Revision of New World Helava Masner &amp; Huggert (Platygastridae, Sceliotrachelinae)

Elijah J. Talamas; Lubomír Masner

doi:10.3897/jhr.53.10217

Monograph

Revision of New World Helava Masner & Huggert (Platygastridae, Sceliotrachelinae)

Elijah J. Talamas^‡, Lubomír Masner^§

‡ USDA/SEL, Smithsonian Institution, Washington, DC, United States of America

§ Agriculture and Agri-Food Canada, Ottawa, Canada

Corresponding author: Elijah J. Talamas ( billy.jenkins@GMAIL.COM )

Academic editor: Matthew Yoder

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.

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

ZooBank: urn:lsid:zoobank.org:pub:13184D63-2047-4F62-A987-B844E6386BCD

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 terminology in which clavomeres are defined by the presence of basiconic sensilla. Our revision 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 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 Australian 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.

Figure 1.

Aphanomerus sp. female (USNMENT00916681), head, mesosoma, metasoma, lateral view. Scale bar in millimeters.

Figures 2–3.

Austromerus grandis, female paratype (USNMENT00916679) 2 head, mesosoma, metasoma, lateral view 3 head, mesosoma, metasoma, ventral view. Scale bars in millimeters.

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 taxonomic 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 Hymenoptera 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 magnification. 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 carbon 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 abbreviations 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

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 encountered 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 metapleuron (Figures 9, 16, 19, 29, 50)	2
–	Foamy structures on lateral propodeum covering area distinctly smaller than hairy metapleuron (Figures 4, 24, 35, 40, 55) or foamy structures absent (Figure 45)	5
2	Lateral pronotum with dorsoventral strip of dense setation posteriorly (Figure 9); notaulus percurrent (Figure 10); male antenna filiform (Figure 13); female antennal clava 3-merous (Figure 2)	H. allomera Masner & Talamas, sp. n.
–	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)	3
3	Upper frons densely setose (Figure 53)	H. samanthae Masner & Talamas, sp. n.
–	Upper frons glabrous or with sparse setae only along inner orbits (Figures 14–15, 21, 32)	4
4	Posterior vertex glabrous or only very sparsely setose (Figures 30, 34)	H. carinata Masner & Talamas, sp. n.
–	Posterior vertex densely setose (Figures 15, 20–21)	H. alticola Masner & Huggert
5	Wings reduced to strips, reaching only to anterior T2, or absent (Figures 36, 46)	6
–	Wings fully developed, exceeding apex of metasoma	7
6	Scutoscutellar sulcus absent (Figure 46); ocelli absent (Figure 46); mesopleuron without transepisternal line (Figure 45)	H. reducta Masner & Talamas, sp. n.
–	Scutoscutellar sulcus present (Figure 36); ocelli present (Figure 36); mesopleuron with transepisternal line (Figure 35)	H. microptera Masner & Talamas, sp. n.
7	Medial S2 distinctly projecting in lateral view in both sexes (Figure 4); mesopleuron without transepisternal line (Figure 4); antenna in male filiform, with A9 and A10 approximated (Figure 8)	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 (Figures 27, 43)	8
8	Notaulus present (Figure 25)	H. aureipes Masner & Talamas, sp. n.
–	Notaulus absent (Figures 41, 56)	9
9	Basal vein (Rs+M) in fore wing absent (Figure 60)	H. simplex Masner & Talamas, sp. n.
–	Basal vein (Rs+M) in fore wing nebulous (Figure 41)	H. pygmea Masner & Talamas, sp. n.

Figures 4–8.

Helava acutiventris 4 female holotype (USNMENT00989201), 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, anterior view 8 male paratype (USNMENT00989202), antenna, ventral view. Scale bars in millimeters.

Figures 9–13.

Helava allomera 9 female holotype (USNMENT00989217), head, mesosoma, metasoma, lateral view 10 female holotype (USNMENT00989217), head, mesosoma, metasoma, dorsal view 11 female holotype (USNMENT00989217), head, anterior view 12 female holotype (USNMENT00989217), antenna, dorsal view 13 male paratype (USNMENT00989218), antenna, dorsal view. Scale bars in millimeters.

Figures 14–18.

Helava 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.

Figures 19–23.

Helava alticola 19 female (USNMENT00989211), 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.

Figures 24–28.

Helava aureipes 24 female holotype (USNMENT00989205), head, mesosoma, metasoma, lateral view 25 female holotype (USNMENT00989205), head, mesosoma, metasoma, dorsal view 26 female holotype (USNMENT00989205), head, anterior view 27 male paratype (USNMENT00989206), antenna, anterior view 28 female holotype (USNMENT00989205), antenna, anterior view. Scale bars in millimeters.

Figures 29–34.

