JHR 86: 101-121 (2021) oe, JOURNAL OF Sermtertensing ern doi: 10.3897/jhr.86.71309 RESEARCH ARTICLE ) Hymenopter a 9 https://jhr.pensoft.net The marrational Sociry of Hymenoperiss, RESEARCH Description of Kavayva, gen. nov., (Chalcidoidea, Eurytomidae) and two new species associated with Guarea (Meliaceae), and a review of New World eurytomids associated with seeds Y. Miles Zhang', Michael W. Gates', Rogerio Silvestre*?, Manuela Scarpa? I Systematic Entomology Laboratory, Agricultural Research Service, U.S. Department of Agriculture, clo Na- tional Museum of Natural History, Smithsonian Institution, RO. Box 37012, MRC-168, Washington, DC 20013-7012, USA 2. Hymenoptera Ecology Laboratory- Hecolab, Universidade Federal da Grande Dourados- UFGD. Rod. Dourados Itahum km 12, Cidade Universitaria, 79804-970, Dourados, Mato Grosso do Sul, Brazil 3 Graduate Program in Entomology and Biodiversity Conservation - UFGD, Dourados, Brazil Corresponding author: Michael W. Gates (Michael.Gates@usda.gov) Academic editor: Petr Jansta | Received 8 July 2021 | Accepted 3 September 2021 | Published 29 October 2021 http://zoobank.org/0B892FB9-A903-44C4-9B5 D-4AF6D/6E48A2 Citation: Zhang YM, Gates MW, Silvestre R, Scarpa M (2021) Description of Kavayva, gen. nov., (Chalcidoidea, Eurytomidae) and two new species associated with Guarea (Meliaceae), and a review of New World eurytomids associated with seeds. Journal of Hymenoptera Research 86: 101-121. https://doi.org/10.3897/jhr.86.7 1309 Abstract Kavayva Zhang, Silvestre & Gates, gen. nov., and two species are described from the Neotropics, Kavayva bodoquenensis Zhang, Silvestre & Gates, sp. nov., and Kavayva davidsmithi, Zhang & Gates, sp. nov. Speci- mens of the new species were collected independently during separate research efforts in Peru and Brazil, reared from the seeds of Guarea F. Allam ex L. (Meliaceae), which represents a new host plant family for Eurytomidae. A differential diagnosis of the New World seed-feeding eurytomids is also provided. Keywords Neotropical region, Phytophagy, seed chalcids Copyright Y.Miles Zhang 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. 102 Y.Miles Zhang et al. / Journal of Hymenoptera Research 86: 101-121 (2021) Introduction The Eurytomidae is one of the smaller family of Chalcidoidea, and the majority of their larvae feed endophytically as seed eaters, gall formers, or as parasitoids of phytophagous in- sects (Lotfalizadeh et al. 2007). Most seed-feeders of cultivated plants are considered as pest insects, which can be accidentally transported to new regions given their cryptic lifestyle. In the Neotropics, three genera have been recorded to be associated with seeds. ‘The most commonly encountered genus is Bephratelloides Girault, which are known seed feeders of Annonaceae (Grissell and Schauff 1990; Grissell and Foster 1996; Chang 1998). Bephratelloides abulus Grissell and Foster was erroneously reported to be associated with Diospyros digyna Jacq. (Ebenaceae), but was later shown to be Prodecatoma diospyri Muesebeck (Castafeda-Vilddzola et al. 2011; Ruiz-Montiel et al. 2021). Although not a major pest, Bephratelloides have been documented in various species of custard apple grown for human consumption, such as atemoya, cherimola and others in Florida (USA), Mexico and Brazil (Pefia and Bennett 1995; Moura et al. 2006; Hernandez-Fuentes et al. 2008; Castafeda-Vildézola et al. 2010). The genus is also commonly intercepted at US ports of entry and submitted to the Systematic Entomology Lab for identification, with 634 specimens submitted over the past 25 years (M. Touchet, pers. comm.). Three species of Bephratelloides were included in Lotfalizadeh et al. (2007) as part of the mor- phological phylogenetic analysis of Eurytominae, and the genus was weakly recovered by homoplastic characters including bilobed clypeus and relatively long postgenal bridge. Prodecatoma Ashmead have been recorded from South America, Africa, and Asia, although the genus is likely not monophyletic (DalMolin et al. 2004). Lotfalizadeh et al. (2007) redefined the genus in a restricted sense