Shaw: Studies of Parasitic Wasps () In Grand Teton National P

STUDIES OF PARASITIC WASPS (BRACONIDAE) IN GRAND TETON NATIONAL PARK +

SCOIT R. SHAW DEPARTMENT OF RENEWABLE RESOURCES UNIVERSITY OF WYOMING + LARAMIE,

+ ABSTRACT + METHODS.

A survey of braconid wasps in GTNP and Sampling of parasitic wasps was conducted south YNP recognizes 71 genera and at least 150 in Grand Teton National Park (GTNP) from 28 July species. Of these, 100 are new distribution records to 15 August 2002. Four Townes-style Malaise and at least nine are new (undescribed) species. The (flight intercept) traps were placed in forested areas species richness of Braconidae in GYE (150 species) near the AMK research station. Trap jars were is substantially greater than the maximum species changed daily, samples sorted, and all Braconidae richness of all herbaceous plants, shrubs, trees, birds specimens were preserved in 95% ethanol. and mammals in lodgepole pine communities of YNP Subsequently, most of these samples were (112 species). This total also substantially exceeds chemically-dried, point-mounted on pins, and labeled the number of braconid species recorded in fire­ for taxonomic analysis. Other sampling methods disturbed habitats of south YNP (109 species). included yellow pan traps, black light, and by-hand collecting with nets and aspirators. Because of the difficulty of placing and maintaining Malaise traps in + INTRODUCTION remote areas, sampling in south Yellowstone National Park (YNP) was restricted to by-hand Braconid wasps (: Braconidae) collecting with nets and aspirators from 30 July to 9 are one of the key groups of beneficial organisms that August. naturally regulate other populations (Shaw & Huddleston 1991 ). Prior to our YNP studies done in 1990 (Lockwood, Shaw & Struttmann 1996), there + RESULTS had been no systematic surveys to assess what braconid species were present in the Greater Thus far, 1,021 braconid specimens have Yellowstone Ecosystem (GYE). No studies have been dry-mounted for study. This represents the bulk been done to assess how GYE braconid faunas of the Malaise trap material, but some smaller change over time. This study is targeted towards samples are still stored in alcohol. From these providing a survey of which braconid species are samples 19 subfamilies of Braconidae are present, and better understanding their roles in the represented: Doryctinae, , Rhyssalinae, GYE. Since braconid wasps affect the population Hormiinae, Rogadinae, Opiinae, Alysiinae, dynamics of many plant-feeding , this research Homolobinae, , Meteorinae, relates to the larger NPS goal of examining rangeland Neoneurinae, Blacinae, Helconinae, Orgilinae, restoration after the 1988 fires. Agathidinae, Aphidiinae, Adeliinae, Cheloninae, and Microgastrinae. Six of these subfamilies (Rhyssalinae, Hormiinae, Homolobinae,

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Neoneurinae, Agathidinae, and Adeliinae) are species are important parasites of bark beetles recorded from GTNP for the first time. Only one (Scolytidae) in western forests (Mason 1978). subfamily expected to occur was not sampled, the Coeloides rufovariegatus (Provancher) has been Ichneutinae. For each of these groups, representative previously recorded from GTNP and Coeloides genera and (if possible) species were determined vancouverensis (D.T.) has been found in YNP and using the identification manual provided by Wharton the Shoshone National Forest (Mason 1978) but et al. (1997) and other sources (cited below). A full neither species was found in this study. listing of species is not feasible because of the lack of modem taxonomic treatments for many groups. Subfamily Rhyssalinae Results for each subfamily group follow below. These minute wasps are gregarious Subfamily Doryctinae ectoparasitoids of microlepidoptera and Coleoptera larvae concealed in plant tissue (Muesebeck 1935). Doryctine species are ectoparasites of wood­ A single female specimen of Oncophanes species boring beetle larvae, especially attacking the beetle near californicus (Ashmead) was collected in GTNP families Bostrichidae, Buprestidae, Cerambycidae, on August 9, 2002. This is the first record for this Lyctidae, and Scolytidae. Consequently, they are subfamily in GTNP. A single female specimen of the normally found in association with dead or damaged same species was collected in YNP on September 7, trees, or firewood. Only five doryctine specimens 1990. Oncophanes californicus was previously were sampled, representing three genera and species known only from California, Oregon, and (all new records for GTNP): Dendrosoter scaber Washington and its host is unknown (Marsh 1979). Muesebeck, Onsira imperator (Haliday), and A closely related species in the eastern United States Spathius species near sequoiae (Ashmead). is a parasite of leaf-mining sawflies in birch Dendrosoter scaber was previously known from (Muesebeck 1935; Marsh 1979). California, Oregon, and Montana (Marsh 1979) but not Wyoming. It is recorded as a parasite of bark Subfamily Hormiinae beetles in Pseudotsuga taxifolia and Pinus monophylla (Muesebeck 1938). Onsira imperator is These minute and poorly studied wasps are widely distributed in Europe and North America but gregarious ectoparasites of concealed it has not been previously recorded in GTNP (Marsh microlepidopterous larvae including Gelechiidae, 1966). Spathius sequoiae is widely distributed in Pyralidae, and Tortricidae (Marsh 1979). During this western North America, and is known to parasitize study 36 specimens of a Hormius species were several bark beetle species in a variety of conifers sampled. The genus Hormius has never been revised (Matthews 1970). for North America, so no identification keys are available. The GTNP species may be Hormius Subfamily Braconinae americanus Ashmead, which was previously recorded from Colorado (Marsh 1979). None has Most braconine species are ectoparasites of previously been reported from GTNP. wood-boring beetle larvae or Lepidoptera larvae in plant tissue. Four braco nine genera and ten species Subfamily Rogadinae were found in GTNP during this study: Atanycolus pilosiventris Shenefelt, species near Rogadines are divided into two tribes, the montowesi Viereck, Bracon nigridorsum Ashmead, Rhysipolini and the Rogadini. · Rhysipolines are Bracon species near nigridorsum Ashmead, Bracon solitary ectoparasites of microlepidopterous leaf­ nitidulus Dalla Torre, Bracon nuperus Cresson, miners of the families Mornphidae and Gracillariidae Bracon palliventris (Provancher), Bracon tychii (Spencer & Whitfield 1999). Rhysipolis decorator (Muesebeck), Coeloides sympitys Mason, and (Haliday) and Rhysipolis platygaster Spencer were laevis (Provancher). Most of these prevously recorded from YNP and another species, species are widely distributed across the northern Rhysipolis stenodes Spencer, from GTNP by Spencer United States and Canada but none had been and Whitfield ( 1999). In this study, Rhysipolis previously recorded from GTNP. Species of platygaster is reported from GTNP for the first time. Atanycolus are parasitic on the wood-boring larvae of Cerambycidae and Buprestidae (Shenefelt 1943). Species of the tribe Rogadini are mostly Bracon species parasitize a wide variety of small solitary endoparasites of macrolepidoptera larvae, Coleoptera and Lepidoptera larvae feeding in plant especially attacking the Noctuoidea, Geometroidea, tissues (Muesebeck 1925; Marsh 1979). Coeloides and Arctioidea (Shaw, Marsh & Fortier 1997; Marsh

