Article. Department of Entomology, University of Illinois at Urbana-Champaign, IL 61801, USA. 3

Zootaxa 2084: 1 49 (2009) www.mapress.com/zootaxa/ Copyright 2009 Magnolia Press Article ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) Review of world Parapanteles Ashmead (Hymenoptera: Braconidae: Microgastrinae), with description of fourteen new Neotropical species and the first description of the final instar larvae A. A. VALERIO 1,3, J. B. WHITFIELD 2 & D.H. JANZEN 4 1 Central American Institute of Biological Research and Conservation (CIBRC). P.O. Box 2398-2050 San Pedro de Montes de Oca, San José, Costa Rica. E-mail: avalerio_13@hotmail.com 2 Department of Entomology, University of Illinois at Urbana-Champaign, IL 61801, USA. jwhitfie@life.uiuc.edu. 3 Corresponding author. Current address: Museum of Biological Biodiversity, Ohio State University, 1315 Kinnear road, Columbus, OH 43212, USA Department of Biology 433 South University Avenue University of Pennsylvania Philadelphia, PA 19104. djanzen@sas.upenn.edu Table of contents Abstract... 2 Introduction... 2 Material and methods... 2 Descriptive Taxonomy... 3 Parapanteles Ashmead... 3 Key to world species of Parapanteles Ashmead (based mainly on females)... 6 Species descriptions... 8 Parapanteles aletiae (Riley)... 8 Parapanteles masoni Austin and Dangerfield... 12 Parapanteles paradoxus (Muesebeck)... 13 Parapanteles rooibos Valerio, Whitfield and Kole... 15 New species descriptions... 15 Parapanteles complexus Valerio and Janzen, n. sp.... 15 Parapanteles continua Valerio and Whitfield, n. sp.... 17 Parapanteles em Valerio and Whitfield, n. sp... 20 Parapanteles lincolnii Valerio & Whitfield, n. sp... 27 Parapanteles mariae Valerio & Whitfield, n. sp.... 28 Parapanteles nephos Valerio & Whitfield, n. sp.... 30 Parapanteles noae Valerio and Whitfield, n. sp... 32 Parapanteles polus Valerio and Whitfield, n. sp... 33 Parapanteles rarus Valerio and Whitfield, n. sp... 35 Parapanteles scotti Valerio and Whitfield, n. sp.... 37 Parapanteles sicpolus Valerio and Whitfield, n. sp.... 38 Parapanteles tesssares Valerio and Whitfield, n. sp.... 40 Parapanteles thrix Valerio and Whitfield, n. sp... 42 Parapanteles tlinea Valerio and Whitfield, n. sp.... 44 Discussion... 46 Acknowledgments... 48 References... 48 Accepted by M. Buffington: 7 Mar. 2009; published: 27 Apr. 2009 1

Abstract Currently only two Neotropical species are known for Parapanteles Ashmead (1900): Parapanteles aletiae (Riley), described from the U.S., Cuba and Puerto Rico, and P. paradoxus (Muesebeck), described from Costa Rica. In the present paper we describe 14 new species from the New World (Parapanteles complexus n.sp., P. continua n.sp., P. em n.sp., P. lincolnii n.sp., P. mariae n.sp., P. nephos n.sp., P. noae n.sp., P. polus n.sp., P. rarus n.sp., P.scotti n.sp., P. sicpolus n.sp., P. tessares n.sp., P. thrix n.sp., P. tlinea n.sp.) and provide an illustrated key for the world species. In addition, we present a synopsis of the ecology of species in the genus using new rearing records and previously published host records. Eighteen species in 11 lepidopteran families are known to be hosts for Parapanteles species, and these hosts feed on plants from 21 families totaling at least 43 species. Key words: Braconidae, Microgastrinae, ecology, taxonomy, Neotropics, new species, host records Introduction The Neotropical microgastrine braconid wasp genus Parapanteles was proposed by Ashmead (1900) to include only the type species, Parapanteles aletiae (Riley). Muesebeck (1922) later synonymized Parapanteles under Apanteles Foerster, but Mason (1981) later resurrected the genus name and the species Parapanteles paradoxus (Muesebeck) in his revision of the genus Apanteles s.l.. Since then only two additional species have been described: P. masoni Austin & Dangerfield from Australia (Austin & Dangerfield 1992), and P. rooibos Valerio, Whitfield & Kole from South Africa (Valerio et al. 2005). Parapanteles has presented some problems for microgastrine generic and tribal classification due to its joint possession of the short Macrolepidoptera Suite ovipositor apparatus generally attributed to Cotesiini and Microplitini, along with possession of a medial propodeal areola characteristic of the Apantelini (Mason 1981). Mason placed Parapanteles among his Cotesiini, proposing a relatively early phylogenetic divergence within that tribe, but Williams (1985) found Mason s placement difficult to reconcile with the existence of other, quite different, apparent phylogenetic transitional forms between Mason s Cotesiini and Microplitini. Since the reanalysis of Mason s phylogenetic work by Walker et al. (1990), Mason s tribal divisions of Microgastrinae have been largely abandoned, but the placement of Parapanteles is still not well established. Recent molecular phylogenetic studies (Mardulyn and Whitfield 1999; Whitfield et al. 2002; Banks and Whitfield 2006) show a relatively close relationship with Hypomicrogaster Ashmead. Taxonomically and biologically, the genus still appears to be poorly known, and the published records for P. aletiae are restricted to unidentified host caterpillars in the families Notodontidae and Noctuidae. According to Muesebeck (1958), P. paradoxus was reared from a species of Hemiceras sp. (Notodontidae) feeding on Inga roussoviana (Fabaceae). More recently, Jacobson (1991) in Perú reared some unidentified Parapanteles wasps from the arctiid larva Carales astur Cramer feeding on Citrus spp. (Rutaceae), broadening the host generic and family spectrum for the genus. In South Africa, P. rooibos is known to attack a geometrid moth, Isturgia exerraria, feeding on rooibos tea plants (Valerio et al. 2005). The diversity of host records suggests a relatively broad host range that may still be poorly sampled. The paucity of host records also reflects the taxonomically uncharacterized composition of the genus in the Neotropics (Mason 1981). The objectives of the present paper are to describe a number of recently reared Parapanteles species from the New World assembled from several large-scale rearing projects, and to present an updated key for the known species. A brief review of host use and geographic distributions is also provided, as are some comments about the taxonomy of the genus. Material and methods While DHJ is a coauthor of the paper, he explicitly prefers to not be a coathor of the species names. 2 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

