromalidae: Pteromalinae)
(Figs. 87–93)
Diagnosis.—
Clypeus shallowly bi-
lobed apically (Fig. 88); pedicel ;2.0X
as long as broad, antennal formula 11263
(Fig. 89); propodeum reticulate-rugose,
with median carina incomplete (Fig. 90);
gaster subcircular to short-ovate (Fig. 91).
Fauna.—
Psychophagus contains the
single Holarctic species, P. omnivorus.
Biology.—
This genus has an incred-
ibly broad host range and may act as a
primary or secondary parasitoid (Noyes
2003) across four insect orders and 22
families. Most hosts are pupae of Lep-
idoptera, Hymenoptera, or Diptera (or
their dipteran or hymenopteran parasit-
oids). There is one record of a single
species of Limacodidae being attacked in
North America among numerous addi-
tional host records (see below).
Literature.—
Aside from original de-
scriptive work (Walker 1835) and treat-
ment by Graham (1969), Noyes (2003)
provides citations of the bulk of the lit-
erature (particularly host records) for this
species.
North American Records:
Psychophagus omnivorus (Walker) (Figs.
87–93): Pteromalus omnivorus Walker,
1835: 204. Lectotype ?, BMNH, de-
signated by Graham (1969: 473) (not
examined).
The sole limacodid host, the exotic M.
flavescens, was reported by Schaffner and
Griswold (1934) and Schaffner (1959).
The former recorded this wasp as often
hyperparasitic (only other parasitoid re-
corded therein was Com.concinnata),
with four to 90 emerging per host and
distributed in northeastern North America.
The latter reported four specimens of
VOLUME 114, NUMBER 1
79

P. omnivorus from Boston, Massachusetts
as a gregarious parasitoid emerging from a
limacodid pupa.
Distribution: Holarctic. In North America,
primarily eastern U.S.A. and Canada.
Limacodidae: M. flavescens.
Other hosts: Primary Parasitoid: Sco-
lytidae (Coleoptera); Diprionidae,
Pamphiliidae, Tenthredinidae (Hy-
menoptera); Arctiidae, Coleophoridae,
Lasiocampidae, Lycaenidae, Lyman-
triidae, Noctuidae, Notodontidae,
Nymphalidae, Oecophoridae, Pieridae,
Pyralidae, Saturniidae, Tortricidae
(Lepidoptera). Hyperparasitoid: Sar-
cophagidae, Tachinidae (Diptera);
Braconidae, Ichneumonidae (Hyme-
noptera) (see Noyes 2003).
Trichogramma Westwood
(Chalcidoidea: Trichogrammatidae)
(Figs. 94, 96)
Diagnosis.—
The family is recognized
by the three-segmented tarsi (Fig. 94) and
fore wing setation arranged in lines (Figs.
96). Trichogramma are recognized by
broad fore wing with sigmoid venation
and distinctive Rs1 setal track (Fig. 96),
among other features not included herein
(Pinto 1998).
Fauna.—
Trichogramma is the largest
genus in the family with ;180 species
worldwide and 68 known from North
America (Pinto 1998). They are used
extensively in biological control programs
of numerous species of pestiferous Lepi-
doptera in natu ral and agro eco systems
(S. Smith 1996). A comprehensive regional
monograph is available (Pinto 1998).
Biology.—
Species of Trichogrammati-
dae are solitary or gregarious parasitoids of
insect eggs in several orders (Hemiptera,
Coleoptera, Lepidoptera, Diptera, etc.). A
single limacodid host is recorded (see be-
low) for North America. Three exralimital
records of Trichogramma spp. (Hoong and
Hoh 1992, Wei 1996) and Tr. papilionis
(Nagarkatti) on Darna pallivitta in Hawaii
(Conant et al. 2006) have been reported.
There are also a few extralimital records of
Trichogrammatoidea thoseae Nagaraja
attacking limacodids in three genera as
summarized by Cock (1987). Most Tricho-
gramma species are primarily solitary or
gregarious endoparasitoids of eggs of Lepi-
doptera. The number successfully com-
pleting development in a single egg de-
pends upon host size and the proximity of
host eggs (Pinto 1998; references therein),
and females inject a developmental ar-
restant during oviposition (Strand 1986).
Literature.—
Keytospecies(Pinto1998).
North American Records:
Trichogramma minutum Riley (Figs.
94, 96): Trichogramma minuta Riley,
1871: 157–158. Neotype ?,USNM,
designated by Pinto et al. (1978)
(examined).
Pinto (1998) considers this species to be a
“complex of populations whose taxonomic
status remains under investigation.” Accord-
ing to Pinto (1998), Peck’s (1963) recogni-
tion of only three Nearctic species of
Trichogramma, and following Girault’s
(1911) interpretation of Tr. min utu m,re-
sulted in recognizing ;150 host species
in 33 families for this species alone.
Given that this vast number of host re-
cords and associated research on Tri-
chogramma species provided minimal
voucher specimen documentation, the
bulk of this information is unusable at
the species level. Thus, the previously
published limacodid rearing record cited
below is suspect. Diagnostic genitalic char-
acters for species separation are reported
in Pinto (1998).
Distribution: U.S.A., east of 110°
longitude (Pinto 1998).
Limacodidae: Monema flavescens
(Collins 1933).
Other hosts: Pinto (1998) recorded,
based on specimens examined and
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80

those reared from field-collected hosts
(not literature records), a total of 49
host records from three orders (Hy-
menoptera (1), Lepidoptera (45), and
Neuroptera (3)). None was from Li-
macodidae.
Trichogramma sp.:
Distribution: Eastern U.S.A., minimally.
Limacodidae: E.delphinii*on Q.alba.
Cotesia Cameron (Ichneumonoidea:
Braconidae: Microgastrinae)
(Figs. 64–69)
Diagnosis.—
The combination of the
following features is diagnostic (see
Whitfield 1997): fore wing with vein r-m
absent so that no small “areolet” (tiny sec-
ond submarginal cell) is present (Fig. 65);
propodeum with rugose sculpture and at
least some indication of a percurrent medial
carina (Fig. 66); anterior two metasomal
terga (and sometimes anterior portions of
third) sculptured and relatively broad and
subrectangular (Fig. 66); ovipositor and
sheaths relatively short, with only a few
small setae at tip of sheath (Fig. 67).
Fauna.—
Cotesia is a huge genus
containing several hundred described
species worldwide, and many more un-
described, although it is best represented
in temperate regions. It is one of the
largest and most economically important
genera of Microgastrinae, with hosts in
many lepidopteran (especially macro-
plepidopteran) families in both natural
and agricultural systems. Most available
revisions of Cotesia are strictly regional,
and Muesebeck (1920) is the most recent
comprehensive North American treat-
ment. Extralimitally, there are two records
of Apanteles spp. recorded from Ph.hip-
parchia Cramer and Ac. (as Episibine)
intensa Dyar (Genty et al. 1978) in the
Neotropical Region; these are most cer-
tainly Cotesia spp. based on the images of
adults and cocoons in Genty et al. (1978).
Biology.—
Most Cotesia species are
gregarious endoparasitoids of macro-
lepidopterans from many families, with
only a few microlepidopterans (e.g., plu-
tellids) among their hosts. A minority of
species develop as solitary parasitoids. Like
other microgastrines, species of Cotesia
typically oviposit into early-instar host lar-
vae, emerging from later instar larvae or
prepupae to spin cocoon(s) on or near the
host, which soon dies. A few species are
capable of ovipositing into eggs (Ruberson
and Whitfield 1996). The immune system
of the host caterpillar is compromised in
part by the introduction, along with the
eggs, of symbiotic polydnaviruses by the
female wasp from her reproductive tract
(Whitfield and Asgari 2003).
Literature.—
Muesebeck (1920) is the
only comprehensive North American treat-
ment of Cotesia.
North American Records:
Cotesia empretiae (Viereck) (Figs. 64–67):
Apanteles empretiae Viereck, 1913:
562. Holotype /, USNM (examined).
Distinctive among Cotesia species in
having the body length <2mm,first
metasomal tergite width subequal an-
teriorly and posteriorly (Fig. 66; most
Cotesia have tergite broader posteri-
orly), the third metasomal tergite with
sculpturing over anterior half or two
thirds, especially medially (Fig. 66),
and the hypopygium sharply angled
apically and with ovipositor sheaths
strongly exserted, often visible in dor-
sal view (Fig. 67). The gregarious lar-
vae emerge to spin (separately) 5–15
whitish to pale tan woolly cocoons
dorsally (sometimes laterally) on the
caterpillars.
Distribution: U.S.A., especially south-
eastern.
Limacodidae: E.delphinii,Li.fasciola,
Pa.chloris,andAc.stimulea (Marsh
1979), Ap.biguttata,andPa .indetermina
(Whitfield et al. 1999).
VOLUME 114, NUMBER 1
81

Cotesia phobetri (Rowher) (Fig. 68):
Apanteles phobetri Rohwer, 1915: 228.
Holotype /, USNM (examined).
This species (Fig. 68) differs from both
Co. empretiae and Co. schaffneri in hav-
ing the third metasomal tergite entirely
unsculptured; from Co. empretiae it also
differsinhavingthefirstmetasomalter-
gite broadening posteriorly, and from Co.
schaffneri it differs in having the hind
tibiae and tarsi almost entirely pale orange
brown. The gregarious pale buff cocoons
are spun separately and positioned dor-
sally on the host caterpillar.
Distribution: Eastern U.S.A. (contribu-
tion from possibly non-conspecific, non-
limacodid rearings cannot be ruled out).
Limacodidae: Ph.pithecium (Rowher
1915).
Other hosts: Halysidota tesselaris Smith
and Abbot (Arctiidae) (Muesebeck
1920) (possibly not conspecific Co.
phobetri). Several other series reared
from Acronicta sp. (Noctuidae) and
other arctiids have also been identified
as this species, perhaps mistakenly.
Cotesia schaffneri (Muesebeck) (Fig. 69):
Apanteles schaffneri Muesebeck, 1931: 4.
Holotype /, USNM (not examined).
This species is unique among the three
Cotesia recorded herein in having the hind
tarsi strongly bicolored: the proximal and
distal tarsomeres being mostly blackish,
whereas the third and fourth (and sometimes
also second) tarsomeres are pale orange
yellow. Unlike in Co.empretiae,thegre-
garious pale buff cocoons are spun sepa-
rately (but appressed together; individually
immobile) on the back of the host cater-
pillar. The parasitized cocoons with the
emergence holes on the side are whitish.
Distribution: Nearctic ranging from
mid-Atlantic states and Texas in U.S.
A.; thus it may be distributed broadly
across the eastern and southern U.S.A.
Limacodidae: Unidentified limacodid
(“cochlidiid”) (Muesebeck 1931),
To.pallida H.-S., To.testacea,Pr.
badia,andE.delphinii (specimens in
the USNM with host remains).
Ascogaster (Ichneumonoidea:
Braconidae: Cheloninae)
(Figs. 48, 60–62)
Diagnosis.—
Carapace without pair
of complete transverse grooves, at most
grooves present laterally but absent dor-
sally (Fig. 48); fore wing vein (RS+M)
a present; eyes glabrous or virtually so
(Fig. 60); ocelli configured in shape of
isosceles triangle (Fig. 61); scutellar disc
sculptured (Fig. 62).
Fauna.—
Described species of As-
cogaster occur in all zoogeographic re-
gions, but only 12 species are known
from the Nearctic (Shaw 1997a). Yu et al.
(2005) listed four species in Ascogaster
placed currently in Leptodrepana Shaw.
As pointed out in Shaw (1997a), richness
is higher in temperate areas than tropical
areas of the New World, but richness is
lower in the Nearctic than in all other
regions except the Neotropical (Yu et al.
2005).
Biology.—
As is the case for all chelo-
nines, species of Ascogaster are solitary
koinobiont egg-larval endoparasitoids of
lepidopterans (Shaw 1997a). Host asso-
ciations reported for Ascogaster include
23 families of Lepidoptera, including
Limacodidae; most hosts are in Tortricidae
(Yu et al. 2005). Host associations reported
for species of Ascogaster in the Nearctic
Region include species in Anthomyiidae,
Clusiidae (Diptera), Curculionidae (Co-
leoptera), Cynipidae, Tenthredinidae
(Hymenoptera), Blastodacnidae, Depres-
sariidae, Gelechiidae, Geometridae, Lima-
codidae, Pyralidae, Tineidae, Tortricidae,
and Yponomeutidae (Lepidoptera) (Yu
et al. 2005). Non-lepidopteran host re-
cords are erroneous since all confirmed
hosts for chelonines are lepidopterans
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82

(Shaw 1997a, S. Shaw pers. comm.). Of
the remaining 138 putative lepidopteran
hosts, only one species of Limacodidae,
M.flavescens, has been reported (Wilcox
1918).
Literature.—
Research relevant to this
study published since Yu et al. (2005) in-
cludes As. quadridentata (Wesmael) reared
from the tortricids Lobesia botrana
(Denis and Schiffermu
¨ller) (Carlos et al.
2006) and Spilonota ocellana (Denis and
Schiffermu
¨ller) (Pluciennik and Olszak
2010). The aforementioned hosts were
reported previously in multiple publica-
tions (Yu et al. 2005). Nazemi et al. (2008)
reported As.quadridentata from the tor-
tricid Pammene amygdalana (Duponchel),
an inquiline of the cynipid Andricus
cecconi Kieffer.
North American Records:
Ascogaster quadridentata (Wesmael)
(Figs. 48, 60–62): Ascogaster quad-
ridentatus Wesmael 1835: 237. Lectotype
/, IRSNB, designated by Shaw (1983)
(not examined).
This species can be differentiated from
all Nearctic species of Ascogaster, except
As. shawi Marsh, using the key in Shaw
(1983). It can be differentiated from As.
shawi using the features that differentiate
As.quadridentata and As.provancheri
Dalla Torre in the key by Shaw (1983).
Distribution: All zoogeographic re-
gions except the Afrotropical, although
it has been introduced into South Africa
and several other countries to control
orchard pests, particularly Cydia po-
monella (Linnaeus). It ranges in the
Nearctic Region from Maine south to
Georgia and west to California (Yu
et al. 2005).
Limacodidae: Monema flavescens re-
ported but host plant not mentioned
(Wilcox 1918). RRK considers this
record possibly valid but has not verified
this identification. In terms of rearing
protocol, Wilcox (1918) mentioned only
that cocoons were “placed in rearing
boxes,” so the record should be con-
firmed through rearing from a host
isolated individually. This host species
is likely no longer present in North
America based on subsequent searches
for it (M. Epstein pers. comm.).
Other hosts: Reported from at least
48 species in six lepidopteran fami-
lies (Yu et al. 2005).
Triraphis Ruthe (Ichneumonoidea:
Braconidae: Rogadinae)
(Figs. 46–47, 49–59)
Diagnosis.—
Third maxillary palpo-
mere not enlarged and flattened, similar
in size and shape to distal palpomeres;
second labial palpomere not enlarged
and vesiculate, similar in size and shape
to distal palpomeres; basal carina of pro-
podeum diverging basally to apically (Figs.
53, 55, 58), not complete to apical margin
as single mesal carina; inner margin of
metatibia with fringe of flattened setae (Fig.
59); metatibial spurs straight and setiferous;
pro- and mesotarsus with tarsomeres 2–4
longer than wide (Figs. 50–51, 56); tarsal
claws with basal lobe or tooth (Fig. 47);
fore wing first submarginal cell uniformly
setose, without glabrous patch; fore wing
vein 1cu-a not angled as obliquely as fore
wing 2CUa (Fig. 56); hind wing M+CU
equal to or longer than 1M (as in Fig.
45); metasoma with T1–T4 not car-
apacelike and covering other terga (Figs.
50–51, 56); T2 without triangular area
anteromesally; transverse groove between
T2+T3 distinct (Figs. 52, 57).
Fauna.—
Described species of Triraphis
are known from the Nearctic, Neotropical,
Oriental, and Palearctic regions (Yu et al.
2005). There are three Nearctic species
currently placed in this genus, including
T.eupoeyiae (Ashmead), new combi-
nation (see below), but most if not
all New World species currently in Rogas
VOLUME 114, NUMBER 1
83

