Robert R. Kula

Potential exposure of a classical biological control agent of the soybean aphid, Aphis glycines, on non-target aphids in North America

In summer 2007, the Asian parasitoid Binodoxys communis (Hymenoptera: Braconidae) was released in North America for control of the exotic soybean aphid, Aphis glycines (Hemiptera: Aphididae). Despite its comparatively narrow host range, releases of B. communis may still constitute a risk to native aphid species. To estimate the risk of exposure of non-target aphids to B. communis, we merged assessments of temporal co-occurrence with projections of spatial overlap between B. communis and three native aphid species, and in-field measurements of the incidence of ecological filters that may protect these aphids from parasitism. Temporal co-occurrence was assessed between A. glycines and native aphids (Aphis asclepiadis, Aphis oestlundi, and Aphis monardae) at four different locations in Minnesota, USA. The degree of temporal overlap depended greatly on location and aphid species, ranging between 0 and 100%. All of the native aphids were tended by multiple species of ants, with overall ant-attendance ranging from 26.1 to 89.6%. During temporal overlap with A. glycines, 53 ± 11% of A. monardae colonies were partly found in flower heads of their host plant, with flowers acting as a physical refuge for this aphid. The extent of geographic overlap between B. communis and native aphids based upon Climex modeling was 17–28% for A. monardae, 13–22% for A. oestlundi, 46–55% for A. asclepiadis and 12–24% for the A. asclepiadis species complex. The estimated overall probability of potential exposure of B. communis on native aphids was relatively low (P = 0.115) for A. oestlundi and high (P = 0.550) for A. asclepiades. Physical and ant-mediated refuges considerably lowered probability of population-level impact on A. monardae, and could lead to substantial reduction of exposure for the other native aphids. These findings are used to make broader statements regarding the ecological safety of current B. communis releases and their potential impact on native aphid species in North America.

Leluthia astigma (Ashmead) (Hymenoptera: Braconidae: Doryctinae) as a Parasitoid of Agrilus planipennis Fairmaire (Coleoptera: Buprestidae: Agrilinae), with an Assessment of Host Associations for Nearctic Species of Leluthia Cameron

Abstract Published host associations are assessed for Leluthia astigma (Ashmead), Leluthia floridensis Marsh, and Leluthia mexicana Cameron, the three known species of Leluthia Cameron in the Nearctic Region. Leluthia astigma is reported as a parasitoid of Agrilus planipennis Fairmaire, emerald ash borer (EAB), infesting Fraxinus americana L., white ash, in Delaware County, Ohio. It is the first species of Leluthia for which a determined species of Agrilus Curtis has been confirmed as a host and the association vouchered. All other hosts reported in the literature for L. astigma require confirmation through rearing from an isolated host or documentation of an unequivocal host-parasitoid association. Leluthia astigma adults reared from parasitoid cocoons collected in Delaware County, Ohio parasitized and produced F1 adults in the laboratory on last-instar EAB larvae collected in Ingham County, Michigan. Parasitism of EAB and other natural history data are reported for L. astigma, including the first records of L. astigma from Kansas and New York.

Hymenopteran Parasitoids Attacking the Invasive Emerald Ash Borer (Coleoptera: Buprestidae) in Western and Central Pennsylvania

ABSTRACT We conducted field surveys of the emerald ash borer (EAB), Agrilus planipennis Fairmaire, and associated larval parasitoids in western and central Pennsylvania (Cranberry Township in Butler County and Granville in Mifflin County) in the spring and fall of 2009. The survey procedure involved destructively debarking sections of the main trunk (bole) of EAB-infested green ash (Fraxinus pennsylvanica Marsh.) trees from the ground to the height of 2 m. Three species of the hymenopteran parasitoids were consistently recovered from EAB larvae observed in both survey sites, including two indigenous species of braconids, Spathius laflammei Provancher (= Spathius benefactor Matthews) and Atanycolus nigropyga Shenefelt and the exotic (accidentally introduced) eupelmid Balcha indica (Mani & Kaul). In addition, there are three unidentified species of hymenopteran parasitoids including two braconids Atanycolus sp. [possibly Atanycolus disputabilis (Cresson)] and Spathius sp. (at the Butler Co. site) and one ichneumonid Dolichomitus sp. (at the Mifflin Co. site). These parasitoids together parasitized 0.5–4.6% and 0.5–1.5% of the sampled EAB hosts at the Butler and Mifflin Co. sites, respectively. Parasitism rate by each species or group of those hymenopteran parasitoids varied between the two survey sites—with parasitism rates being generally higher at the Butler Co. site than at the Mifflin Co. site. Studies are needed to determine if those new associations of North American indigenous braconid parasitoids with EAB may play a complementary role in controlling this invasive pest.