Specimens were identified to genus using the key to genera of microgastrines by Whitfield (1997). Species were identified by comparison of specimens with previous published descriptions as well with type specimens from the U.S. National Museum of Natural History, Washington D.C. (USA). Most morphological terminology used in the species descriptions is that of Huber and Sharkey (1993) and Schuh (1989), except propodeal terms, which follow Townes (1969, Fig. E). The cuticular sculpture terminology follows that of Harris (1979), while the terminology for the wing venation is a variation of the Comstock Needham system used by Sharkey and Wharton (1997, Fig. 15). A series of morphological measurements were used to describe the species (see figures 1 and 2) using a micrometer mounted in a stereomicroscope. The measurements were taken from a maximum of 20 specimens per species when possible. The author of each host species does not appear on the main text, but all authors are given in Table 1. The genus description follows the terminology of Whitfield et al. (2002) to facilitate further comparisons with other previously described genera. Scanning electron micrographs for the type specimens were taken using a Philips XL30 ESEM-FEG electron microscope at the Beckman Microscopy Suite (University of Illinois at Urbana-Champaign). Wing illustrations were traced in Adobe Illustrator 10 after digital photographs were taken using a JVC GC-QX5HD digital still camera mounted on a Leica MZ125 stereomicroscope. Female genitalia were slide-mounted using Euparal mounting media (BioQuip Products) after an overnight immersion in 10% KOH and exposure to 80% and 90% alcohol before final mounting. Genitalia were traced using a micro-slide projector and then inked. Larval characters for Parapanteles species were acquired using the protocol proposed by Wahl (1984) [using lactophenol instead of Nesbitt s solution] for reared cocoons remains, and then larval characters were scored using the approach of Whitfield et al. (2002). The larval morphological terminology follows Mason (1981).The rearing project of D. Janzen and W. Hallwachs (http://janzen.sas.upenn.edu/caterpillars/ database.lasso) at the Area de Conservación Guanacaste (Costa Rica, Guanacaste) is the source for many of the host-plant associations, host records, cocoon material and specimens of the Neotropical species. All Janzen and Hallwachs voucher codes (yy-srnp-xxxxx) refer to the event-based record of finding the caterpillar and rearing it. Individual wasps later treated for DNA barcoding or other individual-specific actions are assigned an additional unique voucher code in the form of DHJPARxxxxxxxx, which can be obtained from the yy-srnp-xxxxx voucher-coded record on the Janzen and Hallwachs web site. All, rearing records are reported as they appeared in Janzen and Hallwachs database at the time of this paper preparation except for the holotype information recently updated; for a current state of the motioned host records please consult the Janzen & Wallwachs database.additional sources are the Jacobson (1991) rearing records from Perú, and some rearing records from the Ozarks in the U.S by Whitfield et al. (1999). Sex ratios reported in this investigation are given with the following format: number of males/ number of females (i.e. 3/4, meaning 3 males and 4 females). Specimens were obtained on loan from the following institutions, whose abbreviations are used as follows: Australian National Insect Collection, Canberra, Australia (ANIC); Illinois Natural History Survey, Urbana, USA (INHS); Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica (INBio) and U.S. National Museum of Natural History, Washington D.C., USA (USNM). Descriptive taxonomy Parapanteles Ashmead Parapanteles Ashmead 1900: 131 Type species: Apanteles aletiae Riley Adult description. Head: Basal antennal placodes 2-ranked and with same pattern of arrangement ventrally and dorsally on midapical segments; glossa truncate. REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 3

Mesosoma: Propleuron with simple (unflanged) lower outer corner; pronotum with two pronotal lateral grooves; epicnemial carina absent; scutellar lunulae lunate in outline, midposterior band of scutellum nitid; metanotum sublateral setae present as distinct tuft; propodeum with anterodorsal area greatly shortened, medial longitudinal carina usually absent but sometimes represented by two anterior parallel carinae or short anterior carina; areola present and complete, transverse carinae present and well defined; hind leg with coxa extending far beyond first metasomal tergum, and tarsal claw with basal tooth; fore telotarsus varying from unmodified to exhibiting set of organized setae ventrally to exhibiting conspicuous hooklike seta ventrolaterally. FIGURE 1. Head morphological measurements used for the species descriptions: A Face width, B Intertentorial pit distance, C Tentorial pit-compound eye distance, D Face width at clypeus edge, E Basal width of mandible, F Malar space, G Clypeus height, H Clypeus width, I Compound eye height, J Head height, K Compound eye width, L Head width, LL Inter ocellar distance, M Ocell-ocular distance, N Face width at upper edge of posterior ocelli, O Torulus-anterior ocelli distance, P Terminal flagellomere length, Q Subterminal flagellomere distance, R Terminal flagellomere width, S Third flagellomere length, T Second flagellomere length and U First flagellomere length. 4 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

FIGURE 2. Wing venation and hind leg morphological measures taken for the species description: a) Forewing: A Pterostigma length, B Pterostigma width, C (RS+M)a length, D 1M length, E 1RS length, F M+CU length, G m-cu length, H 1Cub length, I 1CUa length, J 2r-m length; b) Hind wing: K 1RSa length, L 2M length, LL 1r-m length, M 1M length, N M+CU length, O cu-a length, P 1A length; Hind leg: Q Femur length, R Tibia length and S Femur width. FIGURE 3. Propodeum of Parapanteles rooibos Valerio et al. (A); P. nephos n. sp. first metasomal tergum in dorsal view (B), and P. tlinea n. sp. transversal lineate sculpture on mesopleuron (arrow points at it) (C). REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 5

Metasoma: First metasomal tergum with basal excavation and midapical groove both present, variably shaped from almost parallel-sided to abruptly widening posteriorly, junction with second tergum fused; second metasomal tergum normally nitid except one species with longitudinal lineate sculpture present on the second and third tergum, shape from rectangular to triangular; second and third metasomal terga delineated by a fine distinct suture; third metasomal tergum not carapacelike and normally nitid; hypopygium not formed into longitudinal pleats medially, instead sharply folded and evenly sclerotized; female genitalia with eighth tergite 2 4 times taller than long, second valvifer tall, lorate and not expanded apically, ovipositor sheaths attached below midpoint of second valvifer, sheaths normally short but some species more elongate; ovipositor and sheaths mainly hidden by hypopygium, setation mostly only on distal tip of ovipositor sheaths or on distal 2/3 of ovipositor length; ovipositor compressed laterally, short and normally very thick at base, but some species exhibiting more elongate and slender ovipositor. Male genitalia: Apex of digitus acute and directed dorsally, but its ventral edge strongly convex. Wings: Forewing vein 3RS straight or weakly curved, vein 1M+1Rs weakly angled between 20 and 60, veins r-m and 2A absent; hind wing with cells 1R and 2R of approximately same width, vein 2r-m present, vein 2A absent, vein 2Cu mostly straight and not sinuate, vannal lobe separated from remainder of wing by a sharp notch, convex in shape and with setae dense and conspicuous throughout margin. Larval description. Antenna absent; mandible blade with less than five distal teeth, and apex simple (not divided); larval maxilla with two setae; larval labium with four setae; larval skin densely covered by papulae which bear a central spine. Quick diagnosis. Basal antennal placodes 2-ranked and with same pattern of arrangement ventrally and dorsally on midapical segments; propodeum with areola present (sometimes not so evident); metasomal terga 1-3 often (not always) coarsely sculptured but neither fused, broad in shape nor carapace like; first metasomal tergum with mid distal depression present; hypopygium sharply folded medially (normally) and ovipositor usually short and thick in lateral view; hind wing with vannal lode evenly convex and forewing areolet open distally. Key to world species of Parapanteles Ashmead (based mainly on females): 1 Propodeal areola very inconspicuous but complete (Fig. 3A); propodeum posterior lateral areas and areola nitid, anterior portion of propodeum with smooth confused rugulose sculpture; fore telotarsus with hooklike seta at ventral area; ovipositor short and thick in lateral view; ovipositor sheaths with dense setation constrained at distal 1/3; body mainly black in color as ovipositor sheaths; from Africa... P. rooibos Valerio, Whitfield & Kole - Areola well defined and completely present, sometimes partly obscured by confused rugulose sculpture present on propodeum (Fig. 5A vs. 5D); propodeum otherwise variable in sculpture; fore telotarsus, ovipositor sheaths, ovipositor and body color variable; from the New World or Australia... 2 2 Areola open at anterior edge and propodeum without other sculpture except for the areola, transverse and posterior 1/2 of costular carinae (Fig. 4F); ovipositor shape in lateral view short and thick; fore telotarsus medial edge with a conspicuous hooklike setae (Fig. 4A); ovipositor sheaths short and mainly concealed by hypopygium; head and mesosoma black, metasoma mainly dark brown with the interpleural membrane contrastingly white; from Australia... P. masoni Austin & Dangerfield - Areola normally closed at anterior edge, if open then different in shape from above (Fig. 6C); ovipositor and ovipositor sheaths elongate and relative thin in lateral view (Fig. 10A); fore telotarsus, ovipositor, ovipositor sheaths and body color variable; from the New World... 3 3 Clypeus, labrum and mandibles totally black; basal segments of palpi black, remainder yellowish; propodeum with midlongitudinal carina crossed by semicircular carinae in anterior 1/3 (Fig. 7A); propodeal spiracle present on an elevated and expanded area... P. lincolnii Valerio & Whitfield n. sp. - Without the above combination of characters, especially propodeum lacking semicircular carina crossing anterior 1/3 of midlongitudinal carina; palpi not bicolored; propodeal spiracle variable... 4 4 First metasomal tergum I -shaped and with sparse longitudinal lineate sculpture anterolaterally plus dense and fine rugulose sculpture, remainder of tergum with coarser rugose sculpture, tergum posteriorly with depressed midlongitudinal area with only rugose sculpture, posterior edge with circular depression at each corner (Fig. 14A); head totally black except palpi yellow... P. complexus Valerio & Janzen n. sp. 6 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