2 Shaw: Studies of Parasitic Wasps (Braconidae) In Grand Teton National P 49

& Shaw 2001). Four rogadine species were sampled Exotela have been described, these two are presumed in this study: Aleiodes near perplexus (Gahan), to be new species. Aleiodes near scrutator (Say), and two male specimens each representing a new species of the In this study, seven genera and at least 12 Aleiodes coxa/is species-group. species of tribe Alysiini were found in GTNP: Alysia (3 species), Anisocyrta, Aphaereta, Aspilota (2 Subfamily Opiinae species), Dapsilarthra, Dinotrema ( 2 species), and Phaenocarpa species (2 species). Alysia species Opiines are solitary endoparasites of have been revised by Wharton (1980, 1986, 1988). dipterous larvae with pupation occurring in the fly Alysia species are mostly parasites of fly larvae in puparium (Marsh 1979; Fischer 1977). Two genera carrion or mushrooms (Wharton 1986). Alysia and eight species were found in this study: micans (Viereck) was reported from GTNP and YNP Diachasma probably muleibre (Muesebeck), Opius by Wharton (1988). Alysia brevissima Wharton and blantoni Fischer, Opius cinctus Provancher, Opius Alysia brachycera Thomson are reported from GTNP dimidiatus (Ashmead), Opius hartleyi Fischer, Opius for the first time. Alysia brachycera is a widespread hollisterensis Fischer, Opius kinleyensis Fischer, and Holarctic species but Alysia brevissima was formerly Opius rudis Wesmael. With the exception of Opius known only from Alaska, Yukon Territory, and dimidiatus (Ashmead), all are new records for GTNP. Northwest Territories. Anisocyrta nearctica van Diachasma muleibre (Muesebeck) was known Achterberg is recorded from GTNP and Wyoming for previously from Washington, Oregon, and California the first time. The species has been previously as a parasite of Rhagoletis fruit flies (Marsh 1979). recorded from Colorado, British Columbia, Yukon Opius dimidiatus (Ashmead) is widespread across Territory, and Alaska as a parasite of fly larvae in North America and is recorded as a parasite of mushrooms (van Achterberg 1986). Species of Agrogromyza, Cerodontha, Liriomyza, Phytobia, and Aspilota and Dinotrema are mostly parasites of Phytomyza species (Marsh 1979). Opius rudis phorid fly larvae in decaying organic matter, or less W esmael is recorded as a parasite of Agromyza commonly platypezid fly larvae in mushrooms (van species (Marsh 1979). Opius kinleyensis Fischer and Actherberg 1988). Opius hartleyi Fischer were previously known only from Alaska. Subfamily Ichneutinae

Subfamily Alysiinae Ichneutine braconids are solitary internal parasites of sawfly larvae (Hymenoptera). Although The Alysiinae are all internal parasites of the two species are known to occur in YNP, no larvae of cyclorrhaphous Diptera, completing their ichneutine species were found in GTNP during this development in the fly puparium (Marsh 1979). study. This is probably because adults of this group These wasps are easily distinguished by their unusual are active much earlier in the season. twisted and out-turned mandibles, which is regarded as an adaptation for escaping from the fly puparium. Subfamily Homolobinae Alysiine wasps are classified into two tribes: Dacnusini and Alysiini. Members of the Dacnusini are Homolobine wasps are solitary internal mostly solitary endoparasites of leaf-mining fly parasites of and Geometridae caterpillars. larvae of the family Agromyzidae, and there are often The subfamily is recorded from GTNP for the first very host-specific (Griffiths 1964). Members of the time. One specimen of Homolobus (Apatia) tribe Alysiini, on the other hand, have a much truncator (Say) was sampled by Malaise on 5 broader host range and attack at least 25 different August. H. truncator is recorded as a parasite of families of flies (Wharton 1984). Agrotis, Feltia, Spodoptera and other cutworm-like larvae. The subfamily was revised by van In this study, four genera and at least five Achterberg (1979). species of tribe Dacnusini were found in GTNP: Chorebus, Dacnusa, Epimicta griffithsi Wharton, and Subfamily Macrocentrinae Exotela (2 species). Epimicta griffithsi was previously known only from Texas (Wharton 1994) and Macrocentrines are solitary or gregarious Kansas (Bob Kula, personal communication). endoparasites of concealed-feeding Lepidoptera Identification of most dacnusine species was not larvae. Some gregarious species are known to be possible because of the lack of revisionary studies for polyembryonic, with 8-10 individuals developing this group. Since no North Amercan species of from a single egg. Three species of Macrocentrus