belong in Triraphis (Valerio 2006, Zaldı
´var-
Rivero
´n et al. 2008). Neither Triraphis nor
Rogas has been revised for the New
World; New World richness is estimated
at 100–150 species (Shaw 1997b).
Biology.—
As is the case for all roga-
dines, species of Triraphis are koinobiont
endoparasitoids of lepidopteran larvae and
pupate within the mummified remains of
their hosts (Shaw 1997b). Reported host
associations are larvae in Dalceridae,
Limacodidae, Lycaenidae, Lymantriidae,
Megalopygidae, Riodinidae, Tortricidae,
and Zygaenidae (all Lepidoptera) (Yu
et al. 2005, Zalvidar-Riveron et al. 2008).
A single record from Cynipidae (Hyme-
noptera) is obviously erroneous since all
confirmed hosts for rogadines are lepi-
dopterans (Shaw 1997b, S. Shaw pers.
comm.). Other records of questionable
validity are the Palearctic species T.
tricolor (Wesmael) from the tortricids
Archips rosana L. (Tobias 1976) and Tor -
trix testudinana Hu
¨bner (Reinhard 1865)
and the Oriental species T.sichuanensis
Chen and He from the lymantriid Eu-
proctis bipunctapex (Hampson) (Chen
and He 1997). Genera in Dalceridae,
Lycaenidae, Megalopygidae, and Riodi-
nidae were reported in Zaldı
´var-Rivero
´n
et al. (2008) as containing hosts for spe-
cies of Triraphis; particular parasitoid and
host species were not listed. The basis for
those records (e.g., rearing protocol) is
unknown (i.e., cited as Valerio and Shaw
in prep.), although they were noted as
confirmed by a rogadine specialist (i.e.,
S. R. Shaw, University of Wyoming-
Laramie). Thus, in terms of published
records, species of Triraphis are known
from hosts in two monophyletic groups
(i.e., Limacodidae, Zygaenidae) (Wahlberg
et al. 2005, Niehuis et al. 2006).
Literature.—
Aside from literature
reported in Yu et al. (2005), Zaldı
´var-
Rivero
´n et al. (2004) examined venom
gland morphology for Rogadinae, including
Trirap his, and conducted a cladistic
analysis based on characters of the
venom gland. Quick and Shaw (2005)
discussed mummy characteristics for
Tri rap his;T.tricolor was the only species
mentioned. Valerio (2006) transferred T.
brasiliensis (Sze
´pligeti) and T.mac-
ulipennis (Sze
´pligeti) from Rogas,but
Papp (2004) had already transferred those
species to Triraphis. Valerio (2006) also
indicated T.pulchricornis (Sze
´pligeti) as
a new combination transferred from Rogas,
but Papp (2004) had transferred pul-
chricornis from Rogas to Aleiodes
Wesmael. Additionally, Valerio (2006)
mentioned eight species of Rogas as be-
longing in Aleiodes, but all had been
transferred to Aleiodes previously (Shaw
et al. 1997, Marsh and Shaw 1998, Shaw
et al. 1998, Fortier and Shaw 1999).
Zalvidar-Riveron et al. (2008) carried
out cladistic analyses for Rogadinae,
including Trirap his, based on DNA se-
quence data and assessed the evolution
of host range and mummy characteris-
tics. Kula et al. (2010) reported the first
host records for T.discoideus
North American Records:
Triraphis discoideus (Cresson) (Fig. 46–
47, 49, 56–59): Aleiodes discoideus Cres-
son, 1869: 380. Holotype /, ANSP (not
examined).
The body is irregularly yellow and brown
and the stigma is usually (96.6%) brown
with at most a small yellow area where it
meets vein R1a (Fig. 56).
Distribution: Nearctic Region, rang-
ing from Massachusetts to Florida and
west to Illinois (Yu et al. 2005).
Limacodidae: Reported from 10 spe-
cies of Limacodidae sampled collec-
tively from six hardwood tree species
in the District of Columbia and
Maryland (Table 1) (Kula et al. 2010).
See Kula et al. (2010) for additional
information on the natural history of T.
discoideus.
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84

Other hosts: None reported.
Triraphis eupoeyiae (Ashmead), new
combination (Fig. 50): Pelecystoma
eupoeyiae Ashmead, 1897: 113. Holo-
type /, USNM (examined).
The body is primarily orange but with at
least the lateral lobes of the mesoscutum
brown, and the stigma is yellow with a
brown spot directly above vein r (Fig. 50).
This species fits Triraphis sensu van
Achterberg (1991) except the occipital
carina is complete dorsally, and the frons
is rugose in the holotype. Additionally,
several relative features in the diagnosis
for Triraphis sensu van Achterberg (1991)
are difficult to interpret (e.g., size and
shape of basal lobe of tarsal claw). It is
transferred to Trirap his despite the ex-
ceptions and problems with interpretation
because it shares more features with
species in that genus than species in Ro-
gas sensu van Achterberg (1991).
Distribution: Known only from Flor-
ida prior to this research, but a speci-
men from Cuba is in the USNM.
Limacodidae: Ashmead (1897) reported
the holotype as reared from Ala.slos-
soniae, but a host cadaver is not asso-
ciated with the holotype. RRK considers
this record possibly valid, but it requires
confirmation through rearing from
a host isolated individually. The cadaver
associated with this rearing is deposited
in the USNM. In reporting on the biology
of Ala. slossoniae, Dyar (1897) figured
a larva parasitized by T. eu p oeyia e ,in-
dicating that it becomes, “bright red
(Plate V, fig. 23) and hardens.” (p. 125).
Other hosts: Label data associated with
the specimen from Cuba indicates that
it was reared from Leptotes cassius
(Cramer) (Lycaenidae); host remains
are associated with the specimen.
Triraphis harrisinae (Ashmead) (Fig.
51–55): Rhogas harrisinae Ashmead
(1888) 1889: 632. Holotype ?,USNM
(examined).
Thebodyisentirelyyellowtoorange,and
the fore wing stigma is yellow with at most
a light brown spot distad vein r (Fig. 51).
Distribution: Nearctic and Neotropi-
cal regions, ranging from New Jersey
to Florida west to Missouri and south
to Costa Rica. It also occurs in Cuba
(Yu et al. 2005).
Limacodidae: Harrison (1963) reported
T.harrisinae from Ac.apicalis (Dyar)
on banana; host larvae were isolated in-
dividually, and RRK considers this re-
cord valid. Triraphis harrisinae was not
reared from a congener, Ac. stimulea,du-
ring the course of this research. Marsh
(1979) reported T.harrisinae from Li.
fasciola, but there are no specimens of
T.harrisinae at the USNM purportedly
reared from Li.fascicola.Thisrecord
is possibly valid, but T.harrisinae was
not reared from Li.fasciola during the
course of this research.
Other hosts: Riley and Howard (1890)
first reported it as a parasitoid of Harri-
sina americana (Gue
´rin-Me
´neville)
(Zygaenidae). Additionally, specimens
of T.harrisinae reared from H.amer-
icana and undetermined Acoloithus
Clemens (Zygaenidae) larvae collected
in Florida and Illinois were used to start
a colony at the University of California-
Riverside (Smith et al. 1955). Speci-
mens from that colony were released
in California to control H.metallica
Stretch and apparently established in
San Diego (Clausen 1956). See Kula
et al. (2010) and references in Yu et al.
(2005) for additional information on
the natural history of T.harrisinae.
Mesochorus Gravenhorst
(Ichneumonoidea: Ichneumonidae:
Mesochorinae)
(Figs. 19–22)
Diagnosis.—
Mesochorus can be rec-
ognized by the following combination of
VOLUME 114, NUMBER 1
85

characters: areolet of fore wing closed,
large and rhombic (Fig. 20); spiracle of
metasomal segment 1 near or just behind
middle (Fig. 21); glymma large and deep
(Fig. 21); ovipositor needlelike, about 1.5X
as long as metasomal apical depth and
without dorsal subapical notch (Fig. 22);
female hypopygium prominent and trian-
gular in lateral view (Fig. 22); male gon-
oforceps produced into elongate processes.
Fauna.—
Species of Mesochorus occur
in all zoogeographic regions (Yu et al.
2005), with about 695 species worldwide,
108 of which occur in the Nearctic Re-
gion. Dasch (1971) revised the Nearctic
Mesochorinae.
Biology.—
Most observations and rear-
ing records indicated that mesochorines
are obligate hyperparasitoids of endo-
parasitic Ichneumonoidea (and, rarely,
Tachinidae) which parasitize primary
hosts of larval Lepidoptera, Symphyta,
and Coleoptera, and nymphal and adult
Hemiptera and Psocoptera (Carlson 1979a).
Based upon rearing records, many spe-
cies have a wide host range (Dasch 1971,
Carlson 1979a). Ectoparasitic ichneumo-
nids can be utilized as well, based upon
rearings of Mesochorus from Phytodietus
and ?Oedemopsis hosts by M. R. Shaw
(Gauld and Bolton 1988: 17). Although
there are some records of mesochorines as
primary parasitoids (Dasch 1971, Yu et al.
2005), these are likely specimens reared
from undetected primary parasitoids (Wahl
1993a). The pitfalls and deficiencies of
rearing records were ably discussed by
Shaw (1990: 453–455) and are relevant
to this matter.
Literature.—
The Nearctic Mesochorus
fauna may be identified using Dasch
(1971). Aside from literature reported
in Yu et al. (2005), Reis Fernandes et al.
(2010) reported two undetermined spe-
cies of Mesochorus: one from an un-
determined species of Hypomicrogaster
Ashmead (Braconidae: Microgastrinae)
through an undetermined species of
Olethreutinae (Lepidoptera: Tortricidae),
and the other through an undetermined
species of Noctuidae (Lepidoptera) pre-
sumably parasitized by an undetermined
species of Aleiodes Wesmael (Braconidae:
Rogadinae).
North American Records:
Mesochorus discitergus (Say) (Figs. 19–22):
Cryptus discitergus Say, 1835: 231. Type /
(destroyed, Townes et al. 1965).
There are numerous extremely similar
species of Mesochorus in the Nearctic,
and discitergus cannot be separated from
them by a short diagnosis. Dasch (1971)
divided the 96 species that he treated into
17 species groups with this species
placed in the discitergus group with two
other species: M.americanus Cresson and
M.acuminatus Thompson. Dasch pro-
vided information that may be used to
separate discitergus from other species of
Mesochorus, but the characters are diffi-
cult to interpret by non-experts without
access to a large collection. It remains
necessary to confirm species identifications
by sending them to an expert with access to
a well-curated ichneumonid collection
(e.g.,USNM,AEI,CNC).
Distribution: Worldwide except for
the Australian region (Yu et al. 2005).
Limacodidae: The two specimens of Me.
discitergus in this study emerged (post-
erolaterally) from mummified early to
middle instar larvae of Pa.chloris on Q.
rubra and Pr.badia on F.grandifolia;
the mummies were probably made by a
species of Triraphis based on examina-
tion of montage images of host remains
(JTL). This species has not been re-
ported as a hyperparasitoid through a li-
macodid, nor was it reared from any
other parasitoids in this project.
Other hosts: Mesochorus discitergus
has been reported as a hyperparasitoid
of 31 species of Braconidae and three
species of Ichneumonidae (Yu et al.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
86

2005). Records of Me.discitergus as
a primary parasitoid of lepidopterans
are questionable, as discussed above.
Casinaria Holmgren (Ichneumonoidea:
Ichneumonidae: Campopleginae)
(Fig. 28)
Diagnosis.—
Casinaria mayberecog-
nized by the following combination of
characters: clypeus not separated from
supraclypeal area by distinct groove;
ventral posterior corner of propleuron with
strongly produced, more or less angulate
lobe touching or overlapping pronotum (as
in Fig. 24); areolet of fore wing closed,
obliquely quadrate and petiolate (as in Fig.
26); metasomal segment 1 with the fol-
lowing: spiracle beyond middle, petiole
long and cylindrical in cross-section and
with tergo-sternal suture at midline, T1
without trace of glymma (Fig. 28); meta-
soma laterally compressed; ovipositor not
needlelike, about as long as metasomal
apical depth and with dorsal subapical
notch (Fig. 28).
Fauna.—
Casinaria is distributed world-
wide, with 99 species, 20 of which are
recorded from the Nearctic Region (Yu
et al. 2005). Walley (1947) revised the
Nearctic fauna. In a forthcoming revi-
sion of Costa Rican Campopleginae,
I. D. Gauld recognizes a species-group
within Casinaria consisting of grandis
and two undescribed Costa Rican spe-
cies. The latter parasitize various species
of limacodids in the genera Acharia,
Euclea,Euprosterna, and Natada based
on extensive rearing efforts conducted in
Costa Rica (D. Janzen and W. Hallwachs
pers. comm; see also http://janzen.sas.
upenn.edu). Extralimitally, there is one re-
cord of Casinaria sp. from Ac. (as Sibine)
megasomoides (Walker) (Genty et al.
1978) in the Neotropical Region.
Biology.—
Like other campople-
gines, Casinaria species are koinobiont
endoparasitoids of Lepidoptera, with the
egg deposited in an exposed host larva and
the mature wasp larva emerging before the
host pupates (Jerman and Gauld 1988).
Species in the following families have
been recorded as hosts: Arctiidae, Cram-
bidae, Geometridae, Hesperiidae, Lasio-
campidae, Limacodidae, Megalopygidae,
Notodontidae, Nymphalidae, Oecophoridae,
Olethreutidae, and Zygaenidae (Walley
1947, Finlayson 1975, Short 1978, Carlson
1979a, Jerman and Gauld 1988, Janzen
and Hallwachs 2009).
Literature.—
The Nearctic Casinaria fauna
may be identified using Walley (1947).
Additionally, a species of Casinaria was
reported from Ac.(asSibine)megasomoides
Walker on oil palm in Costa Rica (Mexzo
´n
et al. 1996).
North American Records:
Casinaria grandis (Walley) (Fig. 28):
Walley, 1947: 376. Replacement name for
Casinaria texana Ashmead (1890); Ho-
lotype /, USNM (not examined).
Casinaria grandis may be distinguished
from other Nearctic Casinaria by the fol-
lowing characters: large, broad, flat, and
centrally concave scutellum; petiolate
areolet of the fore wing; gena descen ding
perpendicularly behind ocelli; coarsely
rugosopunctate propodeum without ca-
rinae; and large size (about 15 mm long).
Distribution: Gulf of Mexico (Florida,
Texas, Louisiana; Carlson 1979a). It pre-
sumably extends into northern Mexico.
Limacodidae: Reported from Ac. stim-
ulea (Carlson 1979a); no specimens
could be located in the USNM collec-
tion that would verify this record.
Other hosts: None known.
Hyposoter Fo
¨rster (Ichneumonoidea:
Ichneumonidae: Campopleginae)
(Figs. 23–24, 26–27, 30, 32)
Diagnosis.—
Hyposoter may be rec-
ognized by the following combination of
VOLUME 114, NUMBER 1
87