- First metasomal tergum not I -shaped; variable in sculpture but never as described above; head variable in pattern of color... 5 5 Third metasomal tergum with lineate sculpture mixed with confused colliculate sculpture in triangular area that can cover the anterior 1/3 of tergum; legs yellow, except tarsomeres and distal tip of femur fuscous; first metasomal tergum with lineate sculpture present basally; propodeum totally covered by conspicuous areolate-rugose sculpture (Fig. 6A)... P. scotti Valerio & Whitfield n. sp. - Third metasomal tergum without sculpture, or if sculpture present then very sparse and limited to anterior edge; leg color normally yellow and brown but never with hind femur mostly yellow; without lineate sculpture on basal area of first metasomal tergum; propodeum variable in sculpture... 6 6 First metasomal tergum strongly narrowed posteriorly; second metasomal tergum exhibiting a sharply triangular shape in dorsal view (Fig. 8B); ovipositor strongly elongate and narrow in lateral view; ovipositor sheaths with few long widely spaced setae (Fig. 9B)... P. rarus Valerio & Whitfield n. sp. - First metasomal tergum never conspicuously narrowed to posterior edge, usually wider at posterior edge in dorsal view; ovipositor sheaths variable in setation and shape... 7 7 Propodeum exhibiting only the areola, transverse and anteromedial longitudinal carinae, remainder nitid (Fig. 6C); body color reddish brown or light brown with yellow... 8 - Propodeal sculpture not limited to major carinae as described above, with some kind of sculpture anterolaterally or posterolaterally and/or in areolar area (similar to Fig. 7B, 7D); body dark brown or with a combination of brown with yellow colors... 11 8 Second metasomal tergum nitid (Fig. 6D, 11D), pronotum with lateral grooves deeply impressed; body color mainly reddish brown with legs yellow... 9 - Second metasomal tergum with sculpture present over most of surface (Fig. 11 A&E), pronotum with lateral grooves variable; body color also variable... 10 9 First metasomal tergum mostly sculptured (Fig. 6D); ovipositor short and thick and barely exerted from hypopygium (Fig. 6B); ovipositor sheaths with short dense setae on distal 1/5 (Fig. 6B); wings with faint yellowish color... P. aletiae (Riley) - First metasomal tergum essentially nitid (Fig. 3B); ovipositor elongate and strongly exerted from hypopygium; ovipositor sheaths with sparse long setae area covering at least 1/2 of their lengths; palpi, tegula and pterostigma yellowish; wings slightly infuscate... P. nephos Valerio & Whitfield n. sp. 10 Fore telotarsus with a long and thin hooklike seta medioventrally (similar to Fig. 4G); body dark brown and yellow; scrobal areas shallow and widening distally; pronotum with lateral grooves present but not conspicuously impressed... P. noae Valerio & Whitfield n. sp. - Fore telotarsus with a set of 4 setae ventrodistally (similar to Fig. 4B); body totally reddish brown, except anterior and distal portions of leg yellowish; antennal scrobal areas deep at basal area but narrowing distally; pronotum lateral grooves deeply impressed... P. continua Valerio & Whitfield n. sp. 11 Mesopleuron with confused rugulose sculpture on upper half of anterior edge, remainder of anterior edge with transverse lineate sculpture (Fig. 3C); propodeum with areola divided horizontally by a thick and sinuate transverse carina (Fig. 5E)... P. tlinea Valerio & Whitfield n. sp. - Mesopleuron with punctate sculpture on anterior edge, never with transverse lineate sculpture; areola variable, if divided horizontally then by more than one horizontal carina mixed with rugulose sculpture (Fig. 5A, 7B)... 12 12 Telotarsus of all legs with four thick setae on distal 1/3 (Fig. 4B, 4C); ovipositor thickened at base but elongate in lateral view (Fig. 9F, 10C)... 13 - Telotarsus without a set of four setae on distal 1/3 of ventral side, or if setae present, not arranged as described above (Fig. 4D, 4H); ovipositor thick and short in shape (Fig. 9D, 10E)... 14 13 Ovipositor sheaths elongate in lateral view with setation covering at least distal 1/3 (Fig. 10C); first metasomal segment with areolate-rugulose sculpture throughout (Fig. 11A); face midlongitudinal area with less dense punctuate sculpture than remainder and triangular shaped elevated area present; body length greater than 2.8 mm...... P. sicpolus Valerio & Whitfield n. sp. - Ovipositor sheaths with setation limited to distal 1/5; first metasomal tergum with midlongitudinal area bearing confused rugulose sculpture over 2/3 of its length, remainder tergum with confused foveate sculpture except basal 1/5 with sparse confused and smooth rugulose sculpture; face with punctate sculpture except medial 1/3 almost nitid, without any triangular shaped elevated area; body length less than 2.7 mm... P. tessares Valerio & Whitfield n. sp. 14 Ovipositor relatively slender in lateral view (Fig. 9A, 10D); hypopygium long, between 0.48-0.75 mm; ovipositor sheaths elongate and covered by sparse long setae on most of distal 1/3... 15 - Ovipositor broad in lateral view (Fig. 9C, 10E); hypopygium sort in length, not as described above; ovipositor sheaths with setation only at distal tip... 16 15 First metasomal tergum almost perfectly rectangular in dorsal view (Fig. 8F); areolar area covered with sculpture throughout; fore and mid leg mostly yellow; sternauli devoid of sculpture... P. polus Valerio & Whitfield n. sp. - First metasomal tergum wider posteriorly than anteriorly (Fig. 8A); areolar area mostly nitid, without sculpture; fore REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 7

and mid leg yellow and dark brown in color; sternauli with transverse lineate sculpture throughout...... P. thrix Valerio & Whitfield n. sp. 16 Fore telotarsus with a long and narrow hooklike seta ventrally (Fig. 4H); labrum usually black in color (95%); axilla through mesonotum with series of well defined ridges; ovipositor sheaths broad in lateral view and with setae almost throughout (Fig. 9D); body size between 2.35 2.50 mm, very stout in lateral view...... P. paradoxus (Muesebeck) Mason. - Fore telotarsus and ovipositor sheaths variable in shape and setation; labrum variable in color but never black; axilla through mesonotum mainly nitid except at anterior lunular edge (with short and thick ridges), remainder of axilla nitid; ovipositor sheaths setation located on distal 1/3... 17 17 Fore telotarsus with two thick straight setae ventrally (Fig. 4D); ovipositor sheaths setation sparsely and shape of sheaths in lateral view slender (Fig. 10E); labrum brownish yellow... P. mariae Valerio & Janzen n. sp. - Fore telotarsus with one long narrow hooklike seta ventrally (Fig. 2H); ovipositor sheaths with dense setation and with broad shape in lateral view (Fig. 9C); labrum reddish-orange... P. em Valerio & Whitfield n. sp. Species descriptions Parapanteles aletiae (Riley) Figs. 4G, 5C, 6B, 6D Apanteles aletiae Riley, 1869, 4: 296, 298, 306. Parapanteles aletiae, Mason, 1981, 115: 102. Female. Body length = 1.60 2.10 mm. Body color: Legs (except hind coxae anterior 1/2 and tarsal claws) light yellow, as is most of metasoma (except terga, posterior area of metasoma), ovipositor, tegula, labrum, palpi, scape (except distal edge light brownish yellow), pedicel, propleuron anterior edge and posterolateral flange; remainder of antenna brownish yellow; ocelli yellowish; remainder of body light brown as on hind telotarsus and tarsal claws. Wings hyaline, forewing veins translucent except pterostigma and C+SC+R with very light yellow color; hind wing veins translucent. Head. Head height/width = 1.27; compound eye height/width = 1.60; intertentorial pit distance = 0.15 0.18 mm; tentorial pit distance/distance tentorial pit to compound eye = 3.00 3.75; width of face at dorsal clypeal edge = 0.30 mm; clypeus width/height = 4.00 4.25; vertex width/distance between anterior ocelli and edge of torulus = 2.06 2.28; length of first flagellomere = 0.18 mm; first flagellomere length/width = 2.80; length of first flagellomere/length of second flagellomere = 1.00; length of first flagellomere/length of third flagellomere = 1.00; terminal flagellomere length = 0.11 mm; terminal flagellomere length/penultimate flagellomere length = 1.80; terminal flagellomere length/width = 2.25; malar space height/basal width of mandible = 0.86; ocell-ocular distance/lateral ocelli distance = 1.40 1.50. Clypeus with fine punctulate sculpture; face with dense fine punctate sculpture mixed with confused, upper keel present without strong cristate; frons with scrobal areas nitid and shallow near toruli, frons laterally with fine punctate sculpture, distal area more glabrous; vertex with dense punctulate sculpture; temple and gena with dense, shallow and confused punctate sculpture extending to basal 2/3 of postgena near junction area with gena; remainder of postgena nitid. Mesosoma. Mesosoma length = 0.90 0.92 mm; mesosoma length/width = 1.36 1.48; mesosoma height = 0.61 0.64 mm. Propleuron with fine punctate sculpture throughout except nitid anterolateral 1/3; pronotum laterally with anterior 1/3 well impressed and essentially nitid except for two short transverse ridges, ventrolateral groove crossed by short transverse ridges throughout its length, dorsolateral groove also well impressed but transverse ridges only present over distal 1/4; area between grooves nitid, dorsal edge 1/5 and ventral edge 1/3 of posterior edge height; mesonotum with dense punctate sculpture that almost reaches the scutellar groove; scutellar groove with 12 small costulae, medial ones more clearly defined and larger than lateral ones; scutellum with large but shallow and scattered punctations laterally, medial area essentially nitid, scutellum lateral area with narrow costulate sculpture which become narrower and elongated towards 8 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

