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(Hymenoptera, Diptera) Associated with Limacodidae (Lepidoptera) in North America, with a Key to Genera

(Hymenoptera, Diptera) Associated with Limacodidae (Lepidoptera) in North America, with a Key to Genera

PROC. ENTOMOL. SOC. WASH. 114(1), 2012, pp. 24–110

REVIEW OF AND (, DIPTERA) ASSOCIATED WITH () IN NORTH AMERICA, WITH A KEY TO GENERA

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

(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 [email protected], RRK [email protected], DRS dave. [email protected]); (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 [email protected], TMS [email protected]); (JEO) Canadian National Collection of , and Agri-Food Canada, 960 Carling Avenue, Ottawa, Ontario, Canada K1A 0C6 (e-mail: [email protected].); (DBW) American Entomological Institute, 3005 SW 56th Ave., Gainesville, Florida 32608 U.S.A. (e-mail: [email protected]); (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. [email protected])

Abstract.—Hymenopteran and dipteran of (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 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 ) 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): ([1]; Hymenoptera: Chalcidoidea), (3; Chalcidoidea), ([1]; Chalcidoidea), (1; Chalcidoidea), (3 [1]; Hymenoptera: Ichneumo- noidea), (7 [3]; ), (1; Hymenoptera: ), (2; Hymenoptera: Trigonaloidea), ([1]; Diptera: Asilioidea), and (3; ). 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): li- macodis (Walker) (1), U. pristis (Townsend) (5), Austrophorocera spp. (6), Ceraphron sp. (1), Alveoplectrus lilli Gates (1), Playplectrus americana (Girault) (10), VOLUME 114, NUMBER 1 25 crassicornis (Thomson) (1), (1), 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,istransferredfrom (Hymenoptera: Braconidae). Key Words: hyperparasitoid, parasitic, slug moth , , Acrolyta, Alveoplectrus, Ascogaster, Austrophorocera, Baryceros, , Ceraphron, , , , , Hyposoter, Isa, Isdromas, Isochaetes, , Lysibia, Mesochorus, , Orthogonalys, , , Pediobius, , Platyplectrus, , Psychophagus, , , Tortricidea, Trichogramma, Triraphis, Uramya DOI: 10.4289/0013-8797.114.1.24

Slug moth caterpillars (Lepidoptera: that facilitates their capture. Outbreaks Limacodidae) are mostly polyphagous of North American limacodids are re- external foliage feeders on a broad array ported rarely, but many tropical species of deciduous trees and shrubs in eastern are important agricultural pests, partic- North America (Wagner 2005, Lill 2008). ularly in banana and palm plantations Their broad diets, rather distinctive larval in South America and Southeast Asia morphologies, and often bright coloration (Ostmark 1974, Cock et al. 1987, Godfray make them interesting subjects for eco- and Chan 1990). Because many species logical and behavioral studies (e.g., Lill of Limacodidae possess stinging spines et al. 2006, Murphy et al. 2010). Unlike during all or a portion of their larval de- many more mobile caterpillars, limaco- velopment (Epstein 1988), the larvae can did larvae tend to remain on an individual be a nuisance when they occur in planta- host for the duration of their de- tions (Ostmark 1974) or on ornamental velopment: typically about two months, palms near tourist attractions (e.g., Conant going through numerous instars (7–12) et al. 2002). Their protracted larval de- depending on the species, with some velopment period exposes the caterpillars species requiring more than 100 days to attack from a diverse assemblage of to complete development (J. Lill and larval and larval-pupal parasitoids. How- S. Murphy pers. obs.). Larvae tend to ever, research to-date on host-parasitoid occur in low densities (< 1caterpillar/ interactions in Limacodidae has focused m2 foliage; Lill et al. 2006, Lill 2008) almost entirely on biocontrol of pest li- and can be highly cryptic, especially in macodids in tropical agricultural systems early instars, but they produce a rather (e.g., Cock et al. 1978) with the notable distinctive pattern of feeding damage exception of the massive caterpillar rear- ing database compiled for tropical forests of Area de Conservacio´nGuanacastein *Accepted by John W. Brown Costa Rica (Janzen and Hallwachs 2009). 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

By contrast, the hymenopteran/dipteran focused on Limacodidae (as mentioned parasitoid community that attacks North above). American limacodids has been described Tachinidae and parasitoid Hymenoptera only anecdotally, and concerted rearing are diverse in forest canopy communi- efforts are mostly lacking (but see Le ties in North America with new host- Corff and Marquis 1999 and Stireman parasitoid associations being recorded with and Singer 2003a, b for examples where some frequency (Butler 1993, Strazanac limacodids were reared as part of larger et al. 2001, Petrice et al. 2004). Bio- community sampling). logically, tachinids are recorded from In general, dipteran (i.e., Tachinidae) 15 orders (Wood 1987), whereas and hymenopteran parasitoids are essen- parasitoid Hymenoptera have been docu- tial in regulating native forest macrolep- mented from 16 (LaSalle 1993). As a idoptera and are reasonably well studied group, parasitoid Diptera have been (LaSalle 1993, McCullough et al. 1999). viewed as secondary to hymenopterans Conversely, there is little information as effective parasitoids (Askew 1971, regarding bombyliid parasitoids (i.e., in Quicke 1997); however, both groups Systropus) and their roles in regulating are abundant in North American forests native forest macrolepidoptera, likely where limacodids occur most commonly. due to their relative rarity compared to North American hymenopteran and dip- parasitic hymenopterans and tachinids. teran parasitoid-host associations have Most hymenopteran parasitoids of mac- been cataloged by Krombein et al. (1979) rolepidoptera belong to the superfamilies and Arnaud (1978), respectively. A world Ichneumonoidea and Chalcidoidea, and catalog to Tachinidae is in preparation species of Tachinidae are the most com- (JEO), and databases are available for mon dipteran parasitoids of microlepi- Ichneumonoidea (Yu et al. 2005) and doptera (Askew 1971, Quicke 1997). Two Chalcidoidea (Noyes 2003). Recent large- recent studies in North America have scale rearings of forest macrolepidoptera focused on Tachinidae and Hymenoptera by Butler (1993) documented 115 new specifically and their host relationships in hymenopteran host associations, and eastern North American forests (Strazanac Strazanac et al. (2001) reported 60 new et al. 2001, Petrice et al. 2004). Most tachinid host associations. research on parasitoids of native forest Many of the tachinid and hymenop- macrolepidoptera has focused on out- teran parasitoids reported herein are break species such as fall webworm known to attack lepidopterans (or their (Hyphantria cunea Drury; Arctiidae), primary parasitoids) more generally, but (Malacosoma several specialize on Limacodidae, and americanum (F.); Lasiocampidae) (Kulman others are facultative (see Pediobius; 1965, Morris 1976, Witter and Kulman Peck 1985) or obligatory hyperparasitoids 1979), and gypsy moth (Lymantria (see Conura Spinola; Delvare 1992). dispar (L.); Lymantriidae) (Barbosa et al. Given their specialization on Limacodidae, 1975, Elkinton and Liebhold 1990 and many of these taxa, particularly certain references therein). Few studies have Chalcidoidea such as Platyplectrus Ferrie`re documented parasitoids in less abundant/ and Alveoplectrus Wijesekara and Schauff, non-outbreak forest macrolepidoptera (in- were encountered infrequently in North cluding limacodids) (Schaffner and Griswold America (Wijesekara and Schauff 1997). 1934, northeasternU.S.A.; Raizenne 1952, This can be ascribed to the lack of major Ontario, Canada), and fewer still have limacodid pests in North America, the VOLUME 114, NUMBER 1 27 difficulty of locating/handling their larvae, collection date, and host plant. In addition and the lack of financial resources avail- to these wild-caught larvae, we also con- able to study non-outbreak pests. Most ducted a series of field experiments as part of the parasitoids documented as a re- of a different project examining tri-trophic sult of the rearing portion of this study interactions in Limacodidae that involved parasitize early limacodid instars. placing out “sentinel” larvae on each of the The purpose of this study is to review six common host described above the parasitoids associated with limacodids at the Little Bennett Regional Park site. in America north of Mexico, including These larvae came from laboratory colo- introduced limacodids (i.e., nies established through a combination of flavescens Walker) and exotic parasitoids larval and adult (ex ovo)collections;thus, recorded from limacodids elsewhere that larvae were unparasitized when placed in could potentially attack limacodids in the field. These experimental larvae were North America. We assemble these vari- left exposed to parasitoid attack in the field ous records herein and add host-parasitoid for one to several weeks depending on the associations generated from six years of experiment and then brought back to the intensive rearing of larval limacodids in lab for rearing. the greater Washington, DC metropolitan Collected larvae were reared individu- area (Table 1). ally in 16 oz. clear plastic deli containers containing a disk of moistened filter paper MATERIALS AND METHODS to prevent host foliage from drying out. Collection and Rearing Fresh leaves from the various host plants Beginning in mid- to late-June of each were replaced as needed (typically twice summer of 2004–2009, we searched per week). Larvae showing signs of para- manually for limacodid larvae on the sitism were checked routinely for parasit- undersides of leaves of common woody oid larvae/pupae, and emerging adults were trees at the following field sites near either placed in 95% ethanol (Hymenop- Washington, DC: Little Bennett Regional tera) or frozen (Diptera) for later mounting/ Park, Plummers Island (Montgomery pinning and identification. One of County, MD), Patuxent National Wildlife tachinid flies (Austrophorocera spp.) exclu- Refuge (Prince George’s County, MD), sively contains larval-pupal parasitoids, so Rock Creek Park (Washington, DC), and adults eclosed from overwintering cocoons the U.S. National Arboretum (Washington, the following year (pupae were kept in an DC). More than a dozen host plants have environmental chamber in moist peat moss yielded limacodids, but most of our efforts during the winter months and then exposed were focused on searching six common to spring conditions to induce fly pupation host plants used by most species of and emergence). In addition, several of Limacodidae: American beech (Fagus the hymenopteran parasitoids reared from grandifolia Ehrh.), white oak (Quercus limacodid hosts late in the season dia- alba L.), northern red oak (Quercus rubra paused as pupae and emerged the follow- L.), black gum (Nyssa sylvatica Marsh.), ing summer. black cherry (Prunus serotina Ehrh.), and pignut hickory (Carya glabra Miller). Hymenopteran Parasitoid Additional host plant species were sam- Preparation and Imaging pled less intensively (Table 1). For each Specimens in ethanol were dehydrated wild-caught , we recorded the species, through increasing concentrations of 28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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 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 VOLUME 114, NUMBER 1 29

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 virginiana Co. empretiae 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 30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Continued.

Limacodid Species Plant Species Parasitoid Species Notes

Isa/Natada Nyssa sylvatica Al. lilli Isochaetes Fagus grandifolia U. pristis* beutenmuelleri O. pulchella Hyperparasitoid of U. pristis Quercus rubra U. pristis* 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 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 Quercus rubra Austrophorocera sp.* Only egg observed T. discoideus I. lycaenae* Mesochorus discitergus Hyperparasitoid of Triraphis; unconfirmed Phobetron pithecium Fagus grandifolia Pl. americana* 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* VOLUME 114, NUMBER 1 31

Table 1. Continued.

Limacodid Species Plant Species Parasitoid Species Notes

Quercus rubra A. imitator* Pl. americana* T. discoideus 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

ethanol to hexamethyldisilazane (HMDS) data to a desktop computer. The program (Heraty and Hawks 1998) before point- or Cartograph 5.6.0 (Microvision Instru- card-mounting. Images of specimens were ments, ) was used to merge an produced by scanning electron micros- image series (typically representing 15– copy (SEM) and an EntoVision Imaging 30 focal planes) into a single in-focus, Suite. Card- and point-mounted specimens composite image. Lighting was achieved were examined using stereomicroscopes using techniques summarized in Buffington with 10X or 25X oculars and fiber optic et al. (2005), Kerr et al. (2008), and light sources. Mylar film was used to dif- Buffington and Gates (2008). fuse glare from fiber optic light sources to Several images (see below) were ob- reduce glare from the specimens. Scan- tained with a Visionary Digital imaging ning electron microscope (SEM) images station. The station consists of an Infin- were taken with an Amray 1810 (LaB6 ity Optics K2 long distance microscope source). Specimens were affixed to 12.7 X affixed to a Canon EOS 40D digital SLR 3.2 mm Leica/Cambridge aluminum SEM camera. Lighting was provided by a Dyna- stubs with carbon adhesive tabs (Electron lite M2000er power pack and Microptics Microscopy Sciences, #77825-12). Stub- ML1000 light box. Image capture soft- mounted specimens were sputter coated ware is Visionary Digital proprietary ap- using a Cressington Scientific 108 Auto plication with images saved as TIF with with gold from at least three differ- the RAW conversion occurring in Adobe ent angles to ensure complete coverage Photoshop Lightroom 1.4. Image stacks (;20–30 nm coating). Wings were re- were montaged with Helicon Focus moved and slide-mounted in polyvinyl 4.2.1 for images of Conura nortonii alcohol prior to imaging. Wing and (Cresson), C. immaculata (Cresson), habitus images were captured using an Orthogonalys pulchella (Cresson), and EntoVision Imaging Suite, which includes Taeniogonalos gundlachii (Cresson), afirewireJVCKY-753CCDdigitalcam- as well as habitus shots of Psychophagus era mounted on a Leica M16 zoom lens omnivorus (Walker), Platyplectrus ameri- via a Leica z-step microscope stand. Ad- cana (Girault), and Pediobius crassicornis ditionally, a GT-Vision Lw11057C-SCI (Thomson). digital camera attached to a Leica DMRB Final image plates for hymenopteran compound scope was used to feed image and dipteran figures were prepared using 32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

InDesign CS4. Figures 14–17, previously the posterior ocelli; ocular ocellar line unpublished, were used with the permis- (OOL), the shortest distance between the sion of D. M. Wood (Agriculture and lateral margin of the posterior ocellus Agri-Food Canada, Ottawa). Figures 38 and the eye orbit; marginal vein, the and 43–44 were reproduced from Townes length coincident with the leading fore (1970) with permission. wing edge to the base of the stigmal vein; stigmal vein, the length between its Diptera Imaging base on the marginal vein and its apex; The image of Systropus macer Loew postmarginal vein, the length from the was captured with a Nikon Coolpix 8800 base of the stigmal vein to its apex on and adapters through the ocular of a the leading fore wing edge. Mesosomal Leica MZ9.5 stereoscope. Tachinid images and metasomal sclerites were measured were taken with a Canon EOS 40D digital dorsally along the midline. The use of SLR camera mounted on a Kaiser RS1 “[ ]” in descriptions denotes structures copy stand. A Canon EF 100 mm f/2.8 that are not visible in the specimens upon macro lens was used for full body images, which the description is based (observed and a Canon MP-E 65 mm 1–5X macro via SEM), whereas their use in the ma- lens was used for images of body struc- terial examined section refer to author tures. A ring light consisting of 80 LEDs notes. For braconids metasomal terga 1, 2, 3, etc. are abbreviated as T1, T2, T3; and covered with a reflective dome antennal flagellomeres are abbreviated provided the lighting. Image stacks were as F1, F2, F3, etc. The ovipositor of montaged using Syncroscopy’s Auto- ichneumonids was measured from the Montage, and the resultant images received structure’s base (observed or inferred) further treatment in Adobe Photoshop to its apex. The ovipositor sheaths must CS4. sometimes be separated by a fine needle to expose the ovipositor valves. The junc- Terminology ture of the occipital and hypostomal carina Hymenopteran terminology for sur- above the mandibular base is measured face sculpture follows Harris (1979) and in posterolateral view; it is sometimes for morphology follows Wahl (1993a), necessary to remove the head to properly Gibson (1997), Sharkey and Wharton measure. Dipteran terminology follows (1997), and Deans et al. (2010). Several McAlpine (1981). measurements for chalcidoids were taken, The portion of the key pertaining to including the following: body length, in Chalcidoidea is based upon the keys in lateral view from the anterior projection Gibson et al. (1997). That for Braconidae is of the face to the tip of the metasoma; based on the keys of Sharkey (1997) and head width through an imaginary line Whitfield (1997). That for Ichneumonidae connecting the farthest lateral projection is based on the key of Wahl (1993b). The of the eyes; head height through an diagnosis for Cotesia Cameron is based on imaginary line from the vertex to the the key in Whitfield (1997); the diagnosis clypeal margin bisecting both the median for Ascogaster Wesmael is based on the ocellus and the distance between the tor- key in Shaw (1997a). The diagnosis for uli; malar space, in lateral view between Triraphis Ruthe is based on the diagnoses for the ventral margin of the eye and lateral Triraphis and Rogas Nees in van Achterberg margin of the oral fossa; posterior ocellar (1991), as well as the keys in van Achterberg line (POL), the shortest distance between (1991) and Shaw (1997b). VOLUME 114, NUMBER 1 33

All binominals for hosts and parasitoids Canada), USNM (National Museum of are reported in their current nomenclatural Natural History, Smithsonian Institution, combinations. Although this is not Washington, DC, U.S.A.), and ZMUC necessarily straightforward in any single (Zoologisk Museum, Copenhagen, publication, we cite the most recent/ Denmark). comprehensive nomenclatural authorities where required. For Limacodidae, nomen- RESULTS AND DISCUSSION clatural references include: Fletcher and Key to Primary and Secondary Nye (1982), Davis (1983), and Becker and Parasitoids Known From Limacodids Epstein (1995). Host records from the lit- in North America erature for Tachinidae list the current combination, as well as the combinations 1. Hind wing modified as haltere (Fig. 1), hamuli absent, mouthparts sponging (Fig. 1) under which the hosts and tachinids were ...... 2(Diptera) originally cited. New host records result- 1’ Hind wing not modified as haltere, hamuli ing from the rearing efforts of JTL, SMM present (Figs. 92–93), mouthparts mandib- and TMS for a particular parasitoid are ulate (e.g., Figs. 39, 82) ...... denoted with * in the text and Table 1...... 5(Hymenoptera) Those newly reported herein from other 2. (1) Abdomen narrow and elongate, > 2.0X ‡ as long as broad and swollen apically sources are denoted with .Wereportnew (Fig. 3). Wings shorter than abdomen. host records at the parasitoid species level Antennae and proboscis much longer than only rather than higher taxonomic levels. head ...... Thus, if a parasitoid species is newly re- ..Systropus (macer Loew; Bombyliidae) corded for a particular host genus or spe- 2’ Abdomen stout, less than 2.0X as long as broad (Figs. 1, 11). Wings longer than ab- cies, we report it as new even if congeneric domen. Antennae and proboscis at most as parasitoids were previously recorded from long as head ...... 3 (Tachinidae) that host genus or species. 3. (2’) Facial ridge bare except for a few small Abbreviations for collections are as and decumbent setae on lower third or less follows: AEI (American Entomological (Fig. 2). Metathoracic spiracle fringed with plumose hairs of aboutequallengthalong Institute, Gainesville, Florida, U.S.A.), both anterior and posterior edges, leaving a ANSP (Academy of Natural Sciences, V-shaped middorsal opening (Fig. 4). Pros- Philadelphia, Pennsylvania, U.S.A.), BMNH ternumbare(Fig.6)...... (The Natural History Museum, London, Uramya (pristis (Walker), limacodis (Townsend)) United Kingdom), CNC (Canadian 3’ Facial ridge with row of stout setae on lower National Collection of Insects, Ottawa, one-half or more (Fig. 9). Metathoracic spiracle with posterior lappet much larger Ontario, Canada), IRSNB (Institut Royal than anterior one (Fig. 5). Prosternum haired des Sciences Naturelles de Belgique, laterally (Fig. 7)...... 4 Brussels, ), MZLU (Zoological 4. (3’) Ocellar seta vestigial or absent (Fig. 9). Museum, Lund University, Lund, Sweden), Abdominal terga 3 and 4 each with 1 pair of MCZ (Museum of Comparative Zoology, median discal setae. Bend of vein M obtuse (Fig. 8). Female with sickle-shaped, pierc- Harvard University, Cambridge, Massa- ing ovipositor (Fig. 8) ...... chusetts, U.S.A.), MSUC (Michigan State ...... Compsilura (concinnata (Meigen)) University, East Lansing, Michigan, 4’ Ocellar seta well developed, similar in size U.S.A.), NHMW (Naturhistorisches to outer vertical seta (Fig. 10). Abdominal Museum Wien, Vienna, ), SEMK terga 3 and 4 without median discal setae (Fig. 11). Bend of vein M almost a right angle (Snow Entomological Museum, Univer- (Fig. 11). Female with short, non-piercing sity of Kansas, Lawrence, Kansas, U.S. ovipositor (Fig. 11) ...... A.), ULQC (Universite´ Laval, Quebec, ..Austrophorocera (cocciphila (Aldrich and 34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Webber), coccyx (Aldrich and Webber), A. main area of propleuron (Fig. 25). Sternaulus einaris (Smith), A. imitator (Aldrich and of mesopleuron present and reaching middle Webber), A. n. sp.) coxa (Fig. 31). Clypeus separated from su- 5 (1’) Fore wing venation complete, with at praclypeal area by groove (Fig. 33). Areolet least 2 closed cells (Figs. 18, 28, 34) . . . . . of fore wing open and pentagonal (Fig. 34)...... 6 (Trigonaloidea, Ichneumonoidea) ...... 11 5’ Fore wing venation reduced, with fewer 10. (9) Petiole of first metasomal segment than 2 closed cells (Figs. 77–78, 96, 113) long and cylindrical in cross-section; .....17(Chalcidoidea,Ceraphronoidea) midpoint of petiole with tergo-sternal 6. (5) Fore wing with veins C and SC+R suture at midline; T1 without trace of glymma touching/fused, costal cell absent (Figs. (Fig. 28) . . . .Casinaria (grandis Walley) 63, 65). Metasomal sterna less strongly 10’ Petiole of first metasomal segment shorter sclerotized than terga (Figs. 19, 23, 27, and quadrate in cross-section; midpoint of 29) ...... 7 (Ichneumonoidea) petiole with tergo-sternal suture close to 6’ Fore wing with veins C and SC+R separate, ventral margin; T1 with glymma present as costal cell present (Fig. 18). Metasomal sterna pitlike impression (Fig. 27) ...... and terga equally sclerotized (Figs. 70–73) ...... Hyposoter (fugitivus (Say)) ...... 16(Trigonaloidea) 11. (9’) Body color (excluding legs) black with 7. (6) Fore wing with vein 2m-cu present white markings (Fig. 29). Mesosoma with (Fig. 28, 35) ...... 8(Ichneumonidae) coarse punctures (Fig. 37). Dorsal margin 7’ Fore wing with vein 2m-cu absent (Fig. of pronotum with strong swelling at dorsal 56) ...... 14 (Braconidae) end of epomia (Fig. 37). Fore wing 5.0–8.5 8. (7) Areolet of fore wing closed, large and mm long ...... rhombic (Figs. 20, 26). Ovipositor delicate, ...... Baryceros (texanus (Ashmead)) needlelike, sheaths thick and rigid (Figs. 19, 11’ Body color (excluding legs) ranging from 22). Female hypopygium prominent and tri- uniformly black/dark brown to having angular in lateral view (Fig. 22). Male gon- brownish red areas on T2–T3 (Figs. 34–36). oforceps produced into elongate process. Mesosoma with punctures ranging from Spiracle of metasomal segment 1 near or just fine to absent. Dorsal margin of pronotum behind middle, glymma large and deep without strong swelling at dorsal end of (Fig. 21) ...... epomia. Fore wing 1.9–3.8 mm long...... Mesochorus (discitergus (Say)) ...... 12 8’ Areolet of fore wing open (vein 3rs-m ab- 12. (11’) Apical 0.3 of clypeus turned inward sent) or closed, if closed then cell obliquely at 90° (Fig. 43). Vein 2-Cu of hind wing quadrate and petiolate (Figs. 26, 34). basally incomplete ...... Ovipositor always stouter than above, sheaths ....Lysibia (mandibularis (Provancher)) thin, often curved if ovipositor ; 2.0X as 12’ Apical 0.3 of clypeus flat, not turned in- long as metasomal apical depth. Female ward (Fig. 44). Vein 2-Cu of hind wing hypopygium small and quadrate in lateral complete ...... 13 view (Fig. 23). Male gonoforceps not 13. (12’) Occipital and hypostomal carinae produced. Spiracle of metasomal segment meeting at mandibular base (Fig. 41) . . . 1beyondmiddle,glymmasmallorabsent Acrolyta (nigricapitata (Cook and Davis)) (Fig.27) ...... 9 13’ Occipital carina meeting hypostomal ca- 9. (8’) Ventral posterior corner of pro- rina above mandibular base, juncture sep- pleuron with strongly produced, more or arated from base by about 0.2X basal less angulate lobe touching or overlapping mandibular width (Fig. 42)...... pronotum (Fig. 24). Sternaulus about 0.3X ...... Isdromas (lycaenae (Howard)) as long as mesopleuron (Fig. 30). Clypeus 14. (7’) Labrum visible through gap between not separated from supraclypeal area by ventral margin of clypeus and mandibles distinct groove (Fig. 32). Areolet of fore and concave (as in Fig. 39) (cyclostome wing closed, obliquely quadrate and peti- Braconidae). Epicnemial carina present olate (Fig. 26) ...... 10 (Fig. 47); median carina present on T1 and 9’ Ventral posterior corner of propleuron not usually extending posteriorly from dorsal developed as distinct lobe, not angulate, at carinae; T2 striate to striate-rugose (Figs. most with weak groove delimiting it from 52, 57); fore wing vein 1m-cu basad or in VOLUME 114, NUMBER 1 35