characters: clypeus not separated from
supraclypeal area by distinct groove (Fig.
32); ventral posterior corner of propleuron
with strongly produced, more or less an-
gulate lobe touching or overlapping pro-
notum (Fig. 24); areolet of fore wing
closed, obliquely quadrate and petiolate
(Fig. 26); metasomal segment 1 with the
following: spiracle beyond middle, petiole
short and quadrate in cross-section and
with tergo-sternal suture close to ventral
margin, T1 with glymma present as pitlike
impression (Fig. 27); metasoma laterally
compressed; ovipositor not needlelike,
about as long as metasomal apical depth
and with dorsal subapical notch (Fig. 23).
Fauna.—
Species of Hyposoter occur in
all zoogeographic regions. Yu et al. (2005)
listed 117 valid species worldwide and 29
for the Nearctic Region. Horstmann (2008)
described one new species from the Pale-
arctic Region, and Horstmann (2009)
transferred Hyposoter koentzeii (Kiss),
known from Hungary, from Omorgus
Fo
¨rster. Hyposoter bellus (Cresson),
known from Cuba, was transferred from
Xanthocampoplex Morley in Gauld and
Ferna
´ndez-Triana (2010). Viereck
(1925, 1926) provided keys that include
most species known from the Nearctic
Region, but a single key to species of
Hyposoter in the Nearctic Region has
not been published. Further, the keys in
Viereck (1925, 1926) use characters
that are of doubtful utility for species
diagnostics (see specific information
below).
Biology.—
Hyposoter spp. are koino-
biont endoparasitoids of ditrysian lepi-
dopterans, including some species of
Rhopalocera. Egg deposition is in early to
middle instar larvae, and the wasp larva
usually emerges from the penultimate
instar of the host larva. The wasp spins an
ovoid cocoon near or under the host cat-
erpillar’s remains. The cocoon is some-
times completely covered by the host’s
cuticle; this is not, however, mummifi-
cation as found in rogadine braconids
(see the discussion of rogadine mummi-
fication in Shaw (2006)) in which the host
mummy remains intact due to internal
pupation by the parasitoid.
Literature.—
Aside from literature re-
ported in Yu et al. (2005), Marconato
et al. (2008) reported an undetermined
Hyposoter from two geometrids (Lepi-
doptera). Reis Fernandes et al. (2010)
reported two undetermined species of
Hyposoter from two geometrids (Lepi-
doptera). There are no reliable or com-
prehensive treatments of the taxonomy
of the Nearctic Hyposoter fauna.
North American Records:
Hyposoter fugitivus (Say) (Figs. 23–24,
26–27, 30, 32): Banchus fugitivus Say,
1835: 247. Lectotype /, USNM, desig-
nated by Cushman (1925) (not found in
USNM collection).
Besides the 29 described Nearctic Hy-
posoter species there are numerous un-
described species; in the absence of a
formal revision, meaningful diagnosis of
fugitivus may be misleading. The char-
acters presented in the key, in conjunc-
tion with an association with a rearing
from a limacodid host in the Nearctic
Region, provide the best information at
this point to attempt to assign a species
name. As with other ichneumonids (e.g.,
Mesochorus) reported herein, final identi-
fication should be made by an ichneumo-
nid expert with access to a large col-
lectionsuchasUSNM,AEI,orCNC.
Distribution: Transcontinental in the
Nearctic (except possibly absent in the
southwestern U.S.A.) (Carlson 1979a).
Recorded from Brazil (Costa Lima
1962), but not recognized in the Neo-
tropical catalog of Townes and Townes
(1966) nor is it present in Costa Rica (I.
Gauld et al. unpubl.).
Limacodidae: Isa textua on Acer ne-
gundo L. and Salix nigra (Marsh)
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(Barbosa and Caldas 2004). It was
reared in this study from Pr.badia*
on Q. alba. The Hyposoter cocoon is
completely covered by the skin of the
host caterpillar. This is the only de-
scribed species of Hyposoter known to
attack limacodids; several undescribed
Costa Rican species have been reared
from species of Acharia,Euclea, Na-
tada,andPara s a based on extensive
caterpillar rearing effort in Costa Rica
(D. Janzen and W. Hallwachs pers.
comm.).
Other hosts: It has been reported from
30 other lepidopteran species in 12
families (Yu et al. 2005); a new record
from this study was its rearing from
Meg.crispata on Q. alba.
Baryceros Gravenhorst
(Ichneumonoidea: Ichneumonidae:
Cryptinae)
(Figs. 29, 37–38)
Diagnosis.—
Baryceros mayberecog-
nized by the following combination of
characters: clypeus separated from su-
praclypeal area by distinct groove; ventral
posterior corner of propleuron not devel-
oped as distinct lobe, at most with weak
groove delimiting it from main area of
propleuron (Fig. 37); dorsal margin of
pronotum with strong swelling at dorsal
end of epomia (Fig. 37); areolet of fore
wing open (vein 3rs-m absent) and pen-
tagonal (Fig. 29); mesosoma with coarse
punctures (Fig. 37); spiracle of metasomal
segment 1 beyond middle; glymma absent
(Fig. 29); metasoma cylindrical/dorsoven-
trally compressed; ovipositor not needlelike,
about 2.0X as long as metasomal apical
depth and without dorsal subapical notch
(Fig. 38); body (excluding legs) black
with white markings (Fig. 29).
Fauna.—
Baryceros is a New World
genus, with 32 described species; five are
found in the Nearctic (Yu et al. 2005).
Townes and Townes (1962) provide a key
to the Nearctic fauna.
Biology.—
Baryceros is placed in the
subtribe Baryceratina, with all known
host records indicating that the genera of
this subtribe are idiobiont ectoparasitoids
of Limacodidae (Townes 1970). Three
species of Baryceros have been reared.
Baryceros eucleidis (Blanchard) has been
reared from Ph.hipparchia (Cramer) and
Ac. (as Sibine)nesea (Stoll), and B. sibine
(Cameron) has been reared from Ac. (as
S.) trimaculata (Sepp) and Ac. (as S.)ne-
sea (Townes and Townes 1966). Bar-
yceros texanus is discussed below. A suite
of morphological characters appears to be
adaptations to attaching limacodid co-
coons. One such character is the posses-
sion of prominent dorsal and ventral teeth
on the ovipositor apex (Fig. 29); charac-
terized by Gauld (1984) as “corkscrew-
like,” this specialized structure is used to
penetrate the hard limacodid cocoon.
Literature.—
Townes and Townes
(1962) remains the definitive taxonomic
treatment of the Nearctic fauna.
North American Records:
Baryceros texanus (Ashmead) Figs. 29,
39–38: Crypturus texanus Ashmead,
1890: 413. Holotype ?, USNM (not
examined).
Baryceros texanus can be differentiated
from other Nearctic Baryceros by the
black and white tergites and the single
large white spot on the ventral portion of
the mesopleuron (Fig. 29).
Distribution: North America, in U.S.A.
from Maryland to Florida, west to
Michigan and eastern Texas. Recorded
from Sonora, Mexico and is presumably
more widely distributed (Townes and
Townes 1962 Carlson 1979a).
Limacodidae: Ala. slossoniae from
southern Florida (Ashmead 1897, Dyar
1897). The USNM collection has a reared
specimen from Stock Island (Monroe
Co.), Florida; label data state that the
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89

host was a “Lep. larva” on a leaf of
Conocarpus erecta L. (Combretaceae).
Other hosts: None known.
Lysibia Fo
¨rster (Ichneumonoidea:
Ichneumonidae: Cryptinae)
(Figs. 25, 35, 43)
Diagnosis.—
Lysibia may be recog-
nized by the following combination of
characters: clypeus separated from su-
praclypeal area by distinct groove, apical
0.3 turned inward at 90°angle (Fig. 43);
occipital carina meeting hypostomal ca-
rina above mandibular base (as in Fig. 41);
ventral posterior corner of propleuron not
developed as distinct lobe, at most with
weak groove delimiting it from main area
of propleuron (Fig. 25); dorsal margin of
pronotum without strong swelling at dor-
sal end of epomia (Fig. 35); areolet of fore
wing open (vein 3rs-m absent) and pen-
tagonal; vein 2-Cu of hind wing basally
incomplete (Fig. 35); mesosoma with
punctures ranging from fine to absent;
spiracle of metasomal segment 1 beyond
middle; glymma absent; metasoma cy-
lindrical/dorsoventrally compressed; ovi-
positor not needlelike, about 1.4X as long
as metasomal apical depth and without
dorsal subapical notch. Body color (ex-
cluding legs) black to dark brown, with
eastern Nearctic specimens often having
T2–T7 brown and with brownish red
areas on T2–T3 (Fig. 35).
Fauna.—
Lysibia is found worldwide
except for Africa and Australia. There
are nine described species, with two
(mandibularis (Provancher) and tenax
Townes) present in the Nearctic. They
may be identified using the key to world
species in Townes (1983).
Biology.—
Like the majority of spe-
cies in the Phygadeuontini, Lysibia spp.
are idiobiont ectoparasitoids of cocooned
hosts. They are apparently pseudohyperp-
arasitoids, specializing in microgastrine
cocoons (Townes 1970), including Co.
glomerata L. (Harvey et al. 2004); Yu
et al. (2005) list 33 microgastrine species
as hosts. Yu et al. (2005) also list 26
species of Lepidoptera in 14 families as
hosts. While some of these host records
may be correct (see discussion under
Isdromas), many likely are the result of
faulty record keeping (Shaw 1990). A
series of recent papers have been pub-
lished on the biology and ecology of
Ly.nana (Gravenhorst) (Harvey et al.
2004; 2006; 2009a, b) attacking Co.
glomerata.
Literature.—
Townes (1983) remains the
definitive taxonomic treatment of the
Nearctic fauna.
North American Records:
Lysibia mandibularis (Provancher) (Figs.
25, 35, 43): Hemiteles madibularis
Provancher, 1875: 315. Holotype /,
ULQC (not examined).
Lysibia mandibularis has the punctures of
tergites 2–3 centrally sparse, separated by
1.5–2.5X the length of the setae; Ly.
tenax has the tergal punctures evenly
distributed and separated by about 0.8X
the length of the setae.
Distribution: Transcontinental in the
Nearctic (Townes 1983).
Limacodidae: The specimens in this
study were reared from cocoons of an
undetermined gregarious microgastrine
parasitizing Ac.stimulea on Lindera
benzoin (L.) Blume.
Other hosts: Yu et al. (2005) list seven
species of microgastrines as hosts; four
species of Lepidoptera are also listed,
but these records are probably errone-
ous as noted above.
Acrolyta Fo
¨rster (Ichneumonoidea:
Ichneumonidae: Cryptinae)
(Figs. 36, 41)
Diagnosis.—
Acrolyta may be recog-
nized by the following combination of
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90

characters: clypeus separated from su-
praclypeal area by distinct groove, apical
0.3 flat and not turned inward (as in Fig.
44); occipital carina meeting hypostomal
carina at mandibular base (Fig. 41); ven-
tral posterior corner of propleuron not
developed as distinct lobe, at most with
weak groove delimiting it from main area
of propleuron (as in Fig. 25); dorsal mar-
gin of pronotum without strong swelling
at dorsal end of epomia (Fig. 36); areolet
of fore wing open (vein 3rs-m absent) and
pentagonal; vein 2-Cu of hind wing com-
plete; mesosoma with punctures ranging
from fine to absent; spiracle of metasomal
segment 1 beyond middle; glymma ab-
sent; metasoma cylindrical/dorsoventrally
compressed; ovipositor not needlelike,
1.6–1.8X as long as metasomal apical
depth and without dorsal subapical notch.
Body color of females (excluding legs)
ranges from uniformly black/dark brown
to having extensive brownish red areas on
T2–T4; males have metasomal coloration
ranging from having brownish red re-
stricted to the apices of T2–T3, to having
brownish red on the apex of T2, all of T3,
and the basal area of T4 (Fig. 36).
Fauna.—
Acrolyta is found in the Afro-
tropical, Palaearctic, Nearctic, and Orien-
tal regions. There are 25 described species,
with the following four species in the
Nearctic: alticola (Ashmead), mesochori
Ashmead, nigricapitata (Cook and Davis),
and washingtonensis (Cushman) (Yu et al.
2005). No keys are available to separate
the species.
Biology.—
Acrolyta species are inferred
idiobiont ectoparasitoids of cocooned hosts,
as are other species in the Phygadeuontini
(Gauld 1984). Townes (1970) stated that
“[p]robably the usual hosts are cocoons
of Braconidae and Ichneumonidae.” Yu
et al. (2005) list 28 species of braconids
as hosts; 13 lepidopteran species in 10
families are also listed (as well as a spe-
cies of Cynipidae!). While some of the
Lepidoptera may be hosts (see discussion
under Isdromas), it is likely that these re-
cords are due to faulty observations and
lax record keeping of the sort discussed by
Shaw (1990). Harvey et al. (2009b) re-
ported on the biology of Acrolyta nens
Hartig attacking Co.glomerata.
Literature.—
There are no reliable or
comprehensive treatments of the taxon-
omy of the Nearctic Acrolyta fauna.
North American Records:
Acrolyta nigricapitata (Cook and Davis)
(Figs. 36, 41): Ischnoceros nigricapitatus
Cook and Davis, 1891: 11. Holotype /,
MSUC (not examined).
Acrolyta nigricapitata can be distin-
guished from other Nearctic Acrolyta by
the following characters: tergite 3 with
fine longitudinal striae covering most of
surface (tergite 3 without striae); fore
and middle coxae light brownish red
(fore and middle coxae white); tergite 3
completely brownish red (tergite 3 dark
brown/fuscous, ranging from completely
so to having anterior and posterior mar-
gins narrowly brownish red).
Distribution: Northeastern U.S.A. and
southeastern Canada to the Pacific
Coast (California and Washington)
(Gauld 1984, Yu et al. 2005).
Limacodidae: Acrolyta nigricapitata
is recorded as parasitizing Ac. stimulea
in Ashmead (1896: 209), where the
only information is as follows: “De-
scribed from two specimens labeled
No. 295o, reared May 14, 1883, from
Empretiae stimulea.” It is unlikely to
be a primary parasitoid.
Other hosts: Yu et al. (2005) list three
braconid species as hosts; a noctuid,
Pseudaletia unipunctata (Haworth) (=
Mythimna unipunctata), is also listed, but
this record also stems from Ashmead
(1896: 210) with only the following
entry: “Described from one specimen
labeled “parasite on Leucania uni-
punctata, June, 1880.” The USNM
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91

collection has three nigricapitata spec-
imens with host records/remains (R.
Kula pers. comm.): (1) a specimen
without host remains but label data in-
dicating Ap.melanoscelus (Ratzeburg)
as the host, (2) a specimen with a
mounted cocoon and label data iden-
tifying it as Ap. melanoscelus, and (3)
a specimen with an associated cocoon
(probably braconid) and no associated
information.
Isdromas Foerster (Ichneumonoidea:
Ichneumonidae: Cryptinae)
(Figs. 31, 33–34, 42, 44)
Diagnosis.—
Isdromas may be recog-
nized by the following combination of
characters: clypeus separated from supra-
clypeal area by distinct groove, apical 0.3
flat and not turned inward (Fig. 44); oc-
cipital carina meeting hypostomal carina
above mandibular base (Fig. 42); ventral
posterior corner of propleuron not devel-
oped as distinct lobe, at most with weak
groove delimiting it from main area of
propleuron (as in Fig. 25); dorsal margin
of pronotum without strong swelling at
dorsal end of epomia (Fig. 34); areolet of
fore wing open (vein 3rs-m absent) and
pentagonal (Fig. 34); vein 2-Cu of hind
wing complete (Fig. 34); mesosoma with
punctures ranging from fine to absent;
spiracle of metasomal segment 1 be-
yond middle; glymma absent (Figs. 34);
metasoma cylindrical/dorsoventrally com-
pressed; ovipositor not needlelike, about
3.0X as long as metasomal apical depth
and without dorsal subapical notch. Body
color (excluding legs) usually black; T2–
T7 may vary from black to dark brown
(Fig. 34).
Fauna.—
Isdromas contains eight de-
scribed species worldwide with one spe-
cies reported in the Nearctic (Yu et al.
2005). Although Townes (1970) estimated
that the world fauna consists of at least 60
species, Is.lycaenae (Howard) appears to
be the sole Nearctic representative.
Biology.—
As with most Cryptinae,
Acrolyta,Isdromas, and Lysibia are idi-
obiont ectoparasitoids of cocooned hosts
(Gauld 1984). Gauld (1988) points out
that many species of Phygadeuontini
“are facultative hyperparasitoids and use
as hosts whatever is in a particular cocoon
be it the cocoon maker or one of its par-
asitoids.” This is particularly true for the
subtribe Acrolytina, to which these gen-
era belong, due to the habit of parasitizing
ichneumonid and braconid cocoons
(Gauld 1995). This then leads to consid-
eration of two different modes of hyper-
parasitism (Shaw and Askew 1976): (1)
true hyperparasitism in which the hyper-
parasitoid develops on a primary parasit-
oid living on or within the primary’s host
and (2) pseudohyperparasitism, which
involves the pseudohyperparasitoid at-
tacking the primary parasitoid after it
has destroyed its host.
This background puts a necessary
perspective on the rearing records of I.
lycaenae. The species has been reported
as reared from five species of Braconi-
dae and two species of Ichneumonidae
on various lepidopteran hosts (Yu et al.
2005); this would thus be pseudohy-
perparasitism. Isdromas lycaenae has also
been reported from nine species of Lepi-
doptera in seven families (Yu et al. 2005);
these could either be instances of primary
parasitism or utilization of overlooked
primary parasitoids. De Santis (1987) and
Lourenca
˜o et al. (1989) reported Is.mon-
terai (Costa Lima), the only other species
of Isdromas for which host use is known,
as a likely pseudohyperparasitoid of Prota-
panteles dalosoma (De Santis) (Braconidae:
Microgastrinae) on Anacraga citrinopsis
Dyar (Lepidoptera: Dalceridae).
Literature.—
The genus may be iden-
tified using Townes (1983); lycaenae is the
only Nearctic representative.
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92