Revision of New World Chaenusa Haliday sensu lato (Hymenoptera: Braconidae: Alysiinae), with New Species, Synonymies, Hosts, and Distribution Records

Abstract The New World species of Chaenusa Haliday sensu lato are revised, and a diagnosis is provided for Chaenusa s.l. Four new species from North America, Chaenusa trumani Kula, Chaenusa virgili Kula, Chaenusa whartoni Kula, and Chaenusa woolleyi Kula, and two new species from South America, Chaenusa hirsutissima Kula and Chaenusa ireneae Kula, are described. Chaenusa americana (Riegel), new combination and Chaenusa saxicola (Riegel), new combination are transferred from Chorebidea Viereck. Chaenusa bergi (Riegel), revised combination is returned from Chorebidella Riegel. Chorebidea bessae Riegel is a new synonym of Cha. americana, and Chorebidea mcclurei Riegel is a new synonym of Cha. saxicola. Chaenusa anticostae Riegel and Chaenusa illinae Riegel are new synonyms of Chaenusa quadriceps (Ashmead). Chaenusa americana, Cha. bergi, Chaenusa californica Riegel, Cha. quadriceps, Chaenusa rossi Riegel, and Cha. saxicola are redescribed. Host and spatiotemporal distribution data are provided, including new host and distribution records. The status of several holotypes is discussed, and a key to the New World species of Chaenusa s.l. is provided.

Review of Parasitoid Wasps and Flies (Hymenoptera, Diptera) Associated with Limacodidae (Lepidoptera) in North America, with a Key to Genera

Abstract. Hymenopteran and dipteran parasitoids of slug moth caterpillars (Lepidoptera: Limacodidae) from North America are reviewed, and an illustrated key to 23 genera is presented. Limacodid surveys and rearing were conducted during the summer months of 2004–2009 as part of research on the ecology and natural history of Limacodidae in the mid-Atlantic region of the U.S.A. Parasitoid rearing involved a combination of collecting naturally occurring larvae in the field (at least 14 host species) and placing out large numbers of “sentinel” larvae derived from laboratory colonies of three host species. Species in the following families are documented from limacodids in North America as primary or secondary parasitoids (number of genera for each family in parentheses; number of genera included in key but not reared through this research in brackets): Chalcididae ([1]; Hymenoptera: Chalcidoidea), Eulophidae (3; Chalcidoidea), Pteromalidae ([1]; Chalcidoidea), Trichogrammatidae (1; Chalcidoidea), Braconidae (3 [1]; Hymenoptera: Ichneumonoidea), Ichneumonidae (7 [3]; Ichneumonoidea), Ceraphronidae (1; Hymenoptera: Ceraphronoidea), Trigonalidae (2; Hymenoptera: Trigonaloidea), Bombyliidae ([1]; Diptera: Asilioidea), and Tachinidae (3; Oestroidea). We recovered 20 of 28 genera known to attack limacodids in North America. Records discerned through rearing in the mid-Atlantic region are augmented with previously published host-parasitoid relationships for Limacodidae in North America north of Mexico. New records are reported for the following parasitoids (total new records in parentheses): Uramya limacodis (Walker) (1), U. pristis (Townsend) (5), Austrophorocera spp. (6), Ceraphron sp. (1), Alveoplectrus lilli Gates (1), Playplectrus americana (Girault) (10), Pediobius crassicornis (Thomson) (1), Trichogramma (1), Mesochorus discitergus (Say) (1), Hyposoter fugitivus (Say) (1), and Isdromas lycaenae (Howard) (5). The male of Platyplectrus americana (Hymenoptera: Eulophidae) is redescribed, and the female is described for the first time. Incidental and miscellaneous host-parasitoid associations are discussed, and it is concluded that most of these records are likely parasitoids of contaminants accidentally introduced during the limacodid rearing process. Triraphis eupoeyiae (Ashmead), new combination, is transferred from Rogas (Hymenoptera: Braconidae).

Revision of the genus Pseudapanteles (Hymenoptera, Braconidae, Microgastrinae), with emphasis on the species in Area de Conservación Guanacaste, northwestern Costa Rica.