to contain only the Neotropical phy- tophagous species, which is supported by the following morphological characteristics: lower face strigose with a median carina continued on intertorular space; intertorular space raised into a broadly laminate and discoid projection continuing dorsally on the scrobal depression; and prepectus with subventral carinae distinctly diverging anterior- ly. This group now contains gall formers and inquilines on Araceae, Dipterocarpaceae, Ebenaceae, Fabaceae, Liliaceae, Myrtaceae, Meliaceae, Rubiaceae, Sabiaceae and Vita- ceae (DalMolin et al. 2004; Lotfalizadeh et al. 2007; Ruiz-Montiel et al. 2021). Paradecatoma Masi is a small genus restricted to Afrotropical region, with a single described species Paradecatoma bannensis Masi from the pyrene/drupe of Cordia africana Lam. (Boraginaceae) (Yirgu and Delvare 2019), and at least three undescribed species asso- ciated with seeds of Combretum glutinosum Perr. ex DC. and Terminalia macroptera Guill. & Perr. (Combretaceae) (Lotfalizadeh et al. 2007). All four species of Paradecatoma were included in the Lotfalizadeh et al. (2007) study, and the genus was weakly recovered with the following characters: narrow intertorular space and strongly raised margin on antennal toruli, lateral foraminal plate not delimited, and subforaminal bridge with vestigial median strip. Eurytoma werauhia Gates & Cascante-Marin (2004) is a phytophagous Neotropical species associated with floral buds of Werauhia gladioliflora (Wendl.) (Bromeliaceae), Lot- falizadeh et al. (2007) suggested that E. werauhia probably belongs to Paradecatoma Masi, which would greatly expand the biogeographic range of the genus. However, ongoing phylogenomic analysis suggests E. werauhia is actually a new genus (Zhang et al. in prep). Description of Kavayva and two new species associated with Guarea 103 Table |. Morphological comparison of comparison of New World Eurytomidae associated with seeds. Bepratellotdes ne Intertorular | minute, rounded projection] projection thin, long and | projection thin, long and no projection space with tip truncate narrow, pointy narrow, pointy Frontal wider ventrally, slightly wider ventrally, slightly wider ventrally, slightly wider dorsally, narrowing depression broadening towards broadening towards toruli, broadening towards towards toruli, anterior toruli, anterior ocellus not | anterior ocellus partially | toruli, anterior ocellus not ocellus not included included included included Ventral plaque no in some males no in all males, some females Forewing stigmated not stigmated not stigmated thickened in males, stigmated Procoxa striate-rugose smooth, with lamella Propodeum _| coarsely sculptured laterally, | coarsely sculptured laterally, | broad, flat, median furrow broad, flat, smooth broad, median furrow broad, no median furrow | areolate, slightly narrowing present or absent The research presented here is based on specimens collected independently dur- ing separate research efforts in Peru, Panama, and Brazil, reared from the seeds of Guarea F. Allam ex L. (Meliaceae), which represents a new host plant family for Eu- rytomidae (Fig. 1). The goal of this study is to describe the new genus Kavayva and the two new species, and provide an overview of the generic concepts of New World seed-feeding eurytomids. Materials and methods Field collection Guarea kunthiana A. Juss. (Meliaceae), is a perennial tree of secondary to late climax communities. Its height reaches up to 20 m and its diameter up to 60 cm, and it occurs commonly in semi-deciduous forests in Central and South American (Pennington and Clarkson 2013). The common name in Brazil is “Figo do Mato” (Lorenzi 2002). The unisexual flowers secrete nectar and are pollinated by Coleoptera and Lepidoptera, with seed dispersal by birds and rodents (Wenny 1999). Flowering occurs from November to December, but may occur sporadically throughout the year (Souza et al. 2002). Fruits were collected in a semi-deciduous Atlantic Forest (Fig. 2) at Serra da Bo- doquena, Bonito, Mato Grosso do Sul, Brazil, near the Taquaral river (21°06'56'S, 