North American Records:
Isdromas lycaenae (Howard) (Figs. 31,
33–34, 42, 44) Hemiteles lycaenae
Howard, 1889: 1880. Holotype /,
USNM (examined).
Isdromas lycaenae is the sole Nearctic
representative of this genus and may be
recognized by the apical 0.3 of clypeus
turned inward at 90°(Fig. 43), vein 2-Cu
of hind wing basally incomplete, and the
dark brown body with paler legs (Fig. 34).
Distribution: In the Nearctic, Is.lycae-
nae is found from Ontario south to the
Gulf of Mexico and west to Texas and
Kansas. In the Neotropics it is found from
Honduras to Argentina (Yu et al. 2005).
Limacodidae: Isdromas lycaenae was
reared from mummified early to
middle instar larvae of Ac.stimulea*
on Q.rubra;I.textula*on Q. rubra;N.
nasoni*on F.grandifolia,P.serotina,
and Q.rubra;Pa.chloris*on Q. rubra;
and Pr.badia*on F. grandifolia (Table
1). The mummies were probably made
by a species of Triraphis based upon
images of the host remains (JTL).
Other hosts: Yu et al. (2005) listed
records of Is.lycaenae from five spe-
cies of Braconidae and two species of
Ichneumonidae; they also record it
from nine lepidopteran species in seven
families. Records of Is.lycaenae as
a primary parasitoid are questionable
for the reasons discussed above.
Orthogonalys Schulz (Trigonaloidea
Trigonalidae: Orthogonalinae)
(Figs. 70–71)
Diagnosis.—
Antenna black with white
or light yellow band in middle; propo-
deum usually with light markings; meta-
soma thin, without punctures, usually
entirely orange; without female arma-
ture; male antenna without tyloids.
Fauna.—
Carmean and Kimsey (1998)
listed 11 species worldwide, one from
eastern North America, one from South
America, and the others from India,
Vietnam, Japan, Taiwan, Madagascar, and
southern Africa. An additional species was
described from eastern North America by
Smith and Stocks (2005). Carmean and
Kimsey (1998) gave a discussion of the
genus.
Biology.—
Although some species are
collected rather commonly, very little is
known about hosts and biology. Seasonal
flight activity of O. pulchella in eastern
North America was recorded in D. Smith
(1996). Carlson (1979b) reported O.
pulchella from a tachinid parasitizing an
undetermined species of Lepidoptera.
Benoit (1951) recorded O. hova Bischoff
from a limacodid moth in Madagascar.
The first verified host records were re-
ported by Murphy et al. (2009) from
tachinids parasitizing limacodids, noctuids,
and megalopygids. The female deposits
numerous eggs on leaves of angiosperms.
The eggs are ingested by caterpillars
feeding on the leaves, eclosion occurs in
the gut of the caterpillar, and the first in-
star bores through the gut wall. Further
development depends on parasitization of
the caterpillar by a parasitoid, in this case
a tachinid (Murphy et al. 2009). The other
Nearctic species, O. bella Smith and
Stocks, is known from a single speci-
men from Great Smoky Mountains
National Park, Tennessee.
Literature.—
Townes (1956) revised the
North American Trigonalidae and sepa-
rated Orthogonalys from other genera.
The genus was treated in D. Smith (1996),
and seasonal flight data were given for
O. pulchella. Carmean and Kimsey (1998)
gave a diagnosis and description and
provided the known distribution and bi-
ology of the genus. Murphy et al. (2009)
recorded hosts for O. pulchella.
North American Records:
Orthogonalys pulchella (Cresson) (Figs.
70–71): Trigonalys pulchellus Cresson,
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93

1867: 351. Holotype ?,ANSP(not
examined).
This species and O. bella are the only
species of Orthogonalys in North and
Central America. They are illustrated and
separated by Smith and Stocks (2005).
Orthogonalys bella, known only from a
female, is almost entirely black, has
antennal segments 7–9 white, smaller
eyes slightly diverging below and far
removed from the posterior margin of
the head, tarsal claws with both teeth
subequal in size, and the sheath in lat-
eral view broadly rounded at its apex.
Figures 58 and 59 show the typical color
of O. pulchella. There is some variation
in the amount of black on the mesosoma
and dorsum of the metasoma.
Distribution: Southeastern Canada,
eastern U.S.A., Mexico.
Limacodidae: Associated with primary
parasitoids of Iso. beutenmuelleri and
I. textual. Most records are from tach-
inids parasitizing I.textula feeding on
Q. prinus and Q. rubra.
Other hosts: Uramya pristis parasitiz-
ing Meg.crispata (Megalopygidae),
Isochaetes beutenmuelleri,I.textula,
and Acrocnicta increta (Noctuidae)
(Murphy et al. 2009). Recorded para-
sitizing Com. concinnata through
Actias luna (L.) (Kellogg et al. 2003).
Taeniogonalos Schulz (Trigonaloidea
Trigonalidae: Trigonalinae)
(Figs. 18, 72–73)
Diagnosis.—
Antenna uniformly yellow
brown or brown becoming darker apically,
never banded (Figs. 72–73); metasoma
stout, with punctures, black with trans-
verse yellow stripes; female with armature
(Fig. 72); male antenna with tyloids (ventral
sensory structures).
Fauna.—
Taeniogonalos is the most
widely distributed of all trigonalid genera.
Carmean and Kimsey (1998) listed 34
species worldwide from the Nearctic, Neo-
tropical, eastern Palearctic, Australasian,
Oriental, and Afrotropical regions. Most
are from eastern Asia and South America.
Only one species, Taeniogonalos gund-
lachii, occurs in eastern North America.
Biology.—
Species of this genus have
been reared from a variety of tachinid
and ichneumonid parasitoids of species
of Pyralidae, Noctuidae, Saturniidae,
Megalopygidae, and Arctiidae. A summary
of their biology and hosts was given by D.
Smith (1996) and Carmean and Kimsey
(1998). Krauth and Williams (2006)
recorded T. gundlachii from an arctiid,
Euchaetes egle (Drury), in Wisconsin.
Additionally, there is one record from a
tachinid parasitizing a detritivore tipulid,
Tipula ?flavoumbrosa (Gelhaus 1987). The
African Taeniogonalos maynei Benoit was
reared from pupae of Latoia albipunctata
Holland (Limacodidae) in which it pre-
sumably was parasitizing a primary par-
asitoid (Benoit 1950). In Australia, T.
maculata (Smith) and T. venatoria Riek
have been reared as primary parasitoids
from pergid sawfly hosts including Per-
gagrapta condei Benson, Perga dorsalis
Leach, P. affinis Kirby, and Pseudoperga
belinda (Kirby) (Raff 1934; Carne 1969;
Weinstein and Austin 1991, 1995, 1996).
Oviposition and general biology are sim-
ilar to that described for Orthogonalys.
Literature.—
Townes (1956) separated
the genus (as Poecilogonalos)fromother
North American genera. The genus was
also separated from other eastern North
American trigonalids and the seasonal
activity of its species recorded in D. Smith
(1996).
North American Records:
Taeniogonalos gundlachii (Cresson) (Figs.
18, 72–73): Poecilogonalos gundlachii
Cresson, 1865: 10. Holotype ?, ANSP
(not examined).
This is the only species of Taeniogonalos in
eastern North America. Two other species
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occur in southern Mexico and Central
America. The typical color of eastern
North and Central American specimens is
shown in Figs. 72–73. Specimens from
Cuba are more extensively yellow, espe-
cially the head and mesosoma, whereas
some specimens from the southern tier
of states from Florida to Louisiana are
somewhat intermediate in color. Carmean
and Kimsey (1998) considered all as a single
widespread, variable species. In the literature
prior to Carmean and Kimsey (1998), the
eastern North American species is cited as
Poecilogonalos costalis (Cresson).
Distribution: Southeastern Canada,
eastern U.S.A. from Massachusetts to
Florida west to Wisconsin, Ohio, Loui-
siana; Costa Rica, Cuba.
Limacodidae: Associated with primary
parasitoids of E.delphinii.
Other hosts: One specimen reared from
a tachinid, probably Com.concinnata
parasitizing E.delphinii feeding on
Nyssa sylvatica. Recorded parasitiz-
ing Com. concinnata through Actias
luna (L.) (Kellogg et al. 2003).
Questionable/Invalid/Miscellaneous
Parasitoid Records (North America)
Tachinidae
The following are single records that
require confirmation, or in the case of
Chaetexorista javana, involve an intro-
duced species that is probably no longer
extant in North America. Some records
may be based on misidentifications or
represent rare or accidental host records.
(1) Carcelia spp. (Exoristinae: Eryciini):
Adoneta spinuloides (Brauer and Ber-
genstamm 1895, as Chaetolyga sp.), Ph.
pithecium (Patton 1958, ex “hag-moth”,
“Phobetron pithecium?”). Both records
cited by Arnaud (1978).
(2) Chaetexorista javana Brauer and
Bergenstamm, 1895 (Exoristinae:
Exoristini):
Monema flavescens (numerous records,
see Arnaud 1978). Chaetexorista javana
was introduced into Massachusetts from
Japan in 1929 and 1930 for control of the
Oriental moth, M. flavescens (Dowden
1946). The introduction was successful,
but Ch. javana has not been reported for
many years and is probably now extinct in
North America. Similarly, M. flavescens is
likely extinct in North America also based
on subsequent searches (M. Epstein pers.
comm.).
(3) Hyphantrophaga virilis (Aldrich and
Webber, 1924) (Exoristinae: Goniini):
Euclea delphinii (Coquillett 1897, as
Exorista blanda (Osten Sacken) ex E.
cippus (Cramer)). The original record by
Coquillett (1897) was later cited by
Aldrich and Webber (1924, as Zenillia
blanda blanda (Osten Sacken)), Sellers
(1943, as Z.virilis), and Arnaud (1978, as
Eusisyropa virilis).
(4) Lixophaga mediocris Aldrich, 1925
(Exoristinae: Blondeliini):
Monema flavescens (Brimley 1938,
“reared from twig infested by Oriental
moth”; Arnaud 1978 [citing Brimley
record]).
(5) Neomintho celeris (Townsend, 1919)
(Tachininae: Euthelairini):
Packardia ceanothi Dyar (Raizenne
1952, as Eupelecotheca celer; Arnaud
1978 [citing Raizenne record, also as
Eupelecotheca celer]).
Hymenoptera
(Note: None of the taxa listed below
appear in Table 1)
(1) Closterocerus cinctipennis Ashmead
(Eulophidae: Entedoninae):
Associated with Ac.stimulea on Q.
rubra (no host remains), but this species
is possibly the result of a contaminant
introduced during rearing as they are
known parasitoids of microlepidopteran,
VOLUME 114, NUMBER 1
95

buprestid, and agromyzid leafminers; eggs
of Chrysomelidae and Tenthredinidae;
and “Diptera on Catalpa” (Hansson
1994).
(2) Neochrysocharis diastatae (Howard)
(Eulophidae: Entedoninae):
Associated most frequently with lepi-
dopteran and dipteran leaf/stem miners
(Hansson 1995). Five specimens are as-
sociated with E.delphinii on Q.alba,but
the limacodid remains offer no indication
of parasitization (e.g., emergence holes).
We suspect that a leaf miner parasitized
by Neo. diastatae was introduced during
the rearing of the larva.
(3) Horismenus sp. (Eulophidae:
Entedoninae):
A single male is associated with Ac.
stimulea on Q. rubra (no host remains)
but became infected with fungus after
emergence, obscuring morphological fea-
tures. It was run through Burks’ (1971)
key to Nearctic species (even though key
is written for females) and compared with
males of species near which it keyed. It
is not conspecific with males of Ho. la-
trodecti Burks or Ho. carolinensis Burks.
Two extralimital species from the Neo-
tropics, Ho. clavicornis Cameron and Ho.
hipparchia Cameron, attack Ph.hip-
parchia in Guyana (De Santis 1979).
(4) Aroplectrus dimerus Lin (Eulophi-
dae: Eulophinae):
This species is known from the Phil-
ippines, Taiwan, China, and India and
recorded from Pa. bicolor (Singh et al.
1988) and Penthocrates sp. (Fry 1989). It
was released in Hawaii beginning in
2010 as part of a biological control
program against the invasive limacodid
Darna pallivitta (Moore) (HDOA 2010,
2011; Conant et al. 2011). Although
neither D. pallivitta nor Ar. dimerus are
currently known from the U.S.A., it is
highly likely that the limacodid will be
introduced into California given its his-
tory of interception at ports of entry
(CDFA 2005, 2006). Were this to hap-
pen, it is possible that A. dimerus would
be introduced also, either accidentally
or, later, deliberately.
(5) Brachymeria euploeae (Westwood)
(Chalcididae: Chalcidinae):
Recorded from the U.S.A. (but not
attacking a limacodid) and as a parasit-
oid on Chalcoscelis albiguttata Snellen
in Bhutan (as Butjan) and Maluku
Islands (as Moluccas) (Thompson 1954).
Chalcoscelis albiguttata does not occur in
the U.S.A. and has never been introduced
(M. Epstein pers. comm.). The recorded
distribution for Br. euploeae is in eastern
and southeastern Asia and India with the
lone U.S.A. record (Thompson 1954)
standing out as the single record for the
Western Hemisphere. Other host records
and associated references are presented in
Cock (1987). The Thompson (1954) record
indicates that Br. euploeae was introduced
into the U.S.A. from Japan to control
Ostrinia (as Pyrausta)nubilalis (Hu
¨bner).
All of the limacodid hosts recorded for Br.
euploeae are distributed in eastern and
southeastern Asia and India.
(6) Brachymeria lasus Walker (Chalci-
didae: Chalcidinae):
This species is native to the Oriental and
Australasian regions but has been intro-
duced into the U.S.A. several times to con-
trol the gypsy moth (L.dispar; Weseloh
and Anderson 1982, ROBO Database
1981–1985), fall webworm (Hyphantria
cunea Drury; ROBO Database 1981–
1985), and range caterpillar (Hemileuca
oliviae Cockerell; ROBO Database 1981–
1985). Known limacodid hosts include
Thosea cinereomarginata Banks and Th.
sinensis Walker from the Philippines
(Baltazar 1965). This species has a broad
host range on small lepidopteran pupae
and is facultatively hyperparasitic through
hymenopterans and dipterans (Cock 1987).
(7) Conura immaculata (Cresson)
(Chalcididae: Chalcidinae):
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
96

This species is primarily Neotropical
but occurs in southern U.S.A., although
it is not associated with Limacodidae
there. It has been recorded parasitizing
N.subpectinata Dyar and Euprosterna
elaea (as elaeasa) Dyar (Delvare 1992),
primarily Neotropical species, in that re-
gion. It is also known as a hyperparasitoid
on Ichneumonoidea (Delvare 1992), and
Burks (1979) recorded it from Meteorus
sp. (Braconidae) from Cameron County,
Texas. The latter specimens and parasitoid
pupal remains are housed in the USNM.
Lingren (1977) records it as a hyperpara-
sitoid of Campoletis sonorensis (Ca-
meron) through Heliothis virescens (F.) on
cotton in Brownsville, Texas.
(8) Conura lasnierii Gue
´rin-Me
´neville
(Chalcididae: Chalcidinae):
This chalcidid is recorded from Florida
in the U.S.A. but is not known from a
limacodid host. It has been recorded from
Leucophobetron argentiflua Hu
¨bner in
Cuba (Gahan 1934). The moth has never
been intentionally introduced into the U.
S.A. (M. Epstein pers. comm.). Addi-
tionally, Ala. slossoniae, a common spe-
cies in Florida, is closely related to the
monotypic Leucophobetron and perhaps
congeneric (M. Epstein pers. comm.).
(9) Trichopria sp. (Diapriidae: Dia-
priinae):
A single male is associated with E.
delphinii on Q. alba and is likely the
result of contamination during the rear-
ing process (see Neochrysocharis dia-
statae, above). Species of Diapriinae are
recorded as parasitoids of orthorhaphous
(soldier flies, flower flies, etc.) and cy-
clorhaphous (tachinid flies, muscoid flies,
etc.) Diptera (www.diapriid.org/public/
site/diapriid/home). The larval cadaver
associated with this specimen was thor-
oughly dissected in search of a tachinid
puparium that would have likely served
as the host in this instance, but none was
discovered. There was no evidence of any
type of emergence from the cadaver
during the pre-dissection inspection.
A
CKNOWLEDGMENTS
We thank Steve Lingafelter, Thomas
Henry, John Brown (Systematic Ento-
mology Laboratory [SEL], PSI, ARS,
USDA), and Paul Hanson (University of
Costa Rica, San Jose
´, Costa Rica) for
their critical reviews of and suggestions
on this manuscript. We also thank Scott
Whittaker (SEM Lab Manager of the
Scanning Electron Microscopy Lab,
Smithsonian Institution, National Museum
Natural History) for stub preparation and
SEM access. A special thanks to Marc
Epstein (California Department of Food
and Agriculture) for his thorough review
of the manuscript, including pointing out
omissions and limacodid nomenclatural
issues. Thanks to Jerrett McCormick
(SEL), Taina Litwak (SEL), Lynette Gates
(spouse, MWG), Matt Buffington (SEL),
and Terry Nuhn (SEL) for consultation,
information entry, and specimen imaging.
Istva
´nMiko
´and Matt Yoder (North
Carolina State University, Raleigh, North
Carolina) provided consultation on Cera-
phron and Trichopria, respectively. Andy
Deans (North Carolina State University,
Raleigh, North Carolina) permitted the
reproduction of two images of Ceraphron
(Figs. 63–64). Neal Evenhuis (Bishop
Museum, Honolulu, Hawaii) vetted the
section on Systropus and provided the im-
age of S.macer. Funding for the rearing
portion of this project was provided by
NSF-DEB 0643438 to JTL. The fourth au-
thor (JEO) is grateful to D. M. Wood (Ag-
riculture and Agri-Food Canada, Ottawa)
for sharing his knowledge of the tachinid
flies treated in this paper and for allowing
the use of his unpublished illustrations of
Austrophorocera male genitalia in Figs.
14–17. The fourth author (JEO) also
thanks S. J. Henderson (Agriculture and
VOLUME 114, NUMBER 1
97