Abstract Pseudapanteles is a moderately diverse genus of Microgastrinae parasitoid wasps (Hymenoptera: Braconidae), endemic to the New World and with the vast majority of its species (including many undescribed) in the Neotropical region. We describe here 25 new species from Area de Conservación Guanacaste (ACG), northwestern Costa Rica, based on 400 studied specimens. A key to all 36 known species of Pseudapanteles is provided (except for Pseudapanteles brunneus, only known from a single male), and species are placed in three newly created species-groups. Host records are known for only 25% of the species; most are solitary parasitoids of the caterpillars of several families of small Lepidoptera (Crambidae, Elachistidae, Gelechiidae, Incurvariidae, Sesiidae, Tineidae). DNA barcodes (part of the CO1 gene) were obtained for 30 species (83%), and provide a start for future study of the genus beyond ACG. Brief descriptions (generated by Lucid 3.5 software) and extensive illustrations are provided for all species. The following new taxonomic and nomenclatural acts are proposed: Pseudapanteles moerens (Nixon, 1965), comb. n., Pseudapanteles brunneus Ashmead, 1900, comb. rev., a lectotype is designated for Pseudapanteles ruficollis (Cameron, 1911), and the following 25 species nova of Pseudapanteles (all authored by Fernández-Triana and Whitfield): alfiopivai, alvaroumanai, analorenaguevarae, carlosespinachi, carlosrodriguezi, christianafigueresae, hernanbravoi, jorgerodriguezi, josefigueresi, laurachinchillae, luisguillermosolisi, margaritapenonae, mariobozai, mariocarvajali, maureenballesteroae, munifigueresae, oscarariasi, ottonsolisi, pedroleoni, raulsolorzanoi, renecastroi, rodrigogamezi, rosemarykarpinskiae, soniapicadoae, teofilodelatorrei.

Phylogenomic Analysis of Ants, Bees and Stinging Wasps: Improved Taxon Sampling Enhances Understanding of Hymenopteran Evolution

The importance of taxon sampling in phylogenetic accuracy is a topic of active debate. We investigated the role of taxon sampling in causing incongruent results between two recent phylogenomic studies of stinging wasps (Hymenoptera: Aculeata), a diverse lineage that includes ants, bees and the majority of eusocial insects. Using target enrichment of ultraconserved element (UCE) loci, we assembled the largest aculeate phylogenomic data set to date, sampling 854 loci from 187 taxa, including 30 out of 31 aculeate families, and a diversity of parasitoid outgroups. We analyzed the complete matrix using multiple analytical approaches, and also performed a series of taxon inclusion/exclusion experiments, in which we analyzed taxon sets identical to and slightly modified from the previous phylogenomic studies. Our results provide a highly supported phylogeny for virtually all aculeate lineages sampled, supporting ants as sister to Apoidea (bees+apoid wasps), bees as sister to Philanthinae+Pemphredoninae (lineages within a paraphyletic Crabronidae), Melittidae as sister to remaining bees, and paraphyly of cuckoo wasps (Chrysidoidea). Our divergence dating analyses estimate ages for aculeate lineages in close concordance with the fossil record. Our analyses also demonstrate that outgroup choice and taxon evenness can fundamentally impact topology and clade support in phylogenomic inference.

DORYCTINAE (HYMENOPTERA: BRACONIDAE) OF KONZA PRAIRIE EXCLUDING SPECIES OF HETEROSPILUS HALIDAY

Abstract. The results of a survey of Doryctinae (Hymenoptera: Braconidae), excluding species of Heterospilus Haliday, at Konza Prairie near Manhattan, Kansas are reported. Eleven sites representing prairie and woodland/wetland areas, including gallery forest, were sampled in 2001 and 2005 using Malaise and canopy traps. Topographic trap placement included lowland, midslope, and upland areas. Twentyfour species were collected, including 18 in 2001 and 16 in 2005. Twenty-three of the 24 species were collected from the woodland/wetland sites, including 19 from gallery forest sites; five of the 24 species were collected from the prairie sites. Rhaconotus fasciatus (Ashmead) was the most abundant species in 2001 (n = 13); Callihormius bifasciatus (Ashmead) and Coiba jeffersoni Kula were the most abundant species in 2005 (n = 10). The following new species are described: Doryctes xanthogaster Kula and Marsh, Doryctes xanthosoma Kula and Marsh, Doryctinus zolnerowichi Kula and Marsh, and Pambolidea dollari Kula and Marsh. Glyptocolastes caryae (Ashmead), status revised is removed from synonymy with Glyptocolastes rugulosus (Cresson), the type species for Doryctinus Roman. The latter species is transferred to Acrophasmus Enderlein, resulting in Acrophasmus as a new synonym of Doryctinus. Synonymy of the aforementioned genera results in the following nomenclatural changes: Doryctinus amazonicus (Roman), new combination; Doryctinus arizonensis (Marsh), new combination; Doryctinus atriventris (Cresson), new combination; Doryctinus butleri (Marsh), new combination; Doryctinus costaricensis (Marsh), new combination; Doryctinus erugatus (Marsh), new combination; Doryctinus exilis (Enderlein), new combination; Doryctinus ferrugineus (Marsh), new combination; Doryctinus gauldi (Marsh), new combination; Doryctinus immigrans (Beardsley), new combination; Doryctinus maeandrius (Enderlein), new combination; Doryctinus marshi Greenbaum, revised combination; Doryctinus rubronotum (Marsh), new combination; Doryctinus rugulosus (Cresson), revised combination; Doryctinus scobiciae (Marsh), new combination; and Doryctinus secundus (Muesebeck and Walkley), new combination. Doryctes infuscus Marsh is a new synonym of Doryctes rufipes (Provancher), Pioscelus wichitus (Viereck) is a new synonym of Pioscelus borealis (Ashmead), and Rhaconotus graciliformis (Viereck) is a new synonym of R. fasciatus. The following species are first records for Kansas: C. bifasciatus, Dendrosoter sulcatus Muesebeck, D. rufipes, D. ferrugineus, Ecphylus hypothenemi Ashmead, Ecphylus rohweri Muesebeck, Ontsira mellipes (Ashmead), and Rhaconotus canadensis Marsh.