FIGURE 4. Parapanteles masoni Austin & Dangerfield fore telotarsus (A) and pronotum in dorsal view (F); fore telotarsus structure of P. tessares n. sp. (B), P. sicpolus n. sp. (C), P. mariae n. sp. (D), P. scotti n. sp. (E), P. aletiae (Riley) (G), P. paradoxus (Muesebeck) (H), and P. rarus n. sp. (I). The arrows in these figures point to the different setae on the fore telotarsus. REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 9

FIGURE 5. Propodeum of Parapanteles tessares n. sp. (A), P. sicpolus n. sp. (B), P. aletiae (Riley) (C), P. em n. sp. (D), P. tlinea n. sp. (E), and P. continua n. sp. (F). posterior edge; axilla through mesonotum mainly nitid except for a series of transverse narrow ridges on anterior edge of lunulae; metanotum subrectangular and nitid; axilla through metanotum with short transverse ridges on distal edge, remainder nitid; propodeum with anteromedial longitudinal and transverse, areolar and posterior part of costular carinae cristate and well defined, interior of areola posteriorly with sparse light rugulose sculpture, anterolateral areas with punctate sculpture, remainder of propodeum nitid; mesopleuron with anterior 1/4 punctate, sternaulus forming an elongate depression, dorsal edge with anterior 1/3 nitid, remainder with well defined, dense and fine scrobiculate sculpture, posterior edge nitid, remainder of 10 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

mesopleuron nitid; metapleuron with dorsal and posterior edge bearing dense narrow transverse ridges, nearby with shallow and confused punctate sculpture, small medial pit present, pleural suture present, remainder of metapleuron nitid. FIGURE 6. Parapanteles scotti n. sp. propodeum (A); P. aletiae (Riley) female genitalia (B) and first metasomal tergum in dorsal view (D); P. nephos n. sp. propodeum (C) and wings (F), and P. rooibos Valerio et al. female genitalia (E). Legs. Hind femur length = 0.50 0.60 mm; hind femur length/width = 3.69 5.00; hind tibia length/hind femur length = 1.12 1.13. Fore telotarsus with one large elongate hooklike seta set on ventral area located on a small lobe at mid length of telotarsus, ventral area below hooklike seta bare and concave in shape, in lateral view the REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 11

telotarsus slightly bent dorsally, shorter in length than basitarsus; hind telotarsus in dorsal view slightly bent and with an elongate setae on anterior 1/3. Wings. Forewing length = 2.30 2.45 mm; 1RS length = 0.09 0.10 mm; 1CUa length/1cub length = 0.81 0.86; length RS+Ma = 0.35 mm; length M+CU = 0.75 0.78 mm; 1M length/m-cu length = 1.86 2.00; pterostigma length/height = 1.25 1.54. Hind wing: 1M length = 0.34 0.35 mm; 1M length/2m length = 1.69 1.75; 1M length/m+cu length = 1.33 1.50; length r-m/length cu-a = 0.75 0.88; 1RSa length/2r-m = 1.25; 1A length = 0.20 0.21 mm. Metasoma. First tergum basal width = 0.24 0.26 mm; first tergum length/distal width = 1.39 1.5; second tergum length/distal width = 0.27 0.30; third tergum length/distal width = 0.37 0.38; hypopygium length = 0.32 0.34 mm. First metasomal tergum parallel-sided and carinate at edges throughout its length, surface with confused aciculo-rugose sculpture except on basal 1/4 which is nitid; second tergite mainly nitid except for very sparse punctate sculpture on anterior edge of tergite; remainder of tergite nitid; ovipositor short and thick at base, barely exserted from hypopygium; ovipositor sheaths with dense setation over distal 0.2. Variation. The following color variation was observed: scape and pedicel totally brownish yellow or with basal 1/2 light yellow and remainder brownish yellow; forewing with pterostigma and C+SC+R and all hind wing veins with a very faint yellowish color, not white as described in Riley (1869). Material examined. Holotype: female, USNM type number 2771, 2340, Hubblard s (Maybe Hubbard s?) No. 71c. under the name Apanteles aletiae Riley. Paratypes, two females, USNM with type number 1171, 2340, Hubbard s No 71(c). One female identified by Gahan as Apanteles aletiae, P.[uerto] R.[ico], Camuy, Acc330-1922, 5 Oct. 1922, G.N. Wolcott collection. Holotype and paratypes are at USNM. The type specimen is missing the head. Comments. The rearing data for P. aletiae are very meager; the known hosts are larvae of noctuid and notodontid, with only family level of identification at this time. The known distribution is the southeastern United States (Mason 1981, p. 104). According to Riley (1869), the specimens from the type series were reared from Aletia argillacea (currently Alabama argillacea (Hubner), cotton worm) collected from August to October in Florida and Alabama. The type specimen has the head and hind legs missing; the paratypes are in good condition but are totally covered with fungal hyphae. Parapanteles masoni Austin and Dangerfield Figs. 4A, 4F Parapanteles masoni Austin & Dangerfield, 1992: 6: 45 47. Female (Additional descriptive features to Austin and Dangerfield s original description). Mesosoma. Propleuron with anterior edge nitid and remainder with punctate sculpture throughout its length; pronotum lateral area with anterior 1/3 with deep longitudinal and nitid grove, dorsal lateral groove with few smooth transversal ridges across its width and lower lateral groove but the latter with fewer and more thick than those present at dorsal groove, area between lateral grooves with nitid, dorsal edge with posterior edge with dense and confuse punctate sculpture, dorsal edge 1/4 and ventral edge 0.4 posterior edge height; scutellar groove with 11 narrow and well defined costulae; scutellum with big and confused punctate sculpture at anterior lateral 2/3 of its length, medial area with very sparse punctate sculpture an essentially nitid, lateral area with big and well defined costulate sculpture slightly narrower at anterior 1/5; axilla through mesonotum with transversal ridges that reach across 1/2 of its width, transversal ridges are divided at mid length by one carina, remainder axilla nitid; metanotum mainly nitid and subrectangular in shape, mid posterior costulae with two lateral depressions at posterior 1/2 and with remainder of area with few smooth rugulose sculpture. 12 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

