JHR 87: 81-113 (2021) gee JOURNAL OF “revered ovensccese our doi: 10.3897/hr’87.58368 MONOGRAPH ) Hymenoptera https://jhr.pensoft.net The Imarasional Society of Hymenopeeriss, RESEARCH Revision of the Australian genus Alfredella Masner & Huggert (Hymenoptera, Platygastridae, Sceliotrachelinae) Zachary Lahey', Elijah Talamas’, Lubomir Masner?, Norman F. Johnson! | Department of Evolution, Ecology, and Organismal Biology, The Ohio State University, 1315 Kinnear Road, Columbus, Ohio 43212, USA 2. Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Division of Plant Industry, 1911 SW 34” St. Gainesville, Florida 32608, USA 3 Agriculture and Agri-Food Canada, K.W. Neatby Building, Ottawa, Ontario KIA 0C6, Canada Corresponding author: Zachary Lahey (lahey. 18@osu.edu) Academiceditor: Matthew Yoder | Received 18 October 2020 | Accepted 23 February 2021 | Published 23 December 2021 http://zoobank. org/6DB1F7F4-7402-4B7D-92C0-32F97800EF00 Citation: Lahey Z, Talamas E, Masner L, Johnson NF (2021) Revision of the Australian genus A/fredella Masner & Huggert (Hymenoptera, Platygastridae, Sceliotrachelinae). In: Lahey Z, Talamas E (Eds) Advances in the Systematics of Platygastroidea III. Journal of Hymenoptera Research 87: 81-113. https://doi.org/10.3897/jhr.87.58368 Abstract The genus Alfredella Masner & Huggert is revised. Alfredella tasmanica Masner & Huggert is redescribed, Al. teres (Buhl), comb. nov. is transferred to Al/fredella from Amitus Haldeman, and Al. auriel Lahey, sp. nov. (New South Wales, Tasmania) and Al mephisto Lahey, sp. nov. (Western Australia) are described as new. The genus Masnerium Polaszek, syn. nov. is treated as a junior synonym of Amitus, and its type spe- cies, M. wellsae Polaszek, is transferred to Amitus as Am. wellsae (Polaszek), comb. nov. The relationship between A/fredella and morphologically similar genera is discussed, and a key is provided to distinguish between Aleyroctonus Masner & Huggert, Alfredella, and Amitus. Keywords Aleyroctonus, Amitus, endemic, Platygastroidea, Tasmania, taxonomy Introduction The subfamily Sceliotrachelinae contains a diverse assemblage of genera, most of which contain a small number of described species. Sceliotracheline wasps are minute in size, infrequently collected, and are most diverse in regions where they have been studied Copyright Zachary Lahey et al. 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. 82 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 81-113 (2021) the least. Masner and Huggert (1989) made a significant contribution to the study of these wasps in their seminal treatise of the subfamily and described 13 new monotypic genera, most of which are known only from the Southern Hemisphere. In particular, the landmasses that Australasia comprises (Australia, Melanesia, New Zealand, and Tasmania) are major centers of platygastrid diversity (Masner and Huggert 1989). Nineteen of the 28 described genera of Sceliotrachelinae are found throughout this region, eight of which are endemic to either Australia or New Zealand. This research is conducted as part of an effort to revise the fauna of Sceliotracheli- nae, with priority given to monotypic genera (Talamas and Masner 2016; Lahey et al. 2019b, c). Our purpose is to revise the species-level taxonomy of Alfredella Masner & Huggert, update its generic concept, and compare the genus to putative closely related taxa. Masner and Huggert (1989) considered Amitus Haldeman to be the sister taxon to Alfredella, a relationship corroborated by a recent phylogenomic analysis of Platygastroidea based on ultraconserved elements (Blaimer et al., unpublished data). Both genera are extremely similar morphologically, and we compared specimens of the two genera to ensure the validity of A/fredella when developing the generic key. During this process, Amitus specimens from around the world were examined, along with new material of the monotypic Masnerium Polaszek. The contributions of the authors are as follows: Z. Lahey: character definition and coding, generic concept development, key development, manuscript preparation, species concept development; E. Talamas: character definition, imaging, manuscript preparation; L. Masner: provision of specimens; and N.E Johnson: project coordina- tion, manuscript preparation. Materials and methods The numbers prefixed with “OSUC”, “USNMENT”, and “ZMUC” are unique identifiers for the individual