Journal of Hymenoptera research 12(1) 2003

S«3 SOCIETy Journal of Hymenoptera Research Volume Number 12, ril 2003 3N #1070-9428 CONTENTS DEANS, A R., J B WHITFIELD, and D H the microgastrine genus Alphomelon JANZEN, D H., A K specificity WALKER, J B JANZEN Taxonomy and natural history Mason (Hymenoptera: Braconidae) WHITFIELD, G DELVARE, and I D GAULD and hyperparasitoids of three new Costa Rican species of of Host- Microplitis Braconidae: Microgastrinae), parasitoids of sphingid Foerster (Hymenoptera: 42 caterpillars HUBER, Review of Chaetomymar Ogloblin, with description of a new species J T Hawaiian Islands (Hymenoptera: Mymaridae) KIMSEY, L S A peculiar new genus of locally in the 77 abundant Australian Thynninae 102 (Hymenoptera: Tiphiidae) LOPEZ PEREZ, M A preliminary list of the Encyrtidae (Hymenoptera: Chalcidoidea) of Cuba, with descriptions of two new species ROIG-ALSINA, A The bee genus D R A synopsis Holmberg (Hymenoptera: Apidae): Holmberg Doeringiella sion of the subgenus Pseudepeolus SMITH, 125 of the sawflies (Hymenoptera: a revi- 136 Symphyta) of America south of the United States: Tenthredinidae (Allantinae) SMITH, D R and D H JANZEN Argidae (Hymenoptera) in Food plants and life 148 histories of sawflies of the family Costa Rica, with descriptions of two new species 193 INTERNATIONAL SOCIETY OF HYMENOPTERISTS Organized 1982; Incorporated 1991 OFFICERS FOR Lynn Kimsey, 2003 President Denis Brothers, President-Elect James B Woolley, Secretary John T Huber, Treasurer E Eric Grissell, Editor Subject Editors Symphyta and Parasitica Biology: Aculeata Mark Shaw Biology: Donald Quicke Systematics: Sydney Cameron Systematics: Wojciech Pulawski All correspondence concerning Society business should be mailed to the appropriate officer at the following addresses: President, Bohart Museum of Entomology, Department of Entomology, A&M University of California, Davis, CA 95616; Secretary, Department of Entomology, Texas University, College Station, Texas 77843; Treasurer, Eastern Cereal & Oilseed Research Centre, Agriculture Canada, K W Neatby Building, Ottawa, Ontario, Canada K1A 0C6; Editor, National Museum of Natural USDA, P.O Box 37012, % Systematic Entomology Laboratory, History CE 520, MRC 168, Washington, D.C 20013-7012 Membership Members shall be persons who have demonstrated interest in the science of entomology Annual dues for members are US$40.00 per year (US$35.00 if paid before February), payable to The International Society of Hymenopterists Requests for membership should be sent to the Treasurer (address above) Information on membership and other details of the Society may be found on the World Wide Web at http://IRIS.biosci.ohio-state.edu/ish Journal The Journal of Hymenoptera Research is published twice a year by the International Society of Hymenopterists, Department of Entomology, Smithsonian Institution, Washington, D.C U.S.A Members in good standing receive the Journal Nonmember subscriptions are 20560-0168, % $60.00 (U.S currency) per year The Society does not exchange its publications for those of other societies Please see inside back cover of this issue for information regarding preparation of manuscripts Statement of Ownership Title of Publication: Journal of Hymenoptera Research Frequency of Issue: Twice a year Location of Office of Publication, Business Office of Publisher of Hymenopterists, Constitution NW, % Department and Owner: International Society and of Entomology, Smithsonian Institution, 10th Washington, D.C 20560-0168, U.S.A % Editor: E Eric Grissell, Systematic National Museum of NatEntomology Laboratory, USDA, ural History, 10th and Constitution NW, D.C 20560-0168 U.S.A Washington, Managing Editor and Known Bondholders or other Security Holders: none This issue was mailed April 2003 J HYM RES Vol 12(1), 2003, pp 1-41 Taxonomy and Natural History of the Microgastrine Genus Alphomelon Mason (Hymenoptera: Braconidae) Andrew R Deans, James (ARD, JBW) Department (DHJ) Department B Whitfield, and Daniel H Janzen of Entomology, University of Illinois, Urbana, IL 61801, email: adeans@life.uiuc.edu, jwhitfie@life.uiuc.edu; of Biology, University of Pennsylvania, Philadelphia, PA 19104, USA, USA, email: djanzen@sas.upenn.edu Abstract —The New World endemic genus Alphomelon Mason (Braconidae: Microgastrinae) is time This revision includes a redescription of the genus, redescriptions