Wing. Hyaline, forewing with veins r and C+SC+R yellowish brown and pterostigma, remainder of veins whitish yellow; hind wing veins whitish yellow. Legs. Fore telotarsus shorter than fore basitarsus, fore telotarsus with a long hooklike setae at internolateral area of it. Material examined. Holotype, female, [Australia], Finniss Riv. Stn., N.T., coll. 17 July 1986, [Col.] C. Wilson, on leaf Mimosa pigra [L. (Fabaceae)]. Paratype: one male with same data as holotype. Comments. All material deposited at ANIC. This is the only known species for the genus Parapanteles to be described and recorded from the Australian continent. Parapanteles paradoxus (Muesebeck) Figs. 4H, 7E, 9D, 11C, 12E Apanteles paradoxus Muesebeck, 1958: 418, 446 448. Parapanteles paradoxus (Muesebeck) Mason, 1981: 104. Female. Body length = 2.35 2.50 mm. Body color: Fore and mid leg yellow (except coxae), hind leg trochanters and basal area of femur and basal tip of tibia, metasomal sterna and pleura (except hypopygium), ovipositor, tegula, ocelli, compound eyes silver; fore and mid coxae honey brown and remainder of hind femur, labrum, propleuron, pronotum, tarsal claws, hypopygium, hind tarsomeres and tibia; palpi whitish yellow; remainder of body dark brown. Wings hyaline, forewing veins light brownish yellow with pterostigma and veins r and 2RS more darker than remainder of veins, also anterior 2/3 of vein 1-1A transparent as basal area of M+CU; hind wing veins light yellow except for veins SC+R and R1 light brownish yellow. Head. Head height/width = 1.28 1.30; compound eye height/width = 1.47 1.55; intertentorial pit distance = 0.18 mm; tentorial pit distance/distance tentorial pit to compound eye = 2.33 2.80; width of face at dorsal clypeal edge = 0.32 0.35 mm; clypeus width/height = 2.42 2.67; vertex width/distance between anterior ocelli and edge of torulus = 2.67 2.76; length of first flagellomere = 0.22 mm; first flagellomere length/width = 1.00; length of first flagellomere/length of second flagellomere = 1.00; length of first flagellomere/length of third flagellomere = 1.00; terminal flagellomere length = 0.13 0.15 mm; terminal flagellomere length/ penultimate flagellomere length = 1.38 1.50; terminal flagellomere length/width = 2.40 2.75; malar space height/basal width of mandible = 1.00; ocell-ocular distance/lateral ocelli distance = 1.25 1.42. Clypeus with shallow and confused punctate sculpture; face with shallow and confused punctate sculpture at lateral areas of it, central area with an elevated medial keel which does not exhibit any punctate sculpture; frons with scrobal areas nitid and very shallow, lateral areas of frons with fine punctate sculpture, distal area almost nitid except for few punctulate sculpture; vertex with few punctulate sculpture except at lateral and distal areas with dense fine punctate sculpture as gena and junction area between gena and postgena; gena with ocular ring nitid; remainder of postgena nitid. Mesosoma. Mesosoma length = 0.98 1.20 mm; mesosoma length/width = 1.20 1.30; mesosoma height = 0.62 0.70 mm. Propleuron with sparse fine punctate sculpture mainly at posterior 1/2 of it, anterior and externolateral areas almost nitid; pronotum with lateral anterior 1/3 with transversal narrow ridges, dorsal and ventral grooves with smooth, confused and dense transversal ridges which are not evident, area between lateral grooves nitid as dorsal and ventral edges, dorsal edge 1/3 and ventral edge 1/5 of posterior edge height; mesonotum with dense punctate sculpture which becomes slightly denser and smaller at midlongitudinal areas of lateral area of distal 1/2 of mesonotum, sculpture almost reaching scutellar groove; scutellar groove deeply impressed and with few transversal division which are not well defined; scutellum with punctate sculpture throughout its length which is more dense at anterior 1/2 of it, lateral areas with well defined and narrow transversal ridges except distal costulae wider and thicker in shape; axilla through mesonotum with transversal ridges across its width and with transversal ridges next to anterior edge of lunulae; metanotum REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 13

very narrow and normally nitid, sometimes exhibits few irregular lineate sculpture next to middistal costula; axilla through metanotum with three transversal short ridges and with two upper longitudinal carinae; propodeum with confused rugose sculpture throughout its length and areolar carinae, costula, midlongitudinal and transversal carinae, anterior lateral areas with few punctate sculpture at medial area, spiracular areas with rugose sculpture too, costula totally present; mesopleuron with anterior edge punctate, posterior edge nitid, dorsal edge with big and few transversal ridges at anterior area of it but remainder of edge with dense and well defined scrobiculate sculpture, sternaulus as a longitudinal depression which is nitid as remainder of mesopleuron; metapleuron mainly nitid, medial pit present an dorsal and posterior edges with transversal ridges, posterior edge with more spaced sculpture and mixed with few confused rugulose sculpture, remainder nitid. Legs. Hind femur length = 0.65 0.68 mm; hind femur length/width = 3.38 3.70; hind tibia length/hind femur length = 1.26 1.27. Telotarsus of fore leg with a hooklike setae at internolateral area of it and not set of 4 setae present at ventrodistal area of telotarsus; hind telotarsus with long setae at internolateral area of it; hind tarsal claw with a basal tooth in one of the claws and the other one simple. Wings. Forewing length = 2.6 2.65 mm; 1RS length = 0.06 0.07 mm; 1CUa length/1cub length = 0.88; length RS+Ma = 0.38 0.40 mm; length M+CU = 0.81 0.84 mm; 1M length/ m-cu length = 2.13 2.14; pterostigma length/height = 1.12 1.18. Hind wing: 1M length = 0.43 0.46 mm; 1M length/2m length = 1.17 1.80; 1M length/m+cu length = 1.23 1.59; length r-m/length cu-a = 0.71 0.81; 1RSa length/2r-m = 0.43 0.46; 1A length = 0.22 0.28 mm. Metasoma. First tergum basal width = 0.98 1.02 mm; first tergum length/distal width = 1.23 1.25; second tergum length/distal width = 0.44 0.45; third tergum length/distal width = 0.32 0.33; hypopygium length = 0.32 0.35 mm. First metasomal tergum with coarsely rugulose sculpture and basal 1/3 with more confused rugulose sculpture present at medial area and less conspicuous coarsely rugulose sculpture at lateral areas; second metasomal tergum with coarsely rugulose sculpture throughout its area, remainder of terga nitid; hypopygium short in length; ovipositor sheaths with distal 1/3 sparsely setose; ovipositor very thick at basal 1/ 2, distal 1/3 normal and thin, as long as 1/5 as hind tibial length; hypopygium slightly divided at posterior edge. Material examined. Holotype, female USNM Type No. 6307, Costa Rica (San José), Paso Ancho, San Sebastián, 13/xii/1936, No. 5060, C.P. Ballou. Other material: Costa Rica, Alajuela, ACG, Sector San Cristóbal, Sendero Vivero, 730 m, 316490 N 384531 E, 27/viii/2001, Col. F. Quesada; (2) Costa Rica, Puntarenas, A.C.L.A.P., P.I.L.A., Buenos Aires, Sabanas Esperanza, 1800 m, 15-23/iii/1999, Col. R. Villalobos, amarilla. LN 337300_570200, #52480; Costa Rica, Buenos Aires, PILA, Sector Altamira, A.C. Amistad, 1150 1400 m, Junio 1994, Col. R. Delgado, LN 331500_571700, #3015. The Costa Rican material is deposited at INBio except holotype. Male. Similar to female. Comments. Sometimes the fore leg telotarsus is darker in color than remaining tarsomeres; also the propleuron and pronotum can be dark brown. Rearing records. The type specimen was reared from Hemiceras sp. (Notodontidae) feeding on Inga roussoviana (Fabaceae). Janzen and Hallwachs specimens that morphologically match P. paradoxus have been reared from larva of Tithraustes seminigrata (Notodontidae) feeding on Chamaedorea tepejilote (Arecaceae); Dunama mexicana (Notodontidae) feeding on Chamaedorea costaricana (Arecaceae); Rosema sp. (Notodontidae) feeding on Inga oerstediana (Fabaceae); Elachistidae larva feeding on Dichapetalum sp. (Dichapetalaceae) and Hemiceras conspirata (Notodontidae) feeding on Inga vera (Fabaceae). According to Muesebeck (1958) P. paradoxus was reared from other unidentified Hemiceras species at Santa Clara (El Salvador) by P.A. Berry. The observed sex ratio was 1/6. Voucher numbers: 00-SRNP-11175, 00-SRNP-8625, 00-SRNP-8626, 01-SRNP-3235, 02-SRNP-431, 02- SRNP-434, 02-SRNP-437, 02-SRNP-1455, 02-SRNP-1532, 03-SRNP-3261, 03-SRNP-3264, 03-SRNP- 3267, 03-SRNP-7519, 03-SRNP-3268, 03-SRNP-3269, 03-SRNP-3265, 03-SRNP-3260, 04-SRNP-55287, and 04-SRNP-55754. However, when these specimens were DNA barcoded and carefully compared with 14 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