line with fore wing vein 2RS (Fig. 56) . . 18. (17’) Hind femur enlarged (< 3.0X as long ...... Triraphis Ruthe (eupoeyiae as broad) (Figs. 79–80). Prepectus reduced (Ashmead), discoideus (Cresson), harri- to a small sclerite along dorsal margin of sinae (Ashmead)) mesopleuron (Fig. 83). Gaster petiolate 14’ Labrum concealed by clypeus or if vis- (Figs. 79–80) ...... ible not concave (Figs. 40 (as in), 60) ....Conura (camescens (Cameron), nig- (non- cyclostome Braconidae). Without ricornis (F.), nortonii (Cresson)) combination of characters found in 18’ Hind femur not enlarged (> 3.0X as long as Triraphis ...... broad). Prepectus larger, triangular (Figs...... , , 15 87, 109). Gaster indistinctly or not petiolate 15. (14’) Occipital carina present, fore wing ...... Eulophidae,Trichogrammatidae, vein RS reaching wing margin as tubular Pteromalidae, 19 vein, T1–T3 fused into carapace cover- 19. (18’) Legs with 5 tarsomeres. Clypeus ing all other terga (Figs. 48, 63). Other shallowly bilobed apically (Fig. 88). characters not in combination found in Pedicel ; 2.0X as long as broad, antennal Cotesia (see below) ...... formula 11263 (Fig. 89). Propodeum re- ticulate-rugose, with median carina in- ..Ascogaster (quadridentata Wesmael) 15’ Occipital carina absent, fore wing vein RS complete (Fig. 90). Gaster subcircular to not reaching wing margin as tubular vein short-ovate (Fig. 91)...... (Fig. 65), T1–T3 variable (Microgastrinae)...... Psychophagus (omnivorus (Walker)) 19’ Legs with 3 or 4 tarsomeres. Other features Fore wing vein r-m absent, areolet ab- not as above ...... sent (Fig. 65). Ovipositor and sheaths . . . . Eulophidae, Trichogrammatidae, 20 short, weakly extending beyond tip of 20. (19’) Legs with 4 tarsomeres. Fore wing hypopygium (Fig. 67). Hypopygium evenly lacking setal tracks (Fig. 113) ...... sclerotized medially. Propodeum with- ...... Eulophidae, 21 out areola, often with distinct medial 20’ Legs with 3 tarsomeres (Fig. 94). Fore longitudinal carina and usually rugose wing with setal tracks, broad with sig- (Fig.66)...... moid venation and distinctive Rs1 setal .. Cotesia (empretiae (Viereck), phobetri track(Fig.96)...... (Rohwer), schaffneri (Rohwer)) ...... Trichogramma (minutum Riley) 16. (6’) black with white or light 21. (20’) Scutellum with one pair of setae (Fig. yellow band at center. Head and mesosoma 95). Submarginal vein with two setae on with black and white pattern. Metasoma dorsal surface. Propodeum with paired and legs mostly orange (Figs. 70–71). submedian carinae that diverge posteriorly Metasoma thin, smooth, impunctate . . . (Fig.95) ...... Orthogonalys (pulchella Cresson) ...... Pediobius (crassicornis (Thomson)) 16’ Antenna yellowish brown without lighter 21’ Scutellum with two pairs of setae (Figs. band at center. Head and body black with 104–105). Submarginal vein with three or yellow markings (Figs. 72–73). Meta- more setae on dorsal surface (Fig. 113). soma black and yellow, legs yellow with Propodeum with single carina, or if with dark brown to black on femur, tibia paired submedian carinae then carinae not dusky in apical third. Metasoma stout, arcuately divergent (Figs. 104–105) ...... punctate...... ,22 ..Taeniogonalos (gundlachii (Cresson)) 22. (21’) Propodeum with two submedian 17. (5’) Fore wing with tubular submarginal carinae (Fig. 104). Scutellum with scu- vein basally on anterior margin, stigmal tellar grooves broad, contiguous with vein curving distally (Fig. 77). Female an- posterior margin (Fig. 104). Hind tibia tenna with 7–8 flagellomeres (Fig. 76), male withonespur...... with 8–9 (Fig. 74). Dorsum flat in lateral ...... Alveoplectrus (lilli Gates) view...... 22’ Propodeum with single median carina ...... Ceraphron Jurine (Ceraphronidae) (Fig. 105). Scutellum with scutellar 17’ Fore wing lacking tubular vein basally on grooves narrow (Fig. 105). Hind tibia with anterior margin (Fig. 78). Other features twospurs...... not as above ...... 18 ....Platyplectrus (americana (Girault)) 36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Diptera although they are documented to oviposit Systropus Weidemann (Bombyliidae: directly onto their hosts by adhering eggs Toxophorinae: Systropodini) to the host’s integument (Genty 1972). In (Fig. 3) at least one instance, oviposition of black eggs is accompanied by rapid movement Diagnosis.—Systropus can be identi- of the abdomen of the fly on the host fied in North America using the key by (Genty 1972). Systropus spp. are specialists Hall (1981) or Kits et al. (2008; Mel- on limacodid larvae and are placed in anderella Cockerell not extant) and can be Toxophorinae, one of only two bomb- distinguished from other North American yliid subfamilies containing true endo- genera (Dolichomyia Weidemann) of Sy- parasitoids (the other is Anthracinae) stropodini by the following characters: (Greathead 1987, Yeates and Greathead pedicel at least 2–3 times longer than wide 1997). Hosts are known only for S. macer (as wide as long in Dolichomyia); first in North America (see below). Extra- flagellomere flattened (cylindrical in limitally, host associations have been re- Dolichomyia); and anal lobe of wing ported from Africa (Hull 1973), Central/ not reduced (extremely reduced in Do- South America (Adams and Yanega 1991, lichomyia). However, Dolichomyia is Aiello 1980, Dyar 1900), and Asia not known from eastern North America. (Greathead 1987). Adults routinely take Athirdgenus,Zaclava Hull, is repre- nectar from flowers (Robertson 1928, sented by four species known only from Painter and Painter 1963) and are com- Australia. Typically, Systropus spp. mimic monly reported from woodland glades sphecid wasps and thus are often elongate, and various habitats in the wet tropics largely orange and black with the abdo- (Painter and Painter 1963, Greathead men long and swollen apically (Fig. 3). 1987). Two species are known from The wings are conspicuously shorter than Australia (Evenhuis and Greathead 1999). the abdomen, and both the antennae and Literature.—Painter and Painter (1963) proboscis are much longer than the head revised the Systropus of North and (Fig. 3). Central America treating 23 species, Fauna.—Systropus is a large genus including three found north of Mexico. containing 154 species most commonly Extralimitally, Enderlein (1926) keyed encountered in tropical areas of all con- 13 species from Southeast Asia, but 48 tinents (Evenhuis and Greathead 1999). species are known from the region today The majority of the species are known (Evenhuis and Greathead 2003). The group from the African and Oriental regions has been cataloged along with a full bibli- (Evenhuis and Greathead 1999). Species ography as part of larger works on Bomb- from other biogeographic regions have yliidae (Evenhuis and Greathead 1999), not been treated via comprehensive tax- and these data are now available on the onomic publications. In North and Cen- Internet (Evenhuis and Greathead 2003). tral America, the 23 described species North American Records: range throughout tropical and subtropical Systropus macer Loew (Fig. 3): Systropus areas (Painter and Painter 1963, Adams macer Loew, 1863: 375. Lectotype /, and Yanega 1991), with three found north MCZ, designated by Evenhuis (1982) (not of Mexico. examined). Biology.—Specific biology related to Systropus macer is diagnosed in the key host location and oviposition behavior of Painter and Painter (1963) by: has not been reported for Systropus spp., swollen part of propleura bright yellow, VOLUME 114, NUMBER 1 37 metasternum solid brown/black, meso- (2009) recorded several undescribed mor- notum with small yellow spots, legs light phospecies of Uramya from Guanacaste brown/black, and the caudal edge of the (Costa Rica); all were reared from 8th sternum notched/undulating. limacodids. Distribution: Northeastern North Biology.—Females of Uramya are America (Painter and Painter 1963). ovolarviparous on lepidopteran larvae. Limacodidae: , Hosts are known for four of the seven sp., Euclea delphinii (Boisduval), species in North America. Two of the Lithacodes fasciola (H. –S.), Proli- four, U. limacodis and U. pristis, attack macodes badia (Adams and Yanega limacodids (Arnaud 1978, see below). 1991), (Boisduval) Uramya halisidotae (Townsend) is most (Epstein 1997). commonly reared from Lophocampa Other hosts: No records. argentata (Packard) (as Halisidota argen- tata;Arctiidae),butthereisasinglerecord Uramya Robineau-Desvoidy from Malacosoma sp. (Lasiocampidae) (Tachinidae: : Uramyini) (Aldrich 1921, Arnaud 1978). Stireman (Figs. 1–2, 4, 6) and Singer (2003a, b) reared U. indita (Walker) from Norape tenera Druce Diagnosis.—Uramya can be identi- (Megalopygidae) in Arizona. fied in North America using the key by Two South American species have Wood (1987) and can be distinguished been reared in Venezuela: U. longa from Compsilura and Austrophorocera (Walker) from Ammalo helops (Cramer) by the following combination of characters: (Arctiidae) and U. sibinivora Guimara˜es ocellar seta weak or absent; facial ridge from Acharia sp. (as Sibine;Limacodidae) bare except for a few small, decumbent (Guimara˜es 1980). Cruttwell (1969) setae on lower third or less (Fig. 2); pros- documented the biology of U. brevicauda ternum bare (Fig. 6); 3 large postsutural Curran, reared from Pareuchaetes pseu- dorsocentral setae; metathoracic spiracle doinsulata Rego Barros (Arctiidae) in fringed with plumose hairs of about equal Trinidad. Eggs are laid directly on a host size along anterior and posterior edges, and hatch immediately; the first instar leaving V-shaped middorsal opening penetrates the host’s integument within (Fig. 4); scutellum with 3 pairs of strong a few seconds. Cruttwell (1969) found marginal setae, including pair of strong, that larvae could develop gregariously in crossed apicals; vein M sharply angled lab-reared hosts, but wild hosts were at bend (Fig. 1); abdomen with 1–2 generally parasitized by single maggots pairs of median discal setae on each of continuously throughout the year in Tri- tergites 3 and 4 (Fig. 1); female ovi- nidad. Schaffner (1959) reported that the positor short and pointed. North American species U. limacodis Fauna.—Uramya is a New World genus and U. pristis develop as solitary parasit- of about 20 described species ranging from oids in their recorded hosts (listed below), Canada to northern Argentina. Twelve have one generation per year, and species were recognized from South overwinter in the pupal stage. All of the America by Guimara˜es (1980), and seven U. pristis reared as part of this study species were recorded from North America followed this pattern. by O’Hara and Wood (2004); however, About 10 morphospecies of Uramya there is no recent revision or key available were reared from five genera of lima- to these species. Janzen and Hallwachs codids (Acharia Hu¨bner, Euclea Hu¨bner, 38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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).

Isochaetes Dyar, Natada Walker, and Literature.—Aldrich (1921) revised the Parasa Moore) by Janzen and Hallwachs Uramya of the Americas under the ge- (2009). neric names Uramya and Pseudeuantha VOLUME 114, NUMBER 1 39

Townsend. Twelve species were treated, portions of the abdomen are typically including only four of the seven species reddish yellow. Uramya limacodis and U. currently recognized from North America. pristis are the only two species of the genus Curran (1930) followed the of in North America that have been reared Aldrich (1921) and provided a key to the from limacodids. They are typically 7–12 three species of Pseudeuantha then known mm long, and only the males are easily from the U.S.A. but excluded without distinguished. In males the long slender comment P. johnsoni.Reinhard(1935) hairs on the anepisternum, from the central described two additional species, one in portion of the sclerite to behind the ane- Pseudeuantha and the other in Uramya. pisternal setae, are black in U. limacodis Sabrosky and Arnaud (1965) reorganized (Fig. 13) and pale in U. pristis (Fig. 12). the species under Anaporia, Pseudeuantha Distribution: Saskatchewan to Nova and Uramya, also adding indita Walker to Scotia, south to Wisconsin and Georgia Pseudeuantha and synonymizing john- (O’Hara and Wood 2004). soni with pristis under Anaporia. Wood Limacodidae: E. delphinii‡ (CNC [North (1987) synonymized Anaporia and Pseu- Burgess Township, Lanark County, deuantha with Uramya and keyed the Ontario, 1973]), Isa textula (H.-S.) genus among other Nearctic genera. This (Schaffner 1959, as Anaporia limacodis classification of seven species of Uramya ex Sisyrosea textula), Limacodes sp. in North America was followed in the (Townsend 1892, as Aporia limaco- catalog by O’Hara and Wood (2004). dis), Li. fasciola (Schaffner 1959, as O’Hara and Wood (2004; catalog) pro- Anaporia limacodis;CNC[NorthBur- vided type and distributional information. gess Township, Lanark County, Ontario, North American Records: 1971]), Tortricidia flexuosa (Grote) Uramya limacodis (Townsend) (Fig. 13): (Aldrich 1921, as Pseudeuantha co- Aporia limacodis Townsend, 1892: 275. quilletti Aldrich; Johnson 1925, as Holotype ?, SEMK (O’Hara and Wood Pseudeuantha coquilletti;Schaffner 2004) (not examined). 1959, as Anaporia limacodis). There is no key to the Uramya of North Uramya pristis (walker) (Figs. 1–2, 4, America, but the species can be readily 6, 12): pristis Walker, 1849: 841. separated into three groups. The first com- Holotype ?, BMNH (O’Hara and Wood prises U. halisidotae and U. aldrichi, 2004) (not examined). which are characterized in the male by See U. limacodis for differential diagnostics. an unusual taillike projection of ab- Distribution: Michigan to Quebec and dominal tergite 5 that extends dorsally New Hampshire, south to Florida; also beyond the male terminalia by about the Arizona (O’Hara and Wood 2004). preceding length of tergite 5. The second Limacodidae: Acharia stimulea group comprises U. indita and U. um- (Clemens) (Aldrich 1921, as Pseudeuan- bratilis. These species are recognized tha pristis ex. Empretia stimulea;Johnson easily by their bi-colored wings in both 1925, as Pseudeuantha pristis ex. Em- sexes: yellow basally and blackish on pretia stimulea), E. delphinii (Coquillett apical half (particularly close to veins). The 1897, as Exorista blanda (Osten Sacken) final three species, U. limacodis, ex Euclea cippus (Cramer); Johnson, U. pristis, and U. rubripes, are more or- 1925, as Pseudeuantha pristis ex Eu- dinary in appearance although the last clea “ceppus”[= cippus] (Cramer)), I. (known only from Florida and Texas) is textula (Coquillett 1897, as Exorista the only species in which the legs and isae Coquillett ex Isa inornata (Grote 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

and Robinson)), Phobetron pithecium (J. from Europe into North America on E. Smith) (Schaffner 1959, as Anaporia multiple occasions beginning in the early pristis). Five additional new limacodid 1900s to control a variety of lepidopteran hosts are reported herein (Table 1): Ad. pests, most notably gypsy moth (L. dispar) spinuloides*, Isochaetes beutenmuelleri*, and browntail moth (Euproctis chrysor- Li. fasciola*, Packardia geminata rhoea (L.)) (Boettner et al. 2000). (Packard)*, Tortricidia sp*. Biology.—Only Com. concinnata has Other hosts: Megalopyge crispata reported host records. This species is (Packard) (Megalopygidae). extremely polyphagous, attacking many Misidentification: The record by species of Lepidoptera in a variety of Coquillett (1897) and cited by Arnaud families and some species of Hymenoptera (1978) of U. pristis (as Macquartia (Symphyta) (Herting 1960, Europe; Arnaud pristis) reared from the arctiid Lopho- 1978, North America; Shima 2006, Japan). campa argentata (as Halisidota ar- Over 180 hosts have been recorded in North gentata)isinerror.Thetachinidreared America, including a single limacodid, M. was U. halisidotae (Aldrich 1921), a flavescens (Schaffner and Griswold 1934, well known parasitoid of L. argentata. Schaffner 1959, repeated in Arnaud 1978; all as Cnidocampa flavescens). Compsilura Bouche´ (Tachinidae: Compsilura species are ovolarvipar- : ) ous, injecting eggs with fully developed (Figs. 5, 7–9) and ready-to-hatch first instars. In Com. concinnata, there is a peculiar adaptation Diagnosis.— known in few other tachinids: the first (Meigen) is the single species of the genus instar occupies the narrow space between in North America. It is typically 7–8 mm the peritrophic membrane and the wall of in length but can be as short as 4 mm. It the midgut and remains there throughout can be identified in North America using most of its three larval instars (Ichiki and the key by Wood (1987) and can be dis- Shima 2003). Bourchier (1991) suggested tinguished from Uramya and Austro- that living in the midgut allows the de- phorocera by the following combination of veloping maggot to evade host immune characters: ocellar seta usually absent; fa- response. cial ridge with row of stout setae on lower The life history of Com. concinnata one-half or more (Fig. 9); prosternum haired is varied according to host species and laterally (Fig. 7); 4 postsutural dorsocentral geographic area. There are two to four setae; metathoracic spiracle with posterior generations per year, often involving al- lappet much larger than anterior one (Fig. ternate hosts throughout the season. In 5); scutellum with 4 pairs of setae, apicals larger hosts, multiple maggots can develop crossed and shorter than laterals, subapicals gregariously. The species overwinters in usually divergent; vein M obtusely-angled the larval stage within a host (Culver 1919, at bend (Fig. 8); abdomen with 1 pair of Schaffner 1959) or possibly in the pupal median discal setae on each of tergites 3 stage away from a host (Tothill 1922). and 4 (Fig. 8); female with short spines Literature.—Shima (1984) diagnosed ventromedially on tergites 3 and 4, and Compsilura and the single Japanese sicklelike piercing ovipositor (Fig. 8). species Com. concinnata in a paper on Fauna.—Compsilura contains four spe- Japanese blondeliine genera cies, mostly of Old World distribution. Robineau-Desvoidy and Compsilura. Compsilura concinnata was introduced Wood (1985) revised and keyed the VOLUME 114, NUMBER 1 41

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). 42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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). VOLUME 114, NUMBER 1 43

Blondeliini of North and Central America from Uramya and Compsilura by the fol- and the West Indies. The genus was keyed lowing combination of characters: ocellar among other Nearctic Tachinidae by seta well developed (Fig. 10); facial ridge Wood (1987) and among other Palearctic with row of stout setae on more than lower Tachinidae by Tschorsnig and Richter one-half (Fig. 10); prosternum haired lat- (1998). O’Hara and Wood (2004) pro- erally (as in Fig. 7); 4 postsutural dorso- vided type and distributional information. central setae; metathoracic spiracle with North American Records: posterior lappet much larger than anterior Compsilura concinnata (Meigen) (Figs. 5, one (as in Fig. 5); scutellum with 4 pairs of 7–9): concinnata Meigen, 1824: setae, apicals crossed and subequal in length 412. Holotype /,NHMW(Herting1972, to laterals; vein M right-angled at bend (Fig. O’Hara et al. 2009) (not examined). 11); abdomen without median discal setae Compsilura concinnata is the most po- on tergites 3 and 4 (Fig. 11); female with lyphagous species of Tachinidae. It has non-piercing ovipositor (Fig. 11). recently been implicated in the decline Fauna.—There are 10 described spe- of native species of saturniid in cies of Austrophorocera in North America New England (Boettner et al. 2000, El- (O’Hara and Wood 2004), and at least 15 kinton and Boettner 2004). Historical species are described from the Neo- evidence suggests it may have success- tropical Region (Wood and Zumbado fully controlled browntail moth, one of 2010), although most of the latter are the invasive pest species it was originally currently assigned to other genera. Janzen introduced against (Elkinton et al. 2006). and Hallwachs (2009) recorded about 20 Distribution: Widespread throughout morphospecies of Austrophorocera in all biogeographic regions of the Old Guanacaste, Costa Rica, of which only World (Shima 1984). Introduced into two are referable to named species. There North America and currently known is no modern key to the New World spe- from most of southern Canada and cies of Austrophorocera. There are five species in the Oriental and Australasian/ western and northeastern U.S.A. Oceanic regions (Crosskey 1976, Cantrell (O’Hara and Wood 2004). The species and Crosskey 1989), with two of those may still be expanding its range in species reaching the Palearctic part of North America. China (O’Hara et al. 2009). Limacodidae: M. flavescens (Webber Biology.—Austrophorocera species are and Schaffner 1926, Schaffner and oviparous. Eggs are laid directly on a host Griswold 1934, Schaffner 1959; all and hatch within a few days; the first in- as Cnidocampa flavescens)andE. star then uses its mouth hooks to burrow delphinii*. into the host. Two or three species have Other hosts: Over 180 species of Lepi- been reared in North America (one nom- doptera and Symphyta in North America inal), and except for one record from (Arnaud 1978, Boettner et al. 2000). a notodontid, the hosts have all belonged to Limacodidae (see below). Janzen and Austrophorocera Townsend Hallwachs (2009) documented 307 (Tachinidae: Exoristinae: ) Austrophorocera specimens (20 - (Figs. 10–11, 14–17) species, two nominal) reared from 14 gen- Diagnosis.—The genus can be identi- era and > 20 species of Limacodidae in fied in North America using the key by Guanacaste. A widespread species in the Wood (1987) and can be distinguished Oriental and Australasian regions, A. 44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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. VOLUME 114, NUMBER 1 45 grandis (Macquart) has been reared from North American Records: several species of Thosea Walker and Dor- Austrophorcera cocciphila (Aldrich and atifera Duncan (Limacodidae) (Crosskey Webber): Phorocera (Parasetigena) 1976, Chadwick and Nikitin1985), as well cocciphila Aldrich and Webber, 1924: as Heliothis sp. () (Cantrell 53. Holotype ?, USNM (O’Hara and 1986). Wood 2004) (not examined). Schaffner (1959) briefly reviewed the Two of the North American species life history of A. coccyx parasitizing two of Austrophorocera, A. cocciphila and limacodid species (see below). He re- A. einaris, are easily distinguished from ported one parasitoid per host, one gen- the other species by having two katepis- eration per year, and larva overwintering ternal setae instead of three. The male in the prepupal stage of the host. Murphy syncercus of these two species is densely et al. (2009) reported on the undescribed setose posteriorly (Figs. 14, 16). The se- A. sp. recorded in the present study, stat- tae are black, and their outer portions ing that it is a larval-pupal parasitoid that have a crinkled appearance. The setae parasitizes the larva and emerges from form a thicker and tighter mat in A. ei- the host’s the following spring. Host naris (Fig. 16) than in A. cocciphila (Fig. pupation appears to be a trigger for de- 14), and the mat in A. einaris is positioned velopment of A. sp., because flies have more centrally on the more elongate syn- repeatedly been observed to eclose from cercus of that species. The syncercus of A. non-diapausing host cocoons in the one einaris in lateral view is distinctly bent North American limacodid species with subapically, compared to slightly curved a partial second generation, E. delphinii. subapically in A. cocciphila. Literature.—Aldrich and Webber Distribution: District of Columbia, (1924) treated seven of the 10 species , Ohio, Tennessee (O’Hara of Austrophorocera recognized currently and Wood 2004). from North America as Phorocera Limacodidae: Ac. stimulea. Robineau-Desvoidy (subgenus Paraseti- Other hosts: None known. gena Brauer and Bergenstamm) in part. Austrophorocera coccyx (Aldrich and Sabrosky and Arnaud (1965) dispersed Webber) (Fig. 15): Phorocera (Para- the species of P.(Parasetigena)among setigena) coccyx Aldrich and Webber, several genera but kept together six 1924: 64. Holotype ?,USNM(O’Hara of the seven Austrophorocera species and Wood 2004) (not examined). in the genus Palpexorista Townsend. Austrophorocera coccyx and A. n. sp. be- These authors added three additional long to a group of seven North American species to Palpexorista that had been Austrophorocera species that have three described in Phorocera after Aldrich katepisternal setae and a patch of short, fine, and Webber (1924). Wood (1987) synon- wavy yellowish hairs within a depressed ymized Palpexorista under Austrophoro- area on the basal portion of the posterior cera, a genus formerly known only from surface of the male syncercus. Two of the the Old World. Thus, the nine species seven species, A. pellecta (Reinhard) assigned to Palpexorista by Sabrosky and (Arizona) and A. tuxedo (Curran), are Arnaud (1965) were treated under Austro- smaller than the rest at about 6–9 mm phorocera by O’Hara and Wood (2004). long; the former has an unusually broad A tenth species was added as a new North parafacial, and the latter is yellowish on American record by O’Hara and Wood the upper portion of the head (fronto- (2004). orbital plate) and the thoracic dorsum. 46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The remaining five species comprise Limacodidae: E. delphinii. A. alba (Townsend), A. coccyx, A. stol- Other hosts: None known. ida (Reinhard), A. virilis (Aldrich and Austrophorcera imitator (Aldrich and Webber), and A. n. sp. These species are Webber): Phorocera (Parasetigena) im- moderately large tachinids (about 10–13 itator Aldrich and Webber, 1924: 63. mm long except for A. n. sp., which is Holotype ?, USNM (O’Hara and Wood 8.5–10.0 mm long) and are grayish black 2004) (not examined). in coloration. They are most easily Austrophorocera imitator and another separated by distinct differences in North American species, A. sulcata their male syncercus: A. alba (and other (Aldrich and Webber), have three kate- forms that may represent undescribed pisternal setae and only black setae species) has the posterior surface of the posteriorly on the male syncercus. The syncercus expanded laterally at mid- setae on the syncercus are shorter and length to form a lobe on either side, and more regular in appearance than in A. the apical portion of the syncercus einaris and A. cocciphila. The syncercus is short and narrow; in A. coccyx (Fig. 15) in both species is slender and evenly the syncercus in posterior view is tapered in posterior view; in lateral view abruptly narrowed at midlength and ta- the apex is slightly clubbed and bent pered to a pointed tip; in A. stolida the posteriorly in A. imitator and tapered to syncercus is gently narrowed at mid- a point and slightly curved anteriorly in length, tapered to a rounded tip, and A. sulcata. The surstylus of A. imitator is bears a tuft of dense black setae basally distinctive among the North American above the area with the short yellow species of the genus in being tightly ap- hairs; in A. virilis the syncercus is smoothly pressed against, and possibly fused with, tapered from base to tip; in A. n. sp. (Fig. the syncercus along its length, giving the 17) the syncercus is broadened on lower appearance on first inspection of being third and abruptly narrowed subapically absent. to a pointed tip. Distribution: Missouri to Massachusetts, Distribution: Ontario to New York, south to Texas and Florida (O’Hara south to Florida, also Arizona and and Wood 2004). Texas (O’Hara and Wood 2004). Limacodidae: Pr. badia. Limacodidae: Ac. stimulea (Aldrich and Other hosts: None known. Webber 1924, as Pho. coccyx;Schaffner Austrophorocera n. sp. (Fig. 17) Murphy 1959, as Pho. longiuscula (Walker)) and et al. 2009 (Austrophorocera n. sp.). Pho. pithecium (Schaffner and Griswold See A. coccyx for differential diagnostics. 1934, as Pho. longiuscula;Schaffner Distribution: Eastern U.S.A. 1959, as Pho. longiuscula). Limacodidae: Reported from “Lima- Other hosts: Datana ministra (Drury) codidae” in Murphy et al. (2009) and (Notodontidae). Li. fasciola* (Table 1). Austrophorcera einaris (Smith): Phor- Austrophorocera “sp. 1”: Stireman and ocera einaris Smith, 1912: 119. Holotype Singer 2003a, b. ?,USNM(O’HaraandWood2004)(not Not examined and characters not known. examined). Distribution: Studied in Arizona. See A. cocciphila for differential diagnostics. Limacodidae: Pa. chloris (H.-S.) Distribution: “Michigan to Quebec and (Stireman and Singer 2003b), Pro. Massachusetts, south to Texas and trigona (Edwards) (Stireman and Singer Florida” (O’Hara and Wood 2004). 2003a, b). VOLUME 114, NUMBER 1 47