Helava carinata 29 female holotype (USNMENT00989215), head, mesosoma, metasoma lateral view 30 female holotype (USNMENT00989215), head and mesosoma, dorsal view 31 male paratype (USNMENT00989216), propodeum and metasoma, dorsal view 32 female holotype (USNMENT00989215), head, anterior view 33 male paratype (USNMENT00989216), antenna, dorsal view 34 male paratype (USNMENT00989216), head and mesosoma, dorsolateral view. Scale bars in millimeters.

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.

Figures 40–44.

Helava pygmea 40 female holotype (USNMENT00989207), head, mesosoma, metasoma, lateral view 41 male paratype (USNMENT00989208), head, mesosoma, metasoma, dorsal view 42 female holotype (USNMENT00989207), head, anterior view 43 male paratype (USNMENT00989208), antenna, dorsal view 44 female holotype (USNMENT00989207), antenna, dorsal view. Scale bars in millimeters.

Figures 45–49.

Helava reducta 45 female holotype (USNMENT00989203), head, mesosoma, metasoma, lateral view 46 female holotype (USNMENT00989203), head, mesosoma, metasoma, dorsal view 47 female holotype (USNMENT00989203), head, anterior view 48 male paratype (USNMENT00989204), antenna, dorsal view 49 female holotype (USNMENT00989203), antenna dorsal view. Scale bars in millimeters.

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 view 54 head and mesosoma, posterolateral view. Scale bars in millimeters.

Figures 55–60.

Helava simplex 55 female holotype (USNMENT00989195) head, mesosoma, metasoma, 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.

Helava acutiventris Masner & Talamas, sp. n.

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 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 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 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: distinctly 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 transepisternal line on the mesopleuron is shared only with H. reducta, which is a starkly different 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: Bío-Bío Reg., Ñuble Prov., Termas Rd., 60km SE Chillán, 1300m, 7.XII–19.XI.1985, flight intercept trap, S. Peck & J. Peck, USNMENT00989201 (deposited in CNCI). Paratypes: CHILE: 12 females, 25 males, CNC424698–424733, USNMENT00989202 (CNCI).

Comments

The diagnostic shape of S2 is found in both males and females, leading us to believe that this is not an adaptation for housing the retracted ovipositor system, as can be found in some species of Synopeas Förster and Platygaster Latreille.

Helava allomera Masner & Talamas, sp. n.

http://zoobank.org/F786DD15-F5B5-41CB-8910-479F75142D03

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: continuous 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 T2: absent. 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 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 Helava, this is the only species with a dorsoventral band of dense setae along the posterior margin of the lateral pronotum (Figure 9).

Etymology

The epithet “allomera” is given to this species in reference to the unusual 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: Araucanía Reg., Malleco Prov., 1200m, 37.809°S 73.016°W, Nahuelbuta National Park, 9.I–12.I.2000, Malaise trap, D. Webb & D. Yeates, USNMENT00989217 (deposited in CNCI). Paratypes: (58 females, 30 males) ARGENTINA: 7 females, 2 males, CNC424981–424985, 424993–424994, 425022, 425031 (CNCI). CHILE: 51 females, 28 males, CNC424946–424980, 424986–424992, 424995–425021, 425023–425030, 425032, 494556, USNMENT00989218 (CNCI).

Helava alticola Masner & Huggert

http://bioguid.osu.edu/xbiod_concepts/12334

Figures 14–18, 19–23

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. 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 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, 494622–494633, 494663–494671, USNMENT00989213–00989214 (CNCI); USNMENT00989943 (USNM).ECUADOR: 37 females, 17 males, CNC494567–494568, 494571–494592, 494594–494621, 494672–494673 (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, 494754–494765, 494767–494771, 494773–494781, 494784, 494794–494795, 494800–494803, 494806, 494807,494809, 494810, USNMENT00989211 (CNCI). ECUADOR: 10 females, 35 males, CNC494593, 494704, 494706–494724, 494726, 494749–494752, 494766, 494772, 494778, 494783, 494786, 494788–494793, 494796–494799, 494804, 494805,494808, 494811 (CNCI). VENEZUELA: 11 females, 14 males, CNC494725, 494729–494748, 494779, 494782, 494787, USNMENT00989212 (CNCI).

Helava aureipes Masner & Talamas, sp. n.

http://zoobank.org/96D177A2-D794-49FF-9B6C-54CBAAF05BCA

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. 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: 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 species 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 golden colour of the legs.

Link to distribution map

http://hol.osu.edu/map-large.html?id=354390

Material examined

Holotype, female: CHILE: Araucanía Reg., Malleco Prov., 1200m, 37.809°S 73.016°W, Nahuelbuta National Park, 9.I–12.I.2000, Malaise trap, D. Webb & D. Yeates, USNMENT00989205 (deposited in CNCI). Paratypes: CHILE: 25 females, 88 males, CNC424741, 425033–425101, 494411–494451, 494813, USNMENT00989206 (CNCI).

