ore JHR 53; |—24 (20 I 6) JOURNAL COR. Ppeeteiewed open atontiouryat doi: 10.3897/jhr.53.10217 (ME Hymenopter a http://jhr.pensoft.net The international Society of Hymenoptersts. RESEARCH Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) Elijah J. Talamas', Lubomir Masner’ I Systematic Entomology Laboratory, USDA/ARS clo USNM, Smithsonian Institution, Washington, D.C. 20560, U.S.A. 2. Agriculture and Agri-Food Canada, K.W. Neatby Building, Ottawa, Ontario KIA 0C6, Canada Corresponding author: Elijah J. Talamas (elijah.talamas@ars.usda.gov) Academic editor: /. Yoder | Received 17 August 2016 | Accepted 31 October 2016 | Published 19 December 2016 Attp://zoobank.org/13184D63-2047-4F62-A98 7-B844E6386BCD Citation: Talamas EJ, Masner L (2016) Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae). Journal of Hymenoptera Research 53: 1-24. https://doi.org/10.3897/jhr.53.10217 Abstract Nine new species of Helava are described: H. acutiventris sp. n., H. allomera sp. n., H. aureipes sp. n.., H. carinata sp. n., H. microptera sp. n., H. pygmea sp. n., H. reducta sp. n., H. simplex sp. n., and H. samanthae sp. n., and Helava alticola Masner & Huggert is redescribed. New characters are presented to supplement the generic description of Masner and Huggert (1989) and the genus is diagnosed from similar genera in Sceliotrachelinae: Aphanomerus Dodd and Austromerus Masner & Huggert. Keywords Parasitoid, taxonomy, Platygastroidea Introduction Helava was described by Masner and Huggert in their 1989 treatment of world genera of Sceliotrachelinae. Masner and Huggert separated Helava from Austromerus on the basis of “clavate” antennae in males. We here replace “clavate” with the term “clubbed” to describe the apically enlarged antennomeres in males to maintain strict use of termi- nology in which clavomeres are defined by the presence of basiconic sensilla. Our revi- sion of Helava reveals that the male antenna is filiform in two species, H. acutiventris and H. allomera, and thus Helava and Austromerus are separable only by the form of Copyright Elijah J. Talamas, Lubomir Masner. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 2 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) the clava in females: compact in Helava and with articulated clavomeres in Austromerus (compare Figures 2 and 12; see also figures 168-173 in Masner and Huggert (1989)). Helava is also morphologically very close to Aphanomerus, from which Masner and Huggert (1989) separated Helava by the dense setation on T1—T2 and presence of propodeal foamy structures (compare Figures 1 and 20). In the concept of Masner and Huggert (1989), Helava is found in South America, Tasmania and continental Australia, a distribution consistent with a Gondwanan origin. The limits of Helava, Aphanomerus and Austromerus have become blurred following this revision and testing hypotheses about the phylogeography of the genus will require a better understanding of relationships between these three genera. We refrain from describing the Austral- ian species of Helava until species-level revision of Aphanomerus and Austromerus are conducted to provide a full grasp of the morphological diversity of their constituent species. Currently no host data are known for Helava. Materials and methods The numbers prefixed with “CNC” or “OSUC ” are unique identifiers for the individual specimens (note the blank space after some acronyms). Details of the data associated with these specimens may be accessed at the following link: http://purl.oclc.org/NET/ hymenoptera/hol, and entering the identifier in the form. Persistent URIs for each taxo- nomic concept were minted by xBio:D in accordance with best practices recommended by Hagedorn et al. (2013). Morphological terms were matched to concepts in the Hy- menoptera Anatomy Ontology (Yoder et al. 2010) using the text analyzer function. A table of morphological terms and URI links is provided in Suppl. material 1. Photographs were captured with a Z16 Leica lens with a JVC KY-F75U digital camera using Cartograph software. Single montage images were produced from image stacks with the program CombineZP. In some cases, multiple montage images were stitched together in Photoshop to produce larger images at high resolution and mag- nification. Full resolution images are archived at the image database at The Ohio State University (http://purl.oclc.org/NET/hymenoptera/specimage). Scanning electron micrographs were produced with a Hitachi TM300 Tabletop Microscope. The specimen was disarticulated with a minuten probe and forceps and mounted on 12 mm slotted aluminum mounting stub (EMS Cat. #75220) using car- bon adhesive tabs (EMS Cat. #77825-12) by means of a fine paint brush and sputter coated with approximately 70 nm of gold/palladium. This work is based on specimens deposited in the following repositories with ab- breviations used in the text: ANIC Australian National Insect Collection, Canberra City, Australia CNCI = Canadian National Collection of Insects, Ottawa, Canada OSUC_ C.A. Triplehorn Collection, The Ohio State University, USA USNM_ Smithsonian National Museum of Natural History, Washington, DC, USA Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) 3 Figure |. Aphanomerus sp. female (USNMENT00916681), head, mesosoma, metasoma, lateral view. Scale bar in millimeters. Figures 2-3. Austromerus grandis, female paratype (USNMENT00916679) 2 head, mesosoma, meta- soma, lateral view 3 head, mesosoma, metasoma, ventral view. Scale bars in millimeters. 