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50 were found in GTNP during this study. M. species. Euphoriella pacifica Muesebeck and incompletus Muesebeck is recorded from G TNP for Microctonus species are newly recorded fromYNP the first time. One female specimen of Macrocentrus (based on material from the 1990 study). A key to species near clypeatus Muesebeck shows unusual New World species of Centistes was provided by variation, and may be a new species. The North Shaw (1995). Centistes agilus is recorded as a American species of Macrocentrus were revised by parasite of anthicid beetles of the genus Notoxus. C. Muesebeck ( 1932). A modem revision of the genus laevis is recorded as a parasite of carabid ground is currently in progress by Ken Ahlstrom. According beetles of the genus Amara. Euphoriella was revised to Ahlstrom the single male specimen from GTNP is by Loan and New (1971). They are parasites of a new species near Macrocentrus innuitorum Walley. nymphal Psocidae. F alcosyntretus was revised by Papp and Shaw (2000). The host of Falcosyntretus Subfamily Meteorinae muesebecki is not known, but other syntretines parasitize adult ichneumonid wasps or adult Meteorine braconids are mostly solitary (a bumblebees. Microctonus is a large genus few are gregarious) internal parasites of Lepidoptera parasitizing various beetles. The genus is in need of or (less commonly) Coleoptera larvae. The most revision, and species-level identifications are difficult frequently attacked hosts are caterpillars of without host-associated material. Myiocephalus Noctuidae, Geometridae, Tortricidae, and Arctiidae hoops is a holarctic species that has previously been (Huddleston 1980). The hosts of Meteorus species found in south YNP. Its host is not known but it has attacking beetles are mostly species living in wood, tree been associated with adult Formica ants (Shaw bark, and bracket fungi, especially Cerambycidae, 1985). Peristenus is a large genus that parasitizes Tenebrionidae, Scolytidae, Cryptophagidae, nymphs of mirid plant bugs (Shaw 1985). Melandryidae, and Ciidae (Huddleston 1983). Most Meteorus species produce very distinctive cocoons Subfamily Neoneurinae suspended by a long single thread of silk (Zitani and Shaw 2003). The most recent revision for North Neoneurine braconids are thought to be American species is that of Muesebeck (1923) and internal parasites in the abdomen of adult worker the group is very much in need of modem studies. ants, but information on their biology and behavior is Four meteorine species were found in this study and sparse (Shaw 1992, 1993; Poinar & Miller 2002). all are new records for GTNP: Zele reticu/atus Three neoneurine species are recorded from GTNP (Muesebeck), Meteorus species near angustipennis for the first time: Elasmosoma vigilans Cockerell, Muesebeck, Meteorus species near dimidiatus Neoneurus mantis Shaw and Neoneurus species near Cresson, Meteorus species near rubens (Nees). Z. masneri Shaw. These were from Malaise trap reticulatus is a parasite of geometrid caterpillars of samples, but on 12 August two additional individuals the genus Semiothisa (Marsh 1979). Two specimens of Neoneurus were found in a spider web. The genus of M. near angustipennis were found indoors in the Elasmosoma was revised by Huddleston (1976). The Lawrence House. Although they might have been North America species of Neoneurus were revised by attracted to lights, they were females and they Shaw (1992). appeared to be searching the walls. This raises the possibility that this, or other beetle-associated A new species of Elasmosoma near braconid species, might be used as indicator species pergandei was discovered in south YNP on 30 July for beetle activity in structural wood of historic near the South Gate Ranger Station and in areas just buildings of the park system. south of the park entrance. These wasps were observed to be active from 30 July to 9 August. They Subfamily Euphorinae were observed hovering near nests of Formica obscuripes mound-building ants between 9am and Euphorine braconids are mostly solitary (a noon on sunny days. They were less active on cooler few are gregarious) internal parasites of various adult days and during cloudy weather. Samples were and nymphal insects, including beetles, bark lice obtained by aspirator. A manuscript describing and (Psocidae ), worker bees, and worker ants (Shaw naming this new species is in preparation. 1985; Shaw & Huddleston 1991). The following genera and species are recorded from GTNP for the Subfamily Blacinae first time: Centistes (Syrrhizus) agilus (Cresson), Centistes (Ancylocentris) /aevis (Cresson), These minute and poorly studied wasps are Falcosyntretus muesebecki Papp & Shaw, thought to be internal parasites of beetle larvae in Myiocephalus hoops W esmael, and Peristenus damp decaying vegetation. Males of some species

4 Shaw: Studies of Parasitic Wasps (Braconidae) In Grand Teton National P 51

form mating swanns. One species, Blacus (Blacus) a parasite of the tortricid moth Grapholitha caduceus van Achterberg, is recorded from the GYE conversana W alsingham in Oregon. for the first time. The Blacinae were revised by van Achterberg (1988b). Subfamily Aphidiinae