First Host Records for the Nearctic Species Triraphis discoideus (Hymenoptera: Braconidae: Rogadinae)

ABSTRACT: Limacodid larvae were collected from 2004 – 2007 on leaves of the following host plants in the District of Columbia and Maryland: Carya glabra, pignut hickory; Quercus alba, white oak; Quercus rubra, northern red oak; Nyssa sylvatica, black gum; Prunus serotina, black cherry; and Fagus grandifolia, American beech. Field-collected larvae were brought to the laboratory where they were kept in isolation for the purpose of rearing parasitoids. The rogadine braconid Triraphis discoideus (Cresson) was reared from the following host-host plant combinations: Acharia stimulea (Clemens), Adoneta spinuloides (Herrich-Schäffer), Lithacodes fasciola (Herrich-Schäffer), and Parasa chloris (Moore) on red oak; Euclea delphinii (Boisduval) on black gum and red oak; Isa textula (Herrich-Schäffer) on American beech, red oak, and white oak; Natada nasoni (Grote) on American beech, black gum, red oak, and white oak; Prolimacodes badia Hübner on black cherry and red oak; and an undetermined species of Tortricidia Packard on American beech. Host use was previously unknown for T. discoideus. Host use and phenology are discussed for Triraphis harrisinae (Ashmead), the only other described species of Triraphis in the Nearctic Region.

Parasitoid Wasps (Hymenoptera: Apocrita) Associated with Sagittaria latifolia Willd. and Sagittaria platyphylla (Engelm.) J. G. sm. (Alismatales: Alismataceae) in the Nearctic Region

Abstract. The results of a survey of parasitoid wasps associated with herbivorous insects on Sagittaria latifolia Willd. and Sagittaria platyphylla (Engelm.) J. G. Sm. (Alismatales: Alismataceae) in the Nearctic Region are reported. The following 10 wasp species were reared from insects on S. platyphylla: Gonatocerus sp., Mestocharis tropicalis Yoshimoto, Syntomopus americanus Ashmead (all Chalcidoidea), Hexacola websteri (Crawford), Rhoptromeris sp. (both Cynipoidea), Alabagrus texanus (Cresson), Bracon sp. nr. mellitor Say, Cotesia diacrisiae (Gahan), Enicospilus glabratus (Say), and Opius (Tolbia) hoffmanni (Fischer) (all Ichneumonoidea). Three wasp species, S. americanus, Chaenusa psillosae Kula (Ichneumonoidea), and B. sp. nr. mellitor, were reared from insects on S. latifolia. All species except C. psillosae are reported as associated with a species of Sagittaria for the first time. Mesochorus discitergus Say is reported as a hyperparasitoid of C. diacrisiae for the first time and is also reported as associated with a species of Sagittaria for the first time. Opius (Tolbia) hoffmanni is reported as a parasitoid of Hydrellia sp. prob. nobilis (Loew); this is the first record of O. (T.) hoffmanni from a species of Hydrellia Robineau-Desvoidy (Diptera: Ephydridae). Mestocharis tropicalis, S. americanus, and O. (T.) hoffmanni are reported from Louisiana for the first time. Syntomopus americanus and O. (T.) hoffmanni are reported from Texas for the first time.