4 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) Abbreviations and characters annotated in the figures apc anterior pronotal patch (Figure 16, 50) apS2 —_ anterior setal patch on S2 (Figures 2, 17) apT1 anterior setal patch on T2 (Figure 17) fp foamy structure on propodeum (Figures 16, 18, 25) fS1 foamy structure on S1 (Figure 17) hoc hyperoccipital carina (Figures 34, 54) mfp mesofurcal pit (Figure 14) mkT1 = median keel on T1 (Figure 17) mkT2 = median keel on T2 (Figures 10, 25) not notaulus (Figures 10, 25) ppc posterior pronotal patch (Figure 9) sss scutoscutellar sulcus (Figure 36) tel transepisternal line (Figures 3, 35) Diagnosis of Helava In the process of coding characters for potential use in species delimitation we encoun- tered a small number of new characters shared between all New World species of Helava. We here present a generic diagnosis based on these characters and those presented by Masner and Huggert (1989): Antennal formula 10-10. Clava compact in females. Epomium absent. Fore wing with tubular submarginal vein terminating in a truncate knob. Ventral rim of pronotum forming lamella flanking procoxa. Mesopleural carina absent. Setation of axillar area present. Setation of mesoscutellum along posterior and lateral margins dense. 1st trochanter longest, particularly on metatrochanter. Setation of coxae dense. Tibial spur formula 1-2-2. Setation of laterotergites present. Sculpture of tergites absent. T2 with narrow strip of dense setation along anterior margin. Sculpture of sternites absent. Felt fields on S2 present. Key to species (males and females) 1 Foamy structures on lateral propodeum covering area larger than visible part Of metapletiton:(Bieures. 95 Ler. 294 SO) sh. sata isan tebeeteeedae te eerels 2 - Foamy structures on lateral propodeum covering area distinctly smaller than hairy metapleuron (Figures 4, 24, 35, 40, 55) or foamy structures absent CPICUiee Eo) cteneas ceo Mette tert sete eerie ty Rete cnetip i tcE Bec xy ah oactu at 5 2, Lateral pronotum with dorsoventral strip of dense setation posteriorly (Fig- ure 9); notaulus percurrent (Figure 10); male antenna filiform (Figure 13); female antennal:clava 3-merous' (igure: 2) éicsscsefovsesedcoacacdaneionacsessccutectiacacates A feo ae Le, Ae nh H. allomera Masner & Talamas, sp. n. Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) 5 — Lateral pronotum without dense setation posteriorly (Figures 19, 29, 50); notaulus absent (Figures 20, 30, 51); male antenna clubbed (Figures 22, 33); female antennal clava 4-merous (Figures 23, 32).....scscssssscssesseeseeeseeseeseeees 3 3 Wypper tronsdenselyceros]e CPE 59 Jes tstactcteabepelacseuresthostasdgutosteseidelouerees ie eektdves riba Menaeeor belbee aote hike vibe enti 9 H. samanthae Masner & Talamas, sp. n. - Upper frons glabrous or with sparse setae only along inner orbits (Figures NAS aie ODA oe aresteh elation blllh ona ien weiivesdee Sadlese oA Be crt SoS g0h dud Gast 4 4 Posterior vertex glabrous or only very sparsely setose (Figures 30, 34)........... oh nt ae et epee net eee H. carinata Masner & Talamas, sp. n. — Posterior vertex-densely setose (Figures: 15720=21) scccca.scccsctzcatcesnestesansucecdane UA Mts ath ciate ose tne eees Louie NSE wsakiey De ele H. alticola Masner & Huggert 5 Wings reduced to strips, reaching only to anterior T2, or absent (Figures 36, AG) cee ee ee | eee See eh een, ee he eer ya. aerne On oe en eon ee Teens 6 — Wings fully developed, exceeding apex of metasoMa........icseeeeseeeeeeseeeees 7 6 Scutoscutellar sulcus absent (Figure 46); ocelli absent (Figure 46); mesopleu- ron without transepisternal line (Figure 45) oo... ee eeceseeseeeeeeeeseeeeeeseeeeneees pM Ades 2, Bea Se Mn Nene eyane ts enh cert H. reducta Masner & Talamas, sp. n. = Scutoscutellar sulcus present (Figure 36); ocelli present (Figure 36); meso- pleuron;with transepisternal line (Figure: 35). ccs: (ocsvecmesavendeewrnboswventesvecseestes Lyte Sid PORTS An rr Reimer ein AE H. microptera Masner & Talamas, sp. n. t Medial S2 distinctly projecting in lateral view in both sexes (Figure 4); mes- opleuron without transepisternal line (Figure 4); antenna in male filiform, with Mand ATO approximated: (Figure Get, lice ctosenns cdberonssdelonntovlauuliubn shies s GAR Pte ud, Ny, ee nonlethal eel H. acutiventris Masner & Talamas, sp. n. = S2 evenly convex medially in lateral view (Figures 24, 40, 55); mesopleuron with transepisternal line (Figures 24, 40, 55); antenna clubbed in male (Fig- WH Rat Say Ae te, NN OP Da tas oa ig RE, ck PR tsa acl mares “ha 8 8 Notaulus present (Figure 25).......... H. aureipes Masner & Talamas, sp. n. — Notaulus-absent (Figures 419 56): cscs cccodtsce.ssnseruss i suovaatseocbelpeacseazeatesssau dea: 9 9 Basal vein (Rs+M) in fore wing absent (Figure 60)... eeeeesssecseeeeeeseeeeees