56°38'24"W), and the Boqueirao farm (21°07'31.8"S, 56°43'20.9"W), at altitudes of 582 m and 540 m above sea level, respectively. There were four seasonal samplings in May and December 2015, and in February and May 2016. The fruits were collected from trees in linear transects 1000 meters long, with a perpendicular distance of 5 meters on either side, when allowed by the topography of the area, totaling 10,000 square meters (1 ha). The fruits were collected manually with scissors (Fig. 3), the quantity varied ac- cording to the availability on each plant. They were individually placed, where possi- ble, in plastic pots containing sterilized sand as a substrate for pupal burial, with small holes in the lid for air circulation (Fig. 4). Pots were observed daily and parasitoids that 104 Y.Miles Zhang et al. / Journal of Hymenoptera Research 86: 101-121 (2021) aes tty Lite hs Figure |. Illustration of Kavayva davidsmithi with its host plant, Guarea guidonia (Meliaceae). The upper right shows the emergence holes from the fruit. Illustrated by Taina Litwak. emerged from the fruits kept alive six hours to fix the chromatic patterns, and then placed in ethanol and frozen. Voucher species were incorporated into the Hymenoptera collection (HyMB) of the Museum of Biodiversity (MuBio) of the Federal University of Grande Dourados Description of Kavayva and two new species associated with Guarea 105 Figure 2-6. Collection of Kavayva bodoquenensis 2 semi-deciduous forest along the Taquaral river, Serra da Bodoquena, Brazil 3 fruits of Guarea kuntiana on tree 4 individualized plastic pots with sterilized sand 5 seed damage by K. bodoquenensis 6 pupa of K. bodoquenensis. Photo | by Paulo Robson de Souza 3, 4 by Manuela Scarpa 5, 6 by Bhrenno Trad. 106 Y.Miles Zhang et al. / Journal of Hymenoptera Research 86: 101-121 (2021) (UFGD), Dourados-MS, Brazil. For the species of trees from which fruits are sampled, vouchers were collected, and sent to the Botanic Sector of the UFGD to be identified by Dr. Zefa Valdevina Pereira and incorporated into the MuBio Herbarium. Molecular protocol Specimens were extracted, amplified, and sequenced at the Laboratories of Analyti- cal Biology (LAB) at the Smithsonian Institution’s National Museum of Natural His- tory (NMNH, Washington, DC, USA). A single specimen of K. bodoquenensis was destructively sampled using the DNeasyT'M Tissue Kit protocol (Qiagen, Valencia, CA, USA). Fragments of mtDNA COI were amplified using LCO1490 5'-GGT- CAACAAATCATAAAGATATTGG-3' and HCO2198 5'-TAAACTTCAGGGT- GACCAAAAAATCA-3' (Folmer et al. 1994). PCR was performed using approximate- ly 2 ul DNA extract, 1.25 wL 10x Buffer, 1 pl dNTP, 1 pl of each primer, 1 unit of Taq DNA polymerase (TaKaRa Bio, Mountain View, CA, USA), and purified water for a final volume of 25 pl. Amplicons of COI were generated with an initial denaturation of 1 min at 95 °C, followed by 35 cycles at 95 °C for 15 s, 49 °C for 15 s and 72 °C for 45 s, and a final elongation period of 4 min at 72 °C. Sequencing was conducted using an ABI 3730xl DNA sequencer following the manufacturer's instructions. Con- tigs were assembled and edited using Geneious Prime v2021.1. DNA sequences were then compared with all available sequences in the Basic Local Alignment Search Tool (BLAST) for nucleotides in GenBank. Imaging Ethanol-preserved specimens were dehydrated through increasing concentrations of ethanol, and transferred to hexamethyldisilazane (HMDS) (Heraty and Hawks 1998) before point-mounting. MWG identified the specimens using a Leica M205C ster- eomicroscope with 10X oculars and a Leica LED ring light source for point-mounted specimen observation. We took scanning electron microscope (SEM) images with a Hitachi TM3000 (Tungsten source). Body parts of disarticulated specimens were ad- hered to a 12.7 X 3.2 mm Leica/Cambridge aluminum SEM stub by a carbon adhesive tab (Electron Microscopy Sciences, #77825—12). Stub-mounted specimens were sput- ter coated with gold-palladium using a Cressington Scientific 108 Auto from multi- ple angles to ensure complete coverage (~20—30 nm