of its named species and descriptions of new species with illustrations of diagnostic characters, an illustrated key to Alphomelon species, and a summary of their natural history as parasitoids of revised for the first four of hesperiid caterpillars Alphomelon is a monophyletic group containing seventeen species, which were previously described: Alphomelon disputabile (Ashmead) (Argentina to U.S.A.), A niA talidicida (Wilkinson) (Brazil griceps (Ashmead) (Caribbean islands and surrounding countries), to Costa Rica), and A conformis (Muesebeck) (Venezuela to Costa Rica) Thirteen species are described as new: Alphomelon arecaphile Deans (Brazil to Costa Rica), A brachymacher Deans (Brazil and Peru to Costa Rica), A bromeliphile Deans (Costa Rica to Mexico), A citroloma Deans (Argentina to Costa Rica), A crocostetlius Deans (Brazil to Colombia), A melanoscelis Deans nanosoma Deans (Brazil to Mexico), A panrogetium Deans (Argentina and rhyssocerms Deans (Argentina to Panama), A simpsononim Deans (Paraguay to Costa Rica), A pyrrhogluteum Deans (Argentina), A winnieivertzae Deans (Costa Rica to Canada), and A xestopyga Deans (Costa Rica) (Brazil to Mexico), A Chile), A Distributed throughout most of the ternperate and tropical New World, the members of the genus Alphomelon Mason parasitize skipper larvae (Hesperiidae) and are frequently collected and often reared Although members of this genus are easy to recognize due to the white coloration on their genae (which gives the genus its Ashmead described two species of Al- from the island of phomelon Vincent in the Caribbean as Urogastcr in 1900 St ni- Ashmead and Urogaster disputabilis Ashmead In 1920 Muesebeck synonymized Urogaster under Apanteles Forster and reassigned nigriceps and disputabilis griceps (renamed accordingly disputabile) Since name), identification at the species level has remained difficult owing to the ab- then only two other species have been de- sence of keys, adequate descriptions, and illustrations This revision updates the Guyana (Wilkinson 1931) and Apanteles conformis Muesebeck from Venezuela classification of the species within Alphomelon, and includes a brief discussion of sification of Apanteles led to his creation of its taxonomic history, biology, and distri- bution together with descriptions of thir- new scribed: Apanteles talidicida (Muesebeck the 1958) new genus Wilkinson from Mason's (1981) reclas- Alphomelon comprising putabile, nigriceps, and talidicida dis- Mason apwhen reas- species, redescriptions of four previously named species, illustrations of parently overlooked conformis signing species, but morphology, color diagnostic characters, and a key to patterns, teen cies known to date all spe- belongs and biology suggest in Alphomelon that it also Journal of Hymenoptera Research Collections often contain mens many of Alphomelon, particularly speciif they basitarsus; see Figure 6a,b) to long (the exserted portion as long as the hind tibia) World malaise trap samples However, due to the paucity of microgastrine taxonomists and a lack of il- with varying degrees of curvature More than half the species are equipped with a lustrations or keys these specimens are usually located in the "unsorted" drawers and medium curvature (Fig 6c), including most of the reared species Species also are rich in New lumped with Apanteles This lack of exwith the large inin combination pertise, flux of specimens from Janzen's caterpil- or lar-parasitoid rearing project in north- western Costa Rica (Janzen and Hallwachs 2002, Schauff and Janzen 2001, Burns and Janzen 2001), clarified the need for a tax- onomic revision of Alphomelon NATURAL HISTORY AND DISTRIBUTION OF ALPHOMELON Host records mens in for the few reared speci- major collections indicate that species of Alphomelon invariably parasitize hesperiine skipper larvae (Lepidoptera: Hesperiidae) feeding mostly on monocots There is one doubtful rearing record from a dicot-eating caterpillar: DHJ voucher 98- "typical" ovipositor of medium length with respect to the number of spines (from one to four) on the tarsal claws (Fig differ 3) Since skipper larvae typically live in with silk (Hogue 1993), further exploration of their ovipositors and tarsal claws may provide clues into ovileaf-rolls lined position behavior, host species escape biology, and the host environment This genus occurs only in the New 