Agri-Food Canada, Ottawa) for creating
the montaged images of tachinids and
their structures that appear herein.
USDA is an equal opportunity provider
and employer.
Literature Cited
van Achterberg, C. 1991. Revision of the genera of
the Afrotropical and W. Palaearctical Roga-
dinae Foerster (Hymenoptera: Braconidae).
Zoologische Verhandelingen 273: 1–102.
Adams, J. and D. Yanega. 1991. The lepidopteran
host of a Neotropical bombyliid fly (Lepi-
doptera: Limacodidae, Diptera: Bombyliidae).
Journal of the Kansas Entomological Society
64: 443–444.
Aiello, A. 1980. Systropus columbianus (Diptera:
Bombyllidae [sic]) reared from a larva of
Limacodid moth. Entomological News 91:
89–91.
Aldrich, J. M. 1921. The muscoid genera Pseu-
deuantha and Uramyia (Diptera). Insecutor
Inscitiae Menstruus 9: 83–92.
Aldrich, J. M. 1925. Two new species of the
tachinid genus Lixophaga, with notes and key
(Diptera). Proceedings of the Entomological
Society of Washington 27: 132–136.
Aldrich, J. M. and R. T. Webber. 1924. The North
American species of parasitic two-winged
flies belonging to the genus Phorocera and
related genera. Proceedings of the United
States National Museum 63 (Art. 17) [=No.
2486]: 1–90.
Andre
´, E. 1881. Notes hyme
´nopte
´rologiques. I.
Chalcidites. Annales de la Socie
´te
´Entomolo-
gique de France (6
e
Se
´rie) 1: 332–344.
Arnaud, P. H., Jr. 1978. A host-parasite catalog
of North American Tachinidae (Diptera).
United States Department of Agriculture.
Miscellaneous Publication 1319: 1–860.
Ashmead, W. H. (1888) 1889. Descriptions of
new Braconidae in the collection of the U. S.
National Museum. Proceedings of the United
States National Museum 11: 611–671.
doi:10.5479/si.00963801.11-760.611
Ashmead, W. H. 1890. Description of new Ich-
neumonidae in the collection of the U.S.
National Museum. Proceedings of the United
States National Museum 12: 387–451.
doi:10.5479/si.00963801.12-779.387
Ashmead, W. H. 1893. A monograph of the North
American Proctotrypidae. Bulletin of the
United States National Museum 45: 1–472.
doi:10.5479/si.03629236.45.1
Ashmead, W. H. 1896. Descriptions of new para-
sitic Hymenoptera. Transactions of the Amer-
ican Entomological Society 23: 179–234.
Ashmead, W. H. 1897. Two new parasites from
Eupoeya slossoniae. Canadian Entomologist
29: 113–114. doi:10.4039/Ent29113-5
Ashmead, W. 1904. Classification of the chalcid
flies of the superfamily Chalcidoidea, with
descriptions of new species in the Carnegie
Museum, collected in South America by
Herbert H. Smith. Memoirs of the Carnegie
Museum 1: 225–551.
Askew, R. R. 1971. Parasitic Insects. Heinemann
Educational Books, London, England. 316 pp.
Austin, A. D. 1987a. A review of the Braconidae
(Hymenoptera) that parasitize Limacodidae
in Southeast Asia, particularly those associ-
ated with coconut and oil palm, pp. 139–164.
In M. J. W. Cock, H. C. J. Godfray, and J. D.
Holloways, eds. Slug and Nettle Caterpillars.
The Biology, Taxonomy and Control of the
Limacodidae of Economic Importance on
Palms in South-east Asia. CAB International,
Wallingford, Oxon, United Kingdom. 270 pp.
Austin, A. D. 1987b. Ceraphronidae, pp. 181–
183. In M. J. W. Cock, H. C. J. Godfray, and
J. D. Holloways, eds. Slug and Nettle Cater-
pillars. The Biology, Taxonomy and Control
of the Limacodidae of Economic Importance
on Palms in South-east Asia. CAB Interna-
tional, Wallingford, Oxon, United Kingdom.
270 pp.
Baltazar, C. R. 1965. A new species of ichneumon-
fly and a chalcid wasp parasitic on slug cat-
erpillars in Basilan Island. Philippine Journal
of Plant Industry 30: 109–113.
Barbosa, P. and A. Caldas. 2004. Patterns of
parasitoid-host associations in differently
parasitized macrolepidopteran assemblages
on black willow Salix nigra (Marsh) and box
elder Acer negundo L. Basic and Applied
Ecology 5: 75–85.
Barbosa, P., J. L. Capinera, and E. A. Harrington.
1975. The gypsy moth parasitoid complex in
western Massachusetts: a study of parasitoids
in areas of high and low host density. Envi-
ronmental Entomology 4: 842–846.
Barbosa, P. and A. Caldas. 2004. Patterns of para-
sitoid-host associations in differently parasit-
ized macrolepidopteran assemblages on black
willow Salix nigra (Marsh) and Box Elder
Acer negundo L. Basic and Applied Ecology 5:
75–85.
Becker, V. O. and M. E. Epstein. 1995. Limaco-
didae, pp. 128–132. In J. B. Heppner, ed.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
98

Checklist: Part 2. Hyblaeoidea – Pyraloidea –
Torticoidea. Atlas of Neotropical Lepidoptera.
Scientific Publishers, Gainesville, Florida.
Benoit, P. L. G. 1950. Les Trigonaloidae du Congo
Belge (Hym. – Terebr.). Revue de Zoologie et
de Botanique Africaines 43: 216–221.
Benoit, P. L. G. 1951. La systematique des Trig-
onaloidae (Hym.) ethiopiens. Revue de Zoo-
logie et de Botanique Africaines 44: 141–147.
Boettner, G. H., J. S. Elkinton, and C. J. Boettner.
2000. Effects of a biological control in-
troduction on three nontarget native species
of saturniid moths. Conservation Biology 14:
1798–1806. doi:10.1046/j.1523-1739.2000.
99193.x
Boucˇek, Z. 1992. The New World genera of
Chalcididae. Memoirs of the American En-
tomological Institute 53: 49–118, 443–446.
Boucˇek, Z. and G. Delvare. 1992. The identities
of species described or classified under
Chalcis by J. C. Fabricius. Memoirs of the
American Entomological Institute 53: 29.
Boucˇek, Z. and J. Halstead. 1997. Chalcididae,
pp. 151–164. In G. A. P. Gibson, J. T. Huber,
and J. B. Woolley, eds. Annotated Keys to the
Genera of Nearctic Chalcidoidea (Hyme-
noptera). Research Press. National Research
Council of Canada, Ottawa, Ontario. 794 pp.
Bourchier, R. S. 1991. Growth and development
of Compsilura concinnata (Meigan [sic])
(Diptera: Tachinidae) parasitizing gypsy moth
larvae feeding on tannin diets. Canadian
Entomologist 123: 1047–1055. doi:10.4039/
Ent1231047-5
Brauer, F. and J. E. von Bergenstamm. 1895. Die
Zweiflu
¨gler des Kaiserlichen Museums zu
Wien. VII. Vorarbeiten zu einer Monographie
der Muscaria schizometopa (exclusive An-
thomyidae). Pars IV. F. Tempsky, Wien. 88 pp.
([Also published in 1895. In Denkschriften der
Kaiserlichen Akademie der Wissenschaften.
Wien. Mathematisch-Naturwissenschaftliche
Classe, 61 (1894), 537–624.])
Brimley, C. S. 1938. The Insects of North Carolina.
Being a List of the Insects of North Carolina
and Their Close Relatives. North Carolina De-
partment of Agriculture, Raleigh. 560 pp.
Brues, C. 1906. Notes and descriptions of North
American parasitic Hymenoptera. II. Bulletin
of the Wisconsin Natural History Society 4:
143–152.
Brues, C. 1916. Serphoidea, pp. 529–576. In H. L.
Viereck, ed. Guide to the Insects of Con-
necticut. Part III. The Hymenoptera, or Wasp-
like Insects, of Connecticut. Chalcidoidea.
Bulletin. Connecticut State Geological and
Natural History Survey 22: 1–848.
Buffington, M. L., R. Burks, and L. McNeil.
2005. Advanced techniques for imaging
parasitic Hymenoptera (Insecta). American
Entomologist 51: 50–56.
Buffington, M. L. and M. W. Gates. 2008.
Advanced imaging techniques II: using a
compound microscope for photographing
point-mount specimens. American Entomol-
ogist 54: 222–224.
Burks, B. D. 1940. Revision of the chalcid-flies of
the tribe Chalcidini in America north of
Mexico. Proceedings of the United States
National Museum 88: 237–354. doi:10.5479/
si.00963801.88-3082.237
Burks, B. D. 1966. The North American species
of Pediobius Walker (Hymenoptera: Eulo-
phidae). Proceedings of the Entomological
Society of Washington 68: 33–43.
Burks, B. D. 1971. The Nearctic species of Ho-
rismenus Walker (Hym., Eulophidae). Pro-
ceedings of the Entomological Society of
Washington 73: 68–83.
Burks, B. D. 1979. Torymidae (Agaoninae) and
all other families of Chalcidoidea (excluding
Encyrtidae), pp. 748–749, 768–889, 967–1043.
In K.V. Krombein, P. D. Hurd, Jr., D. R. Smith,
andB.D.Burks,eds.CatalogofHyme-
noptera in America North of Mexico. Vol 1.
Smithsonian Institution Press, Washington,
D.C. 1198 pp.
Butler, L. 1993. Parasitoids associated with the
macrolepidoptera community at Coopers
Rock State Forest, West Virginia: a baseline
study. Proceedings of the Entomological So-
ciety of Washington 95: 504–510.
California Department of Food and Agriculture
(CDFA). 2005. Darna pallivitta,p.21.Cal-
ifornia Plant Pest and Disease Report 22: 1–76.
California Department of Food and Agriculture
(CDFA). 2006. Darna pallivitta, p. 12. California
Plant Pest and Disease Report 23: 1–117.
Cameron, P. 1913. The Hymenoptera of the
Georgetown Museum. Part V. Timehri, Guyana
3: 105–137.
Cantrell, B. K. 1986. An updated host catalogue
for the Australian Tachinidae (Diptera). Jour-
nal of the Australian Entomological Society
25: 255–265. doi:10.1111/j.1440-6055.1986.
tb01112.x
Cantrell, B. K. and R. W. Crosskey. 1989. Family
Tachinidae, pp. 733–784. In N. L. Evenhuis,
ed. Catalog of the Diptera of the Australasian
and Oceanian Regions. Bishop Museum
VOLUME 114, NUMBER 1
99

Special Publication 86. Bishop Museum
Press (Honolulu) and E. J. Brill (Leiden).
1155 pp.
Carlos, C. R., J. R. Costa, C. B. Ta
˜o, F. Alves, and
L. M. Torres. 2006. Natural parasitism of the
grape berry moth, Lobesia botrana (Denis &
Schiffermu
¨ller) at Regia
˜o Demarcada do
Douro. Boletin de Sanidad Vegetal, Plagas
32: 355–362.
Carlson, R. W. 1979a. Family Ichneumonidae,
pp. 315–740. In K. V. Krombein, P. D. Hurd,
Jr., D. R. Smith, and B. D. Burks, eds. Cat-
alog of Hymenoptera in American North of
Mexico. Vol. 1. Smithsonian Institution Press,
Washington, D.C. 1198 pp.
Carlson, R. W. 1979b. Superfamily Trigonaloidea,
pp. 1197–1198. In K. V. Krombein, P. D.
Hurd, Jr., D. R. Smith, and B. D. Burks, eds.
Catalog of Hymenoptera in American North
of Mexico. Vol. 1. Smithsonian Institution
Press, Washington, D.C. 1198 pp.
Carmean, D. and L. Kimsey. 1998. Phylogenetic
revision of the parasitoid wasp family Trig-
onalidae (Hymenoptera). Systematic Ento-
mology 23: 35–76. doi:10.1046/j.1365-3113.
1998.00042.x
Carne, P. B. 1969. On the population dynamics of
the Eucalyptus-defoliating sawfly Perg a a ffi -
nis affinis Kirby (Hymenoptera). Australian
Journal of Zoology 17: 113–141. doi:10.1071/
ZO9690113
Chadwick, C. E. and M. I. Nikitin. 1985. Records
of parasitism by members of the family Ta-
chinidae (Diptera: Tachinidae). Australian
Zoologist 21: 587–598.
Cha
´ves, T. H. A., D. I. A. Dı
´az, and G. R. A.
Bricen
˜o. 1993. Introduction to Venezuela and
biology of Cotesia plutellae Kurdj. (Hyme-
noptera: Braconidae), parasitoid of Plutella
xylostella L. (Lepidoptera: Plutellidae). Manejo
Integrado de Plagas No. 29: 24–27.
Chen, X. and J. He. 1997. Revision of the sub-
family Rogadinae (Hymenoptera: Braconi-
dae) from China. Zoologische Verhandelingen
308: 1–187.
Clausen, C. P. 1956. Biological control of insect
pests in the continental United States. United
States Department of Agriculture Technical
Bulletin No. 1139: 1–151.
Cock, M. J. W. 1987. Chalcidoidea. Notes on the
chalcidoids attacking South-east Asian Li-
macodidae, pp. 165–179. In M. J. W. Cock,
H. C. J. Godfray, and J. D. Holloways, eds.
Slug and Nettle Caterpillars: the Biology,
Taxonomy, and Control of the Limacodidae
of Economic Importance on Palms in South-
east Asia. CAB International, Wallingford,
Oxon, United Kingdom. 270 pp.
Cock,M.J.W.,H.C.J.Godfray,andJ.D.Holloway.
1987. Slug and Nettle Caterpillars: the Biology,
Taxonomy, and Control of the Limacodidae of
Economic Importance on Palms in South-east
Asia. CAB International, Wallingford, United
Kingdom. 270 pp.
Collins, C. 1933. The Oriental moth (Cnido-
campa flavescens Walk.) in Massachusetts
and the work of its newly introduced para-
site. Journal of Economic Entomology 26:
54–57.
Conant, P., A. H. Hara, L. M. Nakahara, and R. A.
Heu. 2002. Nettle caterpillar, Darna palli-
vitta Moore (Lepidoptera: Limacodidae).
New Pest Advisory No. 01–03. Hawaii De-
partment of Agriculture, Honolulu, HI.
Conant, P., A. H. Hara, W. T. Nagamine, C. M.
Kishimoto, and R. A. Heu. 2011. Nettle cat-
erpillar, Darna pallivitta Moore (Lepidop-
tera: Limacodidae). New Pest Advisory No.
01–03, updated. Hawaii Department of Ag-
riculture, Honolulu, HI.
Conant, P., C. K. Hirayama, C. M. Kishimoto,
and A. H. Hara. 2006. Trichogramma papil-
ionis (Nagarkatti), the first recorded Tri-
chogramma species to parasitize eggs in the
family Limacodidae. Proceedings of the Ha-
waiian Entomological Society 38: 133–135.
Cook, A. J. and G. C. Davis. 1891. Kerosene emul-
sion, some new insects. Bulletin of the Michigan
Agricultural Experiment Station 73: 1–16.
Coquillett, D. W. 1897. Revision of the Tachini-
dae of America north of Mexico. A family of
parasitic two-winged insects. United States
Department of Agriculture. Division of En-
tomology. Technical Series 7: 1–156.
da Costa Lima, A. 1962. Insetos do Brasil. 12°
Tomo. Hymeno
´pteros. 2a Parte. Escola Na-
cional de Agronomia (Se
´rie Dida
´ctica N°14),
Rio de Janeiro. 393 pp.
Cresson, E. T. 1865. On the Hymenoptera of
Cuba. Proceedings of the Entomological
Society of Philadelphia 4: 1–200.
Cresson, E. T. 1867. Descriptions of two new spe-
cies of Trigonalys. Proceedings of the Ento-
mological Society of Philadelphia 6: 351–352.
Cresson, E. T. 1869. List of the North American
species of the genus Aleiodes Wesmael.
Transactions of the American Entomological
Society 2: 377–382.
Cresson, E. T. 1872. Synopsis of the North Amer-
ican species belonging to the genera Leucospis,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
100