Stockett Aetna Si sede be eget ane Dattani H. simplex Masner & Talamas, sp. n. — Basal vein (Rs+M) in fore wing nebulous (Figure 41) oo... cece eeeeeeeeeeeees we scisae ele iuslse Rs tle lpn cnsatS ac. H. pygmea Masner & Talamas, sp. n. Helava acutiventris Masner & Talamas, s http://zoobank.org/FFCC0883-2123-4A2E- BC69- 4CCC679C47D5 http://bioguid.osu.edu/xbiod_concepts/354388 Figures 4-8 Description. Female body length: 0.99-1.03 mm (n=10). Male body length: 0.90-— 0.85 mm (n=20). Male antenna: filiform. Number of female clavomeres: 3. Setation of frons anterior to ocellar triangle: present. Setation of vertex posterior to lateral ocel- 6 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) PA he : nn, = O41 % = = Rae = Figures 4-8. Helava acutiventris 4 female holotype (USNMENT0989201), head, mesosoma, metasoma, lateral view 5 female holotype (USNMENT00989201), head and mesosoma, dorsal view 6 male paratype (USNMENT00989201), metasoma, dorsal view 7 female holotype (USNMENT00989201), head, ante- rior view 8 male paratype (USNMENT00989202), antenna, ventral view. Scale bars in millimeters. lus: dense. Hyperoccipital carina: absent. Pronotum in dorsal view: present mostly as lateral shoulders. Dorsoventral band of dense setation on posterior part of lateral pronotum: absent. Setation of pronotal cervical sulcus: sparse to absent. Width of dorsal mesopleuron in lateral view: about equal ventrally and dorsally to 1.5 times as wide ventrally. Longitudinal striation on dorsal mesopleuron: absent. Transepisternal Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) 7 line: absent. Mesofurcal pit: present. Notaulus: absent. Rs+M in fore wing: nebulous. Wings: macropterous. Rs+M in hind wing: nebulous. Color of legs: yellow. Foamy structures of lateral propodeum: smaller than hairy metapleuron. Median tubercule on T2: absent. Setation of anterior T2: continuous across tergal midline. Lateral patch on T2: present. Foamy structures on S1: present at lateral margin of sternite. Transverse felt field on anterior S2: present as transverse strip. Shape of S2 in lateral view: dis- tinctly bulging medially. Diagnosis. The ventral protrusion of S2 in H. acutiventris separates this species from all other species in Helava. In addition to the shape S2, the absence of a transepis- ternal line on the mesopleuron is shared only with H. reducta, which is a starkly dif- ferent species that can be separated by the absence of ocelli and a scutoscutellar sulcus. Etymology. The epithet “acutiventris” is given to this species in reference to the sharp projection on S2 in both sexes. Link to distribution map. http://hol.osu.edu/map-large.html?id=354388 Material examined. Holotype, female: CHILE: Bio-Bio Reg., Nuble Prov., Ter- mas Rd., 60km SE Chillan, 1300m, 7.XII-19.X1.1985, flight intercept trap, S. Peck & J. Peck, USNMENT00989201 (deposited in CNCI). Paratypes: CHILE: 12 fe- males, 25 males, CNC424698—424733, USNMENT00989202 (CNCI). Comments. The diagnostic shape of S2 is found in both males and females, lead- ing us to believe that this is not an adaptation for housing the retracted ovipositor system, as can be found in some species of Synopeas Forster and Platygaster Latreille. Helava allomera Masner & Talamas, sp. n. http://zoobank.org/F786DD 15-F5B5-41 CB-8910-479F75 142D03 http://bioguid.osu.edu/xbiod_concepts/354395 Figures 9-13 Description. Female body length: 0.89-1.33 mm (n=20). Male body length: 0.99— 1.33 mm (n=20). Male antenna: filiform. Number of female clavomeres: 3. Setation of frons anterior to ocellar triangle: absent or sparsely present only along inner orbit of eye. Setation of vertex posterior to lateral ocellus: dense. Hyperoccipital carina: con- tinuous across vertex. Pronotum in dorsal view: present mostly as lateral shoulders. Dorsoventral band of dense setation on posterior part of lateral pronotum: present. Setation of pronotal cervical sulcus: dense. Width of dorsal mesopleuron in lateral view: about equal ventrally and dorsally to 1.5 times as wide ventrally. Longitudinal striation on dorsal mesopleuron: absent. Transepisternal line: present. Mesofurcal pit: present. Notaulus: percurrent. Rs+M in fore wing: nebulous. Wings: macropterous. Rs+M in hind wing: nebulous Color of legs: bright orange yellow. Setation of coxae: dense Foamy structures of lateral propodeum: larger than hairy metapleuron. Median tubercule on T2: present. Setation of anterior T2: interrupted medially. Lateral patch on [2: absent. Foamy structures on S1: present at lateral margin of sternite. Transverse felt field on anterior $2: present. Shape of S2 in lateral view: broadly convex. 8 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) , ep “ty - . Figures 9-12. Helava allomera 9 female holotype (USNMENT00989217), head, mesosoma, meta- soma, lateral view 10 female holotype (USNMENT00989217), head, mesosoma, metasoma, dorsal view II female holotype (USNMENT00989217), head, anterior view 12 female holotype (USN- MENT00989217), antenna, dorsal view 13 male paratype (USNMENT00989218), antenna, dorsal view. Scale bars in millimeters. Diagnosis. Helava allomera can be differentiated from other species in the genus by the combination of the well-developed hyperoccipital carina, percurrent notauli, and foamy structures on the propodeum that are larger than the visible part of the metapleuron in lateral view. Within He/ava, this is the only species with a dorsoven- tral band of dense setae along the posterior margin of the lateral pronotum (Figure 9). Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) 9 Etymology. The epithet “allomera” is given to this species in reference to the unu- sual form and segmentation of antennae in both sexes. Link to distribution map. http://hol.osu.edu/map-large.html?id=354395 Material examined. Holotype, female: CHILE: Araucania Reg., Malleco Prov., 1200m, 37.809°S 73.016°W, Nahuelbuta National Park, 9.I-12.1.2000, Malaise trap, D. Webb & D. Yeates, USNMENT00989217 (deposited in CNCI). Paratypes: (58 fe- males, 30 males) ARGENTINA: 7 females, 2 males, CNC42498 1—424985, 424993— 424994, 425022, 425031 (CNCI).CHILE: 51 females, 28 males, CNC424946- 424980, 424986424992, 424995425021, 425023-425030, 425032, 494556, US- NMENT00989218 (CNCI). Helava alticola Masner & Huggert http://bioguid.osu.edu/xbiod_concepts/12334 Figures 14-23 Helava alticola Masner & Huggert, 1989: 72 (original description. Species list); Vlug 1995: 26 (cataloged, type information). Description. Female body length: 1.00—1.29 mm (n=20). Male body length: 0.91—1.32 mm (n=21). Male antenna: apically clubbed. Number of antennomeres in male club: 4, Number of female clavomeres: 4. Setation of frons anterior to ocellar triangle: absent or sparsely present only along inner orbit of eye. Setation of vertex posterior to lateral ocellus: dense. Hyperoccipital carina: absent; continuous across vertex; indicated by lateral tubercules. Pronotum in dorsal view: present mostly as lateral shoulders; slightly collarlike. Dorsoventral band of dense setation on posterior part of lateral pronotum: absent. Setation of pronotal cervical sulcus: dense. Width of dorsal mesopleuron in lateral view: about equal ventrally and dorsally to 1.5 times as wide ventrally. Longitu- dinal striation on dorsal mesopleuron: absent. Transepisternal line: present. Mesofurcal pit: present. Notaulus: absent. Rs+M in fore wing: nebulous. Wings: macropterous. Rs+M in hind wing: nebulous. Color of legs: brownish, yellowish brown. Foamy struc- tures of lateral propodeum: larger than hairy metapleuron. Median tubercule on T2: absent. Setation of anterior T2: continuous across tergal midline. Lateral patch on T2: present. Foamy structures on S1: present at lateral margin of sternite. Transverse felt field on anterior S2: present. Shape of S2 in lateral view: broadly convex. Diagnosis. Helava alticola can be separated from species with large propodeal foamy structures by the evenly rounded form of S2 and the pattern of setation on the dorsal head: posterior to the ocelli the posterior vertex is densely setose, and anterior to the ocelli the upper frons is glabrous or with sparse setae only along the inner orbit of the eye. Link to distribution map. http://hol.osu.edu/map-large.html?id=12334 Material examined. Holotype, female: COLOMBIA: Colombia, Caldas, 5.IV.1973, CNC494814 (deposited in CNCI). Paratypes: (56 females, 33 males, 1 unsexed) COLOMBIA: 17 females, 15 males, CNC494557-494566, 494622— 10 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) Figures 14-18. Helave alticola, female (USNMENT00989211) 14 head and mesosoma, ventral view 15 head, dorsal view 16 mesosoma, lateral view 17 metasoma, lateral view 18 mesosoma, posterolateral view. Scale bars in millimeters. 494633, 494663-494671, USNMENT00989213-00989214 (CNCI); USN- MENT00989943 (USNM).ECUADOR: 37 females, 17 males, CNC494567— 494568, 494571-494592, 494594494621, 494672494673 (CNCI). PERU: 2 females, CNC494569-494570 (CNCI). Other material: (115 females, 100 males) BOLIVIA: 1 female, 1 male, CNC494753, 494785 (CNCI). CHILE: 1 female, CNC424908 (CNCI). COLOMBIA: 92 females, 50 males, CNC424852-424894, 494506, 494634-494662, 494674, 494675-494703, 494705, 494727-494728, 494754494765, 494767-494771, 494773-494781, 494784, 494794494795, 494800-494803, 494806, 494807, 494809, 494810, USNMENT00989211 (CNCI). ECUADOR: 10 females, 35 males, CNC494593, 494704, 494706494724, Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) 11 Figures 19-23. Helava alticola \9 female (USNMENT989211), head, mesosoma, metasoma, lateral view 20 female holotype (CNC494814), head, mesosoma, metasoma, dorsal view 21 female holotype (CNC494814), head, anterodorsal view 22 male (USNMENT00989212), antenna, dorsal view 23 female (USNMENT00989211), antenna, anterior view. Scale bars in millimeters. 494726, 494749494752, 494766, 494772, 494778, 494783, 494786, 494788— 494793, 494796494799, 494804, 494805, 494808, 494811 (CNCI). VENEZUE- LA: 11 females, 14 males, CNC494725, 494729494748, 494779, 494782, 494787, USNMENT00989212 (CNCI). 12 E.J. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) Helava aureipes Masner & Talamas, sp. n. http://zoobank.org/96D177A2-D794-49 FF-9B6C-54CBAAFO5BCA http://bioguid.osu.edu/xbiod_concepts/354390 Figures 24-28 Description. Female body length: 0.95—1.36 mm (n=20). Male body length: 0.98— 1.10 mm (n=20). Male antenna: apically clubbed. Number of antennomeres in male club: uncertain, 3. Number of female clavomeres: 4. Setation of frons anterior to ocellar triangle: present. Setation of vertex posterior to lateral ocellus: dense. Hyperoccipital carina: absent. Pronotum in dorsal view: present mostly as lateral shoulders. Dorsoven- tral band of dense setation on posterior part of lateral pronotum: absent. Setation of pronotal cervical sulcus: sparse to absent. Width of dorsal mesopleuron in lateral view: about equal ventrally and dorsally to 1.5 times as wide ventrally. Longitudinal striation on dorsal mesopleuron: present. Transepisternal line: present. Mesofurcal pit: present. Notaulus: percurrent. Rs+M in fore wing: nebulous. Wings: macropterous. Rs+M in hind wing: nebulous. Color of legs: yellow. Foamy structures of lateral propodeum: smaller than hairy metapleuron. Median tubercule on T2: present. Setation of anterior T2: interrupted medially. Lateral patch on T2: absent. Foamy structures on S1: absent. Transverse felt field on anterior S2: present. Shape of S2 in lateral view: broadly convex. Diagnosis. Helava aureipes and H. allomera are the only two South American spe- cies with notauli. They can be separated from each other by the transepisternal line, which is absent in H. allomera and present as a distinct groove in H. aureipes. Etymology. The epithet “aureipes” is given to this species in reference to the gold- en colour of the legs. Link to distribution map. http://hol.osu.edu/map-large.html?id=354390 Material examined. Holotype, female: CHILE: Araucania Reg., Malleco Prov., 1200m, 37.809°S 73.016°W, Nahuelbuta National Park, 9.J—12.1.2000, Malaise trap, D. Webb & D. Yeates, UINMENT00989205 (deposited in CNCI). Paratypes: CHILE: 25 females, 88 males, CNC424741, 425033425101, 494411494451, 494813, USNMENT00989206 (CNCD). Helava carinata Masner & Talamas, sp. n. http://zoobank.org/1047FBEA-5089-4F07-87FE-E18A1C5BE9E1 http://bioguid.osu.edu/xbiod_concepts/354394 Figures 29-34 Description. Female body length: 1.00-1.15 mm (n=19). Male body length: 0.94— 1.12 mm (n=20). Male antenna: apically clubbed. Number of antennomeres in male club: 4. Number of female clavomeres: 4. Setation of frons anterior to ocellar triangle: absent or sparsely present only along inner orbit of eye. Setation of vertex posterior to lateral ocellus: very sparse or absent. Hyperoccipital carina: continuous across vertex. Pronotum in dorsal view: present mostly as lateral shoulders. Dorsoventral band of dense setation on posterior part of lateral pronotum: absent. Setation of pronotal cer- Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) il) fee Ugg eaical eae meer sar a ae ee TE Ein i al a es "4 d = tan ies a cy Ps a : = teh Serge Se Rae’ Na hs ae oe * : ee al < a uet= «oe ‘ a ean ae) al | 5 wie ele habcose ak i ‘4 i ~< eh 7 a ie see : _ ee A soma, lateral view 25 female holotype (USNMENT00989205), head, mesosoma, metasoma, dor- sal view 26 female holotype (USNMENT00989205), head, anterior view 27 male paratype (USN- MENT00989206), antenna, anterior view 28 female holotype (USNMENT00989205), antenna, anterior view. Scale bars in millimeters. vical sulcus: dense. Width of dorsal mesopleuron in lateral view: about equal ventrally and dorsally to 1.5 times as wide ventrally. Longitudinal striation on dorsal mesopleu- ron: absent. Transepisternal line: present. Mesofurcal pit: present. Notaulus: absent. Rs+M in fore wing: nebulous. Wings: macropterous. Rs+M in hind wing: nebulous. Color of legs: brownish. Foamy structures of lateral propodeum: larger than hairy metapleuron. Median tubercule on T2: absent. Setation of anterior T2: continuous 14 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) i. a t 32% eer ~~» (33, ee. ~* See , Se at 2 Fy ae >= | J = eee Bae a Figures 29-34. Helava carinata 29 female holotype (USNMENT00989215), head, mesosoma, meta- soma lateral view 30 female holotype (USNMENT00989215), head and mesosoma, dorsal view 31 male paratype (USNMENT00989216), propodeum and metasoma, dorsal view 32 female holotype (USN- MENT00989215), head, anterior view 33 male paratype (USNMENT00989216), antenna, dorsal view 34 male paratype (USNMENT00989216), head and mesosoma, dorsolateral view. Scale bars in millimeters. across tergal midline. Lateral patch on T2: present. Foamy structures on S1: present at lateral margin of sternite. Transverse felt field on anterior S2: present. Shape of S2 in lateral view: broadly convex. Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) 15 Diagnosis. Helava carinata belongs to the cluster of species with large propodeal foamy structures that includes H. alticola, H. allomera, and H. samanthae. It can be separated from H. alticola and H. samanthae by the setation of the upper frons and posterior vertex, which is either absent or very sparse. In H. alticola the upper frons is glabrous or nearly so, and then abruptly setose posterior to the ocelli; the dorsal head in H. samanthae is setose throughout. Helava carinata can be separated from H. allomera by the absence of dense setation on the posterior part of the lateral pronotum (compare Figures 9 and 29). Etymology. The Latin adjectival epithet “carinata” refers carinate vertex of the head. Link to distribution map. [http://hol.osu.edu/map-large.html?id=354394] Material examined. Holotype, female: COLOMBIA: Cundinamarca Dept., Tena Mpio., Laguna Pedro Pala, 2100m, 26.1.1992, Malaise trap, E. E. Palacio, USNMENT00989215 (deposited in CNCI). Paratypes: (48 females, 56 males) CO- LOMBIA: 22 females, 25 males, CNC494452-494462, 494481494485, 49449 1— 494495, 494501, 494504494505, 494519494534, 494537-494541, 494555, USNMENT00989216 (CNCI). ECUADOR: 26 females, 31 males, CNC494463— 494480, 494486-494490, 494496494500, 494502-494503, 494507-494518, 494535494536, 494542494554 (CNCI). Helava microptera Masner & Talamas, sp. n. http://zoobank.org/7 1 FA4BB9-FE32-41AF-8ABF-366AA39BC82A http://bioguid.osu.edu/xbiod_concepts/354386 Figures 35—39 Description. Female body length: 0.98—1.31 mm (n=9). Male body length: 1.11 mm (n=1). Male antenna: apically clubbed. Number of antennomeres in male club: 3. Number of female clavomeres: 4. Setation of frons anterior to ocellar triangle: present. Setation of vertex posterior to lateral ocellus: dense. Hyperoccipital carina: absent. Pro- notum in dorsal view: large, collarlike. Dorsoventral band of dense setation on poste- rior part of lateral pronotum: absent. Setation of pronotal cervical sulcus: sparse to ab- sent. Width of dorsal mesopleuron in lateral view: half as wide dorsally. Longitudinal striation on dorsal mesopleuron: absent. Transepisternal line: present. Mesofurcal pit: uncertain, present. Notaulus: absent. Wings: brachypterous. Color of legs: yellowish brown. Foamy structures of lateral propodeum: smaller than hairy metapleuron. Me- dian tubercule on T2: present. Setation of anterior T2: interrupted medially. Lateral patch on T2: absent. Foamy structures on S1: absent. Transverse felt field on anterior S2: present as transverse strip. Shape of S2 in lateral view: broadly convex. Diagnosis. Helava microptera and H. reducta are the only species without fully developed wings. They are easily separable by the presence of ocelli, a transepisternal line, and a scutoscutellar sulcus in H. microptera, all of which are absent in H. reducta. Etymology. The Greek name “microptera” refers to the small size of the wings in males and females of this species. 16 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) i] we! a 00” ees Figures 35-39. Helava microptera 35 female holotype (USNMENT00989197), head, mesosoma, metasoma, lateral view 36 female holotype (USNMENT00989197), head, mesosoma, metasoma, dorsal view 37 female holotype (USNMENT00989197), head, anterior view 38 male paratype (USNMENT00989198), antenna, dorsal view 39 female holotype (USNMENT00989197), antenna, dorsal view. Scale bars in millimeters. Link to distribution map. http://hol.osu.edu/map-large.html?id=354386 Material examined. Holotype, female: ECUADOR: Napo Prov., paramo, Quito- Baeza Rd., 4200m, 2.1J1.1979, pan trap, W. R. M. Mason, USNMENT00989197 (deposited in CNCI). Paratypes: ECUADOR: 9 females, 2 males, CNC424772, 424774424782, USNMENT00989198 (CNCI). Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) 17 Helava pygmea Masner & Talamas, sp. n. http://zoobank.org/A1 1 DCOE8-4C75-4A4C-B4BD-FC67D0374CFC http://bioguid.osu.edu/xbiod_concepts/35439 1 Figures 40-44 Description. Female body length: 0.94-1.54 mm (n=18). Male body length: 0.85— 1.07 mm (n=7). Male antenna: apically clubbed. Number of antennomeres in male club: 3. Number of female clavomeres: 4. Setation of frons anterior to ocellar triangle: present. Setation of vertex posterior to lateral ocellus: dense. Hyperoccipital carina: absent. Pronotum in dorsal view: present mostly as lateral shoulders. Dorsoventral band of dense setation on posterior part of lateral pronotum: absent. Setation of pro- notal cervical sulcus: uncertain, dense. Width of dorsal mesopleuron in lateral view: about equal ventrally and dorsally to 1.5 times as wide ventrally. Longitudinal striation on dorsal mesopleuron: absent. Transepisternal line: present. Mesofurcal pit: present. Notaulus: absent. Rs+M in fore wing: nebulous. Wings: macropterous. Rs+M in hind wing: nebulous. Color of legs: brownish, yellowish brown. Foamy structures of lat- eral propodeum: smaller than hairy metapleuron. Median tubercule on T2: present. Setation of anterior T2: interrupted medially. Lateral patch on T2: absent. Foamy structures on S1: absent. Transverse felt field on anterior S2: present as transverse strip. Shape of S2 in lateral view: broadly convex. Diagnosis. Helava pygmea is closest to H. simplex, with which it shares the presence of small propodeal foamy structures, fully developed wings, and a mesoscutum without notauli. The only character that reliably separates these species is the form of the basal vein (Rs+M) in the fore wing: darkly pigmented in H. pygmea and absent in H. simplex. Etymology. The species name “pygmea’” refers to the small size of the body in this species. Link to distribution map. http://hol.osu.edu/map-large.html?id=354391 Material examined. Holotype, female: ECUADOR: Napo Prov., below Papal- lacta, 3000m, 17.11.1983, L. Masner, USNMENT00989208 (deposited in CNCI). Paratypes: (20 females, 9 males) CHILE: 1 male, USNMENT00989196 (CNCI). COLOMBIA: 11 females, 4 males, CNC424746, 424748, 424754424756, 424758— 424763, 424768-424771 (CNCI). ECUADOR: 5 females, 3 males, CNC424747, 424749424751, 424764, 424766-424767, USNMENT00989207 (CNCI). VENE- ZUELA: 4 females, 1 male, CNC424752—424753, 424757, 424765, 424942 (CNCI). Helava reducta Masner & Talamas, sp. n. http://zoobank.org/908D245E-2CF2-495B-B711-87CB4D6AB3EE http://bioguid.osu.edu/xbiod_concepts/354389 Figures 45—49 Description. Female body length: 1.01 mm (n=1). Male body length: 1.04 mm (n=1). Male antenna: apically clubbed. Number of antennomeres in male club: 3. Number of 18 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) Figures 40-44. Helava pygmea 40 female holotype (USNMENT00989207), head, mesosoma, metasoma, lateral view 41 male paratype (USNMENT00989208), head, mesosoma, metasoma, dor- sal view 42 female holotype (USNMENT00989207), head, anterior view 43 male paratype (USN- MENT00989208), antenna, dorsal view 44 female holotype (USNMENT00989207), antenna, dorsal view. Scale bars in millimeters. female clavomeres: 4. Setation of frons anterior to ocellar triangle: sparse throughout. Se- tation of vertex posterior to lateral ocellus: very sparse or absent. Hyperoccipital carina: absent. Pronotum in dorsal view: large, collarlike. Dorsoventral band of dense setation on posterior part of lateral pronotum: absent. Setation of pronotal cervical sulcus: sparse to absent. Width of dorsal mesopleuron in lateral view: half as wide dorsally. Longitudi- Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) 19 49 Figures 45-49. Helava reducta 45 female holotype (USNMENT00989203), head, mesosoma, meta- soma, lateral view 46 female holotype (USNMENT00989203), head, mesosoma, metasoma, dor- sal view 47 female holotype (USNMENT00989203), head, anterior view 48 male paratype (USN- MENT00989204), antenna, dorsal view 49 female holotype (USNMENT00989203), antenna dorsal view. Scale bars in millimeters. nal striation on dorsal mesopleuron: absent. Transepisternal line: absent. Mesofurcal pit: present. Notaulus: absent. Wings: brachypterous. Color of legs: yellowish brown. Foamy structures of lateral propodeum: smaller than hairy metapleuron. Median tubercule on T2: absent. Setation of anterior T2: interrupted medially. Lateral patch on T2: absent. Foamy structures on S1: absent. Transverse felt field on anterior S2: present as transverse strip. Shape of S2 in lateral view: broadly convex. 20 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) Diagnosis. Helava reducta can easily be identified by severe microptery, the ab- sence of a scutoscutellar sulcus, and the absence of transepisternal line. Etymology. The Latin adjectival epithet “reducta” is applied to this species for the reduced segmentation of the mesosoma, Link to distribution map. http://hol.osu.edu/map-large.html?id=354389 Material examined. Holotype, female: VENEZUELA: Mérida St., Black La- goon, Sierra Nevada National Park, 3500m, 29.IV.1981, sweeping, L. Masner, USN- MENT 00989203 (deposited in CNCI). Paratypes: VENEZUELA: 1 female, 2 males, CNC424744-424745, USNMENT00989204 (CNCI). Helava samanthae Masner & Talamas, sp. n http://zoobank.org/33DE4823-9B7 1 -4510-AG4E- GE2C2A549E9A http://bioguid.osu.edu/xbiod_concepts/354387 Figures 50-54 Description. Female body length: 0.98—1.26 mm (n=19). Male body length: 0.82—1.32 mm (n=21). Male antenna: apically clubbed. Number of antennomeres in male club: 4. Number of female clavomeres: 4. Setation of frons anterior to ocellar triangle: present. Setation of vertex posterior to lateral ocellus: dense. Hyperoccipital carina: absent; indi- cated by lateral tubercules. Pronotum in dorsal view: present mostly as lateral shoulders. Dorsoventral band of dense setation on posterior part of lateral pronotum: absent. Se- tation of pronotal cervical sulcus: dense. Width of dorsal mesopleuron in lateral view: about equal ventrally and dorsally to 1.5 times as wide ventrally. Longitudinal striation on dorsal mesopleuron: absent. Transepisternal line: present. Mesofurcal pit: present. Notaulus: absent. Rs+M in fore wing: nebulous. Wings: macropterous. Rs+M in hind wing: nebulous. Color of legs: coxae yellow to brown, remaining segments yellow. Foamy structures of lateral propodeum: larger than hairy metapleuron. Median tubercule on T2: absent. Setation of anterior T2: interrupted medially. Lateral patch on T2: present. Foamy structures on S1: present at lateral margin of sternite. Transverse felt field on an- terior S2: present. Shape of S2 in lateral view: broadly convex. Diagnosis. ‘The form of the hyperoccipital carina as two lateral tubercules on the pos- terior vertex separates H. samanthae from all species except H. alticola, in which the form of the hyperoccipital carina is highly variable. These two species can be separated from each other by the setation of the upper frons, which in H. samanthae is densely present, and is sparsely present only along the inner orbits of the