coating). Habitus images were obtained using a Visionary Digital imaging system. The system consists of a Canon EOS 5D Mark II digital SLR camera with a 65 mm macro lens. A Dynalite MP8 power pack and lights provided illumination. Image capture software was Visionary Digital’s proprietary application with images saved as TIF with the RAW conversion occurring in Canon Digital Photo Professional software. Image stacks were mounted with Helicon Focus 6.2.2. Image editing was done in Adobe Photoshop and plate lay- out in Adobe Illustrator. The painting (Fig. 1) was made from pinned and live insect specimens, plant herbarium sheets and photographs. Additional structural details of Description of Kavayva and two new species associated with Guarea 107 the insects were obtained from SEM photographs. ‘The final image was painted using Adobe Photoshop. All species identifications were corroborated by comparison with authoritatively identified specimens in the Smithsonian National Museum of Natural History. Termi- nologies used for surface sculptures follow Harris (1979), while the morphology fol- lows Gibson (1997), Lotfalizadeh et al. (2007), and Gates and Pérez-Lachaud (2012) for adults, and Short (1952), Roskam (1982), and Henneicke et al. (1992) for larvae. Abbreviations for museums are: MUSM — Natural History Museum of the San Mar- cos University, UFGD, Museum of Biodiversity of the Federal University of Grande Dourados, Dourados-MS, Brazil, and USNM — United States National Museum of Natural History, Washington, D.C., USA. Results The K. bodoquenensis adults emerged between May 8—29, 2016. Twenty-eight fruits were collected with a combined weight of 584.30 g (average = 20.87 g). The fruits were soft carmine color, with fibrous texture, without pulp, containing 4-8 almond-shaped seeds (Fig. 5). On average, 25% of the seeds were parasitized. Only one hole per seed was observed to indicate emergence of wasps. A total of 32 individuals emerged from seeds in the laboratory (Fig. 6), 20 females and 12 males, with a sex ratio of 0.6 m/f. An uni- dentified adult female Sesiidae (Lepidoptera) emerged from one of the seeds. The COJ sequence (676 bp, GenBank Accession # MZ483873) did not match any known species. Kavayva Zhang, Silvestre, & Gates, gen. nov. http://zoobank.org/E9334690-685 1-4440-8244-986B9BC45405 Figs 1, 7-27 Type species. Kavayva bodoquenensis Zhang, Silvestre, Gates. Diagnosis. Kavayva can be distinguished from other eurytomid genera by the fol- lowing combination of characters — presence of ventral plaque of scape form a projec- tion on the inner face below the attachment to pedicel in males (Kavayva bodoquenen- sis, Fig. 13) or both sexes (Kavayva davidsmithi, Fig. 26), F1 of antenna cylindrical and not constricted (Fig. 12), presence of deep black line along the malar sulcus (Fig. 25), middle of propodeum completely glabrous and smooth (Fig. 15), and associated with seeds of Guarea (Meliaceae). Description. Female body length 6.5—10 mm, male 6.5-9.4 mm. Color. Mostly yellow, black along malar sulcus, with brown infuscation or black bands on the dorsal mesosoma. Head. Quadrate with rounded corners, 2.4—2.5x as wide as long in dorsal view (Figs 9, 25), areolate-rugose with setae. Lower face weakly strigose, clypeus bilobed, mandible tridentate, supraclypeal area smooth, slightly concave, extending to the toruli Y.Miles Zhang et al. / Journal of Hymenoptera Research 86: 101-121 (2021) Figure 7-8. Lateral habitus of Kavayva bodoquenensis 7 female 8 male. Photos by Cecilia Escobar. Description of Kavayva and two new species associated with Guarea imm Figure 9-13. Kavayva bodoquenensis 9 frontal view of head 10 ventral view of head II dorsal view of head 12 male antenna 13 close up of ventral plaque on antennal scape. 