1981) The highest concen- World (Mason tration of species are in Amazonia and southern Central America where hesperiids are also extremely speciose (Hogue 1993) PHYLOGENETIC PLACEMENT OF ALPHOMELON Mason (1981; also Walker et al 1990) SRNP-4564 (Janzen and Hallwachs 2002) The majority of museum records are from skipper larvae feeding on agricultural used morphological characters to place Alphomelon in the Apanteles genus-group (Apanteles, Papanteles, Alphomelon, and crops such as corn, sugar cane, bananas, and canna Two of these hosts, Lerema sp Dasylagon) within the tribe Apantelini Re- and Calpodes ethlius, are also reported to feed on peanuts (extraordinarily doubtful) and wheat (according to caterpillar food plant records in the USNM), but Alphomelon has never been recorded from skipper larvae on these host plants In contrast to the dearth of museum host records (which often have questionable accuracy), D H Janzen and W Hall- wachs' (2002) caterpillar-parasitoid rearing project in Costa Rica (voucher number information accessible at Janzen and Hallwach's website: (http://janzen.sas.upenn morphology combined with molecular data (Mardulyn and Whitfield 1999, Whitfield et al 2002) maintains this clade with Alpmomelon as the sister group analysis of and/or DolichogenAvailable host information suggests that most species in these other three genera parasitize concealed microlepidoptera to Apanteles, Dasylagon, idea in rolled leaves (Mason 1981) Alphomelon, differs in parasitizing one subfamily of macrolepidoptera that also live in leaf rolls (Mason 1981) however, MATERIAL edu)) adds abundant and continuous new records of host caterpillars and food More than 2000 specimens were borrowed for this revision from the following plants (see Table institutions or individuals: (AEI) American Entomological Institute Collection, 1) Alphomelon species exhibit a diverse array of ovipositor types ranging from short (the exserted portion shorter than the hind FL, USA, David B Wahl; (CNC) Canadian National Collection, Bio- Gainesville, Volume 12, Number 1, 2003 systematics Research Centre, Agriculture Canada, Ottawa, Ontario, Canada, Henri Goulet; (DHJ) Department of Biology, University of Pennsylvania, Philadelphia, PA, USA, D H Janzen (these specimens were deposited in: AEI, CNC, INBIO, INHS, MCZ, TAMU, USNM, UWY); (INBIO) Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica, Carolina Godoy; (INHS) Natural Illinois of posterior scutellar depression; see Fig 1), and petiolar ridge (medial carina on petiole; see Fig 1) Quicke et al (1999) was used for ovipositor structural terms Head width, medial ocellar diameter, lateral ocellar diameter, lateral ocellar line, posterior ocellar line, ocular ocellar line, hind tarsal length, and ovipositor sheath length used by followed those measurements Williams (1988) boldt Biological Resources Research Insti- Only specimens closely matching the holotype in all morphological aspects and /or males from the same series were Bogota, Colombia, Fernando Fernan- designated and labeled as paratypes All (MCZ) Museum specimens examined for this revision were supplied with species determination labels USA, Col(HUMB) Alexander von Hum- History Survey, Champaign, in Favret; tute, IL, Comparative Zoology, Harvard University, Cambridge, dez; MA, USA, of Stefan Cover; (NHM) Museum, London, UK, Mike (TAMU) Texas A and M University, Fit- History ton; Natural College Station, TX, USA, Robert A Wharton; (UCD) University of California at Davis, Davis, CA, USA, Lynn Kimsey; (USNM) United States National Museum, Smithsonian Institution, Washington, DC, USA, David Smith; (UWY) University Wyoming, Laramie, WY, USA, Scott Although the conspicuous white color- ation of the genae helps to define Alphomelon, one should not rely on that character alone for identification Several species of Neotropical Cotesia (personal ob- servation of holotypes for A disputabile, A iiigriceps, and A conformis from USNM A cotype for A talidicida was examined during a visit NHM but AND of were the to DIAGNOSTIC CHARACTERS SPECIES KEY R in this material Shaw Included before the loans were returned not borrowed ME HODS reared museum riodinid collections parasitoids) also and have white patches on the genae, as many Dolichogenidea and other genera in the Old World tropics (Mason 1981; personal observation) Additional characters that distinguish