Smicra and Chalcis. Transactions of the
American Entomological Society 4: 29–60.
Crosskey, R. W. 1976. A taxonomic conspectus
of the Tachinidae (Diptera) of the Oriental
Region. Bulletin of the British Museum
(Natural History). Entomology Supplement
26: 1–357.
Cruttwell, R. E. 1969. The biology and mode of
parasitism of Uromacquartia trinitatis
Thompson (Diptera: Tachinidae). Common-
wealth Institute of Biological Control. Tech-
nical Bulletin 12: 20–28.
Culver, J. J. 1919. A study of Compsilura con-
cinnata, an imported tachinid parasite of the
gypsy moth and the brown-tail moth. Bulletin
of the United States Bureau of Entomology
766: 1–27.
Curran, C. H. 1930. Report on the Diptera col-
lected at the Station for the Study of Insects,
Harriman Interstate Park, N.Y. Bulletin of the
American Museum of Natural History 61:
21–115.
Cushman, R. A. 1925. The synonymy and generic
position of two North American ichneumon-
flies. Proceedings of the Entomological So-
ciety of Washington 27: 164–166.
Dasch, C. 1971. Ichneumon-flies of America
North of Mexico: 6. Subfamily Mesochor-
inae. Memoirs of the American Entomolog-
ical Institute 16: 1–376.
Davis, D. R. 1983. Limacodidae, pp. 66-67. In
R. W. Hodges, ed. Checklist of the Lepi-
doptera of America North of Mexico. E. W.
Classey Ltd and The Wedge Entomological
Research Foundation, London, England.
284 pp.
Deans, A. R., I. Miko, M. J. Yoder, K. S. Seltmann,
and The H. A. O. Consortium. [Wed Mar 17
10:45:36 -0500 2010]. Hymenoptera Glossary:
http://purl.oclc.org/NET/hymontology
Delvare, G. 1992. A reclassification of the Chal-
cidini with a checklist of the New World
species. Memoirs of the American Entomo-
logical Institute 53: 119–441, 443–466.
De Santis, L. 1979. Cata
´logo de los hime
´nopteros
calcidoideos de Ame
´rica al sur de los Estados
Unidos. Publicacio
´n Especial Comisio
´nde
Investigaciones Cientificas Provincia de
Buenos Aires. La Plata, Argentina. 488 pp.
De Santis, L. 1987. Himeno
´pteros parasito
´ides e
Hiperparasito
´ides de Anacraga sp. (Lepi-
doptera, Dalceridae) en Brasil. Revista Bra-
sileira de Entomologia 31: 97–99.
De Santis, L. and P. Fidalgo. 1994. Catalogo de
Himenopteros Calcidoideos. Serie de la
Academia Nacional de Agronomia y Vet-
erinaria No 13: 1–145.
Dessart, P. 1981. Definition de quleques sous-
genres de Ceraphron Jurine, 1807 (Hyme-
noptera, Ceraphronoidea, Ceraphronidae).
Bulletin de l’Institut Royal des Sciences
Naturelles de Belgique 53: 1–23.
Dessart, P. and P. Cancemi. 1986. Tableau di-
chotomique des genres de Ceraphronoidea
(Hymenoptera) avec commentaires et nou-
velles espe
´ces. Frustula Entomologica N. S.
7–8(20–21): 307–372.
Dowden, P. B. 1946. Parasitization of the Oriental
moth (Cnidocampa flavescens (Walk.)) by
Chaetexorista javana B. and B. Annals of the
Entomological Society of America 39: 225–241.
Dyar, H. G. 1897. On the white Eucleidae and the
larva of Calybia slossoniae (Packard). Jour-
nal New York Entomological Society 5: 121–
126 (plate 5).
Dyar, H. G. 1900. Life history of a South
American slug-caterpillar Sibine fusca Stoll.
Entomological News 11: 517–526.
Elkinton, J. S. and G. H. Boettner. 2004. The
effects of Compsilura concinnata, an in-
troduced generalist tachinid, on non-target
species in North America: a cautionary tale, pp.
4–14. In R. G. Van Driesche and R. Reardon,
eds. Assessing Host Ranges for Parasitoids and
Predators Used for Classical Biological Con-
trol: a Guide to Best Practice. United States
Deptartment of Agriculture Forest Health
Technology Enterprise Team, Morgantown,
West Virginia. FHTET-2004–03. 243 pp.
Elkinton, J. S. and A. M. Liebhold. 1990. Population
dynamics of gypsy moth in North America.
Annual Review of Entomology 35: 571–596.
doi:10.1146/annurev.en.35.010190.003035
Elkinton, J. S., D. Parry, and G. H. Boettner.
2006. Implicating an introduced generalist
parasitoid in the invasive browntail moth’s
enigmatic demise. Ecology 87: 2664–2672.
doi:10.1890/0012-9658(2006)87[2664:IAIGPI]
2.0.CO;2
Enderlein, G. 1926. Zur Kenntnis der Bombyliiden-
Subfamilie Systropodinae (Dipt.). Wiener En-
tomologische Zeitung 43: 69–92.
Epstein, M. 1988. An overview of slug caterpillar
moths (Lepidoptera: Limacodidae) with em-
phasis on genera in the New World Parasa
Group. Doctoral thesis, Department of En-
tomology, University of Minnesota, St. Paul,
Minnesota. xi +149 pp.
Epstein, M. 1996. Revision and phylogeny of the
limacodid group families, with evolutionary
VOLUME 114, NUMBER 1
101

studies on slug caterpillars (Lepidoptera: Zyg-
aenoidea). Smithsonian Contributions to Zool-
ogy 582: 1–102. doi:10.5479/si.00810282.582
Epstein, M. 1997. Parasa indetermina (Bois-
duval) (Lepidoptera: Limacodidae), a new
host for Systropus macer Loew (Diptera:
Bombyliidae). Proceedings of the Entomo-
logical Society of Washington 99: 585–586.
Evans, G. A and L. A. Stange. 1997. Parasitoids
associated with the brown citrus aphid, Tox-
optera citricida, in Florida (Insecta: Hyme-
noptera). Florida Department of Agriculture
and Consumer Service, Division of Plant In-
dustry, Entomology Circular No. 384 Gain-
esville, Florida. 5 pp.
Evenhuis, N. L. 1982. Catalog of primary types of
Bombyliidae (Diptera) in the entomological
collections of the Museum of Comparative
Zoology, with designations of lectotypes.
Breviora 469: 1–23.
Evenhuis, N. L. and D. J. Greathead. 1999. World
Catalog of Bee Flies (Diptera: Bombyliidae).
Backhuys Publishers, Leiden. xlviii +756 pp.
Evenhuis, N. L. and D. J. Greathead. 2003. World
Catalog of Bee Flies (Diptera: Bombyliidae)
Web site (http://hbs.bishopmuseum.org/
bombcat/). [accessed May 2011].
Fabricius, J. C. 1798. Supplementum Entomolo-
giae Systematicae. Proft and Storch, Copen-
hagen. 243 pp.
Finlayson, T. 1975. The cephalic structures and
spiracles of final-instar larvae of the sub-
family Campopleginae, tribe Campoplegini
(Hymenoptera: Ichneumonidae). Memoirs of
the Entomological Society of Canada 94: 1–
137. doi:10.4039/entm10794fv
Fletcher, D. S. and I. W. B. Nye. 1982. The Generic
Names of Moths of the World. Vol. 4. British
Museum (Natural History), London. 192 pp.
Fortier, J. C. and S. R. Shaw. 1999. Cladistics of
the Aleiodes lineage of the subfamily Roga-
dinae (Hymenoptera: Braconidae). Journal of
Hymenoptera Research 8: 204–237.
Franklin, H. J. 1919. Seventh report of the cran-
berry substation from 1917 to 1919. Bulletin
of the Massachusetts Agricultural Experi-
ment Station 192: 129.
Fry, J. M. 1989. Natural enemy databank, 1987. A
catalogue of natural enemies of arthropods
derived from records in the CIBC Natural
Enemy Databank, p. 94. CAB International,
Wallingford, Oxford, UK. 185 pp.
Gadd, C. H., W. T. Fonseka, and D. J. W. Rana-
weera. 1946. Parasites of tea nettlegrubs with
special reference to Platyplectrus natadea
Ferrie
`re and Autoplectrus taprobanes Gadd.
Ceylon Journal of Science (B) 23: 81–94.
Gahan, A. B. 1934. Descriptions of some new
species of Chalcidoidea from Cuba and Pu-
erto Rico. Memorias de la Sociedad Cubana
de Historia Natural, Habana 8: 125–134.
Gates, M. and R. Burks. 2003. Hosts and ranges.
Chalcid Forum 25: 5 (newsletter).
Gates, M. W. and T. Stoepler. 2010. A new spe-
cies of Alveoplectrus Wijesekara and Schauff
(Hymenoptera: Chalcidoidea: Eulophidae)
parasitic on Limacodidae (Lepidoptera).
Proceedings of the Entomological Society of
Washington 112: 207–213. doi:10.4289/0013-
8797-112.2.207
Gauld, I. D. 1984. An introduction to the Ich-
neumonidae of Australia; with a contribution
on Metopiinae by M. G. Fitton. British Mu-
seum (Natural History), no. 895: 1–413.
Gauld, I. D. 1988. Evolutionary patterns of host
utilization by ichneumonoid parasitoids (Hy-
menoptera: Ichneumonidae and Braconidae).
Biological Journal of the Linnean Society.
Linnean Society of London 35: 351–377.
doi:10.1111/j.1095-8312.1988.tb00476.x
Gauld, I. D. 1995. 12.1 Ichneumonidae, pp. 390–
431. In P. E. Hanson and I. D. Gauld, eds. The
Hymenoptera of Costa Rica. Oxford Uni-
versity Press, New York.
Gauld, I. and B. Bolton. 1988. The Hymenoptera.
Oxford University Press, London, England.
332 pp.
Gauld, I. and J. Ferna
´ndez-Triana. 2010. Type
condition and generic placement of Cuban
species of Ichneumonidae described by
Cresson and collected by Gundlach. Zootaxa
2394: 41–50.
Gelhaus, J. K. 1987. A detritivore Tipula (Diptera:
Tipulidae) as a secondary host of Poe cil o go-
nalos costalis (Hymenoptera: Trigonalidae).
Entomological News 98: 161–162.
Genty, P. 1972. Morfologı
´a y biologı
´adeSibine
fusca Stoll, Lepidoptero defoliador de la palma
de aceite en Colombia. Ole
´agineux 27: 65–71.
Genty, P., R. Desmier, J. P. Morin, and C. A.
Korytkowski. 1978. Les ravaguers du palmier
a huile en Amerique Latine. Ole
´agineux 33:
325–419.
Gibson, G. A. P. 1997. Morphology and termi-
nology. Chapter 2, pp. 16–44. In G. A. P.
Gibson, J. T. Huber, and J. B. Woolley, eds.
Annotated Keys to the Genera of Nearctic
Chalcidoidea (Hymenoptera). Research Press,
National Research Council of Canada, Ottawa,
Ontario. 764 pp.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
102

Gibson, G. A. P., J. T. Huber, and J. B. Woolley.
1997. Annotated Key to the Genera of
Neartic Chalcidoidea (Hymenoptera). Re-
search Press. National Research Council of
Canada, Ottawa, Ontario. 764 pp.
Girault, A. 1911. On the identity of the most
common species of the family Trichog-
rammatidae (Hymenoptera). Bulletin of the
Wisconsin Natural History Society 9: 135–165.
Girault, A. 1916. Descriptions of miscellaneous
chalcid-flies. Insecutor Inscitiae Menstruus 4:
109–120.
Godfray, H. C. J. and M. S. Chan. 1990. How
insecticides trigger single-stage outbreaks in
tropical pests. Functional Ecology 4: 329–
337. doi:10.2307/2389594
Graham, M. W. R. de V. 1969. The Pteromalidae
of north-western Europe (Hymenoptera:
Chalcidoidea). Bulletin of the British Mu-
seum (Natural History). Historical Series
(Supplement 16): 1–908 (Natural History).
Greathead, D. J. 1987. Bombyliidae. A summary
of the recorded parasitoids of South-east
Asian Limacodidae, pp. 195–196. In M. J. W.
Cock, H. C. J. Godfray, and J. D. Holloways,
eds. Slug and Nettle Caterpillars: the Bi-
ology, Taxonomy, and Control of the Lima-
codidae of Economic Importance on Palms in
South-east Asia. CAB International, Wall-
ingford, Oxon, United Kingdom. 270 pp.
Guimara
˜es, J. H. 1980. Revision of the South
American Uramyini (Diptera, Tachinidae).
Pape
´is Avulsos de Zoologia 33: 191–219.
Hall, J. 1981. Bombyliidae, pp. 589–602. In J. F.
McAlpine, B. V. Peterson, G. E. Shewell, H. J.
Teskey, J. R. Vockeroth, and D. M. Wood, eds.
Manual of Nearctic Diptera, Vol. 1. Agricul-
ture Canada Monograph, Research Branch,
Agriculture Canada, Ottawa, Ontario.
Hansson, C. 1987. New records of Swedish Eu-
lophidae and Pteromalidae (Hymenoptera:
Chalcidoidea), with data on host species.
Entomologisk Tidskrift 108: 168–173.
Hansson, C. 1994. Re-evaluation of the genus
Closterocerus Westwood (Hymenoptera:
Eulophidae), with a revision of the Nearctic
species. Entomologica Scandinavica 25: 1–
25. doi:10.1163/187631294X00018
Hansson, C. 1995. Revision of the Nearctic species
of Neochrysocharis Kurdjumov (Hymenoptera:
Eulophidae). Entomologica Scandinavica 26:
27–46.
Harris, R. 1979. A glossary of surface sculptur-
ing. Occasional Papers in Entomology, No.
28. California State Department of Food and
Agriculture, Sacramento, California, U.S.A.
31 pp.
Harrison, J. O. 1963. The natural enemies of some
banana insect pests in Costa Rica. Journal of
Economic Entomology 56: 282–285.
Harvey, J., L. M. A. Witjes, and R. Wagenaar.
2004. Development of hyperparasitoid wasp
Lysibi a nana (Hymenoptera: Ichneumonidae)
in a multitrophic framework. Environmental
Entomology 33: 1488–1496. doi:10.1603/0046-
225X-33.5.1488
Harvey, J. A., L. E. M. Vet, L. M. A. Witjes, and
T. M. Bezemer. 2006. Remarkable similarity
in body mass of a secondary hyperparasitoid
Lysibia nana and its primary parasitoid host
Cotesia glomerata emerging from cocoons of
comparable size. Archives of Insect Bio-
chemistry and Physiology 61: 170–183.
doi:10.1002/arch.20080
Harvey, J. A., R. Wagenaar, and M. Bexemer.
2009a. Interactions to the fifth trophic level:
secondary and tertiary parasitoid wasps
show extraordinary efficiency in utilizing
host resources. Journal of Animal Ecology
78: 686–692. doi:10.1111/j.1365-2656.2008.
01516.x
Harvey,J.A.,R.Wagenaar,andT.M.Bezemer.
2009b. Life-history traits in closely related
secondary parasitoids sharing the same primary
parasitoid host: evolutionary opportunities and
constraints. Entomologia Experimentalis et
Applicata 132: 155–164. doi:10.1111/j.1570-
7458.2009.00882.x
HDOA. 2010. Natural enemy of stinging cater-
pillar to be released on O’ahu. News Re-
lease10-07. Available from: http://hawaii.gov/
hdoa/news/2010-news-releases/natural-enemy-
of-stinging-caterpillar-to-be-released-on-o-ahu.
[accessed May 15, 2011].
HDOA. 2011. Kaua’i residents asked to report
sightings of stinging caterpillar. News Re-
lease11-11. Available from: http://hawaii.gov/
hdoa/news/news-releases-2011/kaua-i-residents-
asked-to-report-sightings-of-stinging-caterpillar.
[accessed June 8, 2011].
Heraty, J. and D. Hawks. 1998. Hexamethyldisi-
lazane – a chemical alternative for drying
insects. Entomological News 109: 369–374.
Herting, B. 1960. Biologie der westpala
¨arktischen
Raupenfliegen. Dipt., Tachinidae. Monographien
zur Angewandte Entomologie 16: 1–188.
Herting, B. 1972. Die Typenexemplare der von
Meigen (1824–1838) beschriebenen Rau-
penfliegen (Dipt. Tachinidae). Stuttgarter
Beitra
¨ge zur Naturkunde 243: 1–15.
VOLUME 114, NUMBER 1
103