110 Y.Miles Zhang et al. / Journal of Hymenoptera Research 86: 101-121 (2021) lee: p- 18 Figure 14-18. Kavayva bodoquenensis \4 lateral view of male mesosoma I5 dorsal view of male meso- soma 16 ventral view of mesosoma I7 ventral view of male metasoma 18 dorsal habitus of female speci- men collected from Panama. Description of Kavayva and two new species associated with Guarea 111 (Figs 9, 25). Malar sulcus present, incomplete, reaching about 74 of malar space. Malar space glabrous, smooth. Genal carina present. Toruli positioned parallel to the lower ocular line, diameter of torulus 3.3x that of the intertorular space. Intertorular space without projection between antennae (Fig. 9). Scrobal depression deeply excavated, converging ventrally in frontal view. Vertex areolate to umbilicate, anterior ocellus above scrobal depression. Scape with or without ventral plaque (females of Kavayva bodoquen- ensis). Antenna pedicel chalice-shaped, six funicular segments cylindrical with multiple irregular rows of longitudinal sensilla and whorls of setae, much shorter than its bearing segment, clava 2-segmented. Occiput concave, postgenal groove diverging, postgenal lamina present, subforaminal bridge ornamentation faint and inconspicuous (Fig. 10). Wing. Forewing slightly infumated below marginal and stigmal vein, or forming a narrow band that curves slightly proximally and extending half way down the wing (Figs 7, 27). Costal cell, basal cell, and speculum (except for anterior edge) setose. Mesosoma. Mesosoma umbilicate, 1.2—1.7x as long as broad. Notauli complete, shallow. Anterior pronotal carina widely interrupted. Femoral depression of mesopleu- ron weakly striate, mespeimeron smooth and shiny ventrally, bulging laterally (Fig. 14). Dorsellum carinae diverging. Propodeum in lateral view forming a 90° angle with meso- soma, broadly delimited by carinae forming a hexagon with raised lateral corners (Fig. 15). Median furrow of propodeum concave and smooth, bordered laterally by irregular setose cells. Forecoxa without oblique groove (Fig. 16). All femora with distal lamella, forecoxa without oblique groove. Metacoxa bare laterally, metatibia densely setose. Metasoma. Metasoma medially compressed, smooth, Gt4—Gt6 glabrous or setose. Petiole very short and not visible while specimen is intact (Figs 7, 22). Gaster S-shaped in lateral view, ovipositor angled at about 30° dorsad of horizontal axis. Gt4 may be emarginate posteriorly in dorsal view. Male. Color and sculpture as described similar to females. Ventral plaque on scape forming a projection on the inner face below the attachment point to pedicel (Figs 12, 13). Antennomeres with multiple rows of erect setae. Toruli positioned above the lower ocular line. Marginal vein swollen (Figs 8, 23). Gastral petiole striate dorsally, 1.5—1.7x as long as the length of metacoxa, smooth laterally. Etymology. In the Guarani Native American language “Kavayva” means: “wasp of the fruit that gives seeds”. Key to species of Kavayva 1 Mesosoma uniform without black bands (Fig. 18), forewing with secondary wing band (Fig. 7), only males have ventral plaques on the scape..............004 cai Be cyinvanshlaetee aaa naodn a kasokd aM oceans incense Kavayva bodoquenensis sp. nov. — Mesosoma with black bands (Fig. 24), forewing without secondary wing band (Fig. 27), both males and females have ventral plaques on the scape E04 2G) atest Seaton te tc oastlwiedtte Veewcuvce cote ateacestes Kavayva davidsmithi sp. nov. 112 Y.Miles Zhang et al. / Journal of Hymenoptera Research 86: 101-121 (2021) me, nase x ae Was « pis ie Jo 7 See Nee rt Tie beet 8 7 um ete oe eee - = ney = WM tT es f 1S le 5 ee el ea carr ena cana Figure 19-21. Kavayva bodoquenensis larva 19 lateral habitus, abdominal segment (ABS), anal segment (AS), thoracic segment (THS) 20 anterolateral view of head, antennae (An), anterior tentorial pit (At), clypeal setae (Ci), cranial depression (Dfm), interior frontal setae (Fi), superior frontal setae (Fs), anterior genal setae (Ge), hypostomal setae (Hy), labral setae (La) 21 ventral view of head, labium (Lb), mandible (Md), maxilla (Mx). Kavayva bodoquenensis Zhang, Silvestre, & Gates, sp. nov. http://zoobank.org/838C50D8-2 1 F5-4092-AEDA-3E9A2675D005 Figs 7-21 Material examined. Holotype Brazit * [1F]; MS, Bonito, Serra da Bodoquena; 21°06'56"S, 56°38'24" W; 8-29 May 2016; R. Silvestre leg.; ex fruit of Guarea kunthiana; Description of Kavayva and two new species associated with Guarea jas USNMENT01788104. Paratypes Brazit ¢ [9F, 9M]; same information as holotype; R. Silvestre and M. Scarpa leg.; USNMENT01788085-—103 © [8F, 3M]; same infor- mation as holotype; UFGD Hymb00023-CH-—00034-CH. Panama ¢ [3F, 4M]; Ar- raijan; Sep. 1938; J. Zetek leg.; ex. fruit of Guarea guarea; No. 4279, Lot # 88-17223; USNMENT01788078-084. Diagnosis. Kavayva bodoquenensis can be distinguished from K. davidsmithi by the lack of black bands across mesosoma in dorsal view (Fig. 18), and the lack of ventral plaque in females. Description. Holotype female. 