Subfamily Helconinae These minute wasps are solitary internal parasites of aphids. Most species pupate inside the Helconine braconids are mostly solitary mummified body of the host aphid (Shaw & internal parasites of beetle larvae under bark or in Huddleston 1991 ). Eight aphidiine genera and plant tissue (Shaw & Huddleston 1991). Six species were recorded from the GYE during this helconine genera and species are recorded for the study: Acanthocaudus species probably schlingeri first time from the GYE: Aliolus new species, (Muesebeck), Aphidius species, Binodoxys species, Allodorus species probably crassigaster Diaeretiella species, Ephedrus species, Euaphidius (Provancher), Diospilus species, Eubazus species, species, Monoctonus species, and Praon unicum Triaspis pissodis Viereck, and Wroughtonia species Smith. Four of these are new distribution records for probably frigida (Cresson). Most helconine genera GTNP. The North American aphidiines were studied are in need of modem revisions. by Smith (1944) but most genera are in need of modern studies. Johnson (1987) revised the North Subfamily Agathidinae American Praon species. Praon unicum has been recorded as a parasite of at least 13 aphid species, Agathidines are solitary endoparasites of including common aphids on willows. concealed-feeding Lepidoptera larvae. Most species attack caterpillars feeding in concealment in leaf Subfamily Adeliinae rolls, silk webs, flower heads, or plant sterns, such as Coleophoridae, Pyralidae, Tortricidae, and Adeliines are inconspicuous wasps that are Gelechiidae (Muesebeck 1927). Although diverse rarely encountered because of their minute size. They and widely distributed, the subfamily Agathidinae are thought to be solitary internal parasites of leaf­ has not been previously recorded from GTNP. Only mining larvae of nepticulid moths, including species five agathidine specimens were sampled, that are leaf-miners in poplars and willows (Marsh representing two genera and three species (all new 1979). One species, Adelius coloradensis records for GTNP): Agathis tibiator (Provancher), Muesebeck, is recorded from GTNP for the first time. Bassus gibbosus Say, and Bassus nigripes (Cresson). A key for North American Adelius species was All three species are widely distributed across the provided by Muesebeck ( 1922). northern U.S. and Canada. Subfamily Cheloninae Subfamily Orgilinae Chelonine wasps are solitary internal egg­ Orgilines are solitary internal parasites of larval parasites of various Lepidoptera (Shaw & caterpillars feeding in concealed sites, especially Huddleston 1991 ). The chelonine females lays an Pyralidae, Tortricidae, Coleophoridae, Gelechiidae, egg directly into the host egg, but development of the Oecophoridae, and Gracillariidae (Muesebeck 1970; parasite is delayed until the host caterpillar is fully Marsh 1979). Seven orgiline species were found developed and ready to pupate. Most chelonines from the GYE during this study: Orgilus species near attack host caterpillars that feed in concealment, such conflictanae Muesebeck, Orgilus grapholithae as Tortricoidea, Pyraloidea, and Gelechioidea (Shaw Muesebeck, Orgilus species near medicaginis & Huddleston 1991). This study documents two Muesebeck, Orgilus politus Muesebeck , Orgilus genera and at least eleven chelonine species, all new rostratus Muesebeck, Orgilus strigosus Muesebeck, records for the GYE: Ascogaster species near and Orgilus species near utahensis Muesebeck. 0. borealis Shaw, Chelonus (Archichelonus) aculeatus rostratus was previously recorded from GTNP but Ashmead, Chelonus ( Chelonus) muesebecki the others are all new distribution records. Orgilines McComb, Chelonus (Chelonus) (several species), comprised 14% of all braconid specimens sampled by Chelonus (Microchelonus) a/ius McComb, Chelonus Malaise trap, making them one of the most abundant (Microchelonus) near caulicola McComb, Chelonus braconid subfamilies (second only to the (Microchelonus) near disjunctus McComb, Chelonus Microgastrinae ). The genus Orgilus was revised by (Microchelonus) species near eucosmae McComb, Muesebeck ( 1970). The most abundant orgiline Chelonus (Microchelonus) marshi McComb, species in this study, 0. grapholithae, is recorded as Chelonus (Microchelonus) near medicaginis

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McComb, and Chelonus (Microchelonus) Microplitis alaskensis Ashmead, Microplitis phthorimaeae McComb. The genus Ascogaster was autographae Muesebeck, Microplitis laticinctus revised by Shaw (1983). Ascogaster borealis is Muesebeck, Microplitis new species 1, Microplitis recorded as a parasite of the spruce needle miner, new species 2, Microplitis species 3, Microplitis Endothenia albolineana. The genus Chelonus is species 4 (near laticinctus), Microplitis species 5, highly diverse and badly in need of taxonomic study. Microplitis species 6, Pholetesor bedelliae (Viereck), The large-bodied species of the subgenus Chelonus and Sathon neomexicanus (Muesebeck). Apanteles are common and conspicuous, yet the subgenus has aristoteliae and A. fumiferanae both have broad host never been revised and no keys for species are ranges but are both recorded as parasites of the available (Marsh 1979). The smaller species of the spruce budworm, Choristoneura fumiferanae (Marsh genus Microchelonus were revised by McComb 1979). Cotesia gillettei is recorded as a parasite of ( 1968) but identification of species remains difficult. Archips argyrospilus, a common leaf-roller on many McComb's lengthy key includes 111 species, but the broad leaf trees including trembling aspen (Marsh key treats only females and is not illustrated. 1979). The most recent revision for Apanteles is that Interpretation of variation is often difficult. Chelonus of Muesebeck (1920). The genus Apanteles (as (Microchelonus) leptogaster McComb has been formerly defined) was reclassified by Mason ( 1981) and previously recorded near the South Gate of YNP many common species were re-assigned to other (McComb 1968). genera including Cotesia and Glyptapanteles. The annotated checklist provided by Whitfield ( 199 5) is Subfamily Microgastrinae highly useful for checking generic and species combinations in this group. The genus Sathon was The subfamily Microgastrinae is regarded as revised by Williams (1988). the single most important group of parasitoids of Lepidoptera in the world, both in economic terms and in numbers of species (Whitfield 1995). + DISCUSSION Microgastrines are solitary or gregarious internal parasites of various caterpillars. Due to their high This survey of braconid wasps in GTNP and diversity and frequent occurrence of gregarious south YNP recognizes 71 genera and at least 150 parasitism, microgastrines are very abundant in forest species. Of these, 100 are new distribution records habitats and are normally the most commonly and at least nine are new (undescribed) species. The encountered braconid subfamily. This was also true species richness of Braconidae in GYE (150 species) in this study, where microgastrine species comprised is substantially greater than the maximum species more than 54% of all Braconidae sampled. Despite richness of all herbaceous plants, shrubs, trees, birds their abundance and economic importance, many and mammals in lodgepole pine communities of YNP microgastrines remain very difficult to identify. (112 species according to Taylor 1973). This total There are few modem taxonomic revisions for most also substantially exceeds the number of braconid groups~ and most males cannot be identified to species recorded in fire-disturbed habitats of south species level unless they are associated with host­ YNP (109 species recorded by Lockwood, Shaw & reared series of females. The following Struttmann 1996). microgastrine genera and species are recorded from the GYE during this study: Apanteles aristoteliae The high number of new distribution records Viereck, Apanteles species near crassicornis ( 100) is probably a reflection not of scarcity of these (Provancher), Apanteles ensiger (Say), Apanteles species, but rather the scarcity of previous studies, near femurnigrum Provancher, Apanteles fumiferanae the difficulties of sampling microscopic insects, and Viereck, Apanteles species near stagmatophorae, the lack of studies in GYE because of permit Cotesia species near anisotae (Muesebeck), Cotesia restrictions. No checklist of Braconidae species has species near carduicola (Packard), Cotesia species previously been published for the GYE. A checklist near clisiocampae (Ashmead), Cotesia species near of species based on this study is provided below. enypiae (Mason), Cotesia gillettei Baker, Cotesia Most of the new distribution records are for species glomerata (Linnaeus), Cotesia griffini (Viereck), that are widely distributed in the northwestern United Cotesia xylina (Say), Cotesia yakutatensis States, Canada, or Alaska. Some species recorded (Ashmead), Dolichogenidea near miantonomoi were previously known from Alaska ( Opius Viereck, Diolcogaster fascetosa (Weed), kinleyensis, Opius hartleyi, Alysia brevissima). Glyptapanteles near compressiventris (Muesebeck), Several circumpolar (Holarctic) species are also Microgaster femoralamericanus Shenefelt, present ( Onsira imperator, Aleiodes unipunctator, Microgaster species near peroneae Walley, Alysia brachycera, Homolobus truncator, Meteorus