Hymenoptera of Ceraphron)waspublishedbyDessart and Cancemi (1986). Ceraphron Jurine North American Records: (Ceraphronoidea: Ceraphronidae) Ceraphron sp. (Figs. 74–77): Diagnostics (Figs. 74–77) unavailable for North American species. Diagnosis.—Vertex rounded (Fig. 74), Distribution: This cosmopolitan genus ocellar triangle small, distant from the is widespread in North America. eyes; median furrow present on the mes- Limacodidae: Hyperparasitoid that oscutum (Fig. 75) (Dessart and Cancemi emerged from a primary parasitoid 1986), and dorsum of the latter flat in lat- (possibly Pl. americana* or Al. lilli*) eral view (Austin 1987b). Adult speci- of Ac. stimulea on Q. rubra. mens of Ceraphron,particularlymales, Other hosts: Unknown for this species. are difficult to identify to species due to their simplified genitalia (I. Miko´ pers. Conura Spinola (Chalcidoidea: comm.) and the dearth and age of pub- Chalcididae: ) lished information for the Nearctic Re- (Figs. 78–86) gion (Ashmead 1904, Brues 1906). Diagnosis.—Conura may be recognized Fauna.—There is little known about by postmarginal vein longer than stigmal the diversity of Ceraphron; no compre- vein (Fig. 78); gaster distinctly petiolate hensive treatments have been published regionally or globally. Twenty-four spe- (Figs. 79–80); and propodeal spiracle cies were recorded from North America subvertical to nearly longitudinal. by Brues (1906). Fauna.—Conura is a large genus of Biology.—Species of Ceraphron are ; 296 species restricted to the New commonly considered hyperparasitic on World, primarily the Neotropical Region primary hymenopteran and dipteran par- (283 species), with only 12 species in the asitoids based largely on records from the Nearctic (Noyes 2003). Species have been related genus Aphanogmus (recorded from recovered as far north as Alaska and as far Ichneumonidae and Braconidae by Austin south as Chile and Argentina (Noyes (1987b)). Other species of Ceraphron have 2003). For a discussion of the classification been reared as hyperparasitoids (through of Conura and related genera, see Delvare Hymenoptera) from both Aphididae (Evans (1992). Of the known species from the and Stange 1997) and Cecidomyiidae Nearctic, two (C. nigricornis, C. nortonii) (Franklin 1919), among other hosts. are specialists on Limacodidae and belong Literature.—Austin (1987b) recorded to the nigricornis group, whose members asingleCeraphron sp. hyperparasitoid exclusively attack limacodids. The third from one limacodid rearing from Southeast (C. camescens)isapparentlypolyphagous, Asia. Brues (1906) published a key to also known as a secondary parasitoid of Nearctic Ceraphron and followed this Hymenoptera through Lepidoptera (see with a key to genera and species of Cera- below), and belongs to the immaculata phronidae (Ceraphronoidea) in Connecticut group. Delvare (1992) recorded the biology (Brues 1916). Ashmead (1893) reported on of this group as hyperparasitoids of ich- the genera and species of Ceraphronidae neumonoids through lepidopterans. The known in North America, and Dessart members of this group are recognized by (1981) proposed species groups of Cera- the superficial malar sulcus (Fig. 82), phron.Themostrecentkeytotheworld interantennal projection a weak carina, genera of Ceraphronoidea (incl. subgenera shallow scrobal depression (Fig. 81), and 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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). VOLUME 114, NUMBER 1 49

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). 50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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. VOLUME 114, NUMBER 1 51

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. 52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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 = 1 mm. Figs. 37–38. Baryceros texanus, female. 37, Mesosoma, lateral (arrow: epomia); 38, Ovipositor tip. reticulo-rugose to rugose mesoscutum. (Figs. 84–85), and a moderately deep Burks (1940) defined the nigricornis group scrobal depression with slightly cari- based on deep and coarse body sculpture, nate margins. Delvare (1992) refined the the presence of conspicuous long pilosity definition of this group by noting notauli VOLUME 114, NUMBER 1 53

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). 54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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. VOLUME 114, NUMBER 1 55

Figs. 50–51. Species of Trirhaphis. 50, Trirhaphis eupoeyiae, female, habitus; 51, T. harrisinae, female, habitus. Scales = 1.0 mm. 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 52–55. Trirhaphis harrisinae, female. 52, Metasoma, anterodorsal (arrows, median carina); 53, Mesosoma, dosal; 54, Mesosoma, lateral, scale = 0.25 mm; 55, Propodeum. VOLUME 114, NUMBER 1 57

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). deep, broad, and crenulate (Fig. 84); pro- Biology.—Generally, species of Conura notum with recurved lamina anteriorly are documented as parasitoids of larvae (Fig. 85); and mesopleuron with ventral and pupae of different insects, mostly of carina (Fig. 83). Lepidoptera. However, some species of 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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). VOLUME 114, NUMBER 1 59

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). 60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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. VOLUME 114, NUMBER 1 61

Figs. 70–71. Orthogonalys pulchella. 70, Habitus, female; 71, Habitus, male. Scales = 4.0 mm.

Conura are recorded from Coleoptera, smaller species of Conura and Brachy- Hymenoptera, Diptera, and , meria are gregarious. and others are known secondary para- Literature.—Boucˇek (1992; key to sitoids of ichneumonoids and tachinids genera) revised the New World genera, through lepidopterans (Delvare 1992, Delvare treated the New World Boucˇek and Halstead 1997). Three spe- (1992; key to species groups of Conura; cies that occur in North America are side group keyed to species), and Burks known to attack limacodids, two of (1940) offered an earlier treatment of these exclusively (see below). Known Chalcidini, with keys to genera, species biologies indicate chalcidid host special- groups, and species. ization as solitary parasitoids, but some North American Records: 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 72–73. Taeniogonalos gundlachii. 72, Habitus, female (arrow: armature); 73, Habitus, male. Scales = 2.0 mm.

Conura nigricornis (F.): nigricornis This species belongs to the rarely col- F., 1798: 243. Lectotype /,ZMUC,des- lected nigricornis group (Burks 1940), ignated by Boucˇek and Delvare (1992) which contains 14 nominal (at least 16 (not examined). undescribed) species, all of which are VOLUME 114, NUMBER 1 63

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,forewing,scale= 1.0 mm. known solitary parasitoids of Limaco- frenal carina is present and not fused didae and Megalopygidae (Delvare with the apical margin of the scutellum. 1992). This species is yellow with black The propodeum has distinct lateral teeth maculae on the body as reported by (fig. 42 in Boucˇek and Delvare 1992). Boucˇ ek and Delvare (1992: 29), the Burks (1940) separates C. nigricornis scutellum has a median groove, and the from C. nortonii by the presence of five 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 79–80. Species of Conura. 79, Conura nortonii, male, habitus. 80, Conura immaculata, fe- male, habitus. Scales = 2.0 mm. to six hamuli on the hind wing and the Distribution: South America (Brazil, presence of an inner tooth on the hind Guyana) and eastern U.S.A. femur as compared to C. nortonii (see Limacodidae: Adoneta sp. (Boucˇek and below). Delvare 1992), Ad. spinuloides (Peck 1963, VOLUME 114, NUMBER 1 65

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).

Herting 1976, Burks 1979), Apoda sp. Conura nortonii (Cresson) (Figs. 78– (Peck 1963, Burks 1979, Boucˇek and 79, 83–86): Smicra nortonii Cresson, Delvare 1992), Parasa sp. (Boucˇek and 1872: 45. Neotype ?,USNM,des- Delvare 1992), Pa. indetermina (Hert- ignated by Delvare (1992: 239) ing 1976, Burks 1979). (examined). 66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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. VOLUME 114, NUMBER 1 67

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). 68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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.

This species also belongs to the nigricornis USNM is more orange brown (perhaps group. Schaffner (1959) reported that two killed with cyanide) than yellow. Com- specimens, one per host, were reared pared to C. nigricornis,thepropodeum from Ph. pithecium cocoons collected in lacks lateral teeth and the scutellum Bolton, Connecticut. The base color of lacks a median groove, although there is this species is yellow with black mac- minute lamina apically (Burks 1940). ulation, although the neotype in the Conura nortonii has three hamuli on the VOLUME 114, NUMBER 1 69 hind wing and lacks the inner tooth on the Alveoplectrus Wijesekara and Schauff hind femur. (Chalcidoidea: Eulophidae: Eulophinae) Distribution: Eastern U.S.A., west to (Fig. 104) Texas (Burks 1940). Diagnosis.—Alveoplectrus may be re- Limacodidae: Apoda sp. (Peck 1963, cognized by the presence of two sub- Burks 1979), Ph. pithecium (Schaffner median carina on the propodeum (Fig. 1959, Peck 1963, Herting 1976, Burks 104), scutellum with lateral grooves broad 1979), Pr. badia (Herting 1976, Burks and continuous posteriorly (Fig. 104), 1979). dorsellum produced medially as cuplike Conura camescens Delvare: Ceratosmicra carina (Fig. 104), and hind tibia with flavescens Cameron, 1913: 114. Conura single elongate spur. The related genus camescens is a replacement name for Platyplectrus (see below) has a single Conura flavescens (Cameron 1913) nec median carina on propodeum, scutellum (Andre´ 1881). Lectotype ?,BMNH, designated by Delvare (1992: 217) (not with narrow lateral grooves discontinuous examined). posteriorly, dorsellum not produced as This species belongs to the immaculata triangular point posteromedially, and hind subgroup of the immaculata group tibia with two elongate spurs (> 0.5X (Delvare 1992). Delvare (1992) reported length of hind tarsus). this species from the U.S.A. without fur- Fauna.—Alveoplectrus is a small ge- ther locality information. A specimen nus of four species restricted to the from Venezuela in the USNM identified New World, primarily the eastern Nearctic by Delvare has label data indicating and Neotropical regions (Wijesekara and (Joh.) as a host [possibly Schauff 1997, Gates and Stoepler 2010). on primary parasitoid]. Two additional Species of Alveoplectrus were previously specimens, also from Venezuela, indicate placed in Euplectrus (Ashmead 1904) Alabama argillacea (Hu¨bner), “par. by or Euplectromorpha (see Schauff et al. Rogas.” This species is reported as a pale 1997). The published distribution for Al- brownish yellow with areas of the face, veoplectrus includes Brazil, Costa Rica, pronotum, mesoscutal sutures, and scu- and the U.S.A. (Florida) (Wijesekara and tellum in basal half more yellow in color Schauff 1997). For a discussion of the (Cameron 1913). The ; 13 teeth of the classification of Alveoplectrus and re- hind femur are black, the basal tooth lated genera, see Wijesekara and Schauff triangular with the apical teeth closely (1997). appressed. Biology.—Known biologies indicate Distribution: New World. (Delvare 1992). host specialization as solitary ectopar- Limacodidae: Acharia sp. (as Sibine; asitoids of larvae of Limacodidae. One hyperparasitoid of Cotesia sp.) (Delvare species, Al. floridanus Wijesekara and 1992). Schauff, is recorded from label data for Other hosts: Prodenia eridania undetermined species in USNM indicat- (Cramer) (Noctuidae); Plutella xy- ing Neotropical Euprosterna elaea (as lostella (L.) (); Apanteles elaesa)DyarandEuprosterna sp. (Li- sp., Cotesia sp., Co. plutellae (Kurd- macodidae) as hosts. jumov), and Rogas sp. (Braconidae) North American Records: (De Santis 1979, De Santis and Fidalgo Alveoplectrus floridanus Wijesekara and 1984, Delvare 1992, Cha´vez et al. Schauff: Wijesekara and Schauff, 1997: 1993). 105. Holotype /, USNM (examined). 70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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).

This species is differentiated from Al. (compare with Gates and Stoepler 2010: lilli by the absence of a short, transverse fig. 2). carina beneath the anterior ocellus and Distribution: U.S.A. (Florida) (Wijesekara the presence of a transepimeral sulcus and Schauff 1997). VOLUME 114, NUMBER 1 71

Limacodidae: Ala. slossoniae (Packard) Australia and eastern Europe (Noyes (Wijesekara and Schauff 1997). 2003). Alveoplectrus lilli Gates (Fig. 104): Gates Biology.—Known biologies indicate and Stoepler 2010: 209. Holotype /, host specialization as solitary or gre- USNM (examined). garious ectoparasitoids of larvae of This species is differentiated from Al. flor- Limacodidae. Laboratory studies of the idanus by the presence of a short, trans- extralimital Pl. natadae Ferrie`re (Gadd verse carina beneath the anterior ocellus et al. 1946) revealed this species engaged and the absence of a transepimeral sul- in host feeding once for every four eggs cus (figured in Gates and Stoepler 2010: laid, and a host used for host feeding was fig. 2). never parasitized. Oviposition occurs on Distribution: Eastern U.S.A. (Gates the lateral surfaces below any tubercles of and Stoepler 2010). the host after the host is paralyzed with a Limacodidae: I. textula, Li. fasciola, sting. The egg, positioned externally, is held E. delphinii (Gates and Stoepler 2010), in place by a pedicel within the host’s body. and Ad. spinuloides*. In nature, this species usually lays one or rarely lays two eggs per host with only one Platyplectrus Ferrie`re (Chalcidoidea: larva typically reaching maturity. However, Eulophidae: Eulophinae) in the present study, the number of eggs (Figs. 105–113) per host appears to be size-dependent, with larger hosts (or later instars) yielding Diagnosis.—Platyplectrus is most as many as 14 mature larvae. A newly likely to be confused with Alveoplectrus eclosed larva migrates to the venter of the (among parasitoids of limacodids in North host’s body where feeding begins imme- America) and Euplectrus,althoughspe- diately and results in the death of the host. cies in the latter genus are not known from Pupation occurs under the host with the limacodids in North America. Species of host cadaver providing a protective cover- Platyplectrus may be recognized by the ing (Gadd et al. 1946, M. Gates pers. obs.). following combination of features: single Literature.—Only eight species have median carina on propodeum (Fig. 105), associated host records, with five of those scutellum with narrow lateral grooves dis- records being from limacodids. The continuous posteriorly (Fig. 105), dorsel- widespread Pl. laeviscuta (Thomson) lum not produced (Fig. 105), and hind tibia ranges from Sweden and China south to with two elongate spurs. Alveoplectrus has Australia and is cited from Stigmella rufi- two submedian carinae on propodeum capitella (Haworth) (Nepticulidae) (Noyes (Fig. 104), scutellum with lateral grooves 2003), but we find no record of it in the broad and continuous posteriorly (Fig. original paper (Hansson 1987). Platyplec- 104), dorsellum produced medially (Fig. trus chlorocephalus (Nees) is recorded 104), and hind tibia with single elongate from a zygaenid ( pruni (Denis spur. Euplectrus,asidefromitshostpref- and Schiffermu¨ller)), and this is perhaps erences, is most similar to Platyplectrus in unsurprising given the relationship be- overall coloration and habitus but lacks tween and Limacodidae scutellar grooves. (Epstein 1996, Wahlberg et al. 2005, Fauna.—One species is known from Niehuis et al. 2006). The third non-limacodid North America of 46 described worldwide. record published by Zhu and Huang Highest species richness is in southeastern (2004) was based on label data: Cifuna Asia, but several species are known from locuples Wlkr (Lymantriidae). 72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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. VOLUME 114, NUMBER 1 73

Figs. 111–113. Platyplectrus americana, female. 111, Antenna; 112, Fore wing; 113, Antenna, male.

Redescription of Platyplectrus americana brown—dorsal half flagellum, tegula, (Figs. 105–113): metafemur in apical half except extreme Diagnosis.—Platyplectrus americana is apex golden, pretarsus, gaster apically, lat- the sole representative of the genus in erally, and ventrally, wing veins; white— North America and thus will not be con- basal three tarsomeres. fused with congeners. Platyplectrus can be Head: 1.7X as broad as high, clypeal differentiated from related eulophid genera region transversely impressed along mar- by the generic characters outlined above. gin, smooth, straight ventrally; malar sul- Note.—Girault (1916) described this cus fine, slightly curved, malar space 0.53X species from a single male. The original eye height; mandibles reduced, not touching description is short, and additional mor- medially; toruli with upper margin posi- phological details of the male require tioned one-half torular diameter above documentation, as given below. The fe- lower ocular line, separated by ; 2torular male is newly described. diameters (Fig. 107). Scrobal depression Description.—Female. Body length: smooth and shallow, shaped as an inverted 2.4 mm (1.2–3.5 mm; n = 20). Color: V. Antenna (Fig. 111) with scape reaching Black except for the following golden— three-quarters distance to anterior ocellus; scape, flagellum in ventral half, fore coxa ratio of scape (minus radicle):pedicel: in apical third, legs, apical tarsomere, and anellus: F1:F2: F3:F4:club as 27:8:1:16: basal half gaster dorsomedially (Fig. 106); 14:12:12:18; funicular segments with 2 74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON irregular rows of interdigitated longitu- sally; Gt1 depressed anterodorsally near dinal sensilla and evenly setose; clava petiole occupying about one-third the apparently bisegmented with fusion com- length of gaster; remaining terga and syn- plete. Ratio of lateral ocellus:OOL:POL tergum smooth; [petiole transverse in dor- as 7:7:14. [Head posteriorly lacking post- sal view, smooth; lateral carinae on petiole occitpital carina, hypostomal carinae pres- body meeting ventrally as slightly pointed ent ventrally but faint (Fig. 108).] collar]; measurements of gastral terga Mesosoma (Fig. 105): 1.3X as long as along midline as 30:9:9:7:5:7:8. broad; midlobe of mesoscutum 0.9X as Male. Body length: 1.5 mm (1.2–2.8 mm; long as broad; scutellum 1.0X as long n = 5). Color: As described for female, as broad; notauli fine, complete; midlobe but with legs and venter of gaster paler, of mesoscutum shallowly bilobate post- whitish. Sculpture as described for fe- eromedially; scutellum shallowly bilobate male. Antenna with funicular segments anteromedially, sublateral grooves mesad (Fig. 112) minutely pedicellate, each with scutellar setae, extending from scutoscu- multiple rows of suberect setae and about tellar suture to near posterior edge scu- 1.0X as long as width of segment; ratio of tellum; scutoscutellar suture obliterated scape (minus radicle):pedicel:anellus: medially; axilla smooth with few setae F1:F2:F3:F4:F5:club as 16:7:1:8:8:8:8:15; posteromedially; lateral surface of pre- scape with ventral plaque not apparent. pectus triangular, smooth; [subventral ca- Gastral petiole in lateral view roughly rinae of prepectus steplike, prepectus with cylindrical, roughly trapezoidal in dorsal ovate depression ventromedially] (Fig. 109). view, length about 0.8X as long as Mesepimeron smooth, femoral depression greatest width. asigmoidalsulcus,smalllineardepression Variation.—Most variation occurs in posteromedially. Mesepisternum smooth coloration and size. Smaller male spec- (Figs. 109–110), convex, defined poste- imens (< 1.8 mm; range = 1.1–1.8 mm) riorly by carina, discrimen medially be- tend toward a browner coloration rather tween mesofurcal pit and anterior margin than black (especially the head and mesopectus. Dorsellum smooth with post- mesosoma) with brownish maculation eromedial truncation, subequal in length on the femora only a faint suffusion with propodeum medially, slightly pro- (especially fore and midfemora) and the jecting over propodeum medially and fore femur may be entirely golden/whitish excavated submedially (Fig. 105). Meta- whereas the hind femur may approach pleuron and lateral areas of propodeum being entirely dark brown. The whitish (Fig. 109) smooth; callus setose; propo- maculation dorsally on the gaster may be deum smooth with elevated median carina confined only to that area but can extend broadest anteriorly and posteriorly broadly laterally and ventrally. Two males are joined to nucha, with fine rugulosity; almost golden in body color. For females flangelike carina extending laterally from (range = 1.3–2.3 mm), the coloration nucha toward metasomal foramen and spi- patterns seen for males based on size seems racle; spiracle about one-third its greatest to hold true. One specimen is medium diameter from dorsellum. All coxae im- brown with white legs, except for brown bricate, smoother laterally. Fore wing with suffusion on the hind femur; the antenna ratio of marginal vein:postmarginal vein: is whitish. For both sexes, there are ex- stigmal vein as 64:45:23 (Fig. 113). ceptions to the size/color trend but typically Metasoma: Smooth; petiole ; 2.0X only at the smaller size range (dark, small as broad as long, finely sculptured dor- specimens). VOLUME 114, NUMBER 1 75

Type Material.—20471 [red label, ex Acharia stimulea on Quercus alba handwritten]; Euplectromorpha americana (1/ USNM); same locality, collected ?,Type,Girault[handwritten](USNM, 25.VII.2007, emerged 21.VIII.2007, J. examined). Lill et al. 07-319, ex Acharia stimulea on Material Examined.—DISTRICT OF Prunus serotina (2/?USNM); same COLUMBIA: Rock Creek Park, col- data, 07-316, (1/ USNM); same data, lected 9.VIII.2005, J. Lill et al. 05-607, 07-315, (2/ USNM); same locality, ex Acharia stimulea on Robinia sp. (1/ collected 25.VII.2007, J. Lill et al. 07- USNM); same locality, collected 12. 340, ex Acharia stimulea on Quercus VII.2006, emerged 23.VII.2006, J. Lill, rubra (1/ USNM); same data, 07-339, et al., 06-281, ex Limacodidae on Carya (1/ USNM); same locality, collected glabra (? USNM); same locality, col- 25.VII.2007, emerged 21.VIII.2007, J. lected 23.VII.2006, emerged 14.VIII.2006, Lill et al. 07-338, ex Acharia stimulea on J. Lill, et al. 06-313, ex Lithacodes fasciola Quercus rubra (1/ USNM); same lo- on Nyssa sylvatica (/?USNM); same cality, collected 9.VII.2008, emerged 25. locality, collected 14.VIII.2006, emerged VII.2008, J. Lill et al. 08-152, ex Lith- 21.VIII.2006, J. Lill et al. 06-326, ex Pro- acodes fasciola on Fagus grandifolia limacodes badia on Quercus alba (/ (1/ USNM); same locality, collected 9. USNM); same locality, collected 13. VII.2008, emerged 18.VII.2008, J. Lill VIII.2006, emerged 3.IX.2006, J. Lill et al., et al. 08-155, ex Euclea delphinii on 06-336, ex Acharia stimulea on Acer sac- Quercus alba (1/ USNM); same local- charinum (/ USNM); same locality, col- ity, collected 10.VII.2008, emerged 25. lected 20.VIII.2006, emerged 3.IX.2006, J. VII.2008, J. Lill et al. 08-175, ex Euclea Lill et al. 06-387, ex Euclea delphinii on delphinii on Fagus grandifolia (1/ Nyssa sylvatica (10/ USNM); same lo- USNM); same data, 08-175 (1/ USNM); cality, collected 20.VIII.2006, J. Lill et al., same data, 08-176, (1/ USNM); same 06-389, ex Adoneta spinuloides on Nyssa data, 08-177, (1/ USNM); same data, 08- sylvatica (6/ USNM); same locality, col- 178 (1/ USNM); same locality, collected lected 25.VIII.2006, J. Lill et al., 06-408, 10.VII. 2008, emerged 28.VII.2008, J. ex Lithacodes fasciola on Nyssa sylvatica Lill et al. 08-186, ex Euclea delphinii on (3/ USNM); same locality, collected 26. Carya glabra (1/ USNM); same locality, VI.2007, emerged 17.VII.2007, J. Lill et al. collected 10.VII.2008, emerged 21. 07-120, ex Lithacodes fasciola on Quercus VII.2008, J. Lill et al. 08-187, ex Euclea rubra (/ USNM); same data, but collected delphinii on Carya glabra (1/ USNM); 30.VI.2007, emerged 17.VII.2007, 07-140, same locality, collected 11.VII.2008, ex Lithacodes fasciola on Nyssa sylvatica emerged 25.VII.2008, J. Lill et al. 08- (? USNM); same data, 07-143, ex 200, ex Euclea delphinii on Quercus ru- Limacodidae sp. on Fagus grandifolia (? bra (1/ USNM); same locality, collected USNM); same locality, collected 30. 14.VII.2008, emerged 28.VII.2008, J. VI.2007, emerged 17.VII.2007, J. Lill Lill et al. 08-217, ex Euclea delphinii on et al., 07-144, ex Limacodidae sp. on Nyssa sylvatica (1/ USNM); same lo- Fagus grandifolia (? USNM); same cality, collected 19.VII.2007, emerged data, 07-148 (/ USNM); same data, 07- 21.VIII.2007, J. Lill et al., 07-260, ex 149 (? USNM); same data, 07-152 (? Lithacodes fasciola on Quercus rubra USNM). MARYLAND: Montgomery (3/ USNM); same locality, collected 19. Co.: Little Bennett Regional Park, col- VII.2007, emerged 4.VIII.2007, J. Lill lected 25.VII.2007, J. Lill et al. 07-320, et al., 07-292, ex Phobetron pithecium on 76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fagus grandifolia (3? USNM); same (2/ USNM); same locality, emerged 2. data, 07-293 (? USNM); same data, 07- VIII.2007, 07-996, ex Euclea delphinii on 295 (2? USNM); same locality, collected Prunus serotina,(3/?USNM); same 25.VII.2007, emerged 21.VIII.2007, J. Lill data, 07-995, (? USNM); same data, 07- et al. 07-307, ex Euclea delphinii on Pru- 997, (1/ USNM); same locality, [No nus serotina (3/ USNM); same locality, date], 07-993, ex Euclea delphinii on collected 25.VII.2007, J. Lill et al. 07-351, Nyssa sylvatica (2/ 2? USNM); same ex Acharia stimulea on Nyssa sylvatica data, emerged 2.VIII.2007, 07-999, ex (1/ USNM); same data, 07-346, ex Euclea delphinii on Quercus rubra, (1/ Acharia stimulea on Carya glabra (1/ USNM); same data, 07-994, ex Euclea USNM); same data, 07-343 (2/ USNM); delphinii on Fagus grandifolia (2/ same data, 07-345 (2/ 2? USNM); USNM); same data, 07-991, ex Acharia same data, 07-349, ex Acharia stimulea stimulea on Quercus alba (1/ USNM); on Nyssa sylvatica (1? USNM); same same locality, collected 3.VIII.2007, data, 07-354 (4/?USNM); same lo- emerged 21.VIII.2007, J. Lill et al. 07- cality and date, emerged 21.VIII.2007, J. 526, ex Acharia stimulea on Fagus gran- Lill et al. 07-363, ex Lithacodes fasciola difolia (1/ USNM); same data, 07-522, on Fagus grandifolia (/?USNM); ex Acharia stimulea on Prunus serotina same data, 07-367, ex Acharia stimulea (1/ USNM); same locality, collected 5. on Fagus grandifolia (/?USNM); VIII.2008, emerged 18.VIII.2008, J. Lill same data [no emergence date], 07-368 et al. 08-632, ex Acharia stimulea on (1/ USNM); same locality, collected 1. Nyssa sylvatica (3/ USNM); same lo- VIII.2007, J. Lill et al. 07-466, ex Acha- cality, collected 31.VII.2008, emerged 13. ria stimulea on Quercus alba (2/? VIII.2008, J. Lill et al. 08-527, ex Acharia USNM); same data, 07-467 (2/ USNM); stimulea on Nyssa sylvatica (1/ USNM); same data, emerged 21.VIII.2007, 07-470 same locality, collected 18.VII.2007, (/?USNM); same data [no emergence emerged 2.VIII.2007, J. Lill et al. 07-998, date], 07-471, ex Acharia stimulea on ex Euclea delphinii on Quercus rubra (1/ Quercus rubra (1/ USNM); same data, USNM); same locality, 23.VIII. 2007, 07-489, ex Adoneta spinuloides on Carya emerged 1.VI.2008, J. Lill et al. 07-672, ex glabra (? USNM); same locality, col- Prolimacodes badia on Quercus rubra lected 27.VII.2007, J. Lill et al. 07-499, (1/ USNM); same locality, collected 3. ex Euclea delphinii on Quercus rubra VIII. 2007, emerged 21.VIII.2007, J. Lill (1/ USNM); same locality, collected 3. et al. 07-524, ex Acharia stimulea on Fa- VIII.2007, emerged 21.VIII.2007, J. Lill gus grandifolia (1/ USNM); same local- et al. 07-558, ex Prolimacodes badia on ity, collected 14.VII.2008, emerged 1. Fagus grandifolia (1/ USNM); same VIII.2008, J. Lill et al. 08-226, ex Euclea data, 07-561 (? USNM); same data, 07- delphinii on Nyssa sylvatica (/? 563 (? USNM); same data, 07-564 (1/ USNM); same data, 08-234 (2/ USNM); USNM); same locality, collected 14. same locality, collected 28.VII.2008, VIII.2007, emerged 1.VI.2008, J. Lill et al. emerged 11.VIII. 2008, J. Lill et al. 08- 07-581, same host (1/ USNM); same lo- 328, ex Acharia stimulea on Quercus cality, collected 31.VII.2008, emerged 11. rubra (3/ USNM); same locality and VII.2008, J. Lill et al. 08-528, ex Acharia collection date, emerged 19.VIII.2008, J. stimulea on Nyssa sylvatica (3/ USNM); Lill et al. 08-326, ex Acharia stimulea on same data but emerged 26.VIII.2008, 08- Nyssa sylvatica (4/ USNM); same lo- 536, ex Acharia stimulea on Carya glabra cality, collected 31.VII.2008, emerged 18. VOLUME 114, NUMBER 1 77