6.5 mm in length. Color. Yellow except malar sulcus, supraclypeal area, Gt3-syntergum of the meta- soma (except for pairs of yellow patches dorsad of Gt4 and Gt5), proximal half of fem- ora and tibiae, wing veins, wing bands near basal setal line and marginal vein brown, edge of mandible, setae on head and mesosoma black and eyes pinkish red (Fig. 7). Head. Quadrate with rounded corners, 1.2x as wide as high in frontal view, 2.5x as wide as long in dorsal view, areolate-rugose with setae (Fig. 11). Lower face weakly strigose, clypeus bilobed, mandible tridentate, supraclypeal area smooth, extending to the toruli. Malar sulcus present, incomplete, reaching to 74 of malar space. Malar space glabrous, smooth. Genal carina present, smooth. Toruli positioned above lower ocular line about 1/3 of the eye length, diameter of torulus 3.3x that of the intertorular space. Scrobal depression deeply excavated, converging ventrally in frontal view (Fig. 9). Vertex areolate-umbilicate, anterior ocellus above scrobal depression, ratios of POL: 0OL:LOL 4:5:1. Scape without ventral plaque. Ratio of scape (minus radicle):pedicel:anellus:F1:F2:F3:F4:F5:F6:club as 14:3.3:1:6:6:6:5.3:4:7, pedicel chalice-shaped, funicular segments cylindrical with mulkti- ple irregular rows of longitudinal sensilla whorls of setae, much shorter than its bearing seg- ment, clava 2-segmented. Occiput concave, postgenal groove diverging, postgenal lamina present, subforaminal bridge ornamentation faint and inconspicuous (Fig. 10). Wing. Forewing infumated below marginal and stigmal vein, band narrow, curving slightly proximally, and extending half way down the wing. Basal and costal setal line also infumated. Ratio of marginal vein:postmarginal vein:stigmal vein as 1.5:1 (Fig. 7). Mesosoma. Mesosoma umbilicate, 1.2x as long as broad. Notauli complete, shallow (Fig. 18). Anterior pronotal carina widely interrupted. Femoral depression of mesopleuron weakly striate, mesepisternum smooth (Fig. 14). Dorsellum carinae diverging. Propodeum in lateral view forming a 90° angle with mesosoma, concave and smooth medially, bordered laterally by irregular, ridged, setose cells (Fig. 15). All femora with distal lamella. Metasoma. Metasoma medially compressed, smooth, Gt4-syntergum setose. Peti- ole very short and not visible while specimen is intact (Fig. 7). Gaster S-shaped in lat- eral view, ovipositor angled at about 30° dorsad to horizontal axis. Gt4 not emarginate posteriorly in dorsal view. Male. 6.5 mm. Scrobal depression black, ventral half of body whitish-yellow, wing vein amber, otherwise color and sculpture as described for females. Ventral plaque on scape forming a projection on the inner face below the attachment point to the pedicle (Figs 12, 13). Antennomeres with multiple rows of erect setae. Gastral petiole striate dorsally, 1.5x as long as the length to metacoxa, smooth laterally (Fig. 17). 