6 Shaw: Studies of Parasitic Wasps (Braconidae) In Grand Teton National P 53

rubens, Myiocephalus hoops, Cotesia glomerata). provided critical assistance with setting and The community of braconid species appears to be maintaining Malaise traps and other field assistance. dominated by boreal elements, many of which probably dispersed into the GYE from the north, along the corridor of the Rocky Mountains. + LITERATURE CITED

Although the high species richness of Achterberg, C. van. 1979. A revision of the Braconidae is perhaps not unexpected, it is still rather subfamily Zelinae auct. (Hymenoptera, remarkable that so many previously unrecorded Braconidae). Tidschrift voor Entomologie species could be sampled during such a short time 122:241-479. frame (28 July to 15 August), mostly at one sampling site (AMK ranch), mostly by one sampling method Achterberg, C. van. 1986. The Holarctic genus (Malaise trap), and during a severe drought season. It Anisocyta Foerster (Hymenoptera: should be kept in mind that sampling done earlier in Braconidae: Alysiinae ). Zoologische the season, in plant communities other than mature Mededelingen 60:285-297. lodgepole pine, at other locations, and using a variety of other sampling methods would likely yield many other Achterberg, C. van. 1988. The genera of the braconid species not recorded here. Host-rearing Aspilota-group and some descriptions of studies are very much needed to establish host ranges fungicolous Alysiini from the Netherlands for braconid species in the GYE and to obtain (Hymenoptera: Braconidae: Alysiinae ). conspecific series to examine variation. Given these Zoologische Verhandelingen 24 7:3-88. considerations, it seems reasonable to suppose that the total richness of Braconidae species in the GYE is Acterberg, C. van. 1988b. Revision of the subfamily probably much greater than the 150 species recorded Blacinae Foerster (Hymenoptera, here, and might easily exceed 200 species. Braconidae). Zoologische Verhandelingen 249:1-324. It is also interesting to observe that the discovery of new insect species reflects not so much Fischer, M. 1977. Hymenoptera: Braconidae the rarity of these organisms, but rather the lack of (Opiinae II - Amerika). Das Tierreich 96: 1- previous sampling and revisionary research for GYE 1001. (in German) forest Braconidae. For example, although Microplitis was one of the most abundant and easily sampled Griffiths, G.C.D. 1964. The Alysiinae (Hym. genera in Malaise samples, the most common Braconidae) parasites of the Agromyzidae Microplitis species in these samples were (Diptera). I. General questions of undescribed species (Jim Whitfield, personal taxonomy, biology, and evolution. Beitrage communication). A new species of Elasmosoma was zur Entomologie 14:823-914. discovered literally within minutes of entering YNP, and was found to be rather common for several days. Huddleston, T. 1976. A revision of Elasmosoma Ruthe (Hymenoptera, Braconidae) with two new species from Mongolia. Annales + ACKNOWLEDGMENTS His tori co-Naturales M usei N ationalis Hungarici 68:215-225. Jim Whitfield (University of Illinois) provided species identifications for Microplitis and Huddleston, T. 1980. A revision of the western Pholetesor. Kenneth Ahlstrom (North Carolina Palearctic species of the genus Meteorus Department of Agriculture) provided species (Hymenoptera: Braconidae). Bulletin of the identifications for Macrocentrus. Robert Kula British Museum (Natural History) 41: 1-58. (University of Kansas) provided the identification for Epimicta griffithsi. Christie Hendrix (YNP) and Huddleston, T. 1983. Meteorus (Hymenoptera: Nicki Tippets (GTNP) assisted with providing park Braconidae) of Australia and New Guinea. collecting permits. Rick Mossman (YNP ranger) Systematic Entomology 8:393-420. pointed out the location of ant colonies in south YNP, crucial to the discovery of Elasmosoma. This Johnson, J.W. 1987. A revision of the species of research was funded by a 2002 grant from the UW­ Praon in North America north of Mexico NPS research station and Macintyre-Stennis project (Hymenoptera: Aphidiidae). The Canadian number WY0-362-02. My son, Michael Shaw, Entomologist 119:999-1025.