VIII.2008, J. Lill et al. 08-525, same host et al. 05-899, ex Lithacodes fasciola on (1/ USNM); same locality, collected 31. Fagus grandifolia (3/?USNM); same VII. 2008, emerged 11.VII.2008, J. Lill locality, collected 6.IX.2005, emerged et al. 08-510, ex Acharia stimulea on Pru- 10.VI.2006, J. Lill et al. 05-916, ex nus serotina (3/?USNM). Plummers Lithacodes fasciola on Fagus grandifolia Island, collected 18.VIII.2006, J. Lill et al. (2? USNM (1 pupa)); same data, but 06-ak, ex Acharia stimulea on Asimina emerged 11.VI.2006, 05-921 (2/ USNM); triloba (/ USNM); same data, but same locality, collected 11.VII.2006, J. Lill emerged 25.VIII.2006, J. Lill et al. 06-am et al. 06-252, ex Lithacodes fasciola on (6/ USNM); same data, but emerged 28. Fagus grandifolia (? USNM); same data, VIII.2006, 06-ab, ex Acharia stimulea on emerged 31.VII.2006, 06-260 (/ USNM); Quercus alba (/ USNM; 6 pupae beneath same data, 06-268 (/ USNM); same data, host, 4 pupae on leaf); same locality, col- emerged 1.VIII.2006, 06-274, ex Limaco- lected 18.VIII.2006, J. Lill et al. 06-aq-V, didae on Carya glabra (/ USNM); ex Acharia stimulea on Quercus rubra collected 30.VIII.2007, emerged 1. (? USNM); same data, but emerged 3. IX.2007, J. Lill et al. 07-841, ex Proli- IX.2006, 06-ai, ex Acharia stimulea on macodes badia on Fagus grandifolia, Asimina triloba (? USNM; specimen (1/ USNM); same locality, collected 8. damaged); same data [no emergence date], VIII.2007, emerged 16. X .2007, J. Lill 06-ae-II, ex Acharia stimulea on Asimina et al. 07-611, ex Lithacodes fasciola on triloba (/?USNM); same data, 06-an- Quercus alba,(/?USNM); same II, ex. Acharia stimulea on Quercus data, no emergence, 07-608, same host, rubra (/ USNM); collected 3.VII.2007, (4/?USNM); same locality, collected emerged 17.VII.2007, J. Lill et al. 07- 14.IX. 2007, emerged 16.X.2007 J. Lill 221, ex Lithacodes fasciola on Carya et al. 07-957, ex Isa textula on Quercus glabra (/ USNM); same data, 07-222 alba (2/?USNM). (? USNM). Prince Georges County: North American Records: Patuxent NWR, collected 21.VII.2005, Platyplectrus americana (Girault) (Figs. emerged 9.VIII.2005, J. Lill et al. 05- 105–113): Euplectromorpha americana 501, ex Lithacodes fasciola on Fagus Girault, 1917: 114. Holotype ?,USNM grandifolia,(2/?USNM); same lo- (examined). cality, collected 17.VIII.2005, emerged 1. Based on the significant new rearing IX.2005, J. Lill et al. 05-730, ex Pack- data (Table 1), Platyplectrus (and most ardia geminata on Fagus grandifolia,(/ eulophid parasitoids) preferentially attack USNM); same locality, collected 17. early limacodid instars. Platyplectrus VIII.2005, emerged 27.VIII.2005, J. Lill americana is a solitary/gregarious ecto- et al. 05-742, ex Lithacodes fasciola on parasitoid with a single emerging Fagus grandifolia,(/ USNM); same lo- from a host 41% of the time, two emerging cality, collected 17.VIII.2005, emerged 26% of the time, three (12%), four (10%), 30.VIII.2005, J. Lill et al. 05-761, ex five (4%), six (4%), and greater than six Lithacodes fasciola on Fagus grandifolia, (4%). The position for larval feeding is as (2/ USNM (5 pupae)); same locality, previously described for Pl. natadae (Gadd collected 17.VIII.2005, emerged 10. et al. 1946). Pupation occurs beneath the VI.2006, J. Lill et al. 05-779, ex Proli- host cadaver (as in Fig. 98; cadaver re- macodes badia on Fagus grandifolia moved), and the pupa and cadaver are (/ USNM); same locality, collected 1. adhered to the substrate with the cadaver IX.2005, emerged 20.IX.2005, J. Lill providing protection for the parasitoid 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pupa. No in situ observations for ovipo- Literature.—Burks (1966) published a sition are yet available for Pl. americana. key to 23 species known from North Distribution: Eastern U.S.A. America, 24 were treated in a catalog Limacodidae: Previously reported from (Burks 1979), and Peck (1985) provided Packardia geminata (Gates and Burks akeyto32speciesfromNorthAmerica. 2003). Newly recorded hosts (Table 1) North American Records: include: Ac. stimulea*, Ad. spinuloides*, Pediobius crassicornis (Thomson) (Figs. E. delphinii*, I. textula*, Li. fasciola*, 95, 97–103): Pleurotropis crassicornis N. nasoni*, Ph. pithecium*, Pr. badia*, Thomson, 1878: 255. Holotype /, and Tortricidia sp.* MZLU (not examined). This Holarctic species has a broad host Pediobius Walker (Chalcidoidea: range, but surprisingly, it had not been Eulophidae: Entedoninae) reported as associated with limacodids (Figs. 95, 97–103) previously (Noyes 2003). It has, however, been reported previously from primary Diagnosis.—The following combina- parasitoids (Ichneumonidae: H. fugiti- tion of features is diagnostic: propodeum vus,Tachinidae:Com. concinnata)that medially with two subparallel carinae (Fig. are reported from limacodids herein. It 95), with distinct plicae; petiole ventrally is most commonly recovered as a pri- pointed; notauli incomplete (Fig. 101); mary (Lepidoptera) or secondary para- antenna with three funiculars and two sitoid of hymenopterans through lepi- claval segments (Figs. 102–103). dopterans. From the rearing results Fauna.—Pediobius is a large genus of reported herein, Pe. crassicornis com- 277 species worldwide with 36 known monly parasitize Pl. americana (Fig. 98). from the Nearctic (Noyes 2003). New When attacking Pl. americana, adult Pe. World species of Pediobius range from crassicornis emerge from the anterior end central Canada to Argentina. Extralimitally, of the pupa and escape the limacodid Cock (1987) summarized the Pediobius as- cadaver laterally or dorsally, depending sociated with limacodids in Southeast Asia: upon the degree of adherence of the li- Pe. imbreus (Walker) (as detrimentosus macodid cadaver to the substrate. In the (Gahan)), Pe. elasmi (Ashmead), and Pe. latter case, circular emergence holes are ptychomyiae (Ferrie`re). They are recorded chewed through the dorsum of the li- from a tachinid, Apanteles (Ap. sp. and Ap. macodid when the margins of the cadaver parasae;Braconidae),andPa. philepida are tightly appressed to the substrate. It is Holloway (Limacodidae), respectively. possible that Pe. crassicornis also parasit- Noyes (2003) summarized additional izes Al. lilli.Thereareafewspecimens host records including primary lepi- associated only with the remains of lima- dopteran hosts and secondary dipteran/ codid or other species of primary parasit- hymenopteran hosts. oid (ichneumonoids), but these records Biology.—Most commonly associated are suspect due to the nature of the with species of Lepidoptera, Coleoptera, emergence hole or the lack of pupal Diptera, and Hymenoptera as primary or remnants reminiscent of Pe. crassi- secondary parasitoids (Peck 1985). Some cornis. In the former instance, the re- species are known from spider egg sacs mains of Ac. stimulea are mummified where they may act as secondary parasitoids and a very large emergence hole is (Peck 1985). Limacodid host associations in chewed posterolaterally, roughly equal to North America are newly reported herein. 4–5 body widths of Pe. crassicornis, and VOLUME 114, NUMBER 1 79 no pupal remnants are visible inside. In additional taxa (see species treatment the latter (same limacodid host), micro- above). gastrine cocoons associated with Pe. crassicornis are empty and appear to have Psychophagus Thomson (Chalcidoidea: emerged normally, but no primary host Pteromalidae: Pteromalinae) remains are associated with them. There (Figs. 87–93) is a series of Co. schaffneri in the USNM Diagnosis.—Clypeus shallowly bi- (no label data) bearing a handwritten lobed apically (Fig. 88); pedicel ; 2.0X identification of, “On Tortricidia testacea as long as broad, antennal formula 11263 Dyar ‘97” that has several larvae with (Fig. 89); propodeum reticulate-rugose, cocoons of Cotesia adhered. A series of with median carina incomplete (Fig. 90); five (formerly seven as two are missing) gaster subcircular to short-ovate (Fig. 91). co-mounted Pe. crassicornis appear to Fauna.—Psychophagus contains the correspond with 10 cocoons of Cotesia, single Holarctic species, P. omnivorus. eight of which bear emergence holes lat- Biology.—This genus has an incred- erally and contain pupal remains. Ped- ibly broad host range and may act as a iobius crassicornis can be separated from primary or secondary parasitoid (Noyes other Pediobius by the following combi- 2003) across four orders and 22 nation of characters: occipital carina pres- families. Most hosts are pupae of Lep- ent, frons coarsely reticulate (Fig. 100), idoptera, Hymenoptera, or Diptera (or male with three funiculars (Fig. 103), their dipteran or hymenopteran parasit- scutellum smooth medially (Fig. 95), oids). There is one record of a single basal gastral tergum alutaceous. species of Limacodidae being attacked in Distribution: Widespread across North North America among numerous addi- America from the west to east coasts tional host records (see below). into Canada in the north. Also found in Literature.—Aside from original de- northern Europe and Japan (Peck scriptive work (Walker 1835) and treat- 1985, Noyes 2003). ment by Graham (1969), Noyes (2003) Limacodidae: Associated with the fol- provides citations of the bulk of the lit- lowing species but appears to be acting as a hyperparasitoid in most instances: erature (particularly host records) for this Ac. stimulea, Ad. spinuloides, N. nasoni, species. E. delphinii, I. textula,andLi. fasciola. North American Records: Natada nasoni may serve as a host in Psychophagus omnivorus (Walker) (Figs. one instance as the larva is mummified 87–93): Pteromalus omnivorus Walker, and an emergence hole is visible post- 1835: 204. Lectotype ?, BMNH, de- erolaterally, roughly equal to two body signated by Graham (1969: 473) (not widths of Pe. crassicornis,andpupal examined). remnants are visible inside. The sole limacodid host, the exotic M. Other hosts: The numerous hosts are flavescens, was reported by Schaffner and listed by Noyes (2003) and span 22 Griswold (1934) and Schaffner (1959). families across the following four insect The former recorded this wasp as often orders: Diptera (2), Coleoptera (2), hyperparasitic (only other parasitoid re- Lepidoptera (10), and Hymenoptera corded therein was Com. concinnata), (8). Herein it is newly recorded as a with four to 90 emerging per host and parasitoid of Pl. americana*, al- distributed in northeastern North America. though it is possible that it is attacking The latter reported four specimens of 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

P. omnivorus from Boston, Massachusetts Hoh 1992, Wei 1996) and Tr. papilionis as a gregarious parasitoid emerging from a (Nagarkatti) on pallivitta in Hawaii limacodid pupa. (Conant et al. 2006) have been reported. Distribution: Holarctic. In North America, There are also a few extralimital records of primarily eastern U.S.A. and Canada. Trichogrammatoidea thoseae Nagaraja Limacodidae: M. flavescens. attacking limacodids in three genera as Other hosts: Primary Parasitoid: Sco- summarized by Cock (1987). Most Tricho- lytidae (Coleoptera); , gramma species are primarily solitary or , (Hy- gregarious endoparasitoids of eggs of Lepi- menoptera); Arctiidae, Coleophoridae, doptera. The number successfully com- Lasiocampidae, , Lyman- pleting development in a single egg de- triidae, Noctuidae, Notodontidae, pends upon host size and the proximity of Nymphalidae, Oecophoridae, Pieridae, host eggs (Pinto 1998; references therein), , Saturniidae, and females inject a developmental ar- (Lepidoptera). Hyperparasitoid: Sar- restant during oviposition (Strand 1986). cophagidae, Tachinidae (Diptera); Literature.—Key to species (Pinto 1998). Braconidae, Ichneumonidae (Hyme- North American Records: noptera) (see Noyes 2003). Trichogramma minutum Riley (Figs. 94, 96): Trichogramma minuta Riley, Trichogramma Westwood 1871: 157–158. Neotype ?,USNM, (Chalcidoidea: Trichogrammatidae) designated by Pinto et al. (1978) (Figs. 94, 96) (examined). Pinto (1998) considers this species to be a Diagnosis.—The family is recognized “complex of populations whose taxonomic by the three-segmented tarsi (Fig. 94) and status remains under investigation.” Accord- fore wing setation arranged in lines (Figs. ing to Pinto (1998), Peck’s (1963) recogni- 96). Trichogramma are recognized by tion of only three Nearctic species of broad fore wing with sigmoid venation Trichogramma, and following Girault’s and distinctive Rs1 setal track (Fig. 96), (1911) interpretation of Tr. minutum, re- among other features not included herein sulted in recognizing ; 150 host species (Pinto 1998). in 33 families for this species alone. Fauna.—Trichogramma is the largest Given that this vast number of host re- genus in the family with ; 180 species cords and associated research on Tri- worldwide and 68 known from North chogramma species provided minimal America (Pinto 1998). They are used voucher specimen documentation, the extensively in biological control programs bulk of this information is unusable at of numerous species of pestiferous Lepi- the species level. Thus, the previously doptera in natural and agroecosystems published limacodid rearing record cited (S. Smith 1996). A comprehensive regional below is suspect. Diagnostic genitalic char- monograph is available (Pinto 1998). acters for species separation are reported Biology.—Species of Trichogrammati- in Pinto (1998). dae are solitary or gregarious parasitoids of Distribution: U.S.A., east of 110° insect eggs in several orders (, longitude (Pinto 1998). Coleoptera, Lepidoptera, Diptera, etc.). A Limacodidae: Monema flavescens single limacodid host is recorded (see be- (Collins 1933). low) for North America. Three exralimital Other hosts: Pinto (1998) recorded, records of Trichogramma spp. (Hoong and based on specimens examined and VOLUME 114, NUMBER 1 81

those reared from field-collected hosts Biology.—Most Cotesia species are (not literature records), a total of 49 gregarious endoparasitoids of macro- host records from three orders (Hy- lepidopterans from many families, with menoptera (1), Lepidoptera (45), and only a few microlepidopterans (e.g., plu- Neuroptera (3)). None was from Li- tellids) among their hosts. A minority of macodidae. species develop as solitary parasitoids. Like Trichogramma sp.: other microgastrines, species of Cotesia Distribution: Eastern U.S.A., minimally. typically oviposit into early-instar host lar- Limacodidae: E. delphinii* on Q. alba. vae, emerging from later instar larvae or prepupae to spin cocoon(s) on or near the Cotesia Cameron (Ichneumonoidea: host, which soon dies. A few species are Braconidae: Microgastrinae) capable of ovipositing into eggs (Ruberson (Figs. 64–69) and Whitfield 1996). The immune system of the host caterpillar is compromised in Diagnosis.—The combination of the part by the introduction, along with the following features is diagnostic (see eggs, of symbiotic by the Whitfield 1997): fore wing with vein r-m female wasp from her reproductive tract absent so that no small “areolet” (tiny sec- (Whitfield and Asgari 2003). ond submarginal cell) is present (Fig. 65); Literature.—Muesebeck (1920) is the propodeum with rugose sculpture and at only comprehensive North American treat- least some indication of a percurrent medial ment of Cotesia. carina (Fig. 66); anterior two metasomal North American Records: terga (and sometimes anterior portions of Cotesia empretiae (Viereck) (Figs. 64–67): third) sculptured and relatively broad and Apanteles empretiae Viereck, 1913: subrectangular (Fig. 66); ovipositor and 562. Holotype /, USNM (examined). sheaths relatively short, with only a few Distinctive among Cotesia species in small setae at tip of sheath (Fig. 67). having the body length < 2mm,first Fauna.—Cotesia is a huge genus metasomal tergite width subequal an- containing several hundred described teriorly and posteriorly (Fig. 66; most species worldwide, and many more un- Cotesia have tergite broader posteri- described, although it is best represented orly), the third metasomal tergite with in temperate regions. It is one of the sculpturing over anterior half or two largest and most economically important thirds, especially medially (Fig. 66), genera of Microgastrinae, with hosts in and the hypopygium sharply angled many lepidopteran (especially macro- apically and with ovipositor sheaths plepidopteran) families in both natural strongly exserted, often visible in dor- and agricultural systems. Most available sal view (Fig. 67). The gregarious lar- revisions of Cotesia are strictly regional, vae emerge to spin (separately) 5–15 and Muesebeck (1920) is the most recent whitish to pale tan woolly cocoons comprehensive North American treat- dorsally (sometimes laterally) on the ment. Extralimitally, there are two records caterpillars. of Apanteles spp. recorded from Ph. hip- Distribution: U.S.A., especially south- parchia Cramer and Ac. (as Episibine) eastern. intensa Dyar (Genty et al. 1978) in the Limacodidae: E. delphinii, Li. fasciola, Neotropical Region; these are most cer- Pa. chloris,andAc. stimulea (Marsh tainly Cotesia spp. based on the images of 1979), Ap. biguttata,andPa. indetermina adults and cocoons in Genty et al. (1978). (Whitfield et al. 1999). 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Cotesia phobetri (Rowher) (Fig. 68): To. pallida H.-S., To. testacea, Pr. Apanteles phobetri Rohwer, 1915: 228. badia, and E. delphinii (specimens in Holotype /, USNM (examined). the USNM with host remains). This species (Fig. 68) differs from both Co. empretiae and Co. schaffneri in hav- Ascogaster (Ichneumonoidea: ing the third metasomal tergite entirely Braconidae: Cheloninae) unsculptured; from Co. empretiae it also (Figs. 48, 60–62) differs in having the first metasomal ter- Diagnosis.—Carapace without pair gite broadening posteriorly, and from Co. of complete transverse grooves, at most schaffneri it differs in having the hind grooves present laterally but absent dor- tibiae and tarsi almost entirely pale orange sally (Fig. 48); fore wing vein (RS+M) brown. The gregarious pale buff cocoons a present; eyes glabrous or virtually so are spun separately and positioned dor- (Fig. 60); ocelli configured in shape of sally on the host caterpillar. isosceles triangle (Fig. 61); scutellar disc Distribution: Eastern U.S.A. (contribu- sculptured (Fig. 62). tion from possibly non-conspecific, non- Fauna.—Described species of As- limacodid rearings cannot be ruled out). cogaster occur in all zoogeographic re- Limacodidae: Ph. pithecium (Rowher gions, but only 12 species are known 1915). from the Nearctic (Shaw 1997a). Yu et al. Other hosts: Halysidota tesselaris Smith (2005) listed four species in Ascogaster and Abbot (Arctiidae) (Muesebeck placed currently in Leptodrepana Shaw. 1920) (possibly not conspecific Co. phobetri). Several other series reared As pointed out in Shaw (1997a), richness from Acronicta sp. (Noctuidae) and is higher in temperate areas than tropical other arctiids have also been identified areas of the New World, but richness is as this species, perhaps mistakenly. lower in the Nearctic than in all other Cotesia schaffneri (Muesebeck) (Fig. 69): regions except the Neotropical (Yu et al. Apanteles schaffneri Muesebeck, 1931: 4. 2005). Holotype /,USNM(notexamined). Biology.—As is the case for all chelo- This species is unique among the three nines, species of Ascogaster are solitary Cotesia recorded herein in having the hind koinobiont egg-larval endoparasitoids of tarsi strongly bicolored: the proximal and lepidopterans (Shaw 1997a). Host asso- distal tarsomeres being mostly blackish, ciations reported for Ascogaster include whereas the third and fourth (and sometimes 23 families of Lepidoptera, including also second) tarsomeres are pale orange Limacodidae; most hosts are in Tortricidae yellow. Unlike in Co. empretiae,thegre- (Yu et al. 2005). Host associations reported garious pale buff cocoons are spun sepa- for species of Ascogaster in the Nearctic rately (but appressed together; individually Region include species in Anthomyiidae, immobile) on the back of the host cater- Clusiidae (Diptera), Curculionidae (Co- pillar. The parasitized cocoons with the leoptera), Cynipidae, Tenthredinidae emergence holes on the side are whitish. (Hymenoptera), Blastodacnidae, Depres- Distribution: Nearctic ranging from sariidae, Gelechiidae, Geometridae, Lima- mid-Atlantic states and Texas in U.S. codidae, Pyralidae, Tineidae, Tortricidae, A.; thus it may be distributed broadly and Yponomeutidae (Lepidoptera) (Yu across the eastern and southern U.S.A. et al. 2005). Non-lepidopteran host re- Limacodidae: Unidentified limacodid cords are erroneous since all confirmed (“cochlidiid”) (Muesebeck 1931), hosts for chelonines are lepidopterans VOLUME 114, NUMBER 1 83