specimens. Details of the data associated with these specimens may be accessed at https://mbd-db.osu.edu/ by entering the unique speci- men identifier (e.g., OSUC 666417) in the form (note the blank space after some acronyms). The following terms and abbreviations are used to describe the morphological structures discussed in the text — sensillar formula of clavomeres: distribution of pap- illary sensilla (PS) on the ventral clavomeres of the female (Yang et al. 2016), with the segment interval followed by the number of PS per segment (e.g., A10—A8/1-2- 2) (Bin 1981); LOL: lateral ocellar line, shortest distance between the outer margins of the lateral and median ocelli (Masner 1980); OD: ocellar diameter, greatest width of each ocellus; OOL: ocular ocellar line, shortest distance between the inner orbit and outer margin of the lateral ocellus (Masner 1980); POL: posterior ocellar line, shortest distance between the inner margins of the lateral ocelli (Masner 1980); T1, ‘E2.+:. 16: métasomal téreitetls 25 --.6;-5:15S2)....SG6nmetasomal-sternite: I; 2; -....06. Morphological terminology follows Masner and Huggert (1989), Miko et al. (2007), and Lahey et al. (2019a—c). Morphological terms were matched to concepts in the Revision of Alfredella Masner & Huggert 83 Hymenoptera Anatomy Ontology (Yoder et al. 2010) using the text analyzer func- tion. A table of morphological terms and URI links is provided in Suppl. material 1. Morphological character matrices and taxonomic descriptions were generated with vSysLab (https://vsyslab.osu.edu/), a web application that utilizes specimen data from the Hymenoptera Online Database (https://mbd-db.osu.edu/). Morphological charac- ters are in the format of character: character state. Multistate characters are separated by a semicolon (e.g., number of clavomeres: 4; 5). The point map of Alfredella species distributions was created with SimpleMappr (Shorthouse 2010). Nomenclature of the bio- and ecoregions of Australia inhabited by Alfredella species follows that used in the Interim Biogeographic Regionalisation for Australia version 7 (https://www.environment.gov.au/land/nrs/science/ibra/australias- bioregions-maps) (Thackway and Cresswell 1995). Photographs of card- or point-mounted insects were captured using a Macroscopic Solutions Macropod Micro Kit, with optical slices rendered in Helicon Focus. Image stacks were processed with CombineZP to produce single montage images, which were subsequently imported into Adobe Photoshop CC to correct for brightness and con- trast. Most scanning electron micrographs were taken with a Phenom XL G2 Desktop Scanning Electron Microscope, using a eucentric stage and Phenom ProSuite Software. The scanning electron micrographs of Amitus (Figs 57-59, 61) were taken following the methods of Talamas et al. (2016). Collections This work is based on specimens deposited in the following repositories: ANIC Australian National Insect Collection, Canberra, ACT, Australia CNCI Canadian National Collection of Insects, Ottawa, Ontario, Canada NHMUK_ Natural History Museum, London, United Kingdom OSUC C.A. Triplehorn Collection, The Ohio State University, Columbus, Ohio, USA USNM Smithsonian National Museum of Natural History, Washington, DC, USA ZMUC Zoological Museum, Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark Abbreviations and characters annotated in the figures ac acetabular carina (Fig. 58) afp anterior mesofurcal pit (Figs 58, 60) als anterolateral striae on T2 (Figs 3, 50, 61) amp anterior mesosternal pits (Figs 58, 60) apT2 anteromedial pits on T2 (Fig. 61) atp —_ anterior tentorial pit (Fig. 59) auc —axillular carina (Figs 2, 61-63) axu _—_axillula (Figs 61, 62) 84 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 81-113 (2021) cly __ clypeus (Fig. 59) daa dorsal axillar area (Fig. 61) ecc __epiclypeal carina (Fig. 59) fs foamy structures (Figs 3, 61) mas malar sulcus (Fig. 49) mbd = mandible (Fig. 59) mnt metanotal trough (Figs 62, 63) msc mesoscutum (Fig. 61) mshs_ mesoscutal humeral sulcus (Fig. 61) not _ notaulus (Fig. 61) pacp _ postacetabular pits (Figs 58, 60) pssu_ prespecular sulcus (Fig. 33) psu _ posterior mesoscutellar sulcus (Fig. 20) R radial vein (submarginal vein) (Fig. 2) sce _ setation of compound eye (Fig. 2) scu mesoscutellum (Fig. 61) sss scutoscutellar sulcus (Figs 1, 61) tac transaxillar carina (Figs 2, 61-63) tel transepisternal line (Figs 51, 58) tmc —_ transmetanotal carina (Figs 62, 63) ts torular striae (Fig. 59) ty tyloid (Figs 38, 40, 41, 55, 57) Character discussion Anterolateral striae on T2 Alfredella and Amitus have striae that radiate from the anterior margin of T2 as well as the margins of the anteromedial pits (Figs 3, 50, 61). This character is particularly subtle in Al/fredella relative to Amitus and requires the anterior margin of T2 to be free of obstructions such as the wings, hind legs, and detritus. Another Australian genus, Oligomerella Masner & Huggert, also possesses striations on T2; however, this genus has a distinct malar sulcus (Fig. 49) and the 3-merous clava is massive relative to A3— A7. We provide images of this exceptionally rare genus for the first time (Figs 46-49). Claval formula The antennal clava of Alfredella is 4- or 5-merous, with a papillary sensillar formula of 1-2-2-1 (AL auriel), 1-2-2-1-1 (AL teres), or 1-2-2-2-1 (AL. auriel, Al. mephisto, Al. tas- manica). Identifying specimens based on this character alone can be challenging and is best performed when examining the antennae ventrally or at an oblique angle. A small drop of exudate can usually be seen at the tip of each papillary sensillum, which serves as a good indicator of their presence (arrows on Fig. 30). Revision of Alfredella Masner & Huggert 85 Shape of clypeus The shape of the ventral margin of the clypeus differs between species of Alfredella. In Al tasmanica, the ventral clypeal margin is rounded (Fig. 44), whereas it is slightly more acuminate in Al mephisto (Fig. 43). The clypeus is distinctly acuminate in A/ auriel and Al. teres; however, these two taxa differ by the setation of the clypeal area: sparse in Al. auriel (Fig. 42) and dense in Al. teres (Fig. 43). Tyloid Males of three of the four A/fredella species possess a noticeable tyloid on A4. The shape of the tyloid is plate-like in Al. auriel (Fig. 38) and Al. teres (Fig. 41), a potential syna- pomorphy that supports A/fredella and Amitus as sister groups. The shape of the tyloid is less clear in Al. tasmanica: its presence is indicated by a ventral ridge that extends most of the length of A4 (Fig. 40), but we were not able to acquire images that com- pletely show this character. A tyloid was not observed on A4 of Al. mephisto (Fig. 39), most likely due to the accumulation of detritus on the only specimen available for scanning electron microscopy. Key to Aleyroctonus, Alfredella, and Amitus 1 Tubular R vein of fore wing absent (Fig. 1); epiclypeal carina present (Fig. 59); transaxillar and axillular carinae fused (Figs 1, 61, 62); mesopleuron without prespectlar sulcus Aric 5a nna is. 0s. Meee ok weeeles Amitus Haldeman = Tubular R vein of fore wing present (Fig. 2); transaxillar and axillular carinae separated (Figs 2, 63); mesopleuron with prespecular sulcus indicated dor- 1 A i fe So.) Baa ne A An ee Sear ey a So Ree te eT RRS POS A 2 2 Propodeum without foamy structures (Fig. 2); anterolateral surface of T2 without longitudinal striae (Fig. 2); setation of compound eyes dense (Fig. 2) saus stempatee easunhinsaiaen teas teauasenwashh eaSewasomaents ttha Aleyroctonus Masner & Huggert = Propodeum with foamy structures (Fig. 3); anterolateral surface of T2 with striae radiating from anteromedial pit (Fig. 3); setation of compound eyes shortiornot, distinct (Fic. 3)x. sas fasssetustossnt Alfredella Masner & Huggert tactauc Figure |. Amitus granulosus MacGown & Nebeker, female holotype (USNMENT01059097), head, mesosoma, metasoma, dorsal view. Scale bar in millimeters. 86 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 81-113 (2021) itt = S Ree SS & WY ew Figure 2. Aleyroctonus stanslyi Lahey & Polaszek, female holotype (OSUC 697919), head, mesosoma, metasoma, dorsolateral view. Scale bar in millimeters. Key to species of Alfredella 1 Notauli constricted posteriorly (Figs 7, 29); transepisternal line weakly arched (GREG 5) 2 coe tesco R on DAS i at ME Baie New Aaas ce tuts be 2 Notauli of even width throughout their length or absent (Figs 13, 20); tran- sepisternal line strongly arched medially (Figs 12, 16, 22)... ceeeeeeeees 3 Medial portion of mesoscutum and mesoscutellum setose (Figs 26, 29, 32); surface sculpture of mesoscutum and mesoscutellum composed of large re- Licwlationish( Pies 95t3)) m8... ee mri a, Mes ones Alfredella teres (Buhl) Medial portion of mesoscutum and mesoscutellum glabrous (Figs 5, 7, 10); surface sculpture of mesoscutum and mesoscutellum composed of fine reticu- lations CE iesi, 07 Ab 0)" cere, creme Alfredella auriel Lahey, sp. nov. Notauli parallel (Figs 18, 20, 25); posterior mesoscutellar sulcus absent medi- ally (Fig. 20); ventral margin of clypeus rounded (Fig. 44)... eee eeeeees sian Avs e354 ly aes anes meacies, eran nea Alfredella tasmanica Masner & Huggert Notauli converging posteriorly or notauli absent (Figs 11, 13); posterior mes- oscutellar sulcus complete (Figs 11, 13); ventral margin of clypeus acuminate Crees) Prermron verse ce rnarenc aeons Alfredella mephisto Lahey, sp. nov. Taxonomy Alfredella Masner & Huggert Alfredella Masner & Huggert, 1989: 39 (original description. Type: Alfredella tasman- ica Masner & Huggert, by monotypy and original designation); Vlug 1995: 10 (citation of type species); Lahey, Masner, and Johnson 2019: 69 (keyed). Diagnosis. Alfredella is most similar to Aleyroctonus and Amitus. The combination of a tubular R vein in the fore wing (absent in Amitus) and anterolateral striae on T2 Revision of Alfredella Masner & Huggert 87 (absent in Aleyroctonus) is enough to separate Al/fredella from these genera. Moreover, Alfredella is one of only two platygastroid genera where the antennal clava is com- posed of both articulated (e.g., AG and A7) and ‘compact’ (e.g., A8—A10) clavomeres. Additional diagnostic characters include the presence of malar striae, a 4- or 5-merous antennal clava, a distinct transepisternal line, and the presence of foamy structures on the propodeum. Description. Head. Color of head: black. Shape of head in dorsal view: transverse. Occipital carina: present. Setation of compound eye: present, short. Hyperoccipital carina: absent. Occipital pit: absent. Paraocellar depressions: absent. Preocellar depres- sions: absent. Setation of occiput: present, short. Antennal scrobe: present. Frontal ledge: absent. Sculpture of frons immediately dorsal to toruli: concentrically rugose. Sculpture of upper frons: reticulate. Sculpture of vertex: reticulate. Malar striae: pre- sent. Malar sulcus: absent. Facial striae: absent. Epistomal sulcus: absent. Anteclypeus: undifferentiated from postclypeus. Orientation of mandiblar teeth: transverse. Man- dibular dentition: bidentate. Number of maxillary palpomeres: 1. Number of labial palpomeres: 1. Number of antennomeres in female: 10. Number of antennomeres in male: 9. Number of clavomeres: 4; 5. Sensillar formula of A10—AG: 1-2-2-1-0; 1-2-2- 1-1; 1-2-2-2-1. Condition of AG: articulate. Condition of A7: articulate. Condition of A10—A8, female: fused, sutures present. Length of pedicel: approximately as long as A3+A4. Mesosoma. Epomial carina: present ventrally. Pronotal shoulders: visible in dorsal view, not sharply angled. Sculpture of mesoscutum: reticulate. Anterior admedian line: present. Median mesoscutal line: absent. Notaulus: present. Parapsidal line: present. Mesoscutal humeral sulcus: present as a thin furrow. Netrion: present. Scutoscutellar sulcus: present as a deep, noncrenulate groove. Sculpture of mesoscutellum: reticulate. Shape of mesoscutellum: nearly hexagonal, widest medially. Setation of axillula: dense. Metascutellum: concealed by posterior margin of mesoscutellum. Prespecular sulcus: present. Transepisternal line: present, terminating in anterior and posterior pits. Meso- pleural carina: absent. Metapleural carina: present. Metapleural sulcus: absent. Meta- pleural pit: present. Location of metapleural pit: at anterior margin of metapleuron. Paracoxal sulcus: absent. Setation of plical area: dense. Protibial spur: bifid. Tibial spur formula: 1-2-2. Tarsal formula: 5-5-5. Length of tarsal claws: equal. Foamy structures: present on lateral propodeal carinae and metapleural carina. Metasoma. Number of visible terga, female: 6. Number of visible terga, male: 8. Number of visible sterna: at least 6. Width of laterotergites: short. Sculpture of terga: T2 anterolaterally striate. Laterotergites: present. Laterosternites: absent. Sculpture of nucha: longitudinally foveolate. Shape of T1: transverse. Anterior pits on T2: present as transverse, setose depressions medially. Longest tergite: T2. Sculpture of S2: smooth except for longitudinal striation and reticulation surrounding felt field. Transverse felt field on anterior S2: present. Wings. Color of wings: hyaline. Wing development: macropterous. Length of