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Loan, C.C. and T.R. New. 1971. An account of the Muesebeck, C.F.W. 1920. A revision of the North North American species of Euphoriella American species of ichneumon-flies Ashmead with descriptions of seven new belonging to the genus Apanteles. species (Hymenoptera: Braconidae, Proceedings of the United States National Euphorinae ). Proceedings of the Museum 58:483-576. Entomological Society of Ontario 102:92- 108. Muesebeck, C.F.W. 1922. A revision of the North American ichneumon-flies belonging to the Lockwood, J.A., S.R. Shaw, and J.M. Struttmann. subfamilies Neoneurinae and 1996. Biodiversity of wasp species (Insecta: Microgastrinae. Proceedings of the United Hymenoptera) in burned and unburned States National Museum 61:1-76. habitats of Yellowstone National Park, Wyoming, USA. Journal of Hymenoptera Muesebeck, C.F.W. 1923. A revision of the North Research 5:1-15. American species of ichneumon-flies belonging to the genus Meteorus Haliday. Marsh, P.M. 1966. The Nearctic Doryctinae, II. Proceedings of the United States National The genus Doryctodes Hellen Museum 63:1-44. (Hymenoptera: Braconidae). Muesebeck, C.F.W. 1925. A revision of the Marsh, P.M. 1979. Family Braconidae, pp. 144-313, parasitic wasps of the genus Microbracon In: Catalog of Hymenoptera in America occurring in America North of Mexico. north of Mexico, Krombein et al. (eds.), Proceedings of the United States National Smithsonian Institution Press, Washington Museum 67:1-85. D.C. Muesebeck, C.F.W. 1927. A rev1s10n of the Marsh, P.M. and S.R. Shaw. 2001. Revision of parasitic wasps of the subfamily Braconinae North American Aleiodes (Part 6): the occurring in America North of Mexico. gasterator and unipunctator species-groups. Proceedings of the United States National Proceedings of the Entomological Society of Museum 69:1-73. Washington 103:291-307. Muesebeck, C.F.W. 1932. Revision of the Nearctic Mason, W.RM. 1978. A synopsis of the Nearctic ichneumon-flies belonging to the genus , with revisions of N earctic species Macrocentrus. Proceedings of the United of Coeloides and Myosoma (Hymenoptera: States National Museum 80:1-55. Braconidae ). Canadian Entomologist 110:721-768. Muesebeck, C.F.W. 1935. On the genus Oncophanes Foerster, with descriptions of Mason, W .R.M. 1981. The polyphyletic nature of two related genera (Hymenoptera: Apanteles Foerster (Hymenoptera: Braconidae ). Annals of the Entomological Braconidae): a phylogeny and Society of America 28:241-250. reclassification of Microgastrinae. Memoirs of the Entomological Society of Canada Muesebeck, C.F.W. 1938. The genus Dendrosoter 115:1-147. Wesmael in the United . States (Hymenoptera: Braconidae). Proceedings of Matthews, R.W. 1970. A revision of the genus the Entomological Society of Washington Spathius in America North of Mexico 40:281-287. (Hymenptera, Braconidae). Contributions of the American Entomological Institute 4:1- Muesebeck, C.F.W. 1970. The nearctic species of 86. Orgilus Haliday (Hymenoptera: Braconidae). Smithsonian Contnbutions to Zoology 30: 1- McComb, C.W. 1968. A revision of the Chelonus 104. subgenus Microchelonus in North America north of Mexico (Hymenoptera: Braconidae). University of Maryland Agricultural Experiment Station Bulletin A- 149, 1-148.

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Papp, J. and S.R. Shaw. 2000. A study of the genus Shaw, S.R., P.M. Marsh, and J.C. Fortier. 1997. Falcosyntretus Tobias from the New World Revision of North American Aleiodes (Part 1): with five new species and a key to known the pulchripes species-group in the New species (Hymenoptera: Braconidae: World. Journal of Hymenoptera Research Euphorinae ). Proceedings of the 6:10-35. Entomological Society of Washington 102:634-642. Shenefelt, R.D. 1943. The genus Atanycolus in America North of Mexico. Research Poinar, G. Jr. and J.C. Miller. 2002. First fossil Studies of the State College of Washington record of endoparasitism of adult ants 11:51-163. (Formicidae: Hymenoptera) by Braconidae (Hymenoptera). Annals of the Smith, C.F. 1944. The Aphidiinae of North America Entomological Society of America 95:41- (Braconidae: Hymenoptera). Ohio State 43. University Contributions in Zoology and Entomology 6:1-154. Shaw, M.R. and T. Huddleston. 1991. Classification and biology of braconid wasps Spencer, L. and J.B. Whitfield. 1999. Revision of the (Hymenoptera: Braconidae). Royal Nearctic species of Rhysipolis Forster Entomolgical Society of London, (Hymenoptera: Braconidae). Transactions Handbooks for the Identification of British of the American Entomological Society Insects 7: 1-126. 125:295-324.