many other rearing Parapanteles rearing records, under the protocol for species discovery through barcoding as described by Smith et al (2008) for other microgastrine wasp genera collected in the same inventory, they have been found to be 4 species that will be described and discussed elsewhere, one parasitizing Hemiceras and Rosema, one parasitizing Dunama mexicana, one parasitizing Tithraustes noctiluces and one parasitizing Elachistidae Parapanteles rooibos Valerio, Whitfield and Kole Figs. 3A, 6E, 11D Parapanteles rooibos, Valerio, Whitfield & Kole, 2005, Zootaxa, 855, 1 8. Female. Body length = 2.05 2.40 mm. Body color: Palpi yellowish (except basal segment) as fore femur distal 1/2, fore tibia and tarsomeres, mid leg distal tip of femur and basal 1/5 of mid tibia, hind tibia basal tip, distal half of mandibles and ovipositor; tarsal claws dark brown; composed eyes silver; ocelli dark orange; spurs of hind tibia whitish yellow as very basal pleura of metasoma; remainder of body black as ovipositor sheaths. Wings hyaline; forewing with veins translucent except pterostigma, 2RS, 2M, r and C+SC+R (except basal 1/4 whitish yellow); hind wing veins translucent except basal area of C+SC+R and distal tip of R1 brownish yellow. For a more extensive description of the species see Valerio et al. (2005). Rearing records and biology. Parapanteles rooibos was reared from Isturgia exerraria (Prout) (Geometridae: Ennominae) feeding on the legume Aspalathus linearis (commonly known as the rooibos tea plant) in South Africa. Material examined. Holotype, female, South Africa, Cederberg region, October 2003, Col. M. Kole. Paratypes six females and one male with same data as holotype. Holotype deposited in South African National Collection of Insects (SANC), Pretoria; paratypes in South African National Collection and in U. S. National Museum (USNM). Comments. This is the only known Parapanteles species from the African continent. New species descriptions Parapanteles complexus Valerio and Janzen, n. sp. Figure 14. Male. Body length = 3.65 mm. Body color: Palpi light yellow; fore leg light brownish yellow (except coxae) and mid and hind leg trochanter and trochanters, posterior tip of mid leg, anterior 1/5 of hind femur; anterior tip of hind basitarsus whitish and tibial spurs; ocelli yellow; compound eyes black with few silver lines (at least in the dead specimen); metasomal pleurites below first tergum brownish yellow; remainder of body dark brown. Wings hyaline, forewing veins light brown, hind wing with vein C+CS+R slightly more yellowish than remainder of veins. Head. Head height/width =1.34; compound eye height/width = 1.57; intertentorial pit distance = 0.19 mm; tentorial pit distance/distance tentorial pit to compound eye = 3.00; width of face at dorsal clypeal edge = 0.35 mm; clypeus width/height = 2.57; vertex width/distance between anterior ocelli and edge of torulus = 2.57; length of first flagellomere = 0.26 mm; first flagellomere length/width = 2.33; length of first flagellomere/ length of second flagellomere = 1.00; length of first flagellomere/length of third flagellomere = 0.95; terminal flagellomere length = 0.15 mm; terminal flagellomere length/penultimate flagellomere length = 1.20; terminal REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 15

flagellomere length/width = 3.00; malar space height/basal width of mandible =? ; ocell-ocular distance/ lateral ocelli distance = 1.00. Clypeus with dense and confused fine punctate sculpture; face with areas around compound eyes with confused punctate sculpture, but mid area with bigger and more defined punctate, face keel well defined and nitid extending itself for 2/3 of face; frons with scrobal areas nitid, remainder with fine and shallow punctate sculpture and anterior vertex; temple and remainder of vertex and gena and genapostgena junction area with fine and well impressed punctate sculpture; remainder of postgena nitid; ocular ring nitid. Mesosoma. Mesosoma length = 1.22 mm; mesosoma length/width = 1.34; mesosoma height = 0.80 mm. Propleuron with fine punctate sculpture throughout; pronotum lateral area with anterior 1/3 nitid and dorsal edge, ventral edge and area between scrobal lateral groove, upper and lower lateral groove with few smooth and transversal ridges, distal edge upper 1/2 and posterior 1/5 of dorsal edge punctate, dorsal edge 1/5 and ventral edge 1/3 of posterior edge height; mesonotum with dense and small punctate sculpture which becomes bigger and less dense towards scutellar groove, area in front of scutellar groove nitid; scutellar groove with 12 small but well defined costulae; scutellum with nitid except for very few punctulate sculpture at midlongitudinal area; axilla through mesonotum with few thick and smooth transversal ridges throughout its width, lunulae edge well defined; metanotum very thin and rectangular in shape, with smooth rugulose sculpture, mid distal costulae without sculpture except for two posterior lateral small pits on it; axilla through metanotum with four transversal ridges; remainder nitid; propodeum without transversal and costula carinae, areola with anterior 1/2 less well defined than posterior 1/2 by presence of small and confused rugulose sculpture on its carinae, anterior lateral areas with punctate sculpture, posterior lateral areas with rugulose sculpture and the areola internal area; mesopleuron anterior 1/3 punctate, dorsal edge with well defined and dense scrobiculate sculpture, epistomal furrow with few smooth and thick transversal ridges, remainder of mesopleuron nitid; metapleuron with mid pit present, dorsal edge with few small transversal ridges, posterior edge with three transversal ridges at mid length, area around dorsal edge with punctate sculpture, remainder of metanotum nitid. Legs. Hind femur length = 0.65 mm; hind femur length/width = 2.60; hind tibia length/hind femur length = 1.42. Fore telotarsus shorter than fore basitarsus, fore telotarsus with two four long setae at ventrodistal edge. Wings. Forewing length = 2.95 mm; 1RS length = 0.09 mm; 1CUa length/1cub length = 1.00; length RS+Ma = 0.46 mm; length M+CU = 1.03 mm; 1M length/ m-cu length = 2.18; pterostigma length/height = 1.93. Hind wing: 1M length = 0.44 mm; 1M length/2m length = 2.44; 1M length/m+cu length = 1.07; length r-m/length cu-a =0.79; 1RSa length/2r-m = 1.00; 1A length = 0.30 mm. Metasoma. First tergum basal width = 0.22 mm; first tergum length/distal width = 2.33; second tergum length/distal width = 0.45; third tergum length/distal width = 0.35. First metasomal tergum with anterior 1/2 with few longitudinal lineate sculpture at lateral areas and with a dense and fine rugulose sculpture, remainder of tergum with a bigger and more evident rugose sculpture, tergum posterior 1/2 with a depressed midlongitudinal area with few rugose sculpture on it, posterior edge with an circular shape depression at each corner of it, mid area of tergum slightly constraint; second metasomal tergum with confused and smooth rugulose sculpture; third metasomal tergum nitid as remainder of terga. Material examined. Holotype, male, Costa Rica, Guanacaste Province, Area de Conservacion Guanacaste, Sector Cacao, Sendero Circular, 1185 m, Lat 10.92714, Long -85.46683, 02-SRNP-8383, 01/iv/ 2002. Holotype deposited at INBio. Comments. This species is described from one known male; this is the only known species that exhibits the I shaped first metasomal tergum. See note under P. lincolnii n. sp. for placement of specimen under the genus Parapanteles Rearing records. Voucher number 02-SRNP-8383 reared from "elachjanzen01 Janzen182" in the family Elachistidae, which was feeding on Erythroxylum macrophyllum (Erythroxylaceae). Etymology. Gender, masculine. This species is named after the complex pattern of sculpture observed on the first tergum. 16 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