114 Y.Miles Zhang et al. / Journal of Hymenoptera Research 86: 101-121 (2021) Figure 22-23. Kavayva davidsmithi lateral habitus 22 female 23 male. Photos by Cecilia Escobar. Variation. Size ranges from 3.5—6.5 mm. Color ranges from mostly yellow to mostly black dorsally, mesepisternum can range from smooth to weakly striate. Larva. Head amber-colored and body beige. Body length 4.7 mm; width 1.7 mm. Body C-shaped; 13 segmented (three thoracic, nine abdominal, and one anal segments); Description of Kavayva and two new species associated with Guarea 115 Figure 24-25. Kavayva davidsmithi 24 male dorsal habitus 25 frontal view of head. Photos by Cecilia Escobar. tapering slightly posteriorly; no protuberance on body segments (Fig. 19). Head heavily sclerotized; antennae positioned ventrolaterally on the head, above the mandible, 1.29x as long as broad. Two pairs of superior frontal setae near the cranial depression, two pairs of interior frontal setae around anterior tentorial pits, two pairs of clypeal setae, 116 Y.Miles Zhang et al. / Journal of Hymenoptera Research 86: 101-121 (2021) antennae low and offset laterally, two pairs of genal setae, two pairs of labral setae, and four pairs of hypostomal setae (Fig. 20). Mandible narrowing apically, bidentate, with two pairs of sensilla. Underlip complex flat, with two pairs of setae on the median lobe (labium), and one on the lateral lobe (maxilla) (Fig. 21). Thoracic segments with two dorsal setae, one pair of pleural setae, one pair of lateral setae, and one pair of ventral setae. Abdominal segments with a single pair dorsal, pleural, and ventral setae. Anal seg- ment with one pair of dorsal terminal setae, and ventral terminal setae present. Biology. Associated with seeds of Guarea kunthiana and G. guarea = G. guidonia (Meliaceae). Distribution. Brazil, Panama. Etymology. Named in honor of the Serra da Bodoquena National Park, an envi- ronmental conservation unit in Mato Grosso do Sul, Brazil. Remarks. ‘The specimens collected from Panama are slightly smaller (3.5—-5.5 mm) than those from Brazil (5.5-6.5 mm), and with lighter coloration on the wings and metasoma which could be the result of specimens being older (Fig. 18). We did not find any consistent morphological differences in either sex that reliably separate the Panama specimens from those collected in Brazil. Therefore we chose to group them all within K. bodoquenensis until fresh material can be collected for molecular work. Kavayva davidsmithi Zhang & Gates, sp. nov. http://zoobank.org/5OEAFCB5-FD0D-4FD5-9724-92A30E33 1LED8 Figs 1, 22—27 Material examined. Holotype Peru ¢ [1F]; Manu National Park, Madre de Dios, Es- tacion Bioldgica Villa Carmen; Trail 0; 14 Dec. 2013; 12°53'41"S, 71°24'13"W; 650 m a.s.l.; A. L. Norrbom leg.; ex. seed in fruit of Guarea guidonia; 13-PE-46; MUSM. Para- types Peru ° [1F, 1M]; same information as holotype; USNMENT01788076, 077. Diagnosis. Kavayva davidsmithi can be distinguished from K. bodoquenensis by the light infumation of the forewing and the absence of a secondary wing band on the basal setal line, extensive black bands across mesosoma in dorsal view, and the presence of ventral plaque on both sexes. Description. Holotype female. 10.1 mm in length. Color. Yellow except antennomeres, supraclypeal area light brown, tip of mandible, vertex, anterior half of occiput, malar sulcus, anterior half of dorsal and lateral pronotum, anterior half of midlobe of mesoscutum, lateral lobes of mesoscutum along the notauli, ax- illula, mediodorsal line on scutellum, ventral prepectus black, clypeus, wing vein, forewing below submarginal and marginal vein, femur, tibia amber and —eyes pinkish red (Fig. 22). Head. Quadrate with rounded corners, 1.2x as wide as high in frontal view, 2.4x as wide as long in dorsal view, areolate-rugose with setae (Fig. 25). Lower face weakly strigose, clypeus bilobed, mandible tridentate, supraclypeal area slightly concave and extending to the toruli. Malar sulcus present, incomplete, reaching about %4 of malar space. Malar space glabrous, smooth. Genal carina present. Toruli positioned above the Description of Kavayva and two new species associated with Guarea 117 1mm Figure 26-27. Kavayva davidsmithi 26 male head and antennae 27 female forewing. Photos by Cecilia Escobar. lower ocular line about 1/3 of the eye length, diameter of torulus 3.3x that of the in- tertorular space. Scrobal depression deeply excavated. Vertex areolate, anterior ocellus above scrobal depression, ratios of POL:O0OL:LOL equal to 3:4:1. Ventral plaque on scape forming a projection on the inner face below the attachment point to the pedicle. Ratio of scape (minus radicle):pedicel:anellus:F 1:F2:F3:F4:F5:F6:club as 10:2.7:1:5.7: 6.3:6.3:5.7:5:4.7:6.7, pedicel chalice-shaped, funicular segments with multiple irregu- lar rows of longitudinal sensilla whorls of setae, much shorter than its bearing segment, clava 2-segmented (Fig. 25). Postgenal lamina present. 