(Shaw 1997a, S. Shaw pers. comm.). Of that cocoons were “placed in rearing the remaining 138 putative lepidopteran boxes,” so the record should be con- hosts, only one species of Limacodidae, firmed through rearing from a host M. flavescens, has been reported (Wilcox isolated individually. This host species 1918). is likely no longer present in North Literature.—Research relevant to this America based on subsequent searches study published since Yu et al. (2005) in- for it (M. Epstein pers. comm.). cludes As. quadridentata (Wesmael) reared Other hosts: Reported from at least from the tortricids Lobesia botrana 48 species in six lepidopteran fami- (Denis and Schiffermu¨ller) (Carlos et al. lies (Yu et al. 2005). 2006) and Spilonota ocellana (Denis and Schiffermu¨ller) (Pluciennik and Olszak Triraphis Ruthe (Ichneumonoidea: 2010). The aforementioned hosts were Braconidae: ) reported previously in multiple publica- (Figs. 46–47, 49–59) tions (Yu et al. 2005). Nazemi et al. (2008) Diagnosis.—Third maxillary palpo- reported As. quadridentata from the tor- mere not enlarged and flattened, similar tricid amygdalana (Duponchel), in size and shape to distal palpomeres; an of the cynipid Andricus second labial palpomere not enlarged cecconi Kieffer. and vesiculate, similar in size and shape North American Records: to distal palpomeres; basal carina of pro- Ascogaster quadridentata (Wesmael) podeum diverging basally to apically (Figs. (Figs. 48, 60–62): Ascogaster quad- ridentatus Wesmael 1835: 237. Lectotype 53, 55, 58), not complete to apical margin /,IRSNB,designatedbyShaw(1983) as single mesal carina; inner margin of (not examined). metatibia with fringe of flattened setae (Fig. This species can be differentiated from 59); metatibial spurs straight and setiferous; all Nearctic species of Ascogaster,except pro- and mesotarsus with tarsomeres 2–4 As. shawi Marsh, using the key in Shaw longer than wide (Figs. 50–51, 56); tarsal (1983). It can be differentiated from As. claws with basal lobe or tooth (Fig. 47); shawi using the features that differentiate fore wing first submarginal cell uniformly As. quadridentata and As. provancheri setose, without glabrous patch; fore wing Dalla Torre in the key by Shaw (1983). vein 1cu-a not angled as obliquely as fore Distribution: All zoogeographic re- wing 2CUa (Fig. 56); hind wing M+CU gions except the Afrotropical, although equal to or longer than 1M (as in Fig. it has been introduced into South Africa 45); metasoma with T1–T4 not car- and several other countries to control apacelike and covering other terga (Figs. orchard pests, particularly Cydia po- 50–51, 56); T2 without triangular area monella (Linnaeus). It ranges in the anteromesally; transverse groove between Nearctic Region from Maine south to T2+T3 distinct (Figs. 52, 57). Georgia and west to California (Yu Fauna.—Described species of Triraphis et al. 2005). are known from the Nearctic, Neotropical, Limacodidae: Monema flavescens re- Oriental, and Palearctic regions (Yu et al. ported but host plant not mentioned 2005). There are three Nearctic species (Wilcox 1918). RRK considers this currently placed in this genus, including record possibly valid but has not verified T. eupoeyiae (Ashmead), new combi- this identification. In terms of rearing nation (see below), but most if not protocol, Wilcox (1918) mentioned only all New World species currently in Rogas 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON belong in Triraphis (Valerio 2006, Zaldı´var- Triraphis,andconductedacladistic Rivero´netal.2008).NeitherTriraphis nor analysis based on characters of the Rogas has been revised for the New venom gland. Quick and Shaw (2005) World; New World richness is estimated discussed mummy characteristics for at 100–150 species (Shaw 1997b). Triraphis; T. tricolor was the only species Biology.—As is the case for all roga- mentioned. Valerio (2006) transferred T. dines, species of Triraphis are koinobiont brasiliensis (Sze´pligeti) and T. mac- endoparasitoids of lepidopteran larvae and ulipennis (Sze´pligeti) from Rogas, but pupate within the mummified remains of Papp (2004) had already transferred those their hosts (Shaw 1997b). Reported host species to Triraphis.Valerio(2006)also associations are larvae in , indicated T. pulchricornis (Sze´pligeti) as Limacodidae, Lycaenidae, Lymantriidae, anewcombinationtransferredfromRogas, Megalopygidae, Riodinidae, Tortricidae, but Papp (2004) had transferred pul- and Zygaenidae (all Lepidoptera) (Yu chricornis from Rogas to et al. 2005, Zalvidar-Riveron et al. 2008). Wesmael. Additionally, Valerio (2006) A single record from Cynipidae (Hyme- mentioned eight species of Rogas as be- noptera) is obviously erroneous since all longing in Aleiodes, but all had been confirmed hosts for rogadines are lepi- transferred to Aleiodes previously (Shaw dopterans (Shaw 1997b, S. Shaw pers. et al. 1997, Marsh and Shaw 1998, Shaw comm.). Other records of questionable et al. 1998, Fortier and Shaw 1999). validity are the Palearctic species T. Zalvidar-Riveron et al. (2008) carried tricolor (Wesmael) from the tortricids out cladistic analyses for Rogadinae, rosana L. (Tobias 1976) and Tor- including Triraphis,basedonDNAse- trix testudinana Hu¨bner (Reinhard 1865) quence data and assessed the evolution and the Oriental species T. sichuanensis of host range and mummy characteris- Chen and He from the lymantriid Eu- tics. Kula et al. (2010) reported the first proctis bipunctapex (Hampson) (Chen host records for T. discoideus and He 1997). Genera in Dalceridae, North American Records: Lycaenidae, Megalopygidae, and Riodi- Triraphis discoideus (Cresson) (Fig. 46– nidae were reported in Zaldı´var-Rivero´n 47, 49, 56–59): Aleiodes discoideus Cres- et al. (2008) as containing hosts for spe- son, 1869: 380. Holotype /,ANSP(not cies of Triraphis;particularparasitoidand examined). host species were not listed. The basis for The body is irregularly yellow and brown those records (e.g., rearing protocol) is and the stigma is usually (96.6%) brown unknown (i.e., cited as Valerio and Shaw with at most a small yellow area where it in prep.), although they were noted as meets vein R1a (Fig. 56). confirmed by a rogadine specialist (i.e., Distribution: Nearctic Region, rang- S. R. Shaw, University of Wyoming- ing from Massachusetts to Florida and Laramie). Thus, in terms of published west to Illinois (Yu et al. 2005). records, species of Triraphis are known Limacodidae: Reported from 10 spe- from hosts in two monophyletic groups cies of Limacodidae sampled collec- (i.e., Limacodidae, Zygaenidae) (Wahlberg tively from six hardwood tree species et al. 2005, Niehuis et al. 2006). in the District of Columbia and Literature.—Aside from literature Maryland (Table 1) (Kula et al. 2010). reported in Yu et al. (2005), Zaldı´var- See Kula et al. (2010) for additional Rivero´netal.(2004)examinedvenom information on the natural history of T. gland morphology for Rogadinae, including discoideus. VOLUME 114, NUMBER 1 85

Other hosts: None reported. The body is entirely yellow to orange, and Triraphis eupoeyiae (Ashmead), new the fore wing stigma is yellow with at most combination (Fig. 50): Pelecystoma alightbrownspotdistadveinr(Fig.51). eupoeyiae Ashmead, 1897: 113. Holo- Distribution: Nearctic and Neotropi- type /, USNM (examined). cal regions, ranging from New Jersey The body is primarily orange but with at to Florida west to Missouri and south least the lateral lobes of the mesoscutum to Costa Rica. It also occurs in Cuba brown, and the stigma is yellow with a (Yu et al. 2005). brown spot directly above vein r (Fig. 50). Limacodidae: Harrison (1963) reported This species fits Triraphis sensu van T. harrisinae from Ac. apicalis (Dyar) Achterberg (1991) except the occipital on banana; host larvae were isolated in- carina is complete dorsally, and the frons dividually, and RRK considers this re- is rugose in the holotype. Additionally, cord valid. Triraphis harrisinae was not several relative features in the diagnosis reared from a congener, Ac. stimulea,du- for Triraphis sensu van Achterberg (1991) ring the course of this research. Marsh are difficult to interpret (e.g., size and (1979) reported T. harrisinae from Li. shape of basal lobe of tarsal claw). It is fasciola, but there are no specimens of transferred to Triraphis despite the ex- T. harrisinae at the USNM purportedly ceptions and problems with interpretation reared from Li. fascicola. This record because it shares more features with is possibly valid, but T. harrisinae was species in that genus than species in Ro- not reared from Li. fasciola during the gas sensu van Achterberg (1991). course of this research. Distribution: Known only from Flor- Other hosts: Riley and Howard (1890) ida prior to this research, but a speci- first reported it as a parasitoid of Harri- men from Cuba is in the USNM. sina americana (Gue´rin-Me´neville) Limacodidae: Ashmead (1897) reported (Zygaenidae). Additionally, specimens the holotype as reared from Ala. slos- of T. harrisinae reared from H. amer- soniae,butahostcadaverisnotasso- icana and undetermined Acoloithus ciated with the holotype. RRK considers Clemens (Zygaenidae) larvae collected this record possibly valid, but it requires in Florida and Illinois were used to start confirmation through rearing from acolonyattheUniversityofCalifornia- ahostisolatedindividually.Thecadaver Riverside (Smith et al. 1955). Speci- associated with this rearing is deposited mens from that colony were released in the USNM. In reporting on the biology in California to control H. metallica of Ala. slossoniae,Dyar(1897)figured Stretch and apparently established in alarvaparasitizedbyT. eupoeyiae,in- San Diego (Clausen 1956). See Kula dicating that it becomes, “bright red et al. (2010) and references in Yu et al. (Plate V, fig. 23) and hardens.” (p. 125). Other hosts: Label data associated with (2005) for additional information on the specimen from Cuba indicates that the natural history of T. harrisinae. it was reared from Leptotes cassius Mesochorus Gravenhorst (Cramer) (Lycaenidae); host remains (Ichneumonoidea: Ichneumonidae: are associated with the specimen. ) Triraphis harrisinae (Ashmead) (Fig. (Figs. 19–22) 51–55): Rhogas harrisinae Ashmead (1888) 1889: 632. Holotype ?,USNM Diagnosis.—Mesochorus can be rec- (examined). ognized by the following combination of 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON characters: areolet of fore wing closed, through an undetermined species of large and rhombic (Fig. 20); spiracle of (Lepidoptera: Tortricidae), metasomal segment 1 near or just behind and the other through an undetermined middle (Fig. 21); glymma large and deep species of Noctuidae (Lepidoptera) pre- (Fig. 21); ovipositor needlelike, about 1.5X sumably parasitized by an undetermined as long as metasomal apical depth and species of Aleiodes Wesmael (Braconidae: without dorsal subapical notch (Fig. 22); Rogadinae). female hypopygium prominent and trian- North American Records: gular in lateral view (Fig. 22); male gon- Mesochorus discitergus (Say) (Figs. 19–22): oforceps produced into elongate processes. Cryptus discitergus Say, 1835: 231. Type / Fauna.—Species of Mesochorus occur (destroyed, Townes et al. 1965). in all zoogeographic regions (Yu et al. There are numerous extremely similar 2005), with about 695 species worldwide, species of Mesochorus in the Nearctic, 108 of which occur in the Nearctic Re- and discitergus cannot be separated from gion. Dasch (1971) revised the Nearctic them by a short diagnosis. Dasch (1971) Mesochorinae. divided the 96 species that he treated into Biology.—Most observations and rear- 17 species groups with this species ing records indicated that mesochorines placed in the discitergus group with two are obligate hyperparasitoids of endo- other species: M. americanus Cresson and parasitic Ichneumonoidea (and, rarely, M. acuminatus Thompson. Dasch pro- Tachinidae) which parasitize primary vided information that may be used to hosts of larval Lepidoptera, Symphyta, separate discitergus from other species of and Coleoptera, and nymphal and adult Mesochorus, but the characters are diffi- Hemiptera and Psocoptera (Carlson 1979a). cult to interpret by non-experts without Based upon rearing records, many spe- access to a large collection. It remains cies have a wide host range (Dasch 1971, necessary to confirm species identifications Carlson 1979a). Ectoparasitic ichneumo- by sending them to an expert with access to nids can be utilized as well, based upon awell-curatedichneumonidcollection rearings of Mesochorus from Phytodietus (e.g., USNM, AEI, CNC). and ?Oedemopsis hosts by M. R. Shaw Distribution: Worldwide except for (Gauld and Bolton 1988: 17). Although the Australian region (Yu et al. 2005). there are some records of mesochorines as Limacodidae: The two specimens of Me. primary parasitoids (Dasch 1971, Yu et al. discitergus in this study emerged (post- 2005), these are likely specimens reared erolaterally) from mummified early to from undetected primary parasitoids (Wahl middle instar larvae of Pa. chloris on Q. 1993a). The pitfalls and deficiencies of rubra and Pr. badia on F. grandifolia; rearing records were ably discussed by the mummies were probably made by a Shaw (1990: 453–455) and are relevant species of Triraphis based on examina- to this matter. tion of montage images of host remains Literature.—The Nearctic Mesochorus (JTL). This species has not been re- fauna may be identified using Dasch ported as a hyperparasitoid through a li- (1971). Aside from literature reported macodid, nor was it reared from any in Yu et al. (2005), Reis Fernandes et al. other parasitoids in this project. (2010) reported two undetermined spe- Other hosts: Mesochorus discitergus cies of Mesochorus: one from an un- has been reported as a hyperparasitoid determined species of Hypomicrogaster of 31 species of Braconidae and three Ashmead (Braconidae: Microgastrinae) species of Ichneumonidae (Yu et al. VOLUME 114, NUMBER 1 87

2005). Records of Me. discitergus as endoparasitoids of Lepidoptera, with the a primary parasitoid of lepidopterans egg deposited in an exposed host larva and are questionable, as discussed above. the mature wasp larva emerging before the host pupates (Jerman and Gauld 1988). Casinaria Holmgren (Ichneumonoidea: Species in the following families have Ichneumonidae: ) been recorded as hosts: Arctiidae, Cram- (Fig. 28) bidae, Geometridae, Hesperiidae, Lasio- Diagnosis.—Casinaria may be recog- campidae, Limacodidae, Megalopygidae, nized by the following combination of Notodontidae, Nymphalidae, Oecophoridae, characters: clypeus not separated from Olethreutidae, and Zygaenidae (Walley supraclypeal area by distinct groove; 1947, Finlayson 1975, Short 1978, Carlson ventral posterior corner of propleuron with 1979a, Jerman and Gauld 1988, Janzen strongly produced, more or less angulate and Hallwachs 2009). lobe touching or overlapping pronotum (as Literature.—The Nearctic Casinaria fauna in Fig. 24); areolet of fore wing closed, may be identified using Walley (1947). obliquely quadrate and petiolate (as in Fig. Additionally, a species of Casinaria was 26); metasomal segment 1 with the fol- reported from Ac.(asSibine) megasomoides lowing: spiracle beyond middle, petiole Walker on oil palm in Costa Rica (Mexzo´n long and cylindrical in cross-section and et al. 1996). with tergo-sternal suture at midline, T1 North American Records: without trace of glymma (Fig. 28); meta- Casinaria grandis (Walley) (Fig. 28): soma laterally compressed; ovipositor not Walley, 1947: 376. Replacement name for needlelike, about as long as metasomal Casinaria texana Ashmead (1890); Ho- apical depth and with dorsal subapical lotype /, USNM (not examined). notch (Fig. 28). Casinaria grandis may be distinguished Fauna.—Casinaria is distributed world- from other Nearctic Casinaria by the fol- wide, with 99 species, 20 of which are lowing characters: large, broad, flat, and recorded from the Nearctic Region (Yu centrally concave scutellum; petiolate et al. 2005). Walley (1947) revised the areolet of the fore wing; gena descending Nearctic fauna. In a forthcoming revi- perpendicularly behind ocelli; coarsely sion of Costa Rican Campopleginae, rugosopunctate propodeum without ca- I. D. Gauld recognizes a species-group rinae; and large size (about 15 mm long). within Casinaria consisting of grandis Distribution: Gulf of Mexico (Florida, and two undescribed Costa Rican spe- Texas, Louisiana; Carlson 1979a). It pre- cies. The latter parasitize various species sumably extends into northern Mexico. of limacodids in the genera Acharia, Limacodidae: Reported from Ac. stim- Euclea, Euprosterna, and Natada based ulea (Carlson 1979a); no specimens on extensive rearing efforts conducted in could be located in the USNM collec- Costa Rica (D. Janzen and W. Hallwachs tion that would verify this record. pers. comm; see also http://janzen.sas. Other hosts: None known. upenn.edu). Extralimitally, there is one re- Hyposoter Forster (Ichneumonoidea: cord of Casinaria sp. from Ac. (as Sibine) ¨ Ichneumonidae: Campopleginae) megasomoides (Walker) (Genty et al. (Figs. 23–24, 26–27, 30, 32) 1978) in the Neotropical Region. Biology.—Like other campople- Diagnosis.—Hyposoter may be rec- gines, Casinaria species are koinobiont ognized by the following combination of 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON characters: clypeus not separated from cuticle; this is not, however, mummifi- supraclypeal area by distinct groove (Fig. cation as found in rogadine braconids 32); ventral posterior corner of propleuron (see the discussion of rogadine mummi- with strongly produced, more or less an- fication in Shaw (2006)) in which the host gulate lobe touching or overlapping pro- mummy remains intact due to internal notum (Fig. 24); areolet of fore wing pupation by the parasitoid. closed, obliquely quadrate and petiolate Literature.—Aside from literature re- (Fig. 26); metasomal segment 1 with the ported in Yu et al. (2005), Marconato following: spiracle beyond middle, petiole et al. (2008) reported an undetermined short and quadrate in cross-section and Hyposoter from two geometrids (Lepi- with tergo-sternal suture close to ventral doptera). Reis Fernandes et al. (2010) margin, T1 with glymma present as pitlike reported two undetermined species of impression (Fig. 27); metasoma laterally Hyposoter from two geometrids (Lepi- compressed; ovipositor not needlelike, doptera). There are no reliable or com- about as long as metasomal apical depth prehensive treatments of the taxonomy and with dorsal subapical notch (Fig. 23). of the Nearctic Hyposoter fauna. Fauna.—Species of Hyposoter occur in North American Records: all zoogeographic regions. Yu et al. (2005) Hyposoter fugitivus (Say) (Figs. 23–24, listed 117 valid species worldwide and 29 26–27, 30, 32): Banchus fugitivus Say, for the Nearctic Region. Horstmann (2008) 1835: 247. Lectotype /, USNM, desig- described one new species from the Pale- nated by Cushman (1925) (not found in arctic Region, and Horstmann (2009) USNM collection). transferred Hyposoter koentzeii (Kiss), Besides the 29 described Nearctic Hy- known from , from Omorgus posoter species there are numerous un- Fo¨rster. Hyposoter bellus (Cresson), described species; in the absence of a known from Cuba, was transferred from formal revision, meaningful diagnosis of Xanthocampoplex Morley in Gauld and fugitivus may be misleading. The char- Ferna´ndez-Triana (2010). Viereck acters presented in the key, in conjunc- (1925, 1926) provided keys that include tion with an association with a rearing most species known from the Nearctic from a limacodid host in the Nearctic Region, but a single key to species of Region, provide the best information at Hyposoter in the Nearctic Region has this point to attempt to assign a species not been published. Further, the keys in name. As with other ichneumonids (e.g., Viereck (1925, 1926) use characters Mesochorus)reportedherein,finalidenti- that are of doubtful utility for species fication should be made by an ichneumo- diagnostics (see specific information nid expert with access to a large col- below). lection such as USNM, AEI, or CNC. Biology.—Hyposoter spp. are koino- Distribution: Transcontinental in the biont endoparasitoids of ditrysian lepi- Nearctic (except possibly absent in the dopterans, including some species of southwestern U.S.A.) (Carlson 1979a). Rhopalocera. Egg deposition is in early to Recorded from Brazil (Costa Lima middle instar larvae, and the wasp larva 1962), but not recognized in the Neo- usually emerges from the penultimate tropical catalog of Townes and Townes instar of the host larva. The wasp spins an (1966) nor is it present in Costa Rica (I. ovoid cocoon near or under the host cat- Gauld et al. unpubl.). erpillar’s remains. The cocoon is some- Limacodidae: Isa textua on Acer ne- times completely covered by the host’s gundo L. and Salix nigra (Marsh) VOLUME 114, NUMBER 1 89

(Barbosa and Caldas 2004). It was Townes and Townes (1962) provide a key reared in this study from Pr. badia* to the Nearctic fauna. on Q. alba. The Hyposoter cocoon is Biology.—Baryceros is placed in the completely covered by the skin of the subtribe Baryceratina, with all known host caterpillar. This is the only de- host records indicating that the genera of scribed species of Hyposoter known to this subtribe are idiobiont ectoparasitoids attack limacodids; several undescribed of Limacodidae (Townes 1970). Three Costa Rican species have been reared species of Baryceros have been reared. from species of Acharia, Euclea, Na- Baryceros eucleidis (Blanchard) has been tada, and Parasa based on extensive reared from Ph. hipparchia (Cramer) and caterpillar rearing effort in Costa Rica Ac. (as Sibine) nesea (Stoll), and B. sibine (D. Janzen and W. Hallwachs pers. (Cameron) has been reared from Ac. (as comm.). S.) trimaculata (Sepp) and Ac. (as S.) ne- Other hosts: It has been reported from sea (Townes and Townes 1966). Bar- 30 other lepidopteran species in 12 yceros texanus is discussed below. A suite families (Yu et al. 2005); a new record of morphological characters appears to be from this study was its rearing from adaptations to attaching limacodid co- Meg. crispata on Q. alba. coons. One such character is the posses- sion of prominent dorsal and ventral teeth Baryceros Gravenhorst on the ovipositor apex (Fig. 29); charac- (Ichneumonoidea: Ichneumonidae: terized by Gauld (1984) as “corkscrew- Cryptinae) like,” this specialized structure is used to (Figs. 29, 37–38) penetrate the hard limacodid cocoon. Literature.—Townes and Townes Diagnosis.—Baryceros may be recog- (1962) remains the definitive taxonomic nized by the following combination of treatment of the Nearctic fauna. characters: clypeus separated from su- North American Records: praclypeal area by distinct groove; ventral Baryceros texanus (Ashmead) Figs. 29, posterior corner of propleuron not devel- 39–38: Crypturus texanus Ashmead, oped as distinct lobe, at most with weak 1890: 413. Holotype ?, USNM (not groove delimiting it from main area of examined). propleuron (Fig. 37); dorsal margin of Baryceros texanus can be differentiated pronotum with strong swelling at dorsal from other Nearctic Baryceros by the end of epomia (Fig. 37); areolet of fore black and white tergites and the single wing open (vein 3rs-m absent) and pen- large white spot on the ventral portion of tagonal (Fig. 29); mesosoma with coarse the mesopleuron (Fig. 29). punctures (Fig. 37); spiracle of metasomal Distribution: North America, in U.S.A. segment 1 beyond middle; glymma absent from Maryland to Florida, west to (Fig. 29); metasoma cylindrical/dorsoven- Michigan and eastern Texas. Recorded trally compressed; ovipositor not needlelike, from Sonora, Mexico and is presumably about 2.0X as long as metasomal apical more widely distributed (Townes and depth and without dorsal subapical notch Townes 1962 Carlson 1979a). (Fig. 38); body (excluding legs) black Limacodidae: Ala. slossoniae from with white markings (Fig. 29). southern Florida (Ashmead 1897, Dyar Fauna.—Baryceros is a New World 1897). The USNM collection has a reared genus, with 32 described species; five are specimen from Stock Island (Monroe found in the Nearctic (Yu et al. 2005). Co.), Florida; label data state that the 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

host was a “Lep. larva” on a leaf of cocoons (Townes 1970), including Co. Conocarpus erecta L. (Combretaceae). glomerata L. (Harvey et al. 2004); Yu Other hosts: None known. et al. (2005) list 33 microgastrine species as hosts. Yu et al. (2005) also list 26 Lysibia Fo¨rster (Ichneumonoidea: species of Lepidoptera in 14 families as Ichneumonidae: Cryptinae) hosts. While some of these host records (Figs. 25, 35, 43) may be correct (see discussion under Diagnosis.—Lysibia may be recog- Isdromas), many likely are the result of nized by the following combination of faulty record keeping (Shaw 1990). A characters: clypeus separated from su- series of recent papers have been pub- praclypeal area by distinct groove, apical lished on the biology and ecology of 0.3 turned inward at 90° angle (Fig. 43); Ly. nana (Gravenhorst) (Harvey et al. occipital carina meeting hypostomal ca- 2004; 2006; 2009a, b) attacking Co. rina above mandibular base (as in Fig. 41); glomerata. ventral posterior corner of propleuron not Literature.—Townes (1983) remains the developed as distinct lobe, at most with definitive taxonomic treatment of the weak groove delimiting it from main area Nearctic fauna. of propleuron (Fig. 25); dorsal margin of North American Records: pronotum without strong swelling at dor- Lysibia mandibularis (Provancher) (Figs. sal end of epomia (Fig. 35); areolet of fore 25, 35, 43): Hemiteles madibularis wing open (vein 3rs-m absent) and pen- Provancher, 1875: 315. Holotype /, tagonal; vein 2-Cu of hind wing basally ULQC (not examined). incomplete (Fig. 35); mesosoma with Lysibia mandibularis has the punctures of punctures ranging from fine to absent; tergites 2–3 centrally sparse, separated by spiracle of metasomal segment 1 beyond 1.5–2.5X the length of the setae; Ly. middle; glymma absent; metasoma cy- tenax has the tergal punctures evenly lindrical/dorsoventrally compressed; ovi- distributed and separated by about 0.8X positor not needlelike, about 1.4X as long the length of the setae. as metasomal apical depth and without Distribution: Transcontinental in the dorsal subapical notch. Body color (ex- Nearctic (Townes 1983). cluding legs) black to dark brown, with Limacodidae: The specimens in this eastern Nearctic specimens often having study were reared from cocoons of an T2–T7 brown and with brownish red undetermined gregarious microgastrine areas on T2–T3 (Fig. 35). parasitizing Ac. stimulea on Lindera Fauna.—Lysibia is found worldwide benzoin (L.) Blume. except for Africa and Australia. There Other hosts: Yu et al. (2005) list seven are nine described species, with two species of microgastrines as hosts; four (mandibularis (Provancher) and tenax species of Lepidoptera are also listed, Townes) present in the Nearctic. They but these records are probably errone- may be identified using the key to world ous as noted above. species in Townes (1983). Biology.—Like the majority of spe- Acrolyta Fo¨rster (Ichneumonoidea: cies in the , Lysibia spp. Ichneumonidae: Cryptinae) are idiobiont ectoparasitoids of cocooned (Figs. 36, 41) hosts. They are apparently pseudohyperp- Diagnosis.—Acrolyta may be recog- arasitoids, specializing in microgastrine nized by the following combination of VOLUME 114, NUMBER 1 91 characters: clypeus separated from su- Lepidoptera may be hosts (see discussion praclypeal area by distinct groove, apical under Isdromas), it is likely that these re- 0.3 flat and not turned inward (as in Fig. cords are due to faulty observations and 44); occipital carina meeting hypostomal lax record keeping of the sort discussed by carina at mandibular base (Fig. 41); ven- Shaw (1990). Harvey et al. (2009b) re- tral posterior corner of propleuron not ported on the biology of Acrolyta nens developed as distinct lobe, at most with Hartig attacking Co. glomerata. weak groove delimiting it from main area Literature.—There are no reliable or of propleuron (as in Fig. 25); dorsal mar- comprehensive treatments of the - gin of pronotum without strong swelling omy of the Nearctic Acrolyta fauna. at dorsal end of epomia (Fig. 36); areolet North American Records: of fore wing open (vein 3rs-m absent) and Acrolyta nigricapitata (Cook and Davis) pentagonal; vein 2-Cu of hind wing com- (Figs. 36, 41): Ischnoceros nigricapitatus plete; mesosoma with punctures ranging Cook and Davis, 1891: 11. Holotype /, from fine to absent; spiracle of metasomal MSUC (not examined). segment 1 beyond middle; glymma ab- Acrolyta nigricapitata can be distin- sent; metasoma cylindrical/dorsoventrally guished from other Nearctic Acrolyta by compressed; ovipositor not needlelike, the following characters: tergite 3 with 1.6–1.8X as long as metasomal apical fine longitudinal striae covering most of depth and without dorsal subapical notch. surface (tergite 3 without striae); fore Body color of females (excluding legs) and middle coxae light brownish red ranges from uniformly black/dark brown (fore and middle coxae white); tergite 3 to having extensive brownish red areas on completely brownish red (tergite 3 dark T2–T4; males have metasomal coloration brown/fuscous, ranging from completely ranging from having brownish red re- so to having anterior and posterior mar- stricted to the apices of T2–T3, to having gins narrowly brownish red). brownish red on the apex of T2, all of T3, Distribution: Northeastern U.S.A. and and the basal area of T4 (Fig. 36). southeastern Canada to the Pacific Fauna.—Acrolyta is found in the Afro- Coast (California and Washington) tropical, Palaearctic, Nearctic, and Orien- (Gauld 1984, Yu et al. 2005). tal regions. There are 25 described species, Limacodidae: Acrolyta nigricapitata with the following four species in the is recorded as parasitizing Ac. stimulea Nearctic: alticola (Ashmead), mesochori in Ashmead (1896: 209), where the Ashmead, nigricapitata (Cook and Davis), only information is as follows: “De- and washingtonensis (Cushman) (Yu et al. scribed from two specimens labeled 2005). No keys are available to separate No. 295o, reared May 14, 1883, from the species. Empretiae stimulea.” It is unlikely to Biology.—Acrolyta species are inferred be a primary parasitoid. idiobiont ectoparasitoids of cocooned hosts, Other hosts: Yu et al. (2005) list three as are other species in the Phygadeuontini braconid species as hosts; a noctuid, (Gauld 1984). Townes (1970) stated that Pseudaletia unipunctata (Haworth) (= “[p]robably the usual hosts are cocoons unipunctata), is also listed, but of Braconidae and Ichneumonidae.” Yu this record also stems from Ashmead et al. (2005) list 28 species of braconids (1896: 210) with only the following as hosts; 13 lepidopteran species in 10 entry: “Described from one specimen families are also listed (as well as a spe- labeled “parasite on Leucania uni- cies of Cynipidae!). While some of the punctata,June,1880.”TheUSNM 92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