Parapanteles continua Valerio and Whitfield, n. sp. Figure 5F, 9E, 11F, 12D Female. Body length = 2.35 2.43 mm. Body color: Head light brownish yellow as mesosoma, first and second terga, antenna; fore and mid leg light yellow as hind tibia basal 1/2, base of hind basitarsus leg, mandibles, tarsal claws, palpi; remainder of hind leg brownish yellow; compound eyes silver; ocelli yellow. Wings hyaline; forewing with light yellow veins but C+SC+R and pterostigma bright yellow; hind wing with transparent tubular veins. Head. Head height/width = 1.26 1.33; compound eye height/width = 1.73 1.75; intertentorial pit distance = 0.16 0.18 mm; tentorial pit distance/distance tentorial pit to compound eye = 2.25 2.33 ; width of face at dorsal clypeal edge = 0.31 0.32 mm; clypeus width/height = 2.33 2.68; vertex width/distance between anterior ocelli and edge of torulus = 2.48 2.50; length of first flagellomere = 0.10 0.11 mm; first flagellomere length/width = 2.40 2.80; length of first flagellomere/length of second flagellomere = 1.00 1.07; length of first flagellomere/length of third flagellomere = 1.00 1.16; terminal flagellomere length = 0.10 0.11 mm; terminal flagellomere length/penultimate flagellomere length = 1.28 1.33; terminal flagellomere length/width = 2.00 2.25; malar space height/basal width of mandible = 0.88 1.00; ocell-ocular distance/lateral ocelli distance = 1.66 1.81. Clypeus and face with fine shallow punctate sculpture, midlongitudinal area of face with few shallow and confused foveate sculpture laterally, centrally nitid; frons with scrobal areas nitid, deeply impressed throughout its length and sharply narrowing distally, lateral areas of frons with few inconspicuous punctate sculpture, distal area nitid as anterior area of vertex; vertex conspicuously punctate at lateral areas and gena and postgena at junction area with gena until mid width of it; ocular ring nitid as remainder of postgena. Mesosoma. Mesosoma length = 0.90 0.98 mm; mesosoma length/width = 1.12 1.20; mesosoma height = 0.65 0.7 mm. Propleuron with punctate sculpture as at distal 1/3 of it but upper edge nitid area between punctate sculpture and upper edge with scattered smooth and confused rugulose sculpture which extends until basal 1/3 of propleuron length; pronotum lateral area with anterior 1/2 with smooth shallow and inconspicuous thin transversal ridges, upper groove nitid but area immediately below with few smooth foveate sculpture, ventral groove nitid but distal 1/6 with few inconspicuous and smooth transversal ridges, remainder of area between grooves nitid, upper groove 1/4 and ventral groove 1/5 of posterior edge height; mesonotum with dense punctate sculpture which becomes more evident and slightly less dense towards scutellum groove; scutellum groove with 4 to 6 big costulae, lateral ones bigger than medial ones; scutellum with evident punctate sculpture throughout, lateral areas with costulate sculpture becoming smaller towards distal edge and of approximately same width; axilla through mesonotum with smooth transversal ridges that in some cases reach the mid width of axilla; metanotum with triangular shape in dorsal view, middistal area with one big medial nitid costula; axilla through metanotum with 4 sharp ridges, area between ridges nitid; propodeum with transversal and posterior 1/2 of costulae well cristate as most of areolar carinae (tip of dorsal area less conspicuously present), anterior 1/2 of costula absent, pronotum anterior 1/2 with transversally lineate sculpture present which is more evident and thick next to areolar carinae, remainder nitid; mesopleuron with anterior edge with fine punctate sculpture, dorsal distal 1/2 with dense small scrobiculate sculpture which becomes even smaller towards distal edge, anterior 1/2 of dorsal edge with few spaced and irregularly defined scrobiculate sculpture (anterior area more shallow and less cristate), distal edge nitid, sternaulus as a wide depress area, remainder of mesopleuron nitid; metapleuron with dorsal edge with big and spaced thin transversal ridges, distal edge with ever bigger and more spaced transversal ridges, distal 1/3 with fine confused rugulose sculpture, medial pit present and remainder of metapleuron nitid. Legs. Hind femur length = 0.55 0.62 mm; hind femur length/width = 2.75 2.78; hind tibia length/hind femur length = 1.2 1.24. Fore telotarsus with set of four big and strong setae at mid ventral area of it, shorter in length than basitarsus; hind telotarsus with a set of elongated setae at internal lateral area of it. REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 17

FIGURE 7. Propodeum of Parapanteles lincolnii n. sp. (A), P. polus n. sp. (B), P. noae n. sp. (C), P. thrix n. sp. (D), P. paradoxus (Muesebeck) (E) and P. rarus n. sp. (F). Wings. Forewing length = 2.40 2.45 mm; 1RS length = 0.08 0.1 mm; 1CUa length/1cub length = 0.72 0.78; length RS+Ma = 0.35 0.38 mm; length M+CU = 0.78 0.8 mm; 1M length/ m-cu length = 2.15 2.16; pterostigma length/height = 1.28 1.31. Hind wing: 1M length = 0.40 mm; 1M length/2m length = 1.77; 1M length/m+cu length = 1.52 1.68; length r-m/length cu-a = 0.73 0.80; 1RSa length/2r-m = 1.57 1.67; 1A length = 0.20 0.22 mm. Metasoma. First tergum basal width = 0.19 0.21 mm; first tergum length/distal width = 1.22 1.24; second tergum length/distal width = 0.44 0.53; third tergum length/distal width = 0.47 0.50; hypopygium length = 18 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

0.70 0.78 mm. First metasomal tergum with midlongitudinal area with distal 2/3 with confused smooth rugulose sculpture with mixed with confused foveate sculpture becoming indistinct at junction area, rugose sculpture becoming more cristate and evident on distal 1/4 of tergum, remainder of tergum with shallow confused foveate sculpture; second metasomal terga with fine smooth lineate sculpture mixed with confused smooth foveate sculpture, both of them becoming less evident towards distal area; remainder of terga nitid; ovipositor almost as big as hind tibia length; ovipositor sheaths elongated in lateral view. FIGURES 8. First metasomal tergum in dorsal view for Parapanteles thrix n. sp. (A), P. rarus n. sp. (B), P. scotti n. sp. (C), P. noae n. sp. (D), P. tessares n. sp. (E), and P. polus n. sp. (F). REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 19