fore wing: exceeding apex of metasoma. Marginal cilia of fore wing: present. R of fore wing: present, tubular. Length of fore wing R: approximately 1/3 length of fore wing. Shape of knob of R: truncate, not rounded. Cu of fore wing: nebulous. M of fore wing: 88 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 81-113 (2021) nebulous. M+Cu of fore wing: nebulous. Marginal cilia of hind wing: present. R of hind wing: present, tubular. Distribution. Mainland Australia (Australian Capital Territory, New South Wales, Queensland, South Australia, Western Australia) and the island of Tasmania. The ab- sence of specimens from Victoria is likely due to a lack of collecting. Species descriptions Alfredella auriel Lahey, sp. nov. http://zoobank.org/6A9392CE-6320-4FD7-84C9-FB052D7553BA Figs 3, 5-10, 34, 38, 42 Description. Setation of mandibles: present. Setation of clypeal area: sparse. Shape of clypeus: acuminate. Anterior tentorial pits: small. Facial and malar striae: present, dorsal striations confluent with concentric sculpture of antennal scrobe. Central keel: absent. Setation of interantennal area: extending to dorsal margin of toruli. Sculpture of frons: reticulate dorsally and along inner orbits, reticulations fading medially, supplanted by concentric striations of antennal scrobe, punctation present near ocelli. Position of lateral ocellus: approximately 1 OD from inner margin of compound eye. Interocular space: 1.25x length of compound eye. Occiput directly anterior to anteromedial por- tion of occipital carina: granulate. Sculpture of occiput: reticulate; granulate. Color of antennomeres: yellow-orange; concolorous with legs. Number of papillary sensilla on AO: 0; 1. Number of papillary sensilla on A7: 1. Glabrous patch on lateral propodeal area adjacent to spiracle: present. Shape of mesoscutellum in lateral view: flat. Shape of transepisternal line: weakly arched. Shape of mesoscutellum in dorsal view: pentago- nal. Sculpture of posterior mesoscutellar sulcus: striate. Posterior mesoscutellar sulcus: complete. Setation of mesoscutum: absent posteromedially, dense lateral to notauli. Setation of mesoscutellum: present throughout. Path of notauli: converging posteri- orly. Shape of notaulus: abruptly widening posteriorly. Setation of anteromedial T2: interrupted medially. Number of setae on lateral surface of T3: 3. Number of setae on ic y = 4! z Figure 3. Alfredella auriel Lahey, female (USNMENT01197967), head, mesosoma, metasoma, dorsal view. Scale bar in millimeters. Revision of Alfredella Masner & Huggert 89 oie, Figure 4. Distribution of Alfredella Masner & Huggert across Australia. Multiple specimens collected from the same locality are indicated by a single circle. States and territories are abbreviated as follows: NSW (New South Wales), NT (Northern Territory), QLD (Queensland), SA (South Australia), TAS (Tasmania), VIC (Victoria), WA (Western Australia). lateral surface of T4: 4. Number of setae on lateral surface of T5: 5. Felt field on S2: 0.5 as long as It2. Setation of laterotergites: present on It2—It5. Setation of dorsal surface of hind coxa: absent medially. Color of coxae: yellow-orange. Diagnosis. This species is easily recognized by the shape of the notauli, lack of setae on the mesoscutal midlobe, and sparsely setose clypeal area. Distribution. New South Wales, Tasmania. Inhabited ecoregions (bioregions). Temperate Broadleaf and Mixed Forests (South Eastern Queensland, South East Corner, South Eastern Highlands, Tasmanian Southern Ranges). Etymology. Auriel is the Archangel of Hope from the Diablo video game fran- chise. The epithet is treated as a noun in apposition. Material examined. Holotype, female: Ausrratia: NSW, New England N. P, 1300-1500 m, Feb. 13,1984, L. Masner, s. s., OSUC 698019 (deposited in ANIC). Paratypes: AUSTRALIA: 8 females, 6 males, OSUC 698007—698009, 698011-698012, 698014-698015, 698017-698018, 698020-698024 (CNCI). Additional material: AUSTRALIA: 1 female, UJSNMENT01197967 (USNM). Comments. While most female specimens in the type series have a 4-merous clava, we examined four specimens (OSUC 698007-698009, USNMENT01197967) from Ko- sciuszko National Park (New South Wales) with 5 clavomeres. We treat these specimens under Al. auriel given a lack of additional characters that would warrant new species status. 90 Zachary Lahey et al. / Journal of Hymenoptera Research 87: 81-113 (2021) 3 Be us ®