Alphomelon from other genera with white genae include: fully sclerotized Morphological terms follow those of Sharkey and Wharton (1997) Additional terms include: lunule (Mason 1981), stem- hypopygium, presence of lack of wing areolet, and maticum (Shaw and Huddleston 1991), lateral and anterior scutellar furrows (to distinguish between scutellar furrows in Sharkey and Wharton 1997; see Fig 1), medial scutellar area (area between lateral Color patterns are the most obvious and perhaps easiest characters to use for diagnosis of certain species Conspicuous body segment coloration of orange or yellow helps distinguish between A uigriceps 1), anterior proareas area podeal (propodeal anteriorly left and right of areola; see Fig 1), posterior propodeal area (propodeal areas pos- scutellar furrows; see Fig teriorly left and right of areola; see Fig 1), axillary projections (rugose projections apical to lunules; see Fig 1), lateral metanotal depression (refers to pits left and right propodeum with petiolar ridge, fully carinate areola, (metasoma and mesosoma yellow), A cro(mesonotum orange), A pyrrhogluteum (metasoma yellow), and A simpsonorum (metasomal tergites I, II, and III yellow) The size of white spots on the genae also varies from moderately reduced (A costethus winniewertzae, A paurogenum) to extensive (A bromeliphile, A brachymacher) (see Fig Journal of Hymenoptera Research Table Host and host plant information Alphomelon for Alphomelon species Host Plant Species Host Species Chamaedorea costaricana Carystoides basoches arecaphile Chamaedorea Chamaedorea Neoxeniades scipio 2001 Arecaceae tepejilote Bromelia pinguin Bromeliaceae Aehmaea magdalenae Hesperiinae Carina indica Cannaceae Muesebeck 1958 Hesperiinae Panicum pilosum Poaceae (specimens in INBIO) Saccharum officinarum Poaceae (specimens in USNM) On/:ii latifolia Poaceae (specimens in USNM) (grass) Poaceae Hesperiinae (grass) Poaceae Hesperiinae (sedge) Cyperaceae Cobalopsis sp Oryza latifolia Poaceae Hesperiinae Caima sp Cannaceae conformis Hesperiinae Lerema spp (suger cane) disputabile Cymaenes trebius mgriceps talidicida Janzen and Hallwachs 2001 melanoscelis nanosoma Janzen and Hallwachs 2001 no known host records eitroloma crocostitliiis Janzen and Hallwachs no known host records brachymacher bromeliphile Arecaceae tepejilote Chamaedorea costaricana Synale cyhaxa Reference Host Plant Family Janzen and Hallwachs 2001 Janzen and Hallwachs 2001 (specimens in USNM) Volume Table 12, Number 1, 2003 Continued Host Species Alphotnelon Host Plant Species Maranta arundinacea, Host Plant Family Referenc Marantaceae Janzen and Hallwachs Calathea macrosepela Calpodes ethlius Thalia geniculata 2001 Cannaceae Caiuia indica Maranta arundinacea, Rhinthon cubana Calathea macrosepela Janzen and Hallwachs 2001 Marantaceae Calathea lutea xestopyga Maranta arundinacea Janzen and Hallwachs Marantaceae Saliana fusta Calathea macrosepela 2001 Maranta arundinacea Janzen and Hallwachs Marantaceae Cynea irma Calathea panamensis Quinta cannae 2001 Janzen and Hallwachs Marantaceae Thalia geniculata 2001 Larval characters were not examined in Additional characters include coloraorange or barred with black), wings (smoky or hyaline), cast skins for ovipositor (reddish-orange or yellow), and face (with a light brown spot or completely black) Although these color patterns one species in the genus In the key to species, values given 4) tion of legs (completely appear to be largely diagnostic, one when relying on color characters alone since temperature during development is known to influence adult should use caution appearance in braconids (Liu and Carver 1982) Other diagnostic morphological characters frequently referred to in this revision concern hind wing vein cu-a curvature (see Fig 2), tarsal claw spine number (see Fig 3), ovipositor length and curvature (see Fig 6), lunule shape, tegula shape and color, open gite I propodeal sculpturing (areola or closed anteriorly), metasomal terand II shape, and sculpturing of the patches reduced - (Fig 4f), Mesonotum dark brown to in parentheses refer to the percentage of speci- mens examined exhibiting that particular The phrase "cheek patches" refers to the white spots on genae The current abundance of species in museum collections (the museums mentioned uncharacteristic der Material) is designated as: rare (fewer than 10 total specimens), uncommon (1020 specimens), common (20-100 specimens), or abundant (more than 100 specimens) A brief description of the species' natural history and range follows the species names in the key Since most males (>75%) are difficult to identify unambig- claws with spine (Fig 3a); wings infumated; cheek with white not extending to post-genal part of occiput black; tarsal claws variable; wings variably colored; cheek to species, the uously male specimens NEW WORLD SPECIES OF key is ALPHOMELON tarsal patches variable Dias (1985) describes larval features for fe- KEY TO FEMALE Mesonotum orange-yellow; due to limited availability of most species, but Penteado- based on head and mesoscutum this revision Metasoma and mesosoma yellow-orange A nigriceps (Ashmead) Journal of Hymenoptera Research (Common Solitary parasitoid reared from Calpodes spp on Carina indica and unidentified sp on corn (Zca mays) Distributed throughout Caribbean islands and countries /states surrounding Caribbean and reaching as far north as North Carolina, U.S.A.) - Metasoma black, mesosoma with mesonotum only orange A crocostethus Deans, n sp cane (Saccharum qfficinarum) Solitary parasitoid reared from hesperiid on sugar Distributed from southern Caribbean islands to Colombia, and Brazil) (Uncommon Metasoma with at least lateral tergites petiole rectangular and elongate and hypopygium in some) yellow; portion of hind wing cu-a angled sternites (except (Fig 5a); posterior toward body (Fig 2c); tarsal claws with spines (Fig 3b) - Metasoma tarsal claws entirely brown to black; petiole variable; hind wing cu-a variable; with 1-4 spines - VII black; meentirely yellow except medial Vs of hypopygium and tergum A pyrrhogluteum Deans, n sp sopleuron nitid except for setae (Rare Biology unknown Collected in Argentina only) Metasoma less than 50% yellow; mesopleuron punctate and tergites I— III yellow; petiolar ridge weakly represented; malar space not conA simpsonorum Deans, n sp vex; pale cheek patch not extending to clypeus (Rare Solitary parasitoid reared from Hesperiinae on a grass (Poaceae) in Costa Rica Distributed from Costa Rica to Paraguay) Petiole - Petiole and Tergites I — III black; medial distinctly convex; pale - malar space (80%) A citroloma Deans, n sp petiolar ridge strongly represented; cheek patch extending (Common Biology unknown - Metasoma to clypeus (Fig 4c) Distributed from Costa Rica to Argentina) Exserted portion of ovipositor shorter than hind basitarsus (Fig 6a,b); tarsal claws with spines (Fig 3d); hind wing cu-a strongly curved towards body (Fig 2b); cheek patches extending onto clypeus (Fig 4a,b) (75%) Exserted portion of ovipositor longer than hind basitarsus (Fig 6c,d,e); tarsal claws variable; hind wing cu-a variable; cheek patches variable Ovipositor less than length of basitarsus (Fig 6a), orange in color; ovipositor sheaths expanded apically (Fig 6a); face black (Fig 4b); antennae dark brown A brachytnacher Deans, n sp (Common Biology unknown Distributed from Costa Rica to Ecuador, Peru and Brazil.) Ovipositor longer than Vi length of basitarsus (Fig 6b), yellow-orange in color; ovipositor Vi sheaths not expanded apically (Fig 6b); face with light brown spot (Fig 4a); antennae brown A bromeliphile Deans, n sp (Rare Gregarious parasitoid reared from Neoxeniades scipio on Bromelia pinguin and Achmaea magdalenae (Bromeliaceae) in Costa Rica Distributed from southern Mexico to Costa Rica) light Hind - Hind legs black except basal Vi of tibia orange; body size large (~5mm); tarsal claws with spines (Figure 3d); petiole costate with ridge strongly represented (Figure 5b) A melanoseelis Deans, n sp (Common Solitary parasitoid reared from hesperiines on grasses (Poaceae) in Costa Rica Distributed from southern Mexico to Brazil) 10 femur never completely black; body size variable (85% claws with 1-4 spines; petiole variable leg coloration variable but 1.4X hind basitarsus length (Fig 6e); hind wing cu-a strongly angled at midpoint towards body (Fig 2e); 5mm) body size medium to large (~4A talidicida (Wilkinson) (Common Gregarious parasitoid reared from Talides spp on Heliconia (Heliconiaceae) and Musa to spp (Musaceae) in Costa Rica Distributed throughout Caribbean islands and from Costa Rica Brazil) Petiole with only an inconspicuous depression (Fig 5e,f); ovipositor obviously decurved, ¥z as thick as hind basitarsus, with exerted portion