Variation. Sculpture of first and second tergum very weak, first tergum with anterior 1/2 nitid and medial rugulose sculpture very smooth and weakly defined second tergum with lineate sculpture which is more cristate posteriorly. Material examined. Holotype, female, Costa Rica, Guanacaste Province, Area de Conservacion Guanacaste, Sector Cacao, Sendero Arenales,1000 m, Lat 10.92471, Long -85.46738, 09/ix/1997, Col. R. Moraga, 97-SRNP-1739 Paratypes: 14 females, all same data as holotype. All material, including the holotype, deposited at INBio except for two females deposited at INHS Male. Similar to female. Comments. This species shares with P. tessares the 4 setae at the ventral-distal area of fore telotarsus but can be easily separated from it by the body color, the deeply impressed scrobal areas as well the ovipositor sheaths with long and sparse setae over most of its distal 1/3. Rearing records. Hylesia continua (Saturniidae) feeding on Hampea appendiculata (Malvaceae). Voucher numbers: 97-SRNP-1673, 97-SRNP-1677, 97-SRNP-1678, 97-SRNP-1679, 97-SRNP-1680, 97- SRNP-1685, 97-SRNP-1691, 97-SRNP-1694, 97-SRNP-1696, 97-SRNP-1703, 97-SRNP-1708, 97-SRNP- 1709, 97-SRNP-1710, 97-SRNP-1715, and 97-SRNP-1739. The observed sex ratio is as follow: 6/35. Etymology. Gender, neutral. The name of this species is a random combination of letters without any meaning. Parapanteles em Valerio and Whitfield, n. sp. Figs. 5D, 9C, 11B, 13F Female. Body length = 2.13 2.75 mm. Body color: Metasomal sterna and pleura at basal 1/2 yellow as fore and mid legs (except fore coxae brownish yellow and mid coxae dark brown in combination with tarsomeres more whitish yellow), hind leg trochanter and trochantellus, yellow as basal half of tibia and basal 1/3 of femur, labrum, mandibles, tegula; scape and pedicel brownish yellow as remainder of antenna, remainder of metasomal sterna and pleura and hypopygium; remainder of body dark brown. Wings hyaline, forewing veins light brownish yellow, except basal 1/2 M+CU and basal 3/4 of 1-1A whitish in color, pterostigma slightly darker in color than remainder of veins with the exception of C+SC+R which appears as dark brownish yellow; hind wing with veins whitish yellow. Head. Head height/width = 1.20 1.21; compound eye height/width = 1.55 1.60; intertentorial pit distance = 0.16 0.17 mm; tentorial pit distance/distance tentorial pit to compound eye = 2.25 2.33; width of face at dorsal clypeal edge = 0.30 0.35 mm; clypeus width/height = 2.10 2.33; vertex width/distance between anterior ocelli and edge of torulus = 2.29 2.53; length of first flagellomere = 0.20 0.22 mm; first flagellomere length/width = 3.00 3.20; length of first flagellomere/length of second flagellomere = 1.00-1.06; length of first flagellomere/length of third flagellomere = 1.00-1.60; terminal flagellomere length = 0.12 0.14 mm; terminal flagellomere length/penultimate flagellomere length = 1.22 1.25; terminal flagellomere length/width = 2.5 2.75; malar space height/basal width of mandible = 1.12 1.14; ocell-ocular distance/lateral ocelli distance = 1.33 1.35. Clypeus with dense, fine and confused punctate sculpture; face with shallow punctate sculpture at most of its area except at lateromedial areas nitid and medial area with more confused, shallow and less defined punctate sculpture; frons with scrobal areas nitid, lateral areas with punctate sculpture present, distal area essentially nitid as anterior area of vertex; vertex with few punctate sculpture at distal area, lateral areas with few to dense punctate sculpture present and remainder of gena (except ocular ring nitid); postgena nitid except at junction with gena. 20 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

FIGURE 9. Female genitalia of Parapanteles thrix n. sp. (A), P. rarus n. sp. (B), P. em n. sp. (C), P. paradoxus (Muesebeck) (D), P. continua n. sp. (E), and P. tessares n. sp. (F). Mesosoma. mesosoma length = 0.95 1.15 mm; mesosoma length/width = 1.31 1.44; mesosoma height = 0.60 0.66 mm. Propleuron with fine punctate sculpture throughout its length except at dorsoexternal edge nitid; pronotum lateral area with anterior 1/2 with few big, spaced and smooth transversal ridges, upper lateral groove essentially nitid, ventral groove with smooth, short and confused transversal ridges present, area between ridges nitid except of area next to distal edge with confused fine rugulose sculpture, dorsal edge 1/5 and ventral edge 1/3 of distal edge height; mesonotum with dense and well defined punctate sculpture throughout its length except area next to scutellar groove nitid, posterior sculpture slightly bigger than anterior REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 21

one; scutellar groove with 7 to 9 small and well defined costulae; scutellum with confused punctate sculpture throughout its length, lateral areas with big wide costulae sculpture, distal one 2 times as long as anterior costulae and narrower too; axilla through metanotum with a series of transversal narrow ridges across its width and with two carinae dividing them at middle length; metanotum triangular in shape and nitid; axilla through metanotum with a series of longitudinal ridges, distal edge nitid; propodeum with rugulose sculpture at posterior lateral areas and areola and posterior 1/2 of anterolateral areas, remainder of anterolateral area with few punctate sculpture, costula totally present; mesopleuron with anterior edge and ventral area punctate, sternaulus as an deep nitid depression along its length, dorsal area with anterior area nitid and remainder of edge with small well defined scrobiculate sculpture except junction with distal edge with more confused and bigger costulate like sculpture, remainder of distal edge nitid as remainder of mesopleuron; metapleuron with medial pit present, anterior 1/2 nitid, reminder area with confused rugulose sculpture present and next to dorsal edge, dorsal edge with big and spaced transversal ridges which are well defined, distal edge with fewer and more sinuate transversal ridges than at dorsal edge. FIGURE 10. Female genitalia of Parapanteles tlinea n. sp. (A), P. scotti n. sp. (B), P. sicpolus n. sp. (C), P. polus n. sp. (D), P. mariae n. sp. (E), and P. lincolnii n. sp. (F). 22 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

FIGURE 11. First metasomal tergum in dorsal view for Parapanteles sicpolus n. sp. (A), P. em n. sp. (B), P. paradoxus (Muesebeck) (C), P. rooibos Valerio et al., P. tlinea n. sp. (E), and P. continua n. sp. (F). Legs. Hind femur length = 0.68 0.72 mm; hind femur length/width = 3.22 3.38; hind tibia length/hind femur length = 1.14 1.15. Fore telotarsus with a long and narrow hooklike setae at internolateral area of it, telotarsus shorter in length than basitarsus; hind telotarsus normal in shape and not modified. Wings. Forewing length = 2.30 2.75 mm; 1RS length = 0.06 0.10 mm; 1CUa length/1cub length = 0.87 0.88; length RS+Ma = 0.34 0.43 mm; length M+CU = 0.79 0.85 mm; 1M length/ m-cu length = 2.08 2.10; pterostigma length/height = 1.33 1.44. Hind wing: 1M length = 0.39 0.42 mm; 1M length/2m length = 1.88 2.00; 1M length/m+cu length = 1.42 1.80; length r-m/length cu-a = 0.66 0.77; 1RSa length/ 2r-m = 1.80 2.00; 1A length = 0.16 0.25 mm. REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 23

FIGURE 12. Wing venation of Parapanteles polus n. sp. (A), P. mariae n. sp. (B), P. thrix n. sp. (C), P. continua n. sp. (D), P. paradoxus (Muesebeck) (E), and P. noae n. sp. (F). Metasoma. First tergum basal width = 1.18 0.26 mm; first tergum length/distal width = 0.82 1.30; second tergum length/distal width = 0.40 0.48; third tergum length/distal width = 0.39 0.43; hypopygium length = 0.46 0.60 mm. First metasomal tergum with basal 1/3 nitid at medial area and with few confused rugulose sculpture at lateral areas, remainder of tergum with confused rugulose sculpture; second metasomal tergum with longitudinal lineate sculpture which is less defined and more sinuate at distal edge and medial area; reminder terga nitid. 24 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.

FIGURE 13. Wing venation of Parapanteles aletiae (Riley) (A), P. sicpolus n. sp. (B), P. tlinea n. sp. (C), P. scotti n. sp. (D), P. tessares n. sp. (E), and P. em n. sp. (F). Material examined. Holotype, female, "Costa Rica, Guanacaste, ACG, Sector Santa Rosa, Vado, Río Calera, 10m, 12/vi/2000, Latitude: 10.80274 Longitude: -85.67423, Col. Gusaneros, 00-SRNP-8628" Paratypes: 7 females, (2) same data as holotype. Costa Rica, Guanacaste, ACG, Sector Santa Rosa, Vado Río Calera, 12/vi/2000, Col. Gusaneros. (2) Costa Rica, Guanacaste, ACG, Sector Cacao, Estación Cacao, 1150 m, 323104 N 375725E, 10/iii/2003, F. Quesada. Material deposited at INBio. Male. Similar to female. REVIEW OF PARAPANTELES Zootaxa 2084 2009 Magnolia Press 25

FIGURE 14. Parapanteles complexus n. sp. first metasomal tergum sculpture (A), mesonotum (B), and wing venation (C). 26 Zootaxa 2084 2009 Magnolia Press VALERIO ET AL.