Herting, B. 1976. Lepidoptera, Part 2 (Macro-
lepidoptera). A catalogue of parasites and
predators of terrestrial arthropods. Section A.
Host or Prey/Enemy, 7: 3 Commonwealth
Agricultural Bureaux, Commonwealth In-
stitute of Biological Control.
Hoong, H. W. and C. K. Y. Hoh. 1992. Major
pests of oil palm and their occurrence in Sa-
bah. Planter 68: 193–210.
Horstmann, K. 2003. Revisionen von Schlupfwespen-
Arten VII. Mitteilungen der Mu
¨nchner En-
tomologischen Gesellschaft 93: 25–37.
Horstmann, K. 2008. Neue westpalaarkitsche arten
der Campopleginae (Hymenoptera: Ichneu-
monidae). Zeitschrift der Arveitsgemeinschaft
Oesterreichischer Entomologen. 60: 3–27.
Horstmann, K. 2009. Typenrevisionen der von
Kiss beschriebenen Taxa der Ichneumonidae
III. Verschiedene Unterfamilien (Hymenop-
tera, Ichneumonidae). Linxer Biologische
Beitra
¨ge 41: 673–689.
Howard, L. O. 1889. The hymenopterous para-
sites of North American butterflies, pp. 1869–
1896. In S. H. Scudder, ed. Butterflies of
Eastern United States and Canada, Vol. III.
W. Wheeler, Cambridge, Massachusetts.
Hull, F. 1973. Bee Flies of the World. The Genera
of the Family Bombyliidae. Smithsonian In-
stitution Press, Washington, DC. 687 pp.
Ichiki, R. and H. Shima. 2003. Immature life of
Compsilura concinnata (Meigen) (Diptera:
Tachinidae). Annals of the Entomological
Society of America 96: 161–167. doi:10.
1603/0013-8746(2003)096[0161:ILOCCM]
2.0.CO;2
Janzen, D. H. and W. Hallwachs. 2009. Dynamic
database for an inventory of the macro-
caterpillar fauna, and its food plants and para-
sitoids, of Area de Conservacion Guanacaste
(ACG), northwestern Costa Rica (nn-SRNP-
nnnnn voucher codes). Available at http://janzen.
sas.upenn.edu. [accessed 15-June-2010].
Jerman, E. J. and I. Gauld. 1988. Casinaria,
a paraphyletic ichneumonid genus (Hyme-
noptera), and a revision of the Australian
species. Journal of Natural History 22: 589–
609. doi:10.1080/00222938800770401
Johnson, C. W. 1925. Fauna of New England. 15.
List of the Diptera or two-winged flies. Oc-
casional Papers of the Boston Society of
Natural History 7: 1–326.
Kellogg, S. K., L. S. Fink, and L. P. Brower. 2003.
Parasitism of native luna moths, Actias luna
(L.) (Lepdioptera: Saturniidae) by the in-
troduced Compsilura concinnata (Meigen)
(Diptera: Tachinidae) in central Virginia, and
their hyperparasitism by Trigonalid wasps
(Hymenoptera: Trigonalidae). Environmental
Entomology 32: 1019–1027. doi:10.1603/
0046-225X-32.5.1019
Kerr, P. H., E. M. Fisher, and M. L. Buffington.
2008. Dome lighting for insect imaging under
a microscope. American Entomologist 54:
198–200.
Kits, J. H., S. A. Marshall, and N. L. Evenhuis.
2008. The bee flies (Diptera: Bombyliidae) of
Ontario, with a key to the species of eastern
Canada. Canadian Journal of Arthropod
Identification No. 6, March 06, 2008, available
online at http://www.biology.ualberta.ca/bsc/
ejournal/kme_06/kme_06.html, doi: 10.3752/
cjai.2008.06. [accessed 15-June-2010].
Krauth, S. J. and A. H. Williams. 2006. Notes on
Taeniogonalos gundlachii Hymenoptera:
Trigonalidae) from Wisconsin. Great Lakes
Entomologist 34: 54–58.
Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and
B. D. Burks. 1979. Catalog of Hymenoptera
in America North of Mexico. Vol. 1. Smith-
sonian Institution Press, Washington, D.C.
xvi +1198 pp.
Kula, R. R., J. T. Lill, S. M. Murphy, and T.
Stoepler. (2009) 2010. The first host records
for the Nearctic species Triraphis discoideus
(Hymenoptera: Braconidae: Rogadinae). En-
tomological News 120: 380–386. doi:10.3157/
021.120.0406
Kulman, H. M. 1965. Natural control of eastern tent
caterpillar and notes on its status as a forest pest.
Journal of Economic Entomology 58: 66–70.
La Salle, J. 1993. Parasitic Hymenoptera, bi-
ological control, and biodiversity, pp. 197–
215. In J. La Salle and I. D. Gauld, eds.
Hymenoptera and Biodiversity. CAB In-
ternational, Wallingford, UK. 348 pp.
Le Corff, J. and R. J. Marquis. 1999. Differences
between undestory and canopy in herbivore
community composition and leaf quality for
two oak species in Missouri. Ecological En-
tomology 24: 46–58. doi:10.1046/j.1365-
2311.1999.00174.x
Lill, J. T. 2008. Caterpillar-host plant relationships
recorded from Plummers Island, Maryland
(Insecta: Lepidoptera). Bulletin of the Bi-
ological Society of Washington 15: 75–79.
doi:10.2988/0097-0298(2008)15[75:CPRRFP]
2.0.CO;2
Lill, J. T., R. J. Marquis, R. E. Forkner, J. Le
Corff, N. Holmberg, and N. A. Barber. 2006.
Leaf pubescence affects distribution and
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
104

abundance of generalist slug caterpillars
(Lepidoptera: Limacodidae). Environmental
Entomology 35: 797–806. doi:10.1603/0046-
225X-35.3.797
Lingren, P. D. 1977. Campoletis sonorensis:
maintenance of a population on tobacco
budworms in a field cage. Environmental
Entomology 6: 72–76.
Lourenca
˜o, A. L., L. O. de Carvalho, and D. H. de
Campos Lasca. 1989. Anacraga citrinopsis
Dyar (Lepidoptera: Dalceridae) em Mamoneira
no Estado de Sa
˜o Paulo. Bragantia 48: 109–112.
Marconato, G., M. M. Dias, and A. M. Penteado
Dias. 2008. Larvas de Geometridae (Lepi-
doptera) e seus parasito
´ides, associadas a
Erythroxylum microphyllum St.- Hilaire
(Erythroxylacae). Revista Brasileira de En-
tomologia 52: 296–299. doi:10.1590/S0085-
56262008000200010
Marsh, P. M. 1979. Braconidae, Aphidiidae,
Hybrizontidae, pp. 144–313. In K. V. Krom-
bein, P. D. Hurd, Jr., D. R. Smith, and B. D.
Burks, eds. Catalog of Hymenoptera in
America North of Mexico. Vol. 1. Smithsonian
Institution Press, Washington, D.C. 1198 pp.
Marsh, P. M. and S. R. Shaw. 1998. Revision of
North American Aleiodes Wesmael (Part 3):
the seriatus (Herrich-Schaeffer) species-group
(Hymenoptera: Braconidae: Rogadinae). Pro-
ceedings of the Entomological Society of
Washington 100: 395–408.
McAlpine, J. F. 1981. Morphology and termi-
nology—adults. In J. F. McAlpine, B. V.
Peterson, G. E. Shewell, H. J. Teskey, J. R.
Vockeroth, and D. M. Wood, eds. Manual of
Nearctic Diptera. Volume 1. Agriculture
Canada Monograph 27: 9–63.
McCullough, D. G., S. A. Katovich, D. L. Mahr,
D. D. Neumann, C. S. Sadof, and M. J.
Raupp. 1999. Biological control of insect
pests in forested ecosystems: a manual for
foresters, Christmas tree growers and land-
scapers. Michigan State University Extension
Bulletin E-2679: 1–123.
Meigen, J. W. 1824. Systematische Beschreibung
der bekannten europa
¨ischen zweiflu
¨geligen
Insekten. Vierter Theil. Schulz-Wundermann,
Hamm. xii +428 pp. +pls. 33–41.
Mexzo
´n, R. G., C. M. Chinchilla, and D. Sala-
manca. 1996. The biology of Sibine mega-
somoides Walker (Lepidoptera, Limacodidae):
observations of the pest in oil palm in Costa
Rica. ASD Oil Palm Papers 12: 1–10.
Morris, R. F. 1976. Relation of mortality caused
by parasites to the population density of
Hyphantria cunea. Canadian Entomologist
108: 1291–1294.
Muesebeck, C. F. W. 1920. A revision of the
North American species of ichneumon-flies
belonging to the genus Apanteles. Pro-
ceedings of the United States National Mu-
seum 58: 483–576.
Muesebeck, C. F. W. 1931. Descriptions of a new
genus and eight new species of ichenumon-
flies with taxonomic notes. Proceedings of
the United States National Museum 79 (Art.
16) [=No. 2882]: 1–16.
Murphy, S. M., J. T. Lill, and D. R. Smith. 2009.
A scattershot approach to host location:
uncovering the unique life history of the
trigonalid hyperparasitoid Orthogonalys pul-
chella (Cresson). American Entomologist 55:
82–87.
Murphy, S. M., S. M. Leahy, L. S. Williams, and
J. T. Lill. 2010. Stinging spines protect slug
caterpillars (Limacodidae) from multiple
generalist predators. Behavioral Ecology 21:
153–160. doi:10.1093/beheco/arp166
Nazemi, J., A. A. Talebi, S. E. Sadeghi, G. Me-
lika, and A. Lozan. 2008. Species richness of
oak gall wasps (Hymenoptera: Cynipidae)
and identification of associated inquilines and
parasitoids on two oak species in western
Iran. North-Western Journal of Zoology 4:
189–202.
Niehuis, O., S. H. Yen, C. M. Naumann, and B.
Misof. 2006. Higher phylogeny of zygaenid
moths (Insecta: Lepidoptera) inferred from
nuclear and mitochondrial sequence data and
the evolution of larval cuticular cavities for
chemical defence. Molecular Phylogenetics
and Evolution 39: 812–829. doi:10.1016/j.
ympev.2006.01.007
Noyes, J. S. 2003. Universal Chalcidoidea Data-
base. World Wide Web electronic publication.
www.nhm.ac.uk/entomology/chalcidoids/index.
html. [accessed 15-June-2010].
O’Hara, J. E., H. Shima, and C.-T. Zhang. 2009.
Annotated catalogue of the Tachinidae (Insecta:
Diptera) of China. Zootaxa 2190: 1–236.
O’Hara, J. E. and D. M. Wood. 2004. Catalogue
of the Tachinidae (Diptera) of America north
of Mexico. Memoirs on Entomology, In-
ternational 18: 1–410.
Ostmark, H. E. 1974. Economic insect pests of
bananas. Annual Review of Entomology 19:
161–176. doi:10.1146/annurev.en.19.010174.
001113
Painter, R. and E. Painter. 1963. A review of the
subfamily Systropinae (Diptera: Bombyliidae)
VOLUME 114, NUMBER 1
105

in North America. Journal of the Kansas En-
tomological Society 36: 278–348.
Papp, J. 2004. Type specimens of the braconid
species by Gy. Sze
´pligeti deposited in the
Hungarian Natural History Museum (Hy-
menoptera: Braconidae). Annals Historico-
Naturales Musei Nationalis Hungarici 96:
153–223.
Patton, C. N. 1958. A catalogue of the Larvae-
voridae of Florida. Florida Entomologist 41:
29–39, 77–89. doi:10.2307/3492632
Peck, O. 1963. A catalogue of the Nearctic
Chalcidoidea (Insecta; Hymenoptera). Cana-
dian Entomologist (Supplement) 30: 1–1420.
Peck, O. 1985. The taxonomy of the Nearctic
species of Pediobius (Hymenoptera: Eulo-
phidae), especially Canadian and Alaskan
forms. Canadian Entomologist 117: 647–704.
doi:10.4039/Ent117647-6
Petrice, T. R., J. S. Strazanac, and L. Butler. 2004.
A survey of hymenopteran parasitoids of
forest macrolepidoptera in the central Appa-
lachians. Journal of Economic Entomology
97: 451–459. doi:10.1603/0022-0493-97.2.451
Pinto, J. 1998. Systematics of the North Ameri-
can species of Trichogramma Westwood
(Hymenoptera: Trichogrammatidae). Mem-
oirs of the Entomological Society of Wash-
ington 22: 1–287.
Pinto, J. D., G. R. Platner, and E. R. Oatman.
1978. Clarification of the identity of several
common species of North American Tri-
chogramma (Hymenoptera: Trichogramma-
tidae). Annals of the Entomological Society
of America 71: 169–180.
Pluciennik, Z. and R. W. Olszak. 2010. The role
of parasitoids in limiting the harmfulness of
leafrollers in apple orchards. Journal of Plant
Protection Research 50: 1–8. doi:10.2478/
v10045-010-0001-9
Provancher, L. 1875. Les Ichneumonides de
Que
´bec. Naturaliste Canadien 7: 309–317.
Quicke, D. L. J. 1997. Parasitic Wasps. Chapman
and Hall, New York. 470 pp.
Quicke, D. L. J. and M. R. Shaw. 2005. First host
records for the rogadine genera Rogasodes
Chen and He and Canalirogas van Achter-
berg and Chen (Hymenoptera: Braconidae)
with description of a new species and survey
of mummy types within Rogadinae s. str.
Journal of Natural History 39: 3525–3542.
doi:10.1080/00222930500392782
Raff, J. W. 1934. Observations on sawflies of the
genus Perga , with notes on some reared pri-
mary parsties of the families Trigonalidae,
Ichneumonidae, and Tachinidae. Proceedings
of the Royal Society of Victoria 47: 54–77.
Raizenne, H. 1952. Forest Lepidoptera of south-
ern Ontario and their parasites received and
reared at the Ottawa Forest Insect Survey
Laboratory from 1937 to 1948. Canada De-
partment of Agriculture, Science Service,
Division of Forest Biology, Ottawa. v +277 pp.
Reinhard, H. 1865. Beitra
¨ge zur Kenntnifs einiger
Braconiden-Gattungen. Berliner Entomologische
Zeitschrift 9: 243–267. doi:10.1002/mmnd.
18650090309
Reinhard, H. J. 1935. New genera and species of
American muscoid flies (Tachinidae: Dip-
tera). Annals of the Entomological Society of
America 28: 160–173.
dos Reis Fernandes, L. B., M. M. Dias Filho, M.
A. Fernandes, and A. M. Penteado-Dias.
2010. Ichneumonidae (Hymenoptera) para-
sitoids of Lepidoptera caterpillars feeding on
Croton floribundus Spreng (Euphorbiaceae).
Revista Brasileira de Entomologia 54: 263–
269. doi:10.1590/S0085-56262010000200009
Riley, C. V. and L. O. Howard. 1890. Some of the
bred parasitic Hymenoptera in the National
Collection. Insect Life 2: 348–353.
Robertson, C. 1928. Flowers and Insects. Lists of
Visitors of 453 Flowers. Science Press
Printing Co., Lancaster, PA. 221 pp.
ROBO (Release of Beneficial Organisms) Data-
base. Introduced Parasites and Predators Re-
leased in the U. S. and Territories - by Target
Host/Prey, 1981–1985. Biological Control
Documentation Center, National Invertebrate
Genetic Resources, http://www.ars-grin.gov/
nigrp/menu.html#link1981. [accessed 5-June-
2010].
Rohwer, S. 1915. Descriptions of new species
of Hymenoptera. Proceedings of the United
States National Museum 49: 205–249. doi:
10.5479/si.00963801.2105.205
Ruberson, J. R. and J. B. Whitfield. 1996. Fac-
ultative egg-larval parasitism of the beet ar-
myworm, Spodoptera exigua (Lepidoptera:
Noctuidae) by Cotesia marginiventris (Hy-
menoptera: Braconidae). Florida Entomolo-
gist 79: 296–302. doi:10.2307/3495577
Sabrosky, C. W. and P. H. Arnaud, Jr. 1965. Family
Tachinidae (Larvaevoridae), pp. 961–1108. In
A. Stone, C. W. Sabrosky, W. W. Wirth, R. H.
Foote, and J. R. Coulson, eds. A Catalog of the
Diptera of America North of Mexico. United
States Department of Agriculture. Agriculture
Handbook 276. U.S. Government Printing
Office, Washington D.C. 1696 pp.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
106