Shaw, S.R. 1983. A taxonomic study of nearctic Taylor, D.L. 1973. Some ecological implications of Asco gaster and a description of a new genus forest fire control in Yellowstone National Leptodrepana (Hymenoptera: Braconidae). Park, Wyoming. Ecology 54:1394-1396. Entomography 2:1-54. Wharton, R.A. 1980. Review of the Nearctic Shaw, S.R. 1985. A phylogenetic study of the Alysiini (Hymenoptera, Braconidae) with subfamilies Meteorinae an Euphorinae discussion of the generic relationships (Braconidae). Entomography 3:277-370. within the tribe. University of California Publications in Entomology 88:1-112. Shaw, S.R. 1988. Euphorine phylogeny: the evolution of diversity in host-utilization by Wharton, R.A. 1984. Biology of the Alysiini parasitoid wasps. Ecological Entomology (Hymenoptera: Braconidae), parasitoids of 13:323-335. cyclorrhaphous Diptera. Texas Agricultural Experiment Station Technical Monograph Shaw, S.R. 1992. Seven new North American 11:1-39. species of Neoneurus (Hymenoptera: Braconidae ). Proceedings of the Wharton, R.A. 1986. The braconid genus Alysia Entomological Society of Washington (Hymenoptera: a description of the 94:26-47. subgenera and a revision of the subgenus Alysia. Systematic Entomology 11:453-504. Shaw, S.R. 1993. Observations on the ovipositional behavior of Neoneurus mantis, and ant­ Wharton, R.A. 1988. The braconid genus Alysia parasitoid from Wyoming. Journal of Insect (Hymenoptera): a revision of the subgenus Behavior 6:649-658. Anarcha. Contributions of the American Entomological Institute 25: 1-69. Shaw, S.R. 1995. A new species of Centistes from Brazil (Hymenoptera: Braconidae: Wharton, R.A. 1994. New genera, species, and Euphorinae) paras1t1zmg adults of records of New World Alysiinae Diabrotica (Coleoptera: Chrysomelidae ), (Hymenoptera: Braconidae). Proceedings of with a key to New World species. the Entomological Society of Washington Proceedings of the Entomological Society of 96:630-664. Washington 97:153-160.

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Wharton, R., P. Marsh, and M. Sharkey. 1997. Williams, D.J.M. 1988. Classification, phylogeny Manual of the New World genera of the and zoogeographic studies of species of family Braconidae (Hymenoptera). Special Sathon Mason (Hymenoptera: Braconidae). Publication of the International Society of Quaestiones Entomologicae 24:529-639. Hymenopterists, Number 1, 439 pp. Zitani, N.M. and S.R. Shaw. 2003. From meteors to Whitfield, J.B. 1995. Annotated checklist of death stars: variations on a silk thread Microgastrinae of North America north of (Hymenoptera: Braconidae: Meteorinae. Mexico (Hymenoptera: Braconidae). American Entomologist 48:228-235. Journal of the Kansas Entomological Society 68:245-262.

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CHECKLIST OF BRACONIDAE FOR THE Aleiodes coxalis species-group new species 2 GTNP GYE Subfamily Opiinae Subfamily Doryctinae Diachasma probably muleibre (Muesebeck) GTNP Doryctes fartus (Provancher) GTNP (new record) Doryctes species near fartus (Provancher) YNP Opius blantoni Fischer GTNP (new record) Doryctes species YNP Opius cinctus Provancher GTNP (new record) Dendrosoter scaber Muesebeck GTNP (new record) Opius dimidiatus (Ashmead) GTNP Heterospilus species YNP Opius hartleyi Fischer GTNP (new record) Ontsira imperator (Haliday) GTNP (new record) Opius hollisterensis Fischer GTNP (new record) Spathius canadensis Ashmead YNP Opius kinleyensis Fischer GTNP (new record) Spathius species near sequoiae Ashmead GTNP (new Opius rudis Wesmael GTNP (new record) record) Subfamily Alysiinae Subfamily Braconinae Tribe Dacnusini Atanycolus pilosiventris Shenefelt GTNP (new Chorebus species GTNP record) Dacnusa species GTNP Bracon species near montowesi .Viereck (new Epimicta griffithsi Wharton GTNP (new record) record) Exotela new species 1 GTNP Bracon nigridorsum Ashmead (new record) Exotela new species 2 GTNP Bracon species near nigridorsum Ashmead (new Tribe Alysiini record) Alysia brachycera Thomson GTNP (new record) Bracon nitidulus Dalla Torre (new record) Alysia brevissima Wharton GTNP (new record) Bracon nuperus Cresson (new record) Alysia lucia Haliday YNP Bracon palliventris (Provancher) (new record) Alysia micans (Viereck) GTNP (Wharton 1988) Bracon tychii (Muesebeck) GTNP (new record) A nisocyrta nearctica van Achterberg GTNP (new Coeloides rufovariegatus (Provancher) GTNP record) (Mason 1978) Aphaereta species GTNP Coeloides sympitys Mason YNP, GTNP (new record) Aspilota species 1 GTNP Coeloides vancouverensis (Dalla Torre) YNP Aspilota species 2 GTNP Cyanopterus laevis (Provancher) YNP, GTNP (new Dapsilarthra species GTNP record) Dinotrema species 1 GTNP Dinotrema species 2 GTNP Subfamily Rhyssalinae (new record) Idiasta species YNP Oncophanes species near californicus (Ashmead) Phaenocarpa species 1 GTNP YNP, GTNP (new record) Phaenocarpa species 2 GTNP

Subfamily Hormiinae (new record) Subfamily Ichneutinae Hormius species possibly americanus Ashmead Ichneutes bicolor (Cresson) YNP (new record) YNP, GTNP (new record) Proterops californicus Cresson YNP (new record)

Subfamily Rogadinae Subfamily Homolobinae Tribe Rhysipolini Homolobus (Apatia) truncator (Say) GTNP (new Rhysipolis decorator (Haliday) YNP (Spencer and record) Whitfield 1999) Rhysipolis platygaster Spencer YNP (Spencer and Subfamily Macrocentrinae Whitfield 1999), GTNP (new record) Macrocentrus species near clypeatus Muesebeck Rhysipolis stenodes Spencer GTNP (Spencer and (new record) Whitfield 1999) Macrocentrus. incompletus Muesebeck (new record) Tribe Rogadini Macrocentrus new species near innuitorum Walley Aleiodes near perplexus (Gahan) GTNP, YNP (male only) Aleiodes near scrutator (Say) GTNP, YNP Aleiodes unipunctator (Thunberg) Subfamily Meteorinae =Aleiodes harrimani (Ashmead) (new Zele reticulatus (Muesebeck) GTNP (new record) synonym) GTNP (Marsh & Shaw 2001) Meteorus species near angustipennis Muesebeck Aleiodes coxalis species-group new species 1 GTNP GTNP (possible new species) (new record)