collection has three nigricapitata spec- species, Is. lycaenae (Howard) appears to imens with host records/remains (R. be the sole Nearctic representative. Kula pers. comm.): (1) a specimen Biology.—As with most Cryptinae, without host remains but label data in- Acrolyta, Isdromas, and Lysibia are idi- dicating Ap. melanoscelus (Ratzeburg) obiont ectoparasitoids of cocooned hosts as the host, (2) a specimen with a (Gauld 1984). Gauld (1988) points out mounted cocoon and label data iden- that many species of Phygadeuontini tifying it as Ap. melanoscelus, and (3) “are facultative hyperparasitoids and use a specimen with an associated cocoon as hosts whatever is in a particular cocoon (probably braconid) and no associated be it the cocoon maker or one of its par- information. asitoids.” This is particularly true for the subtribe Acrolytina, to which these gen- Isdromas Foerster (Ichneumonoidea: era belong, due to the habit of parasitizing Ichneumonidae: Cryptinae) ichneumonid and braconid cocoons (Figs. 31, 33–34, 42, 44) (Gauld 1995). This then leads to consid- eration of two different modes of hyper- Diagnosis.—Isdromas may be recog- (Shaw and Askew 1976): (1) nized by the following combination of true hyperparasitism in which the hyper- characters: clypeus separated from supra- parasitoid develops on a primary parasit- clypeal area by distinct groove, apical 0.3 oid living on or within the primary’s host flat and not turned inward (Fig. 44); oc- and (2) pseudohyperparasitism, which cipital carina meeting hypostomal carina involves the pseudohyperparasitoid at- above mandibular base (Fig. 42); ventral tacking the primary parasitoid after it posterior corner of propleuron not devel- has destroyed its host. oped as distinct lobe, at most with weak This background puts a necessary groove delimiting it from main area of perspective on the rearing records of I. propleuron (as in Fig. 25); dorsal margin lycaenae. The species has been reported of pronotum without strong swelling at as reared from five species of Braconi- dorsal end of epomia (Fig. 34); areolet of dae and two species of Ichneumonidae fore wing open (vein 3rs-m absent) and on various lepidopteran hosts (Yu et al. pentagonal (Fig. 34); vein 2-Cu of hind 2005); this would thus be pseudohy- wing complete (Fig. 34); mesosoma with perparasitism. Isdromas lycaenae has also punctures ranging from fine to absent; been reported from nine species of Lepi- spiracle of metasomal segment 1 be- doptera in seven families (Yu et al. 2005); yond middle; glymma absent (Figs. 34); these could either be instances of primary metasoma cylindrical/dorsoventrally com- parasitism or utilization of overlooked pressed; ovipositor not needlelike, about primary parasitoids. De Santis (1987) and 3.0X as long as metasomal apical depth Lourenca˜oetal.(1989)reportedIs. mon- and without dorsal subapical notch. Body terai (Costa Lima), the only other species color (excluding legs) usually black; T2– of Isdromas for which host use is known, T7 may vary from black to dark brown as a likely pseudohyperparasitoid of Prota- (Fig. 34). panteles dalosoma (De Santis) (Braconidae: Fauna.—Isdromas contains eight de- Microgastrinae) on Anacraga citrinopsis scribed species worldwide with one spe- Dyar (Lepidoptera: Dalceridae). cies reported in the Nearctic (Yu et al. Literature.—The genus may be iden- 2005). Although Townes (1970) estimated tified using Townes (1983); lycaenae is the that the world fauna consists of at least 60 only Nearctic representative. VOLUME 114, NUMBER 1 93

North American Records: eastern North America, one from South Isdromas lycaenae (Howard) (Figs. 31, America, and the others from India, 33–34, 42, 44) Hemiteles lycaenae Vietnam, Japan, Taiwan, Madagascar, and Howard, 1889: 1880. Holotype /, southern Africa. An additional species was USNM (examined). described from eastern North America by Isdromas lycaenae is the sole Nearctic Smith and Stocks (2005). Carmean and representative of this genus and may be Kimsey (1998) gave a discussion of the recognized by the apical 0.3 of clypeus genus. turned inward at 90° (Fig. 43), vein 2-Cu Biology.—Although some species are of hind wing basally incomplete, and the collected rather commonly, very little is dark brown body with paler legs (Fig. 34). known about hosts and biology. Seasonal Distribution: In the Nearctic, Is. lycae- flight activity of O. pulchella in eastern nae is found from Ontario south to the North America was recorded in D. Smith Gulf of Mexico and west to Texas and (1996). Carlson (1979b) reported O. Kansas. In the Neotropics it is found from pulchella from a tachinid parasitizing an Honduras to Argentina (Yu et al. 2005). undetermined species of Lepidoptera. Limacodidae: Isdromas lycaenae was Benoit (1951) recorded O. hova Bischoff reared from mummified early to from a limacodid moth in Madagascar. middle instar larvae of Ac. stimulea* The first verified host records were re- on Q. rubra; I. textula* on Q. rubra; N. ported by Murphy et al. (2009) from nasoni* on F. grandifolia, P. serotina, tachinids parasitizing limacodids, noctuids, and Q. rubra; Pa. chloris* on Q. rubra; and megalopygids. The female deposits and Pr. badia* on F. grandifolia (Table numerous eggs on leaves of angiosperms. 1). The mummies were probably made The eggs are ingested by caterpillars by a species of Triraphis based upon feeding on the leaves, eclosion occurs in images of the host remains (JTL). the gut of the caterpillar, and the first in- Other hosts: Yu et al. (2005) listed star bores through the gut wall. Further records of Is. lycaenae from five spe- development depends on parasitization of cies of Braconidae and two species of the caterpillar by a parasitoid, in this case a tachinid (Murphy et al. 2009). The other Ichneumonidae; they also record it Nearctic species, O. bella Smith and from nine lepidopteran species in seven Stocks, is known from a single speci- families. Records of Is. lycaenae as men from Great Smoky Mountains aprimaryparasitoidarequestionable National Park, Tennessee. for the reasons discussed above. Literature.—Townes (1956) revised the North American Trigonalidae and sepa- Orthogonalys Schulz (Trigonaloidea rated Orthogonalys from other genera. Trigonalidae: Orthogonalinae) The genus was treated in D. Smith (1996), (Figs. 70–71) and seasonal flight data were given for Diagnosis.—Antenna black with white O. pulchella.CarmeanandKimsey(1998) or light yellow band in middle; propo- gave a diagnosis and description and deum usually with light markings; meta- provided the known distribution and bi- soma thin, without punctures, usually ology of the genus. Murphy et al. (2009) entirely orange; without female arma- recorded hosts for O. pulchella. ture; male antenna without tyloids. North American Records: Fauna.—Carmean and Kimsey (1998) Orthogonalys pulchella (Cresson) (Figs. listed 11 species worldwide, one from 70–71): Trigonalys pulchellus Cresson, 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

1867: 351. Holotype ?,ANSP(not species worldwide from the Nearctic, Neo- examined). tropical, eastern Palearctic, Australasian, This species and O. bella are the only Oriental, and Afrotropical regions. Most species of Orthogonalys in North and are from eastern Asia and South America. Central America. They are illustrated and Only one species, Taeniogonalos gund- separated by Smith and Stocks (2005). lachii,occursineasternNorthAmerica. Orthogonalys bella, known only from a Biology.—Species of this genus have female, is almost entirely black, has been reared from a variety of tachinid antennal segments 7–9 white, smaller and ichneumonid parasitoids of species eyes slightly diverging below and far of Pyralidae, Noctuidae, Saturniidae, removed from the posterior margin of Megalopygidae, and Arctiidae. A summary the head, tarsal claws with both teeth of their biology and hosts was given by D. subequal in size, and the sheath in lat- Smith (1996) and Carmean and Kimsey eral view broadly rounded at its apex. (1998). Krauth and Williams (2006) Figures 58 and 59 show the typical color recorded T. gundlachii from an arctiid, of O. pulchella. There is some variation Euchaetes egle (Drury), in Wisconsin. in the amount of black on the mesosoma Additionally, there is one record from a and dorsum of the metasoma. tachinid parasitizing a detritivore tipulid, Distribution: Southeastern Canada, Tipula ?flavoumbrosa (Gelhaus 1987). The eastern U.S.A., Mexico. African Taeniogonalos maynei Benoit was Limacodidae: Associated with primary reared from pupae of albipunctata parasitoids of Iso. beutenmuelleri and Holland (Limacodidae) in which it pre- I. textual. Most records are from tach- sumably was parasitizing a primary par- inids parasitizing I. textula feeding on asitoid (Benoit 1950). In Australia, T. Q. prinus and Q. rubra. maculata (Smith) and T. venatoria Riek Other hosts: Uramya pristis parasitiz- have been reared as primary parasitoids ing Meg. crispata (Megalopygidae), from pergid sawfly hosts including Per- Isochaetes beutenmuelleri, I. textula, gagrapta condei Benson, Perga dorsalis Leach, P. affinis Kirby, and Pseudoperga and Acrocnicta increta (Noctuidae) belinda (Kirby) (Raff 1934; Carne 1969; (Murphy et al. 2009). Recorded para- Weinstein and Austin 1991, 1995, 1996). sitizing Com. concinnata through Oviposition and general biology are sim- Actias luna (L.) (Kellogg et al. 2003). ilar to that described for Orthogonalys. Taeniogonalos Schulz (Trigonaloidea Literature.—Townes (1956) separated Trigonalidae: Trigonalinae) the genus (as Poecilogonalos)fromother North American genera. The genus was (Figs. 18, 72–73) also separated from other eastern North Diagnosis.—Antenna uniformly yellow American trigonalids and the seasonal brown or brown becoming darker apically, activity of its species recorded in D. Smith never banded (Figs. 72–73); metasoma (1996). stout, with punctures, black with trans- North American Records: verse yellow stripes; female with armature Taeniogonalos gundlachii (Cresson) (Figs. (Fig. 72); male antenna with tyloids (ventral 18, 72–73): Poecilogonalos gundlachii sensory structures). Cresson, 1865: 10. Holotype ?,ANSP Fauna.—Taeniogonalos is the most (not examined). widely distributed of all trigonalid genera. This is the only species of Taeniogonalos in Carmean and Kimsey (1998) listed 34 eastern North America. Two other species VOLUME 114, NUMBER 1 95 occur in southern Mexico and Central Monema flavescens (numerous records, America. The typical color of eastern see Arnaud 1978). Chaetexorista javana North and Central American specimens is was introduced into Massachusetts from shown in Figs. 72–73. Specimens from Japan in 1929 and 1930 for control of the Cuba are more extensively yellow, espe- Oriental moth, M. flavescens (Dowden cially the head and mesosoma, whereas 1946). The introduction was successful, some specimens from the southern tier but Ch. javana has not been reported for of states from Florida to Louisiana are many years and is probably now extinct in somewhat intermediate in color. Carmean North America. Similarly, M. flavescens is and Kimsey (1998) considered all as a single likely extinct in North America also based widespread, variable species. In the literature on subsequent searches (M. Epstein pers. prior to Carmean and Kimsey (1998), the comm.). eastern North American species is cited as (3) Hyphantrophaga virilis (Aldrich and Poecilogonalos costalis (Cresson). Webber, 1924) (Exoristinae: Goniini): Distribution: Southeastern Canada, Euclea delphinii (Coquillett 1897, as eastern U.S.A. from Massachusetts to Exorista blanda (Osten Sacken) ex E. Florida west to Wisconsin, Ohio, Loui- cippus (Cramer)). The original record by siana; Costa Rica, Cuba. Coquillett (1897) was later cited by Limacodidae: Associated with primary Aldrich and Webber (1924, as Zenillia parasitoids of E. delphinii. Other hosts: One specimen reared from blanda blanda (Osten Sacken)), Sellers atachinid,probablyCom. concinnata (1943, as Z. virilis), and Arnaud (1978, as parasitizing E. delphinii feeding on Eusisyropa virilis). Nyssa sylvatica.Recordedparasitiz- (4) mediocris Aldrich, 1925 ing Com. concinnata through Actias (Exoristinae: Blondeliini): luna (L.) (Kellogg et al. 2003). Monema flavescens (Brimley 1938, “reared from twig infested by Oriental Questionable/Invalid/Miscellaneous moth”; Arnaud 1978 [citing Brimley Parasitoid Records (North America) record]). (5) Neomintho celeris (Townsend, 1919) Tachinidae (: Euthelairini): The following are single records that Packardia ceanothi Dyar (Raizenne require confirmation, or in the case of 1952, as Eupelecotheca celer; Arnaud Chaetexorista javana,involveanintro- 1978 [citing Raizenne record, also as duced species that is probably no longer Eupelecotheca celer]). extant in North America. Some records may be based on misidentifications or Hymenoptera represent rare or accidental host records. (Note: None of the taxa listed below (1) Carcelia spp. (Exoristinae: ): appear in Table 1) Adoneta spinuloides (Brauer and Ber- genstamm 1895, as Chaetolyga sp.), Ph. (1) Closterocerus cinctipennis Ashmead pithecium (Patton 1958, ex “hag-moth”, (Eulophidae: Entedoninae): “Phobetron pithecium?”). Both records Associated with Ac. stimulea on Q. cited by Arnaud (1978). rubra (no host remains), but this species (2) Chaetexorista javana Brauer and is possibly the result of a contaminant Bergenstamm, 1895 (Exoristinae: introduced during rearing as they are Exoristini): known parasitoids of microlepidopteran, 96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON buprestid, and agromyzid leafminers; eggs (CDFA 2005, 2006). Were this to hap- of Chrysomelidae and Tenthredinidae; pen, it is possible that A. dimerus would and “Diptera on Catalpa”(Hansson be introduced also, either accidentally 1994). or, later, deliberately. (2) Neochrysocharis diastatae (Howard) (5) Brachymeria euploeae (Westwood) (Eulophidae: Entedoninae): (Chalcididae: Chalcidinae): Associated most frequently with lepi- Recorded from the U.S.A. (but not dopteran and dipteran leaf/stem miners attacking a limacodid) and as a parasit- (Hansson 1995). Five specimens are as- oid on Chalcoscelis albiguttata Snellen sociated with E. delphinii on Q. alba, but in Bhutan (as Butjan) and Maluku the limacodid remains offer no indication Islands (as Moluccas) (Thompson 1954). of parasitization (e.g., emergence holes). Chalcoscelis albiguttata does not occur in We suspect that a leaf miner parasitized the U.S.A. and has never been introduced by Neo. diastatae was introduced during (M. Epstein pers. comm.). The recorded the rearing of the larva. distribution for Br. euploeae is in eastern (3) Horismenus sp. (Eulophidae: and southeastern Asia and India with the Entedoninae): lone U.S.A. record (Thompson 1954) A single male is associated with Ac. standing out as the single record for the stimulea on Q. rubra (no host remains) Western Hemisphere. Other host records but became infected with fungus after and associated references are presented in emergence, obscuring morphological fea- Cock (1987). The Thompson (1954) record tures. It was run through Burks’ (1971) indicates that Br. euploeae was introduced key to Nearctic species (even though key into the U.S.A. from Japan to control is written for females) and compared with Ostrinia (as Pyrausta) nubilalis (Hu¨bner). males of species near which it keyed. It All of the limacodid hosts recorded for Br. is not conspecific with males of Ho. la- euploeae are distributed in eastern and trodecti Burks or Ho. carolinensis Burks. southeastern Asia and India. Two extralimital species from the Neo- (6) Brachymeria lasus Walker (Chalci- tropics, Ho. clavicornis Cameron and Ho. didae: Chalcidinae): hipparchia Cameron, attack Ph. hip- This species is native to the Oriental and parchia in Guyana (De Santis 1979). Australasian regions but has been intro- (4) Aroplectrus dimerus Lin (Eulophi- duced into the U.S.A. several times to con- dae: Eulophinae): trol the gypsy moth (L. dispar; Weseloh This species is known from the Phil- and Anderson 1982, ROBO Database ippines, Taiwan, China, and India and 1981–1985), fall webworm (Hyphantria recorded from Pa. bicolor (Singh et al. cunea Drury; ROBO Database 1981– 1988) and Penthocrates sp. (Fry 1989). It 1985), and range caterpillar (Hemileuca was released in Hawaii beginning in oliviae Cockerell; ROBO Database 1981– 2010 as part of a biological control 1985). Known limacodid hosts include program against the invasive limacodid Thosea cinereomarginata Banks and Th. (Moore) (HDOA 2010, sinensis Walker from the Philippines 2011; Conant et al. 2011). Although (Baltazar 1965). This species has a broad neither D. pallivitta nor Ar. dimerus are host range on small lepidopteran pupae currently known from the U.S.A., it is and is facultatively hyperparasitic through highly likely that the limacodid will be hymenopterans and dipterans (Cock 1987). introduced into California given its his- (7) Conura immaculata (Cresson) tory of interception at ports of entry (Chalcididae: Chalcidinae): VOLUME 114, NUMBER 1 97

This species is primarily Neotropical type of emergence from the cadaver but occurs in southern U.S.A., although during the pre-dissection inspection. it is not associated with Limacodidae there. It has been recorded parasitizing ACKNOWLEDGMENTS N. subpectinata Dyar and Euprosterna We thank Steve Lingafelter, Thomas elaea (as elaeasa) Dyar (Delvare 1992), Henry, John Brown (Systematic Ento- primarily Neotropical species, in that re- mology Laboratory [SEL], PSI, ARS, gion. It is also known as a hyperparasitoid USDA), and Paul Hanson (University of on Ichneumonoidea (Delvare 1992), and Costa Rica, San Jose´, Costa Rica) for Burks (1979) recorded it from Meteorus their critical reviews of and suggestions sp. (Braconidae) from Cameron County, on this manuscript. We also thank Scott Texas. The latter specimens and parasitoid Whittaker (SEM Lab Manager of the pupal remains are housed in the USNM. Scanning Electron Microscopy Lab, Lingren (1977) records it as a hyperpara- Smithsonian Institution, National Museum sitoid of Campoletis sonorensis (Ca- Natural History) for stub preparation and meron) through Heliothis virescens (F.) on SEM access. A special thanks to Marc cotton in Brownsville, Texas. Epstein (California Department of Food (8) Conura lasnierii Gue´rin-Me´neville and Agriculture) for his thorough review (Chalcididae: Chalcidinae): This chalcidid is recorded from Florida of the manuscript, including pointing out in the U.S.A. but is not known from a omissions and limacodid nomenclatural limacodid host. It has been recorded from issues. Thanks to Jerrett McCormick Leucophobetron argentiflua Hu¨bner in (SEL), Taina Litwak (SEL), Lynette Gates Cuba (Gahan 1934). The moth has never (spouse, MWG), Matt Buffington (SEL), been intentionally introduced into the U. and Terry Nuhn (SEL) for consultation, S.A. (M. Epstein pers. comm.). Addi- information entry, and specimen imaging. tionally, Ala. slossoniae, a common spe- Istva´nMiko´ and Matt Yoder (North cies in Florida, is closely related to the Carolina State University, Raleigh, North monotypic Leucophobetron and perhaps Carolina) provided consultation on Cera- congeneric (M. Epstein pers. comm.). phron and Trichopria,respectively.Andy (9) Trichopria sp. (: Dia- Deans (North Carolina State University, priinae): Raleigh, North Carolina) permitted the A single male is associated with E. reproduction of two images of Ceraphron delphinii on Q. alba and is likely the (Figs. 63–64). Neal Evenhuis (Bishop result of contamination during the rear- Museum, Honolulu, Hawaii) vetted the ing process (see Neochrysocharis dia- section on Systropus and provided the im- statae, above). Species of Diapriinae are age of S. macer.Fundingfortherearing recorded as parasitoids of orthorhaphous portion of this project was provided by (soldier flies, flower flies, etc.) and cy- NSF-DEB 0643438 to JTL. The fourth au- clorhaphous (tachinid flies, muscoid flies, thor (JEO) is grateful to D. M. Wood (Ag- etc.) Diptera (www.diapriid.org/public/ riculture and Agri-Food Canada, Ottawa) site/diapriid/home). The larval cadaver for sharing his knowledge of the tachinid associated with this specimen was thor- flies treated in this paper and for allowing oughly dissected in search of a tachinid the use of his unpublished illustrations of puparium that would have likely served Austrophorocera male genitalia in Figs. as the host in this instance, but none was 14–17. The fourth author (JEO) also discovered. There was no evidence of any thanks S. J. Henderson (Agriculture and 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Agri-Food Canada, Ottawa) for creating Ashmead, W. H. 1896. Descriptions of new para- the montaged images of tachinids and sitic Hymenoptera. Transactions of the Amer- their structures that appear herein. ican Entomological Society 23: 179–234. Ashmead, W. H. 1897. Two new parasites from USDA is an equal opportunity provider Eupoeya slossoniae. Canadian Entomologist and employer. 29: 113–114. doi:10.4039/Ent29113-5 Ashmead, W. 1904. Classification of the chalcid Literature Cited flies of the superfamily Chalcidoidea, with descriptions of new species in the Carnegie van Achterberg, C. 1991. Revision of the genera of Museum, collected in South America by the Afrotropical and W. Palaearctical Roga- Herbert H. Smith. Memoirs of the Carnegie dinae Foerster (Hymenoptera: Braconidae). Museum 1: 225–551. Zoologische Verhandelingen 273: 1–102. Askew, R. R. 1971. Parasitic Insects. Heinemann Adams, J. and D. Yanega. 1991. The lepidopteran Educational Books, London, England. 316 pp. host of a Neotropical bombyliid fly (Lepi- Austin, A. D. 1987a. A review of the Braconidae doptera: Limacodidae, Diptera: Bombyliidae). (Hymenoptera) that parasitize Limacodidae Journal of the Kansas Entomological Society in Southeast Asia, particularly those associ- 64: 443–444. ated with and oil palm, pp. 139–164. Aiello, A. 1980. Systropus columbianus (Diptera: In M. J. W. Cock, H. C. J. Godfray, and J. D. Bombyllidae [sic]) reared from a larva of Holloways, eds. Slug and Nettle Caterpillars. Limacodid moth. Entomological News 91: The Biology, Taxonomy and Control of the 89–91. Limacodidae of Economic Importance on Aldrich, J. M. 1921. The muscoid genera Pseu- Palms in South-east Asia. CAB International, deuantha and Uramyia (Diptera). Insecutor Wallingford, Oxon, United Kingdom. 270 pp. Inscitiae Menstruus 9: 83–92. Austin, A. D. 1987b. Ceraphronidae, pp. 181– Aldrich, J. M. 1925. Two new species of the 183. In M. J. W. Cock, H. C. J. Godfray, and tachinid genus Lixophaga, with notes and key J. D. Holloways, eds. Slug and Nettle Cater- (Diptera). Proceedings of the Entomological pillars. The Biology, Taxonomy and Control Society of Washington 27: 132–136. Aldrich, J. M. and R. T. Webber. 1924. The North of the Limacodidae of Economic Importance American species of parasitic two-winged on Palms in South-east Asia. CAB Interna- flies belonging to the genus Phorocera and tional, Wallingford, Oxon, United Kingdom. related genera. Proceedings of the United 270 pp. States National Museum 63 (Art. 17) [= No. Baltazar, C. R. 1965. A new species of ichneumon- 2486]: 1–90. fly and a parasitic on slug cat- Andre´, E. 1881. Notes hyme´nopte´rologiques. I. erpillars in Basilan Island. Philippine Journal Chalcidites. Annales de la Socie´te´ Entomolo- of Plant Industry 30: 109–113. gique de France (6e Se´rie) 1: 332–344. Barbosa, P. and A. Caldas. 2004. Patterns of Arnaud, P. H., Jr. 1978. A host-parasite catalog parasitoid-host associations in differently of North American Tachinidae (Diptera). parasitized macrolepidopteran assemblages United States Department of Agriculture. on black willow Salix nigra (Marsh) and box Miscellaneous Publication 1319: 1–860. elder Acer negundo L. Basic and Applied Ashmead, W. H. (1888) 1889. Descriptions of Ecology 5: 75–85. new Braconidae in the collection of the U. S. Barbosa, P., J. L. Capinera, and E. A. Harrington. National Museum. Proceedings of the United 1975. The gypsy moth parasitoid complex in States National Museum 11: 611–671. western Massachusetts: a study of parasitoids doi:10.5479/si.00963801.11-760.611 in areas of high and low host density. Envi- Ashmead, W. H. 1890. Description of new Ich- ronmental Entomology 4: 842–846. neumonidae in the collection of the U.S. Barbosa, P. and A. Caldas. 2004. Patterns of para- National Museum. Proceedings of the United sitoid-host associations in differently parasit- States National Museum 12: 387–451. ized macrolepidopteran assemblages on black doi:10.5479/si.00963801.12-779.387 willow Salix nigra (Marsh) and Box Elder Ashmead, W. H. 1893. A monograph of the North Acer negundo L. Basic and Applied Ecology 5: American Proctotrypidae. Bulletin of the 75–85. United States National Museum 45: 1–472. Becker, V. O. and M. E. Epstein. 1995. Limaco- doi:10.5479/si.03629236.45.1 didae, pp. 128–132. In J. B. Heppner, ed. VOLUME 114, NUMBER 1 99