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 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: brownish. Foamy structures 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 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.I.1992, Malaise trap, E. E. Palacio, USNMENT00989215 (deposited in CNCI). Paratypes: (48 females, 56 males) COLOMBIA: 22 females, 25 males, CNC494452–494462, 494481–494485, 494491–494495, 494501, 494504–494505, 494519–494534, 494537–494541, 494555, USNMENT00989216 (CNCI). ECUADOR: 26 females, 31 males, CNC494463–494480, 494486–494490, 494496–494500, 494502–494503, 494507–494518, 494535–494536, 494542–494554 (CNCI).

Helava microptera Masner & Talamas, sp. n.

http://zoobank.org/71FA4BB9-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. 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. 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. 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 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.

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.III.1979, pan trap, W. R. M. Mason, USNMENT00989197 (deposited in CNCI). Paratypes: ECUADOR: 9 females, 2 males, CNC424772, 424774–424782, USNMENT00989198 (CNCI).

Helava pygmea Masner & Talamas, sp. n.

http://zoobank.org/A11DC0E8-4C75-4A4C-B4BD-FC67D0374CFC

http://bioguid.osu.edu/xbiod_concepts/354391

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 pronotal 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 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 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 Papallacta, 3000m, 17.II.1983, L. Masner, USNMENT00989208 (deposited in CNCI). Paratypes: (20 females, 9 males) CHILE: 1 male, USNMENT00989196 (CNCI). COLOMBIA: 11 females, 4 males, CNC424746, 424748, 424754–424756, 424758–424763, 424768–424771 (CNCI). ECUADOR: 5 females, 3 males, CNC424747, 424749–424751, 424764, 424766–424767, USNMENT00989207 (CNCI). VENEZUELA: 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 female clavomeres: 4. Setation of frons anterior to ocellar triangle: sparse throughout. Setation 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. Longitudinal 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.

Diagnosis

Helava reducta can easily be identified by severe microptery, the absence 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 Lagoon, Sierra Nevada National Park, 3500m, 29.IV.1981, sweeping, L. Masner, USNMENT00989203 (deposited in CNCI). Paratypes: VENEZUELA: 1 female, 2 males, CNC424744–424745, USNMENT00989204 (CNCI).

Helava samanthae Masner & Talamas, sp. n.

http://zoobank.org/33DE4823-9B71-4510-A64E-6E2C2A549E9A

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; indicated by lateral tubercules. 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: 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 anterior S2: present. Shape of S2 in lateral view: broadly convex.

Diagnosis

The form of the hyperoccipital carina as two lateral tubercules on the posterior 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 H. alticola.

Etymology

This species is named for Samantha Fitzsimmons Schoenberger to thank her for excellent work performed as part of the Smithsonian Internship Program, 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: Bío-Bío Reg., Nuble Prov., Las Trancas Valley, 1300m–1650m, 14.XII–17.XII.1976, S. Peck & H. Howden, USNMENT00989199 (deposited in CNCI). Paratypes: (86 females, 25 males) ARGENTINA: 4 females, 3 males, CNC424909–424913, 424930–424931 (CNCI). CHILE: 82 females, 22 males, CNC424783–424851, 424895–424906, 424914–424929, 424932–424937, USNMENT00989200 (CNCI).

Helava simplex Masner & Talamas, sp. n.

http://zoobank.org/0B8E8E62-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 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 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: absent. Rs+M in fore wing: absent. Wings: macropterous. Rs+M in hind wing: absent. Color of legs: yellow; yellowish brown. 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 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 absence 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: Araucanía Reg., Malleco Prov., site 649, 14km E Malalcahuello National Reserve, 1570m, 13.IX–31.XII.1982, trap, A. Newton & M. Thayer, USNMENT00989195 (deposited in CNCI). Paratypes: CHILE: 14 females, CNC424734–424740, 424742–424743, 424943–424945, 494812, USNMENT00989190 (CNCI).

Acknowledgments

We thank Smithsonian interns Samantha Fitzsimmons Schoenberger and Collin Schwantes for producing photographs and scanning electron micrographs, respectively, and Norman Johnson (OSUC) for developing and maintaining the Hymenoptera Online Database, Specimage, and vSysLab, which underlie the cybertaxonomy conducted here. This work was made possible by funding from the Systematic Entomology Laboratory and the National Institute of Food and Agriculture–Specialty Crop Research Initiative (USDA–NIFA–SCRI) #2011-51181-30937. The USDA does not endorse any commercial product mentioned in this research. USDA is an equal opportunity provider and employer.

References

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Masner L, Huggert L (1989) World review and keys to genera of the subfamily Inostemmatinae with reassignment of the taxa to the Platygastrinae and Sceliotrachelinae (Hymenoptera: Platygastridae). Memoirs of the Entomological Society of Canada 147: 1–214. doi: 10.4039/entm121147fv

Mikó I, Vilhelmsen L, Johnson NF, Masner L, Pénzes Z (2007) Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa 1571: 1–78.

Yoder MJ, Mikó I, Seltmann K, Bertone MA, Deans AR (2010) A gross anatomy ontology for Hymenoptera. PLoS ONE 5(12): e15991. doi: 10.1371/journal.pone.0015991