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Meteorus species near dimidiatus Cresson GTNP Subfamily Aphidiinae (new record) Acanthocaudus species probably schlingeri Meteorus species near rubens (Nees) GTNP (new (Muesebeck) GTNP (new record) record) Aphidius species GTNP Binodoxys species GTNP (new record) Subfamily Euphorinae Diaeretiella species YNP Centistes (Syrrhizus) agilus (Cresson) GTNP (new Ephedrus species YNP, GTNP (new record) record) Euaphidius species YNP Centistes (Ancylocentris) laevis (Cresson) GTNP Monoctonus species YNP (new record) Praon unicum Smith GTNP (new record) Euphoriella pacifica Muesebeck YNP (new record) Praon species YNP, GTNP Falcosyntretus muesebecki Papp & Shaw GTNP (new record) Subfamily Adeliinae (new record) Microctonus species YNP Adelius coloradensis Muesebeck GTNP (new record) Myiocephalus boops Wesmael YNP, GTNP (new record) Subfamily Cheloninae Peristenus species GTNP (new record) Ascogaster species near borealis Shaw GTNP (new record) Subfamily Neoneurinae (new record) Chelonus (Archichelonus) aculeatus Ashmead YNP Elasmosoma vigilans Cockerell GTNP (new record) (new record) Elasmosoma new species near pergandei YNP (new Che/onus ( Chelonus) muesebecki McComb YNP, record) GTNP (new record) Neoneurus mantis Shaw GTNP (new record) Chelonus ( Chelonus) species YNP, GTNP (several Neoneurus species near masneri Shaw GTNP (new species) record) Chelonus (Microchelonus) a/ius McComb GTNP (new record) Subfamily Blacinae (new record) Chelonus (Microchelonus) near caulicola McComb Blacus (Blacus) caduceus van Achterberg YNP, GTNP (new record) GTNP (new record) Chelonus ( Microchelonus) near d isjunctus M cComb GTNP (new record) Subfamily Helconinae Chelonus (Microchelonus) species near eucosmae Aliolus new species YNP, GTNP (new record) McComb GTNP (possible new species) Allodorus species probably crassigaster (Provancher) Chelonus (Microchelonus) leptogaster McComb YNP (new record) YNP (McComb 1968) Diospilus species GTNP (new record) Chelonus (Microchelonus) marshi McComb GTNP Eubazus species YNP (new record) Triaspis pissodis Viereck YNP, GTNP (new record) Chelonus (Microchelonus) near medicaginis Wroughtonia species probably frigida (Cresson) McComb GTNP (new record) Chelonus (Microchelonus) phthorimaeae McComb Subfamily Agathidinae GTNP (new record) Agathis tibiator (Provancher) (new record) Chelonus (Microchelonus) species YNP, GTNP Bassus gibbosus Say (new record) (several species) Bassus nigripes (Cresson) (new record) Subfamily Microgastrinae Subfamily Orgilinae Apanteles aristoteliae Viereck GTNP (new record) Orgilus species near conjlictanae Muesebeck YNP, Apanteles species near crassicornis (Provancher) GTNP (new record) GTNP (new record) Orgilus grapholithae Muesebeck GTNP (new record) Apanteles ensiger (Say) GTNP (new record) Orgilus species near medicaginis Muesebeck YNP Apanteles near femurnigrum Provancher GTNP (new (new record) record) Orgilus politus Muesebeck GTNP (new record) Apanteles fumiferanae Viereck GTNP (new record) Orgilus rostratus Muesebeck GTNP (Muesebeck Apanteles species near stagmatophorae GTNP (new 1970) record) Orgilus strigosus Muesebeck YNP (new record) Apanteles species GTNP, YNP Orgilus species near utahensis Muesebeck YNP (new Cotesia species near anisotae (Muesebeck) GTNP record) (new record)

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Cotesia species near carduicola (Packard) GTNP (new record) Cotesia species near clisiocampae (Ashmead) GTNP (new record) Cotesia species near enypiae (Mason) GTNP (new record) · Cotesia gillettei Baker GTNP Cotesia glomerata (Linnaeus) GTNP (new record) Cotesia griffini (Viereck) GTNP (new record) Cotesia xylina (Say) GTNP (new record) Cotesia yakutatensis (Ashmead) GTNP (new record) Cotesia species GTNP, YNP Dolichogenidea near miantonomoi Viereck GTNP (new record) Diolcogaster fascetosa (Weed) GTNP, YNP Glyptapanteles near compressiventris (Muesebeck) GTNP (new record) Glyptapanteles species GTNP, YNP Micro gaster femora/american us Shenefelt YNP, GTNP (new record) Micro gaster species near peroneae Walley GTNP (new record) Microplitis alaskensis Ashmead GTNP (new record) Microplitis autographae Muesebeck GTNP (new record) Microplitis laticinctus Muesebeck GTNP, YNP (new record) Microplitis new species 1 GTNP Microplitis new species 2 GTNP, YNP Microplitis species 3 GTNP Microplitis species 4 (near laticinctus) GTNP Microplitis species 5 GTNP Microplitis species 6 (dwarf males) YNP Microplitis species YNP, GTNP Pholetesor bedelliae (Viereck) GTNP (new record) Sathon neomexicanus (Muesebeck) GTNP, YNP

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