Checklist: Part 2. Hyblaeoidea – – Bulletin. Connecticut State Geological and Torticoidea. Atlas of Neotropical Lepidoptera. Natural History Survey 22: 1–848. Scientific Publishers, Gainesville, Florida. Buffington, M. L., R. Burks, and L. McNeil. Benoit, P. L. G. 1950. Les Trigonaloidae du Congo 2005. Advanced techniques for imaging Belge (Hym. – Terebr.). Revue de Zoologie et parasitic Hymenoptera (Insecta). American de Botanique Africaines 43: 216–221. Entomologist 51: 50–56. Benoit, P. L. G. 1951. La systematique des Trig- Buffington, M. L. and M. W. Gates. 2008. onaloidae (Hym.) ethiopiens. Revue de Zoo- Advanced imaging techniques II: using a logie et de Botanique Africaines 44: 141–147. compound microscope for photographing Boettner, G. H., J. S. Elkinton, and C. J. Boettner. point-mount specimens. American Entomol- 2000. Effects of a biological control in- ogist 54: 222–224. troduction on three nontarget native species Burks, B. D. 1940. Revision of the chalcid-flies of of saturniid moths. Conservation Biology 14: the tribe Chalcidini in America north of 1798–1806. doi:10.1046/j.1523-1739.2000. Mexico. Proceedings of the United States 99193.x National Museum 88: 237–354. doi:10.5479/ Boucˇek, Z. 1992. The New World genera of si.00963801.88-3082.237 Chalcididae. Memoirs of the American En- Burks, B. D. 1966. The North American species tomological Institute 53: 49–118, 443–446. of Pediobius Walker (Hymenoptera: Eulo- Boucˇek, Z. and G. Delvare. 1992. The identities phidae). Proceedings of the Entomological of species described or classified under Society of Washington 68: 33–43. Chalcis by J. C. Fabricius. Memoirs of the Burks, B. D. 1971. The Nearctic species of Ho- American Entomological Institute 53: 29. rismenus Walker (Hym., Eulophidae). Pro- Boucˇek, Z. and J. Halstead. 1997. Chalcididae, ceedings of the Entomological Society of pp. 151–164. In G. A. P. Gibson, J. T. Huber, Washington 73: 68–83. and J. B. Woolley, eds. Annotated Keys to the Burks, B. D. 1979. (Agaoninae) and Genera of Nearctic Chalcidoidea (Hyme- all other families of Chalcidoidea (excluding noptera). Research Press. National Research ), pp. 748–749, 768–889, 967–1043. Council of Canada, Ottawa, Ontario. 794 pp. In K.V. Krombein, P. D. Hurd, Jr., D. R. Smith, Bourchier, R. S. 1991. Growth and development and B. D. Burks, eds. Catalog of Hyme- of Compsilura concinnata (Meigan [sic]) noptera in America North of Mexico. Vol 1. (Diptera: Tachinidae) parasitizing gypsy moth Smithsonian Institution Press, Washington, larvae feeding on tannin diets. Canadian D.C. 1198 pp. Entomologist 123: 1047–1055. doi:10.4039/ Butler, L. 1993. Parasitoids associated with the Ent1231047-5 macrolepidoptera community at Coopers Brauer, F. and J. E. von Bergenstamm. 1895. Die Rock State Forest, West Virginia: a baseline Zweiflu¨gler des Kaiserlichen Museums zu study. Proceedings of the Entomological So- Wien. VII. Vorarbeiten zu einer Monographie ciety of Washington 95: 504–510. der Muscaria schizometopa (exclusive An- California Department of Food and Agriculture thomyidae). Pars IV. F. Tempsky, Wien. 88 pp. (CDFA). 2005. Darna pallivitta,p.21.Cal- ([Also published in 1895. In Denkschriften der ifornia Plant Pest and Disease Report 22: 1–76. Kaiserlichen Akademie der Wissenschaften. California Department of Food and Agriculture Wien. Mathematisch-Naturwissenschaftliche (CDFA). 2006. Darna pallivitta,p.12.California Classe, 61 (1894), 537–624.]) Plant Pest and Disease Report 23: 1–117. Brimley, C. S. 1938. The Insects of North Carolina. Cameron, P. 1913. The Hymenoptera of the Being a List of the Insects of North Carolina Georgetown Museum. Part V. Timehri, Guyana and Their Close Relatives. North Carolina De- 3: 105–137. partment of Agriculture, Raleigh. 560 pp. Cantrell, B. K. 1986. An updated host catalogue Brues, C. 1906. Notes and descriptions of North for the Australian Tachinidae (Diptera). Jour- American parasitic Hymenoptera. II. Bulletin nal of the Australian Entomological Society of the Wisconsin Natural History Society 4: 25: 255–265. doi:10.1111/j.1440-6055.1986. 143–152. tb01112.x Brues, C. 1916. Serphoidea, pp. 529–576. In H. L. Cantrell, B. K. and R. W. Crosskey. 1989. Family Viereck, ed. Guide to the Insects of Con- Tachinidae, pp. 733–784. In N. L. Evenhuis, necticut. Part III. The Hymenoptera, or Wasp- ed. Catalog of the Diptera of the Australasian like Insects, of Connecticut. Chalcidoidea. and Oceanian Regions. Bishop Museum 100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Special Publication 86. Bishop Museum of Economic Importance on Palms in South- Press (Honolulu) and E. J. Brill (Leiden). east Asia. CAB International, Wallingford, 1155 pp. Oxon, United Kingdom. 270 pp. Carlos, C. R., J. R. Costa, C. B. Ta˜o, F. Alves, and Cock, M. J. W., H. C. J. Godfray, and J. D. Holloway. L. M. Torres. 2006. Natural parasitism of the 1987. Slug and Nettle Caterpillars: the Biology, grape berry moth, Lobesia botrana (Denis & Taxonomy, and Control of the Limacodidae of Schiffermu¨ller) at Regia˜o Demarcada do Economic Importance on Palms in South-east Douro. Boletin de Sanidad Vegetal, Plagas Asia. CAB International, Wallingford, United 32: 355–362. Kingdom. 270 pp. Carlson, R. W. 1979a. Family Ichneumonidae, Collins, C. 1933. The Oriental moth (Cnido- pp. 315–740. In K. V. Krombein, P. D. Hurd, campa flavescens Walk.) in Massachusetts Jr., D. R. Smith, and B. D. Burks, eds. Cat- and the work of its newly introduced para- alog of Hymenoptera in American North of site. Journal of Economic Entomology 26: Mexico. Vol. 1. Smithsonian Institution Press, 54–57. Washington, D.C. 1198 pp. Conant, P., A. H. Hara, L. M. Nakahara, and R. A. Carlson, R. W. 1979b. Superfamily Trigonaloidea, Heu. 2002. Nettle caterpillar, Darna palli- pp. 1197–1198. In K. V. Krombein, P. D. vitta Moore (Lepidoptera: Limacodidae). Hurd, Jr., D. R. Smith, and B. D. Burks, eds. New Pest Advisory No. 01–03. Hawaii De- Catalog of Hymenoptera in American North partment of Agriculture, Honolulu, HI. of Mexico. Vol. 1. Smithsonian Institution Conant, P., A. H. Hara, W. T. Nagamine, C. M. Press, Washington, D.C. 1198 pp. Kishimoto, and R. A. Heu. 2011. Nettle cat- Carmean, D. and L. Kimsey. 1998. Phylogenetic erpillar, Darna pallivitta Moore (Lepidop- revision of the family Trig- tera: Limacodidae). New Pest Advisory No. onalidae (Hymenoptera). Systematic Ento- 01–03, updated. Hawaii Department of Ag- mology 23: 35–76. doi:10.1046/j.1365-3113. riculture, Honolulu, HI. 1998.00042.x Conant, P., C. K. Hirayama, C. M. Kishimoto, Carne, P. B. 1969. On the population dynamics of and A. H. Hara. 2006. Trichogramma papil- the Eucalyptus-defoliating sawfly Perga affi- ionis (Nagarkatti), the first recorded Tri- nis affinis Kirby (Hymenoptera). Australian chogramma species to parasitize eggs in the Journal of Zoology 17: 113–141. doi:10.1071/ family Limacodidae. Proceedings of the Ha- ZO9690113 waiian Entomological Society 38: 133–135. Chadwick, C. E. and M. I. Nikitin. 1985. Records Cook, A. J. and G. C. Davis. 1891. Kerosene emul- of parasitism by members of the family Ta- sion, some new insects. Bulletin of the Michigan chinidae (Diptera: Tachinidae). Australian Agricultural Experiment Station 73: 1–16. Zoologist 21: 587–598. Coquillett, D. W. 1897. Revision of the Tachini- Cha´ves, T. H. A., D. I. A. Dı´az, and G. R. A. dae of America north of Mexico. A family of Bricen˜o. 1993. Introduction to Venezuela and parasitic two-winged insects. United States biology of Cotesia plutellae Kurdj. (Hyme- Department of Agriculture. Division of En- noptera: Braconidae), parasitoid of Plutella tomology. Technical Series 7: 1–156. xylostella L. (Lepidoptera: Plutellidae). Manejo da Costa Lima, A. 1962. Insetos do Brasil. 12° Integrado de Plagas No. 29: 24–27. Tomo. Hymeno´pteros. 2a Parte. Escola Na- Chen, X. and J. He. 1997. Revision of the sub- cional de Agronomia (Se´rie Dida´ctica N° 14), family Rogadinae (Hymenoptera: Braconi- Rio de Janeiro. 393 pp. dae) from China. Zoologische Verhandelingen Cresson, E. T. 1865. On the Hymenoptera of 308: 1–187. Cuba. Proceedings of the Entomological Clausen, C. P. 1956. Biological control of insect Society of Philadelphia 4: 1–200. pests in the continental United States. United Cresson, E. T. 1867. Descriptions of two new spe- States Department of Agriculture Technical cies of Trigonalys. Proceedings of the Ento- Bulletin No. 1139: 1–151. mological Society of Philadelphia 6: 351–352. Cock, M. J. W. 1987. Chalcidoidea. Notes on the Cresson, E. T. 1869. List of the North American chalcidoids attacking South-east Asian Li- species of the genus Aleiodes Wesmael. macodidae, pp. 165–179. In M. J. W. Cock, Transactions of the American Entomological H. C. J. Godfray, and J. D. Holloways, eds. Society 2: 377–382. Slug and Nettle Caterpillars: the Biology, Cresson, E. T. 1872. Synopsis of the North Amer- Taxonomy, and Control of the Limacodidae ican species belonging to the genera Leucospis, VOLUME 114, NUMBER 1 101

Smicra and Chalcis. Transactions of the Academia Nacional de Agronomia y Vet- American Entomological Society 4: 29–60. erinaria No 13: 1–145. Crosskey, R. W. 1976. A taxonomic conspectus Dessart, P. 1981. Definition de quleques sous- of the Tachinidae (Diptera) of the Oriental genres de Ceraphron Jurine, 1807 (Hyme- Region. Bulletin of the British Museum noptera, Ceraphronoidea, Ceraphronidae). (Natural History). Entomology Supplement Bulletin de l’Institut Royal des Sciences 26: 1–357. Naturelles de Belgique 53: 1–23. Cruttwell, R. E. 1969. The biology and mode of Dessart, P. and P. Cancemi. 1986. Tableau di- parasitism of Uromacquartia trinitatis chotomique des genres de Ceraphronoidea Thompson (Diptera: Tachinidae). Common- (Hymenoptera) avec commentaires et nou- wealth Institute of Biological Control. Tech- velles espe´ces. Frustula Entomologica N. S. nical Bulletin 12: 20–28. 7–8(20–21): 307–372. Culver, J. J. 1919. A study of Compsilura con- Dowden, P. B. 1946. Parasitization of the Oriental cinnata, an imported tachinid parasite of the moth (Cnidocampa flavescens (Walk.)) by gypsy moth and the brown-tail moth. Bulletin Chaetexorista javana B. and B. Annals of the of the United States Bureau of Entomology Entomological Society of America 39: 225–241. 766: 1–27. Dyar, H. G. 1897. On the white Eucleidae and the Curran, C. H. 1930. Report on the Diptera col- larva of Calybia slossoniae (Packard). Jour- lected at the Station for the Study of Insects, nal New York Entomological Society 5: 121– Harriman Interstate Park, N.Y. Bulletin of the 126 (plate 5). American Museum of Natural History 61: Dyar, H. G. 1900. Life history of a South 21–115. American slug-caterpillar Sibine fusca Stoll. Cushman, R. A. 1925. The synonymy and generic Entomological News 11: 517–526. position of two North American ichneumon- Elkinton, J. S. and G. H. Boettner. 2004. The flies. Proceedings of the Entomological So- effects of Compsilura concinnata, an in- ciety of Washington 27: 164–166. troduced generalist tachinid, on non-target Dasch, C. 1971. Ichneumon-flies of America species in North America: a cautionary tale, pp. North of Mexico: 6. Subfamily Mesochor- 4–14. In R. G. Van Driesche and R. Reardon, inae. Memoirs of the American Entomolog- eds. Assessing Host Ranges for Parasitoids and ical Institute 16: 1–376. Predators Used for Classical Biological Con- Davis, D. R. 1983. Limacodidae, pp. 66-67. In trol: a Guide to Best Practice. United States R. W. Hodges, ed. Checklist of the Lepi- Deptartment of Agriculture Forest Health doptera of America North of Mexico. E. W. Technology Enterprise Team, Morgantown, Classey Ltd and The Wedge Entomological West Virginia. FHTET-2004–03. 243 pp. Research Foundation, London, England. Elkinton, J. S. and A. M. Liebhold. 1990. Population 284 pp. dynamics of gypsy moth in North America. Deans, A. R., I. Miko, M. J. Yoder, K. S. Seltmann, Annual Review of Entomology 35: 571–596. and The H. A. O. Consortium. [Wed Mar 17 doi:10.1146/annurev.en.35.010190.003035 10:45:36 -0500 2010]. Hymenoptera Glossary: Elkinton, J. S., D. Parry, and G. H. Boettner. http://purl.oclc.org/NET/hymontology 2006. Implicating an introduced generalist Delvare, G. 1992. A reclassification of the Chal- parasitoid in the invasive browntail moth’s cidini with a checklist of the New World enigmatic demise. Ecology 87: 2664–2672. species. Memoirs of the American Entomo- doi:10.1890/0012-9658(2006)87[2664:IAIGPI] logical Institute 53: 119–441, 443–466. 2.0.CO;2 De Santis, L. 1979. Cata´logo de los hime´nopteros Enderlein, G. 1926. Zur Kenntnis der Bombyliiden- calcidoideos de Ame´rica al sur de los Estados Subfamilie Systropodinae (Dipt.). Wiener En- Unidos. Publicacio´n Especial Comisio´n de tomologische Zeitung 43: 69–92. Investigaciones Cientificas Provincia de Epstein, M. 1988. An overview of slug caterpillar Buenos Aires. La Plata, Argentina. 488 pp. moths (Lepidoptera: Limacodidae) with em- De Santis, L. 1987. Himeno´pteros parasito´ides e phasis on genera in the New World Parasa Hiperparasito´ides de Anacraga sp. (Lepi- Group. Doctoral thesis, Department of En- doptera, Dalceridae) en Brasil. Revista Bra- tomology, University of Minnesota, St. Paul, sileira de Entomologia 31: 97–99. Minnesota. xi + 149 pp. De Santis, L. and P. Fidalgo. 1994. Catalogo de Epstein, M. 1996. Revision and phylogeny of the Himenopteros Calcidoideos. Serie de la limacodid group families, with evolutionary 102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

studies on slug caterpillars (Lepidoptera: Zyg- Ferrie`re and Autoplectrus taprobanes Gadd. aenoidea). Smithsonian Contributions to Zool- Ceylon Journal of Science (B) 23: 81–94. ogy 582: 1–102. doi:10.5479/si.00810282.582 Gahan, A. B. 1934. Descriptions of some new Epstein, M. 1997. Parasa indetermina (Bois- species of Chalcidoidea from Cuba and Pu- duval) (Lepidoptera: Limacodidae), a new erto Rico. Memorias de la Sociedad Cubana host for Systropus macer Loew (Diptera: de Historia Natural, Habana 8: 125–134. Bombyliidae). Proceedings of the Entomo- Gates, M. and R. Burks. 2003. Hosts and ranges. logical Society of Washington 99: 585–586. Chalcid Forum 25: 5 (newsletter). Evans, G. A and L. A. Stange. 1997. Parasitoids Gates, M. W. and T. Stoepler. 2010. A new spe- associated with the brown citrus , Tox- cies of Alveoplectrus Wijesekara and Schauff optera citricida, in Florida (Insecta: Hyme- (Hymenoptera: Chalcidoidea: Eulophidae) noptera). Florida Department of Agriculture parasitic on Limacodidae (Lepidoptera). and Consumer Service, Division of Plant In- Proceedings of the Entomological Society of dustry, Entomology Circular No. 384 Gain- Washington 112: 207–213. doi:10.4289/0013- esville, Florida. 5 pp. 8797-112.2.207 Evenhuis, N. L. 1982. Catalog of primary types of Gauld, I. D. 1984. An introduction to the Ich- Bombyliidae (Diptera) in the entomological neumonidae of Australia; with a contribution collections of the Museum of Comparative on by M. G. Fitton. British Mu- Zoology, with designations of lectotypes. seum (Natural History), no. 895: 1–413. Breviora 469: 1–23. Gauld, I. D. 1988. Evolutionary patterns of host Evenhuis, N. L. and D. J. Greathead. 1999. World utilization by ichneumonoid parasitoids (Hy- Catalog of Flies (Diptera: Bombyliidae). menoptera: Ichneumonidae and Braconidae). Backhuys Publishers, Leiden. xlviii + 756 pp. Biological Journal of the Linnean Society. Evenhuis, N. L. and D. J. Greathead. 2003. World Linnean Society of London 35: 351–377. Catalog of Bee Flies (Diptera: Bombyliidae) doi:10.1111/j.1095-8312.1988.tb00476.x Web site (http://hbs.bishopmuseum.org/ Gauld, I. D. 1995. 12.1 Ichneumonidae, pp. 390– bombcat/). [accessed May 2011]. 431. In P. E. Hanson and I. D. Gauld, eds. The Fabricius, J. C. 1798. Supplementum Entomolo- Hymenoptera of Costa Rica. Oxford Uni- giae Systematicae. Proft and Storch, Copen- versity Press, New York. hagen. 243 pp. Gauld, I. and B. Bolton. 1988. The Hymenoptera. Finlayson, T. 1975. The cephalic structures and Oxford University Press, London, England. spiracles of final-instar larvae of the sub- 332 pp. family Campopleginae, tribe Gauld, I. and J. Ferna´ndez-Triana. 2010. Type (Hymenoptera: Ichneumonidae). Memoirs of condition and generic placement of Cuban the Entomological Society of Canada 94: 1– species of Ichneumonidae described by 137. doi:10.4039/entm10794fv Cresson and collected by Gundlach. Zootaxa Fletcher, D. S. and I. W. B. Nye. 1982. The Generic 2394: 41–50. Names of Moths of the World. Vol. 4. British Gelhaus, J. K. 1987. A detritivore Tipula (Diptera: Museum (Natural History), London. 192 pp. Tipulidae) as a secondary host of Poecilogo- Fortier, J. C. and S. R. Shaw. 1999. Cladistics of nalos costalis (Hymenoptera: Trigonalidae). the Aleiodes lineage of the subfamily Roga- Entomological News 98: 161–162. dinae (Hymenoptera: Braconidae). Journal of Genty, P. 1972. Morfologı´aybiologı´adeSibine Hymenoptera Research 8: 204–237. fusca Stoll, Lepidoptero defoliador de la palma Franklin, H. J. 1919. Seventh report of the cran- de aceite en Colombia. Ole´agineux 27: 65–71. berry substation from 1917 to 1919. Bulletin Genty, P., R. Desmier, J. P. Morin, and C. A. of the Massachusetts Agricultural Experi- Korytkowski. 1978. Les ravaguers du palmier ment Station 192: 129. a huile en Amerique Latine. Ole´agineux 33: Fry, J. M. 1989. Natural enemy databank, 1987. A 325–419. catalogue of natural enemies of Gibson, G. A. P. 1997. Morphology and termi- derived from records in the CIBC Natural nology. Chapter 2, pp. 16–44. In G. A. P. Enemy Databank, p. 94. CAB International, Gibson, J. T. Huber, and J. B. Woolley, eds. Wallingford, Oxford, UK. 185 pp. Annotated Keys to the Genera of Nearctic Gadd, C. H., W. T. Fonseka, and D. J. W. Rana- Chalcidoidea (Hymenoptera). Research Press, weera. 1946. Parasites of tea nettlegrubs with National Research Council of Canada, Ottawa, special reference to Platyplectrus natadea Ontario. 764 pp. VOLUME 114, NUMBER 1 103