Say, T. 1835. Descriptions of new North Ameri-
can Hymenoptera, and observations on some
already described. Boston Journal of Natural
History 1(3): 210–305.
Schaffner, J. V., Jr. 1959. Microlepidoptera and
their parasites reared from field collections in
the northeastern part of the United States.
United States Department Agriculture Mis-
cellaneous Publications 767: 1–97.
Schaffner, J. V., Jr. and F. L. Griswold. 1934.
Macrolepidoptera and their parasites reared
from field collections in the northeastern part
of the United States. United States De-
partment Agriculture Miscellaneous Publi-
cations 188: 1–160.
Schauff, M. E., J. LaSalle, and L. D. Coote. 1997.
Eulophidae, pp. 327–429. In G. A. P. Gibson,
J. T. Huber, and J. B. Woolley, eds. Annotated
Keys to the Genera of Nearctic Chalcidoidea
(Hymenoptera). Research Press. National
Research Council of Canada, Ottawa, On-
tario. 764 pp.
Sellers, W. F. 1943. The Nearctic species of
parasitic flies belonging to Zenillia and
allied genera. Proceedings of the United
States National Museum 93 [=No. 3157]:
1–108.
Sharkey, M. J. 1997. Key to New World sub-
families of the family Braconidae, pp. 39–45.
In R. A. Wharton, P. M. Marsh, and M. J.
Sharkey, eds. Manual of the New World
Genera of the Family Braconidae (Hymenop-
tera). Special Publication No. 1. International
Society of Hymenopterists, Washington, D. C.
Sharkey, M. J. and R. A. Wharton. 1997. Mor-
phology and terminology, pp. 19–37. In R. A.
Wharton, P. M. Marsh, and M. J. Sharkey, eds.
Manual of the New World Genera of the
Family Braconidae (Hymenoptera). Interna-
tional Society of Hymenopterists, Special
Publication No. 1. Washington, D. C. 439 pp.
Shaw, M. R. 1990. Parasitoids of European but-
terflies and their study, pp. 449–479. In O.
Kudrna, ed. Butterflies of Europe, Vol. 2.
Aula-Verlag, Wiesbaden. 557 pp.
Shaw, M. R. and R. R. Askew. 1976. Parasites,
pp. 240–256. In J. Heath, ed. The Moths and
Butterflies of Great Britain and Ireland. 1.
Micropterigidae – Heliozelidae. Blackwell
Scientific Publications and Curwen Press,
London. 343 pp.
Shaw, S. R. 1983. A taxonomic study of the Ne-
arctic Ascogaster and a description of a new
genus Leptodrepana (Hymenoptera: Braco-
nidae). Entomography 2: 1–54.
Shaw, S. R. 1997a. Subfamily Cheloninae, pp.
193–201. In R. A. Wharton, P. M. Marsh,
and M. J. Sharkey, eds. Manual of the New
World Genera of the Family Braconidae
(Hymenoptera). Special Publication No. 1.
International Society of Hymenopterists,
Washington, D. C. 439 pp.
Shaw, S. R. 1997b. Subfamily Rogadinae s.s, pp.
403–412. In R. A. Wharton, P. M. Marsh,
and M. J. Sharkey, eds. Manual of the New
World Genera of the Family Braconidae
(Hymenoptera). Special Publication No. 1.
International Society of Hymenopterists,
Washington, D. C. 439 pp.
Shaw, S. R. 2006. Aleiodes wasps of eastern
forests: a guide to parasitoids and associated
mummified caterpillars. Forest Health Tech-
nology Enterprise Team/USDA/Forest Ser-
vice. Morgantown, West Virginia.121 pp.
Shaw, S. R., P. M. Marsh, and J. C. Fortier. 1997.
Revision of North American Aleiodes Wes-
mael (Part 1): the pulchripes Wesmael spe-
cies-group in the New World (Hymenoptera:
Braconidae, Rogadinae). Journal of Hyme-
noptera Research 6: 10–35.
Shaw, S. R., P. M. Marsh, and J. C. Fortier. 1998.
Revision of North American Aleiodes Wes-
mael (Part 4): the albitibia Herrich-Schaeffer
and praetor Reinhard species-groups (Hy-
menoptera: Braconidae: Rogadinae) in the
New World. Proceedings of the Entomolog-
ical Society of Washington 100: 553–565.
Shima, H. 1984. Study on the tribe Blondeliini
from Japan (Diptera, Tachinidae). V. The
genera Blondelia Robineau-Desvoidy and
Compsilura Bouche
´. Kontyu
ˆ52: 540–552.
Shima, H. 2006. A host-parasite catalog of
Tachinidae (Diptera) of Japan. Makunagi/Acta
Dipterologica, Supplement 2: 1–171.
Short, J. R. T. 1978. The final larval instars of the
Ichneumonidae. Memoirs of the American
Entomological Institute 25: 1–508.
Singh, B. P., R. P. Singh, and V. D. Verma. 1988.
New record of Aroplectrus dimerus Linn. and
Platyplectrus sp. as larval parasite of slug
caterpillar (Parasa bicolor Walk) from U. P.
Farm Science Journal 3: 199–200.
Smith, D. R. 1996. Trigonalyidae (Hymenoptera)
in the eastern United States: seasonal flight
activity, distribution, hosts. Proceedings of
the Entomological Society of Washington 98:
109–118.
Smith, D. R. and I. C. Stocks. 2005. A new trig-
onalid wasp (Hymenoptera: Trigonalidae)
from eastern North America. Proceedings of
VOLUME 114, NUMBER 1
107

the Entomological Society of Washington
107: 530–535.
Smith, H. E. 1912. A contribution on North Amer-
ican dipterology. Proceedings of the Entomo-
logical Society of Washington 14: 118–127.
Smith, O. J., A. G. Diboll, and H. Rosenberger.
1955. Laboratory studies of Pelecystoma
harrisinae (Ashmead), an adventive braconid
parasite of the western grape leaf skeleton-
izer. Annals of the Entomological Society of
America 48: 232–237.
Smith, S. M. 1996. Biological control with Tri-
chogramma: advances, successes, and potential
of their use. Annual Review of Entomology
41: 375–406. doi:10.1146/annurev.en.41.010196.
002111
Stireman III, J. O. and M. S. Singer. 2003a. De-
terminants of parasitoid-host associations: in-
sights from a natural tachinid-lepidopteran
community. Ecology 84: 296–310. doi:10.1890/
0012-9658(2003)084[0296:DOPHAI]2.0.CO;2
Stireman III, J. O. and M. S. Singer. 2003b. What
determines host range in parasitoids? An
analysis of a tachinid parasitoid community.
Oecologia 135: 629–638.
Strand, M. R. 1986. The physiological in-
teractions of parasitoids with their hosts and
their influence on reproductive strategies, pp.
97–136. In J. K. Waage and D. J. Greathead,
eds. Insect Parasitoids. Academic Press,
London, UK. 389 pp.
Strazanac, J. S., C. D. Plaugher, T. R. Petrice, and L.
Butler. 2001. New Tachinidae (Diptera) host
records of eastern North American forest can-
opy Lepidoptera: baseline data in a Bacillus
thuriengiensis variety kurstaki nontarget study.
Journal of Economic Entomology 94: 1128–
1134. doi:10.1603/0022-0493-94.5.1128
Thompson, W. R. 1954. A catalogue of the par-
asites and predators of insect pests. Section 2.
Host parasite catalogue. Part 3. Hosts of the
Hymenoptera (Calliceratid to Evaniid), pp.
191–332. Commonwealth Agricultural Bu-
reaux, Commonwealth Institute of Biological
Control, Ottawa. 141 pp.
Thomson, C. G. 1878. Hymenoptera Scandina-
viae 5. Pteromalus (Svederus) continuatio.
Typis Expressit Haqv. Ohlsson, Lund. 307 pp.
Tobias, V. I. 1976. Braconids of Caucasus.
Nauka, Leningrad. 287 pp. [In Russian]
Tothill, J. D. 1922. The natural control of the fall
webworm (Hyphantria cunea Drury) in Can-
ada together with an account of its several
parasites. Bulletin of the Canada Department
of Agriculture, N. Ser. 3: 1–107 +6 plates.
Townes, H. K. 1956. The Nearctic species of
trigonalid wasps. Proceedings of the United
States National Museum 106: 295–304.
Townes, H. K. 1970. The genera of Ichneumo-
nidae, part 2. Memoirs of the American En-
tomological Institute 12: 1–537.
Townes, H. K. 1983. Revisions of twenty genera
of Gelini (Ichneumonidae). Memoirs of the
American Entomological Institute 35: 1–
281.
Townes, H. and M. Townes. 1962. Ichneumon-
flies of America North of Mexico: 3. Sub-
family Gelinae, tribe Mesostenini. United
States National Museum Bulletin 216: 1–
602. doi:10.5479/si.03629236.216.1-3
Townes, H. and M. Townes. 1966. A catalogue
and reclassification of the Neotropic Ichneu-
monidae. Memoirs of the American Ento-
mological Institute 8: 1–367.
Townes, H., S. Momoi, and M. Townes. 1965. A
catalogue and reclassification of the eastern
Palearctic Ichneumonidae. Memoirs of the
American Entomological Institute 5: i–v +1–
661.
Townsend, C. H. T. 1892. An Aporia bred from
Limacodes sp. Psyche 6: 275–276. doi:10.1155/
1892/17351
Townsend, C. H. T. (1918) 1919. New muscoid
genera, species and synonymy (Diptera).
[concl.] Insecutor Inscitiae Menstruus 6:
157–182.
Tschorsnig, H.-P. and V. A. Richter. 1998. Family
Tachinidae, pp. 691–827. In L. Papp and B.
Darvas, eds. Contributions to a Manual of
Palaearctic Diptera (With Special Reference to
Flies of Economic Importance), Vol. 3. Higher
Brachycera. Science Herald, Budapest. 880 pp.
Valerio, A. A. 2006. Some taxonomic notes on
named Rogas Nees species (Hymenoptera:
Braconidae: Rogadinae) for the New World.
Methods in Ecology and Systematics 1: 37–43.
Viereck, H. L. 1913. Descriptions of ten new
genera and twenty-three new species of ich-
neumon-flies. Proceedings of the United
States National Museum 44: 555–568.
doi:10.5479/si.00963801.1968.555
Viereck, H. L. 1925. A preliminary revision of the
Campopleginae in the Canadian National
Collection, Ottawa. Canadian Entomologist
57: 176–181, 198–204, 223–228, 296–303.
doi:10.4039/Ent57176-7
Viereck, H. L. 1926. A preliminary revision of the
Campopleginae in the Canadian National
Collection, Ottawa. Canadian Entomologist
58: 2–8, 30–38, 71–78, 122–130, 143–149,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
108

176–182, 196–201, 219–225, 257–260, 276–
283. doi:10.4039/Ent582-1
Wagner, D. L. 2005. Caterpillars of Eastern North
America. Princeton University Press, New
Jersey. 496 pp.
Wahl, D. B. 1993a. Cladistics of the genera of
Mesochorinae (Hymenoptera: Ichneumoni-
dae). Systematic Entomology 18: 371–387.
doi:10.1111/j.1365-3113.1993.tb00673.x
Wahl, D. B. 1993b. Family Ichneumonidae, pp.
395–448. In H. Goulet and J. T. Huber, eds.
Hymenoptera of the World: an Identification
Guide to Families. Monograph 1894E. Ag-
riculture Canada Research Branch. Ottawa,
Ontario. 668 pp.
Wahlberg,N.,M.F.Braby,A.V.Z.Brower,R.de
Jong,M.-M.Lee,S.Nylin,N.E.Pierce,F.A.
H.Sperling,R.Vila,A.D.Warren,andE.Za-
kharov. 2005. Synergistic effects of combining
morphological and molecular data in resolving
the phylogeny of butterflies and skippers. Pro-
ceedings of the Royal Society Series B 272:
1577–1586. doi:10.1098/rspb.2005.3124
Walker, F. 1835. Monographia Chalciditum.
(Continued.). Entomological Magazine 3:
182–206.
Walker, F. 1849. List of the specimens of dip-
terous insects in the collection of the British
Museum, pp. 689–1172. Vol. 4. Printed by
order of the Trustees [British Museum],
London. 483 pp.
Walley, G. S. 1947. The genus Casinaria Holmg-
ren in America north of Mexico (Hymen.,
Ichneumonidae). Scientific Agriculture 27:
364–395.
Webber, R. T. and J. V. Schaffner, Jr. 1926. Host
relations of Compsilura concinnata Meigen,
an important tachinid parasite of the gipsy
moth and the brown-tail moth. United States
Department of Agriculture. Department
Bulletin 1363: 1–31.
Wei, C. G. 1985. A preliminary observation on
the bionomics of Miresina banghaasi.
Kunchong Zhishi (Insect Knowledge) 22:
76–78.
Weinstein, P. and A. D. Austin. 1991. The host-
relationships of trigonalyid wasps (Hyme-
noptera: Trigonalyidae), with a review of
their biology and catalogue to world species.
Journal of Natural History 18: 209–214.
Weinstein, P. and A. D. Austin. 1995. Primay
parasitism, development and adult biology
of the wasp Taeniogonalos venatoria Riek
(Hymenoptera: Trigonalyidae). Australian
Journal of Entomology 43: 541–555.
Weinstein, P. and A. D. Austin. 1996. Thely-
toky in Taeniogonalos venatoria Riek
(Hymenoptera: Trigonalyidae), with notes
on its distribution and first description of
males. Australian Journal of Entomology
35: 81–84. doi:10.1111/j.1440-6055.1996.
tb01366.x
Weseloh, R. M. and J. F. Anderson. 1982. Releases
of Brachymeria lasus and Coccygomimus
disparis, two exotic gypsy moth parasitoids, in
Connecticut: habitat preference and over-
wintering potential. Annals of the Entomo-
logical Society of America 75: 46–50.
Wesmael, C. 1835. Monographie des Braconides
de Belgique. Nouveaux Me
´moires de l’Aca-
demie Royale des Sciences et Bellas-lettres
Bruxelles 9: 1–252.
Whitfield, J. B. 1997. Subfamily Microgastrinae,
pp. 333–364. In R. A. Wharton, P. M. Marsh,
and M. J. Sharkey, eds. Identification Manual
to the New World Genera of the Family
Braconidae (Hymenoptera). International
Society of Hymenopterists, Special Publica-
tion 1. Washington, D. C. 439 pp.
Whitfield, J. B. and S. Asgari. 2003. Virus or not?
Phylogenetics of polydnaviruses and their wasp
carriers. Journal of Insect Physiology 49: 397–
405. doi:10.1016/S0022-1910(03)00057-X
Whitfield, J. B., R. J. Marquis, and J. Le Corff.
1999. Host associations of braconid parasit-
oids (Hymenoptera: Braconidae) reared from
Lepidoptera feeding on oaks (Quercus spp.)
in the Missouri Ozarks. Entomological News
110: 225–230.
Wilcox, A. M. 1918. Ascogaster carpocapsae,
a parasite of the Oriental moth. Psyche 25:
17. doi:10.1155/1918/57974.
Wijesekara, A. and M. Schauff. 1997. Two new
genera and three new species of Euplectrini
(Hymenoptera: Eulophidae) from the New
World. Proceedings of the Entomological
Society of Washington 99: 101–109.
Witter, J. A. and H. M. Kulman. 1979. The par-
asite complex of the forest tent caterpillar in
northern Minnesota. Environmental Ento-
mology 8: 723–731.
Wood, D. M. 1985. A taxonomic conspectus of
the Blondeliini of North and Central America
and the West Indies (Diptera: Tachinidae).
Memoirs of the Entomological Society of Can-
ada 132: 1–130. doi:10.4039/entm117132fv
Wood, D. M. 1987. Tachinidae, pp. 1193–1269.
In J. F. McAlpine, B. V. Peterson, G. E.
Shewell, H. J. Teskey, J. R. Vockeroth, and D.
M. Wood, eds. Manual of Nearctic Diptera.
VOLUME 114, NUMBER 1
109

Vol. 2. Agriculture Canada Monograph 28.
iv +675–1332 pp.
Wood, D. M. and M. A. Zumbado. 2010. Tachinidae
(tachinid flies, parasitic flies), pp. 1343–1417. In
B. V. Brown, A. Borkent, J. M. Cumming, D.
M. Wood, N. E. Woodley, and M. A. Zumbado,
eds. Manual of Central American Diptera, Vol.
2. NRC Research Press, Ottawa.
Yeargan, K. V. and S. K. Braman. 1989. Life
history of the hyperparasitoid Mesochorus
discitergus (Hymenoptera: Ichneumonidae)
and tactics used to overcome the defensive
behavior of the green cloverworm (Lepidop-
tera: Noctuidae). Annals of the Entomologi-
cal Society of America 82: 393–398.
Yeates, D. K. and D. J. Greathead. 1997. The
evolutionary pattern of host use in the
Bombyliidae (Diptera): a diverse family of
parasitoid flies. Biological Journal of the
Linnaean Society 60: 149–185. doi:10.1111/
j.1095-8312.1997.tb01490.x
Yu, D. S., K. van Achterberg, and K. Horstmann.
2005. World Ichneumonoidea 2004. Taxon-
omy, Biology, Morphology and Distribution.
DVD/CD. Taxapad, Vancouver.
Zaldı
´var-Rivero
´n, A., M. R. Shaw, A, G. Sa
´ez, M.
Mori, S. A. Belokoblylskij, S. R. Shaw, and
D. L. J. Quicke. 2008. Evolution of the
parasitic wasp subfamily Rogadinae (Braco-
nidae): phylogeny and evolution of lepidop-
teran host ranges and mummy characteristics.
BMC Evolutionary Biology 8: 329. doi:10.1186/
1471-2148-8-329
Zaldı
´var-Rivero
´n,A.,B.Areekul,M.R.Shaw,
and D. L. J. Quicke. 2004. Comparative
morphology of the venom gland apparatus
in the braconid wasp subfamily Rogadinae
(Insecta, Hymenoptera, Braconidae)
and related taxa. Zoologica Scripta 33:
223–237. doi:10.1111/j.0300-3256.2004.
00144.x
Zhu, C. D. and D. W. Huang. 2004. A study of
Platyplectrus Ferrie
`re (Hymenoptera: Eulo-
phidae) in mainland China. Journal of Natu-
ral History 38: 2183–2209. doi:10.1080/
00222930310001618903
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
110