118 Y.Miles Zhang et al. / Journal of Hymenoptera Research 86: 101-121 (2021) Wing. Forewing infumated half way down the wing below the submarginal and marginal vein, not exceeding stigmal vein. Ratio of marginal vein:postmarginal vein:stigmal vein as 2.3:1.2:1 (Fig. 27). Mesosoma. Mesosoma umbilicate, 1.7x as long as broad. Notauli complete, shal- low (Fig. 26). Anterior pronotal carina interrupted. Femoral depression of mesopleu- ron weakly striate, mesepimeron smooth and shiny ventrally, bulging laterally (Fig. 22). Dorsellum carinae diverging. Propodeum in lateral view forming a 90° angle with mesosoma, broadly delimited by carinae forming a hexagon with raised lateral corners. Median furrow of propodeum concave and smooth, bordered laterally by irregular setose cells. All femora with distal lamella. Metasoma. Metasoma medially compressed, smooth, Gt6-syntergum setose. Gaster S-shaped in lateral view, ovipositor angled at about 30° dorsad to horizontal axis (Fig. 22). Gt4 emarginate posteriorly in dorsal view. Male. 9.4 mm. Scrobal depression black, otherwise color and sculpture as described for female (Fig. 23). Antennomeres with multiple rows of erect setae and about 1.4x as long as width of segment (Fig. 26). Gastral petiole length in dorsal view about 2.8x as long as its greatest width, 1.7x as long as the length to metacoxa, smooth (Fig. 23). Variation. The coloration on the vertex and occiput can be confluent or disconnected. Biology. Associated with seeds of Guarea guidonia (Meliaceae). Distribution. Manu National Park, Peru. Etymology. Patronym honoring David Smith for his decades of devotion to Hy- menoptera and improvement of the Smithsonian’s National Insect Collection. Discussion The new genus Kavayva is only found associated with the seeds of Meliaceae, which represent a new plant family association within Eurytomidae, and the fourth genus as- sociated with seeds in the Neotropics. Based on the upcoming phylogenomic study of Eurytomidae (Zhang et al., in prep.), all of these phytophagous genera are only distantly related to each other, meaning that phytophagy has evolved multiple times within Eu- rytomidae. While addressing the evolutionary relationships is beyond the scope of this paper, we provide a brief literature review of the morphological characters in order to dis- tinguish these four genera (Table 1). The ventral plaque that is present in all known spe- cies of Kavayva is also present in some species of Prodecatoma (e.g., P diospyri), although the latter can be easily distinguished by the presence of a large intertorular projection and the hyaline forewing. Bephratelloides have a stigmated wing and a minute intertorular projection similar to Kavayva, but lack the ventral plaque. Finally, Eurytoma werauhia differs from Kavayva in being mostly black in color, and lacks the ventral plaque. Given the morphological conservatism within Eurytomidae, it is not surprising that a combination of morphological characters is needed to distinguish these four gen- era of eurytomids. We hope this study will aid in the discovery of additional Kavayva specimens and records, as their host plant Guarea can be found from northwestern Mexico down to northern Argentina (Pennington and Clarkson 2013). Description of Kavayva and two new species associated with Guarea 119 Acknowledgements We thank Bhrenno Maykon Trad, Vander Carbonari, Vinicius Marques Lopez for helping us in laboratory work/photos, and to Instituto Chico Mendes de Biodiversi- dade-ICMBio from Bonito MS for field collection permissions under SISBIO number 3266-1 date: Oct/06/2011. We would also like to thank Allen Norrbom for providing specimens as bycatch for the USDA Farm Bill project “Enhancement of Fruit Fly Lar- vae Identification and Taxonomy” (Project Coordinator G.J. 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