eye, or entirely absent, in 1. alticola. Etymology. This species is named for Samantha Fitzsimmons Schoenberger to thank her for excellent work performed as part of the Smithsonian Internship Pro- gram, including most of the photographs presented in this monograph. Link to distribution map. http://hol.osu.edu/map-large.html?id=354387 Material examined. Holotype, female: CHILE: Bio-Bio Reg., Nuble Prov., Las Trancas Valley, 1300m—1650m, 14.XII-17.XII.1976, S. Peck & H. Howden, USN- Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) pp) Figures 50-54. Helava samanthae, female holotype (USNMENT00989199) 50 head, mesosoma, metasoma, lateral view 51 head and mesosoma, dorsal view 52 metasoma, dorsal view 53 head, anterior 2 view 54 head and mesosoma, posterolateral view. Scale bars in millimeters. MENT00989199 (deposited in CNCI). Paratypes: (86 females, 25 males) ARGEN- TINA: 4 females, 3 males, CNC424909-4249 13, 424930—424931 (CNCI). CHILE: 82 females, 22 males, CNC424783-424851, 424895424906, 424914424929, 424932424937, USNMENT00989200 (CNCI). 22 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) Ld Rog Ly = i aS — “tan SS Figures 55-60. Helava simplex 55 female holotype (USNMENT00989195) head, mesosoma, metaso- ma, lateral view 56 female holotype (USNMENT00989195) head and mesosoma, dorsal view 57 female holotype (USNMENT00989195) metasoma, dorsal view 58 female holotype (USNMENT00989195) head, anterior view 59 female holotype (USNMENT00989195) antenna, dorsal view 60 female paratype (USNMENT00989190) fore wing, dorsal view. Scale bars in millimeters. 58 % 0.1 t Wf, Vy, ‘ae Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae) 23 Helava simplex Masner & Talamas, sp. n. http://zoobank.org/OB8E8E62-CE4F-4652-9C39-AD31F276E659 http://bioguid.osu.edu/xbiod_concepts/354385 Figures 55—60 Description. Female body length: 0.94—1.24 mm (n=12). Number of female cla- vomeres: 4. Setation of frons anterior to ocellar triangle: present. Setation of vertex posterior to lateral ocellus: dense. Hyperoccipital carina: absent. Pronotum in dor- sal view: present mostly as lateral shoulders. Dorsoventral band of dense setation on posterior part of lateral pronotum: absent. Setation of pronotal cervical sulcus: sparse to absent. Width of dorsal mesopleuron in lateral view: about equal ventrally and dorsally to 1.5 times as wide ventrally. Longitudinal striation on dorsal meso- pleuron: present. Transepisternal line: present. Mesofurcal pit: present. Notaulus: absent. Rs+M in fore wing: absent. Wings: macropterous. Rs+M in hind wing: ab- sent. Color of legs: yellow; yellowish brown. Foamy structures of lateral propodeum: smaller than hairy metapleuron. Median tubercule on T2: present. Setation of ante- rior ['2: interrupted medially. Lateral patch on T2: absent. Foamy structures on S1: absent. Transverse felt field on anterior S2: present as transverse strip. Shape of S2 in lateral view: broadly convex. Diagnosis. Helava simplex is the only macropterous species in the genus without a pigmented basal vein in the fore wing (Figure 60). Etymology. The epithet “simplex” is given to this species in reference to the ab- sence of several character states (notaulus, foamy structures). Link to distribution map. http://hol.osu.edu/map-large.html?id=354385 Material examined. Holotype, female: CHILE: Araucania Reg., Malleco Prov., site 649, 14km E Malalcahuello National Reserve, 1570m, 13.[X—31.XII.1982, trap, A. Newton & M. Thayer, USNMENT00989195 (deposited in CNCI). Paratypes: CHILE: 14 females, CNC424734-424740, 424742424743, 424943424945, 494812, USNMENT00989190 (CNCD). Acknowledgments We thank Smithsonian interns Samantha Fitzsimmons Schoenberger and Collin Schwantes for producing photographs and scanning electron micrographs, respective- ly, and Norman Johnson (OSUC) for developing and maintaining the Hymenoptera Online Database, Specimage, and vSysLab, which underlie the cybertaxonomy con- ducted here. This work was made possible by funding from the Systematic Entomol- ogy Laboratory and the National Institute of Food and Agriculture—Specialty Crop Research Initiative (USDA-NIFA-SCRI) #201 1-51181-30937. The USDA does not endorse any commercial product mentioned in this research. USDA is an equal op- portunity provider and employer. 24 EJ. Talamas & L. Masner / Journal of Hymenoptera Research 53: 1-24 (2016) References Hagedorn G, Catapano T, Giintsch A, Mietchen D, Endresen D, Sierra S, Groom Q, Biserkov J, Glockler F, Morris R (2013) Best practices for stable URIs. http://wiki.pro-ibiosphere. eu/wiki/Best_practices_for_stable_URIs Masner L, Huggert L (1989) World review and keys to genera of the subfamily Inostemmati- nae with reassignment of the taxa to the Platygastrinae and Sceliotrachelinae (Hymenop- tera: Platygastridae). Memoirs of the Entomological Society of Canada 147: 1-214. doi: 10.4039/entm121147fv Miké I, Vilhelmsen L, Johnson NF, Masner L, Pénzes Z (2007) Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa 1571: 1-78. Yoder MJ, Miko I, Seltmann K, Bertone MA, Deans AR (2010) A gross anatomy ontology for Hymenoptera. PLoS ONE 5(12): e15991. doi: 10.137 1/journal.pone.0015991