Gibson, G. A. P., J. T. Huber, and J. B. Woolley. Agriculture, Sacramento, California, U.S.A. 1997. Annotated Key to the Genera of 31 pp. Neartic Chalcidoidea (Hymenoptera). Re- Harrison, J. O. 1963. The natural enemies of some search Press. National Research Council of banana insect pests in Costa Rica. Journal of Canada, Ottawa, Ontario. 764 pp. Economic Entomology 56: 282–285. Girault, A. 1911. On the identity of the most Harvey, J., L. M. A. Witjes, and R. Wagenaar. common species of the family Trichog- 2004. Development of hyperparasitoid wasp rammatidae (Hymenoptera). Bulletin of the (Hymenoptera: Ichneumonidae) Wisconsin Natural History Society 9: 135–165. in a multitrophic framework. Environmental Girault, A. 1916. Descriptions of miscellaneous Entomology 33: 1488–1496. doi:10.1603/0046- chalcid-flies. Insecutor Inscitiae Menstruus 4: 225X-33.5.1488 109–120. Harvey, J. A., L. E. M. Vet, L. M. A. Witjes, and Godfray, H. C. J. and M. S. Chan. 1990. How T. M. Bezemer. 2006. Remarkable similarity trigger single-stage outbreaks in in body mass of a secondary hyperparasitoid tropical pests. Functional Ecology 4: 329– Lysibia nana and its primary parasitoid host 337. doi:10.2307/2389594 emerging from cocoons of Graham, M. W. R. de V. 1969. The Pteromalidae comparable size. Archives of Insect Bio- of north-western Europe (Hymenoptera: chemistry and Physiology 61: 170–183. Chalcidoidea). Bulletin of the British Mu- doi:10.1002/arch.20080 seum (Natural History). Historical Series Harvey, J. A., R. Wagenaar, and M. Bexemer. (Supplement 16): 1–908 (Natural History). 2009a. Interactions to the fifth trophic level: Greathead, D. J. 1987. Bombyliidae. A summary secondary and tertiary parasitoid wasps of the recorded parasitoids of South-east show extraordinary efficiency in utilizing Asian Limacodidae, pp. 195–196. In M. J. W. host resources. Journal of Ecology Cock, H. C. J. Godfray, and J. D. Holloways, 78: 686–692. doi:10.1111/j.1365-2656.2008. eds. Slug and Nettle Caterpillars: the Bi- 01516.x ology, Taxonomy, and Control of the Lima- Harvey, J. A., R. Wagenaar, and T. M. Bezemer. codidae of Economic Importance on Palms in 2009b. Life-history traits in closely related South-east Asia. CAB International, Wall- secondary parasitoids sharing the same primary ingford, Oxon, United Kingdom. 270 pp. parasitoid host: evolutionary opportunities and Guimara˜es, J. H. 1980. Revision of the South constraints. Entomologia Experimentalis et American Uramyini (Diptera, Tachinidae). Applicata 132: 155–164. doi:10.1111/j.1570- Pape´is Avulsos de Zoologia 33: 191–219. 7458.2009.00882.x Hall, J. 1981. Bombyliidae, pp. 589–602. In J. F. HDOA. 2010. Natural enemy of stinging cater- McAlpine, B. V. Peterson, G. E. Shewell, H. J. pillar to be released on O’ahu. News Re- Teskey, J. R. Vockeroth, and D. M. Wood, eds. lease10-07. Available from: http://hawaii.gov/ Manual of Nearctic Diptera, Vol. 1. Agricul- hdoa/news/2010-news-releases/natural-enemy- ture Canada Monograph, Research Branch, of-stinging-caterpillar-to-be-released-on-o-ahu. Agriculture Canada, Ottawa, Ontario. [accessed May 15, 2011]. Hansson, C. 1987. New records of Swedish Eu- HDOA. 2011. Kaua’i residents asked to report lophidae and Pteromalidae (Hymenoptera: sightings of stinging caterpillar. News Re- Chalcidoidea), with data on host species. lease11-11. Available from: http://hawaii.gov/ Entomologisk Tidskrift 108: 168–173. hdoa/news/news-releases-2011/kaua-i-residents- Hansson, C. 1994. Re-evaluation of the genus asked-to-report-sightings-of-stinging-caterpillar. Closterocerus Westwood (Hymenoptera: [accessed June 8, 2011]. Eulophidae), with a revision of the Nearctic Heraty, J. and D. Hawks. 1998. Hexamethyldisi- species. Entomologica Scandinavica 25: 1– lazane – a chemical alternative for drying 25. doi:10.1163/187631294X00018 insects. Entomological News 109: 369–374. Hansson, C. 1995. Revision of the Nearctic species Herting, B. 1960. Biologie der westpala¨arktischen of Neochrysocharis Kurdjumov (Hymenoptera: Raupenfliegen. Dipt., Tachinidae. Monographien Eulophidae). Entomologica Scandinavica 26: zur Angewandte Entomologie 16: 1–188. 27–46. Herting, B. 1972. Die Typenexemplare der von Harris, R. 1979. A glossary of surface sculptur- Meigen (1824–1838) beschriebenen Rau- ing. Occasional Papers in Entomology, No. penfliegen (Dipt. Tachinidae). Stuttgarter 28. California State Department of Food and Beitra¨ge zur Naturkunde 243: 1–15. 104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Herting, B. 1976. Lepidoptera, Part 2 (Macro- (Diptera: Tachinidae) in central Virginia, and lepidoptera). A catalogue of parasites and their hyperparasitism by Trigonalid wasps predators of terrestrial arthropods. Section A. (Hymenoptera: Trigonalidae). Environmental Host or Prey/Enemy, 7: 3 Commonwealth Entomology 32: 1019–1027. doi:10.1603/ Agricultural Bureaux, Commonwealth In- 0046-225X-32.5.1019 stitute of Biological Control. Kerr, P. H., E. M. Fisher, and M. L. Buffington. Hoong, H. W. and C. K. Y. Hoh. 1992. Major 2008. Dome lighting for insect imaging under pests of oil palm and their occurrence in Sa- a microscope. American Entomologist 54: bah. Planter 68: 193–210. 198–200. Horstmann, K. 2003. Revisionen von Schlupfwespen- Kits, J. H., S. A. Marshall, and N. L. Evenhuis. Arten VII. Mitteilungen der Mu¨nchner En- 2008. The bee flies (Diptera: Bombyliidae) of tomologischen Gesellschaft 93: 25–37. Ontario, with a key to the species of eastern Horstmann, K. 2008. Neue westpalaarkitsche arten Canada. Canadian Journal of Arthropod der Campopleginae (Hymenoptera: Ichneu- Identification No. 6, March 06, 2008, available monidae). Zeitschrift der Arveitsgemeinschaft online at http://www.biology.ualberta.ca/bsc/ Oesterreichischer Entomologen. 60: 3–27. ejournal/kme_06/kme_06.html, doi: 10.3752/ Horstmann, K. 2009. Typenrevisionen der von cjai.2008.06. [accessed 15-June-2010]. Kiss beschriebenen Taxa der Ichneumonidae Krauth, S. J. and A. H. Williams. 2006. Notes on III. Verschiedene Unterfamilien (Hymenop- Taeniogonalos gundlachii Hymenoptera: tera, Ichneumonidae). Linxer Biologische Trigonalidae) from Wisconsin. Great Lakes Beitra¨ge 41: 673–689. Entomologist 34: 54–58. Howard, L. O. 1889. The hymenopterous para- Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and sites of North American butterflies, pp. 1869– B. D. Burks. 1979. Catalog of Hymenoptera 1896. In S. H. Scudder, ed. Butterflies of in America North of Mexico. Vol. 1. Smith- Eastern United States and Canada, Vol. III. sonian Institution Press, Washington, D.C. W. Wheeler, Cambridge, Massachusetts. xvi + 1198 pp. Hull, F. 1973. Bee Flies of the World. The Genera Kula, R. R., J. T. Lill, S. M. Murphy, and T. of the Family Bombyliidae. Smithsonian In- Stoepler. (2009) 2010. The first host records stitution Press, Washington, DC. 687 pp. for the Nearctic species Triraphis discoideus Ichiki, R. and H. Shima. 2003. Immature life of (Hymenoptera: Braconidae: Rogadinae). En- Compsilura concinnata (Meigen) (Diptera: tomological News 120: 380–386. doi:10.3157/ Tachinidae). Annals of the Entomological 021.120.0406 Society of America 96: 161–167. doi:10. Kulman, H. M. 1965. Natural control of eastern tent 1603/0013-8746(2003)096[0161:ILOCCM] caterpillar and notes on its status as a forest pest. 2.0.CO;2 Journal of Economic Entomology 58: 66–70. Janzen, D. H. and W. Hallwachs. 2009. Dynamic La Salle, J. 1993. Parasitic Hymenoptera, bi- database for an inventory of the macro- ological control, and , pp. 197– caterpillar fauna, and its food plants and para- 215. In J. La Salle and I. D. Gauld, eds. sitoids, of Area de Conservacion Guanacaste Hymenoptera and Biodiversity. CAB In- (ACG), northwestern Costa Rica (nn-SRNP- ternational, Wallingford, UK. 348 pp. nnnnn voucher codes). Available at http://janzen. Le Corff, J. and R. J. Marquis. 1999. Differences sas.upenn.edu. [accessed 15-June-2010]. between undestory and canopy in herbivore Jerman, E. J. and I. Gauld. 1988. Casinaria, community composition and leaf quality for a paraphyletic ichneumonid genus (Hyme- two oak species in Missouri. Ecological En- noptera), and a revision of the Australian tomology 24: 46–58. doi:10.1046/j.1365- species. Journal of Natural History 22: 589– 2311.1999.00174.x 609. doi:10.1080/00222938800770401 Lill, J. T. 2008. Caterpillar-host plant relationships Johnson, C. W. 1925. Fauna of New England. 15. recorded from Plummers Island, Maryland List of the Diptera or two-winged flies. Oc- (Insecta: Lepidoptera). Bulletin of the Bi- casional Papers of the Boston Society of ological Society of Washington 15: 75–79. Natural History 7: 1–326. doi:10.2988/0097-0298(2008)15[75:CPRRFP] Kellogg, S. K., L. S. Fink, and L. P. Brower. 2003. 2.0.CO;2 Parasitism of native luna moths, Actias luna Lill, J. T., R. J. Marquis, R. E. Forkner, J. Le (L.) (Lepdioptera: Saturniidae) by the in- Corff, N. Holmberg, and N. A. Barber. 2006. troduced Compsilura concinnata (Meigen) Leaf pubescence affects distribution and VOLUME 114, NUMBER 1 105

abundance of generalist slug caterpillars Hyphantria cunea. Canadian Entomologist (Lepidoptera: Limacodidae). Environmental 108: 1291–1294. Entomology 35: 797–806. doi:10.1603/0046- Muesebeck, C. F. W. 1920. A revision of the 225X-35.3.797 North American species of ichneumon-flies Lingren, P. D. 1977. Campoletis sonorensis: belonging to the genus Apanteles. Pro- maintenance of a population on tobacco ceedings of the United States National Mu- budworms in a field cage. Environmental seum 58: 483–576. Entomology 6: 72–76. Muesebeck, C. F. W. 1931. Descriptions of a new Lourenca˜o, A. L., L. O. de Carvalho, and D. H. de genus and eight new species of ichenumon- Campos Lasca. 1989. Anacraga citrinopsis flies with taxonomic notes. Proceedings of Dyar (Lepidoptera: Dalceridae) em Mamoneira the United States National Museum 79 (Art. no Estado de Sa˜oPaulo.Bragantia48:109–112. 16) [= No. 2882]: 1–16. Marconato, G., M. M. Dias, and A. M. Penteado Murphy, S. M., J. T. Lill, and D. R. Smith. 2009. Dias. 2008. de Geometridae (Lepi- Ascattershotapproachtohostlocation: doptera) e seus parasito´ides, associadas a uncovering the unique life history of the Erythroxylum microphyllum St.- Hilaire trigonalid hyperparasitoid Orthogonalys pul- (Erythroxylacae). Revista Brasileira de En- chella (Cresson). American Entomologist 55: tomologia 52: 296–299. doi:10.1590/S0085- 82–87. 56262008000200010 Murphy, S. M., S. M. Leahy, L. S. Williams, and Marsh, P. M. 1979. Braconidae, Aphidiidae, J. T. Lill. 2010. Stinging spines protect slug Hybrizontidae, pp. 144–313. In K. V. Krom- caterpillars (Limacodidae) from multiple bein, P. D. Hurd, Jr., D. R. Smith, and B. D. generalist predators. Behavioral Ecology 21: Burks, eds. Catalog of Hymenoptera in 153–160. doi:10.1093/beheco/arp166 America North of Mexico. Vol. 1. Smithsonian Nazemi, J., A. A. Talebi, S. E. Sadeghi, G. Me- Institution Press, Washington, D.C. 1198 pp. lika, and A. Lozan. 2008. Species richness of Marsh, P. M. and S. R. Shaw. 1998. Revision of oak gall wasps (Hymenoptera: Cynipidae) North American Aleiodes Wesmael (Part 3): and identification of associated and the seriatus (Herrich-Schaeffer) species-group parasitoids on two oak species in western (Hymenoptera: Braconidae: Rogadinae). Pro- Iran. North-Western Journal of Zoology 4: ceedings of the Entomological Society of 189–202. Washington 100: 395–408. Niehuis, O., S. H. Yen, C. M. Naumann, and B. McAlpine, J. F. 1981. Morphology and termi- Misof. 2006. Higher phylogeny of zygaenid nology—adults. In J. F. McAlpine, B. V. moths (Insecta: Lepidoptera) inferred from Peterson, G. E. Shewell, H. J. Teskey, J. R. nuclear and mitochondrial sequence data and Vockeroth, and D. M. Wood, eds. Manual of the evolution of larval cuticular cavities for Nearctic Diptera. Volume 1. Agriculture chemical defence. Molecular Phylogenetics Canada Monograph 27: 9–63. and Evolution 39: 812–829. doi:10.1016/j. McCullough, D. G., S. A. Katovich, D. L. Mahr, ympev.2006.01.007 D. D. Neumann, C. S. Sadof, and M. J. Noyes, J. S. 2003. Universal Chalcidoidea Data- Raupp. 1999. Biological control of insect base. World Wide Web electronic publication. pests in forested ecosystems: a manual for www.nhm.ac.uk/entomology/chalcidoids/index. foresters, Christmas tree growers and land- html. [accessed 15-June-2010]. scapers. Michigan State University Extension O’Hara, J. E., H. Shima, and C.-T. Zhang. 2009. Bulletin E-2679: 1–123. Annotated catalogue of the Tachinidae (Insecta: Meigen, J. W. 1824. Systematische Beschreibung Diptera) of China. Zootaxa 2190: 1–236. der bekannten europa¨ischen zweiflu¨geligen O’Hara, J. E. and D. M. Wood. 2004. Catalogue Insekten. Vierter Theil. Schulz-Wundermann, of the Tachinidae (Diptera) of America north Hamm. xii + 428 pp. + pls. 33–41. of Mexico. Memoirs on Entomology, In- Mexzo´n, R. G., C. M. Chinchilla, and D. Sala- ternational 18: 1–410. manca. 1996. The biology of Sibine mega- Ostmark, H. E. 1974. Economic insect pests of somoides Walker (Lepidoptera, Limacodidae): bananas. Annual Review of Entomology 19: observations of the pest in oil palm in Costa 161–176. doi:10.1146/annurev.en.19.010174. Rica. ASD Oil Palm Papers 12: 1–10. 001113 Morris, R. F. 1976. Relation of mortality caused Painter, R. and E. Painter. 1963. A review of the by parasites to the population density of subfamily Systropinae (Diptera: Bombyliidae) 106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

in North America. Journal of the Kansas En- Ichneumonidae, and Tachinidae. Proceedings tomological Society 36: 278–348. of the Royal Society of Victoria 47: 54–77. Papp, J. 2004. Type specimens of the braconid Raizenne, H. 1952. Forest Lepidoptera of south- species by Gy. Sze´pligeti deposited in the ern Ontario and their parasites received and Hungarian Natural History Museum (Hy- reared at the Ottawa Forest Insect Survey menoptera: Braconidae). Annals Historico- Laboratory from 1937 to 1948. Canada De- Naturales Musei Nationalis Hungarici 96: partment of Agriculture, Science Service, 153–223. Division of Forest Biology, Ottawa. v + 277 pp. Patton, C. N. 1958. A catalogue of the Larvae- Reinhard, H. 1865. Beitra¨ge zur Kenntnifs einiger voridae of Florida. Florida Entomologist 41: Braconiden-Gattungen. Berliner Entomologische 29–39, 77–89. doi:10.2307/3492632 Zeitschrift 9: 243–267. doi:10.1002/mmnd. Peck, O. 1963. A catalogue of the Nearctic 18650090309 Chalcidoidea (Insecta; Hymenoptera). Cana- Reinhard, H. J. 1935. New genera and species of dian Entomologist (Supplement) 30: 1–1420. American muscoid flies (Tachinidae: Dip- Peck, O. 1985. The taxonomy of the Nearctic tera). Annals of the Entomological Society of species of Pediobius (Hymenoptera: Eulo- America 28: 160–173. phidae), especially Canadian and Alaskan dos Reis Fernandes, L. B., M. M. Dias Filho, M. forms. Canadian Entomologist 117: 647–704. A. Fernandes, and A. M. Penteado-Dias. doi:10.4039/Ent117647-6 2010. Ichneumonidae (Hymenoptera) para- Petrice, T. R., J. S. Strazanac, and L. Butler. 2004. sitoids of Lepidoptera caterpillars feeding on A survey of hymenopteran parasitoids of Croton floribundus Spreng (Euphorbiaceae). forest macrolepidoptera in the central Appa- Revista Brasileira de Entomologia 54: 263– lachians. Journal of Economic Entomology 269. doi:10.1590/S0085-56262010000200009 97: 451–459. doi:10.1603/0022-0493-97.2.451 Riley, C. V. and L. O. Howard. 1890. Some of the Pinto, J. 1998. Systematics of the North Ameri- bred parasitic Hymenoptera in the National can species of Trichogramma Westwood Collection. Insect Life 2: 348–353. (Hymenoptera: Trichogrammatidae). Mem- Robertson, C. 1928. Flowers and Insects. Lists of oirs of the Entomological Society of Wash- Visitors of 453 Flowers. Science Press ington 22: 1–287. Printing Co., Lancaster, PA. 221 pp. Pinto, J. D., G. R. Platner, and E. R. Oatman. ROBO (Release of Beneficial ) Data- 1978. Clarification of the identity of several base. Introduced Parasites and Predators Re- common species of North American Tri- leased in the U. S. and Territories - by Target chogramma (Hymenoptera: Trichogramma- Host/Prey, 1981–1985. Biological Control tidae). Annals of the Entomological Society Documentation Center, National Invertebrate of America 71: 169–180. Genetic Resources, http://www.ars-grin.gov/ Pluciennik, Z. and R. W. Olszak. 2010. The role nigrp/menu.html#link1981. [accessed 5-June- of parasitoids in limiting the harmfulness of 2010]. leafrollers in apple orchards. Journal of Plant Rohwer, S. 1915. Descriptions of new species Protection Research 50: 1–8. doi:10.2478/ of Hymenoptera. Proceedings of the United v10045-010-0001-9 States National Museum 49: 205–249. doi: Provancher, L. 1875. Les Ichneumonides de 10.5479/si.00963801.2105.205 Que´bec. Naturaliste Canadien 7: 309–317. Ruberson, J. R. and J. B. Whitfield. 1996. Fac- Quicke, D. L. J. 1997. Parasitic Wasps. Chapman ultative egg-larval parasitism of the beet ar- and Hall, New York. 470 pp. myworm, Spodoptera exigua (Lepidoptera: Quicke, D. L. J. and M. R. Shaw. 2005. First host Noctuidae) by Cotesia marginiventris (Hy- records for the rogadine genera Rogasodes menoptera: Braconidae). Florida Entomolo- Chen and He and Canalirogas van Achter- gist 79: 296–302. doi:10.2307/3495577 berg and Chen (Hymenoptera: Braconidae) Sabrosky, C. W. and P. H. Arnaud, Jr. 1965. Family with description of a new species and survey Tachinidae (Larvaevoridae), pp. 961–1108. In of mummy types within Rogadinae s. str. A. Stone, C. W. Sabrosky, W. W. Wirth, R. H. Journal of Natural History 39: 3525–3542. Foote, and J. R. Coulson, eds. A Catalog of the doi:10.1080/00222930500392782 Diptera of America North of Mexico. United Raff, J. W. 1934. Observations on sawflies of the States Department of Agriculture. Agriculture genus Perga, with notes on some reared pri- Handbook 276. U.S. Government Printing mary parsties of the families Trigonalidae, Office, Washington D.C. 1696 pp. VOLUME 114, NUMBER 1 107

Say, T. 1835. Descriptions of new North Ameri- Shaw, S. R. 1997a. Subfamily Cheloninae, pp. can Hymenoptera, and observations on some 193–201. In R. A. Wharton, P. M. Marsh, already described. Boston Journal of Natural and M. J. Sharkey, eds. Manual of the New History 1(3): 210–305. World Genera of the Family Braconidae Schaffner, J. V., Jr. 1959. and (Hymenoptera). Special Publication No. 1. their parasites reared from field collections in International Society of Hymenopterists, the northeastern part of the United States. Washington, D. C. 439 pp. United States Department Agriculture Mis- Shaw, S. R. 1997b. Subfamily Rogadinae s.s, pp. cellaneous Publications 767: 1–97. 403–412. In R. A. Wharton, P. M. Marsh, Schaffner, J. V., Jr. and F. L. Griswold. 1934. and M. J. Sharkey, eds. Manual of the New Macrolepidoptera and their parasites reared World Genera of the Family Braconidae from field collections in the northeastern part (Hymenoptera). Special Publication No. 1. of the United States. United States De- International Society of Hymenopterists, partment Agriculture Miscellaneous Publi- Washington, D. C. 439 pp. cations 188: 1–160. Shaw, S. R. 2006. Aleiodes wasps of eastern Schauff, M. E., J. LaSalle, and L. D. Coote. 1997. forests: a guide to parasitoids and associated Eulophidae, pp. 327–429. In G. A. P. Gibson, mummified caterpillars. Forest Health Tech- J. T. Huber, and J. B. Woolley, eds. Annotated nology Enterprise Team/USDA/Forest Ser- Keys to the Genera of Nearctic Chalcidoidea vice. Morgantown, West Virginia.121 pp. (Hymenoptera). Research Press. National Shaw, S. R., P. M. Marsh, and J. C. Fortier. 1997. Research Council of Canada, Ottawa, On- Revision of North American Aleiodes Wes- tario. 764 pp. mael (Part 1): the pulchripes Wesmael spe- Sellers, W. F. 1943. The Nearctic species of cies-group in the New World (Hymenoptera: parasitic flies belonging to Zenillia and Braconidae, Rogadinae). Journal of Hyme- allied genera. Proceedings of the United noptera Research 6: 10–35. States National Museum 93 [= No. 3157]: Shaw, S. R., P. M. Marsh, and J. C. Fortier. 1998. 1–108. Revision of North American Aleiodes Wes- Sharkey, M. J. 1997. Key to New World sub- mael (Part 4): the albitibia Herrich-Schaeffer families of the family Braconidae, pp. 39–45. and praetor Reinhard species-groups (Hy- In R. A. Wharton, P. M. Marsh, and M. J. menoptera: Braconidae: Rogadinae) in the Sharkey, eds. Manual of the New World New World. Proceedings of the Entomolog- Genera of the Family Braconidae (Hymenop- ical Society of Washington 100: 553–565. tera). Special Publication No. 1. International Shima, H. 1984. Study on the tribe Blondeliini Society of Hymenopterists, Washington, D. C. from Japan (Diptera, Tachinidae). V. The Sharkey, M. J. and R. A. Wharton. 1997. Mor- genera Blondelia Robineau-Desvoidy and phology and terminology, pp. 19–37. In R. A. Compsilura Bouche´. Kontyuˆ 52: 540–552. Wharton, P. M. Marsh, and M. J. Sharkey, eds. Shima, H. 2006. A host-parasite catalog of Manual of the New World Genera of the Tachinidae (Diptera) of Japan. Makunagi/Acta Family Braconidae (Hymenoptera). Interna- Dipterologica, Supplement 2: 1–171. tional Society of Hymenopterists, Special Short, J. R. T. 1978. The final larval instars of the Publication No. 1. Washington, D. C. 439 pp. Ichneumonidae. Memoirs of the American Shaw, M. R. 1990. Parasitoids of European but- Entomological Institute 25: 1–508. terflies and their study, pp. 449–479. In O. Singh, B. P., R. P. Singh, and V. D. Verma. 1988. Kudrna, ed. Butterflies of Europe, Vol. 2. New record of Aroplectrus dimerus Linn. and Aula-Verlag, Wiesbaden. 557 pp. Platyplectrus sp. as larval parasite of slug Shaw, M. R. and R. R. Askew. 1976. Parasites, caterpillar (Parasa bicolor Walk) from U. P. pp. 240–256. In J. Heath, ed. The Moths and Farm Science Journal 3: 199–200. Butterflies of Great Britain and Ireland. 1. Smith, D. R. 1996. Trigonalyidae (Hymenoptera) – . Blackwell in the eastern United States: seasonal flight Scientific Publications and Curwen Press, activity, distribution, hosts. Proceedings of London. 343 pp. the Entomological Society of Washington 98: Shaw, S. R. 1983. A taxonomic study of the Ne- 109–118. arctic Ascogaster and a description of a new Smith, D. R. and I. C. Stocks. 2005. A new trig- genus Leptodrepana (Hymenoptera: Braco- onalid wasp (Hymenoptera: Trigonalidae) nidae). Entomography 2: 1–54. from eastern North America. Proceedings of 108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

the Entomological Society of Washington Townes, H. K. 1956. The Nearctic species of 107: 530–535. trigonalid wasps. Proceedings of the United Smith, H. E. 1912. A contribution on North Amer- States National Museum 106: 295–304. ican dipterology. Proceedings of the Entomo- Townes, H. K. 1970. The genera of Ichneumo- logical Society of Washington 14: 118–127. nidae, part 2. Memoirs of the American En- Smith, O. J., A. G. Diboll, and H. Rosenberger. tomological Institute 12: 1–537. 1955. Laboratory studies of Pelecystoma Townes, H. K. 1983. Revisions of twenty genera harrisinae (Ashmead), an adventive braconid of Gelini (Ichneumonidae). Memoirs of the parasite of the western grape leaf skeleton- American Entomological Institute 35: 1– izer. Annals of the Entomological Society of 281. America 48: 232–237. Townes, H. and M. Townes. 1962. Ichneumon- Smith, S. M. 1996. Biological control with Tri- flies of America North of Mexico: 3. Sub- chogramma:advances,successes,andpotential family , tribe Mesostenini. United of their use. Annual Review of Entomology States National Museum Bulletin 216: 1– 41: 375–406. doi:10.1146/annurev.en.41.010196. 602. doi:10.5479/si.03629236.216.1-3 002111 Townes, H. and M. Townes. 1966. A catalogue Stireman III, J. O. and M. S. Singer. 2003a. De- and reclassification of the Neotropic Ichneu- terminants of parasitoid-host associations: in- monidae. Memoirs of the American Ento- sights from a natural tachinid-lepidopteran mological Institute 8: 1–367. community. Ecology 84: 296–310. doi:10.1890/ Townes, H., S. Momoi, and M. Townes. 1965. A 0012-9658(2003)084[0296:DOPHAI]2.0.CO;2 catalogue and reclassification of the eastern Stireman III, J. O. and M. S. Singer. 2003b. What Palearctic Ichneumonidae. Memoirs of the determines host range in parasitoids? An American Entomological Institute 5: i–v + 1– analysis of a tachinid parasitoid community. 661. Oecologia 135: 629–638. Townsend, C. H. T. 1892. An Aporia bred from Strand, M. R. 1986. The physiological in- Limacodes sp. Psyche 6: 275–276. doi:10.1155/ teractions of parasitoids with their hosts and 1892/17351 their influence on reproductive strategies, pp. Townsend, C. H. T. (1918) 1919. New muscoid 97–136. In J. K. Waage and D. J. Greathead, genera, species and synonymy (Diptera). eds. Insect Parasitoids. Academic Press, [concl.] Insecutor Inscitiae Menstruus 6: London, UK. 389 pp. 157–182. Strazanac, J. S., C. D. Plaugher, T. R. Petrice, and L. Tschorsnig, H.-P. and V. A. Richter. 1998. Family Butler. 2001. New Tachinidae (Diptera) host Tachinidae, pp. 691–827. In L. Papp and B. records of eastern North American forest can- Darvas, eds. Contributions to a Manual of opy Lepidoptera: baseline data in a Bacillus Palaearctic Diptera (With Special Reference to thuriengiensis variety kurstaki nontarget study. Flies of Economic Importance), Vol. 3. Higher Journal of Economic Entomology 94: 1128– Brachycera. Science Herald, Budapest. 880 pp. 1134. doi:10.1603/0022-0493-94.5.1128 Valerio, A. A. 2006. Some taxonomic notes on Thompson, W. R. 1954. A catalogue of the par- named Rogas Nees species (Hymenoptera: asites and predators of insect pests. Section 2. Braconidae: Rogadinae) for the New World. Host parasite catalogue. Part 3. Hosts of the Methods in Ecology and Systematics 1: 37–43. Hymenoptera (Calliceratid to Evaniid), pp. Viereck, H. L. 1913. Descriptions of ten new 191–332. Commonwealth Agricultural Bu- genera and twenty-three new species of ich- reaux, Commonwealth Institute of Biological neumon-flies. Proceedings of the United Control, Ottawa. 141 pp. States National Museum 44: 555–568. Thomson, C. G. 1878. Hymenoptera Scandina- doi:10.5479/si.00963801.1968.555 viae 5. Pteromalus (Svederus) continuatio. Viereck, H. L. 1925. A preliminary revision of the Typis Expressit Haqv. Ohlsson, Lund. 307 pp. Campopleginae in the Canadian National Tobias, V. I. 1976. Braconids of Caucasus. Collection, Ottawa. Canadian Entomologist Nauka, Leningrad. 287 pp. [In Russian] 57: 176–181, 198–204, 223–228, 296–303. Tothill, J. D. 1922. The natural control of the fall doi:10.4039/Ent57176-7 webworm (Hyphantria cunea Drury) in Can- Viereck, H. L. 1926. A preliminary revision of the ada together with an account of its several Campopleginae in the Canadian National parasites. Bulletin of the Canada Department Collection, Ottawa. Canadian Entomologist of Agriculture, N. Ser. 3: 1–107 + 6 plates. 58: 2–8, 30–38, 71–78, 122–130, 143–149, VOLUME 114, NUMBER 1 109

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

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

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