Two New Species in the Genus Frumenta Busck 1939 (Lepidoptera: Gelechiidae: Gnorimoschemini) with the Discovery of a Culcitula in the Male

Authors: McCarty, Megan E., Adamski, David, Metz, Mark A., and Landry, Jean-François Source: Proceedings of the Entomological Society of Washington, 122(2) : 415-441 Published By: Entomological Society of Washington URL: https://doi.org/10.4289/0013-8797.122.2.415

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TWO NEW SPECIES IN THE GENUS FRUMENTA BUSCK 1939 (LEPIDOPTERA: GELECHIIDAE: GNORIMOSCHEMINI) WITH THE DISCOVERY OF A CULCITULA IN THE MALE

MEGAN E. MCCARTY,DAVID ADAMSKI,MARK A. METZ, AND JEAN-FRANC¸ OIS LANDRY

(MEM, DA) Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (email: [email protected]); (MAM) USDA, ARS, Systematic Entomology Laboratory, Beltsville, Maryland, USA (urn:lsid:zoobank.org:author:221F8E40-12DA-43AD-85B2-E7BFF46C07C3; https://orcid. org/0000-0002-3535-535X); (JFL) Canadian National Collection of Insects, Agriculture and Agri-Food Canada, Ottawa Research and Development Centre, 960 Carling Avenue, Ottawa, Ontario K1A 0C6, Canada (urn:lsid:zoobank.org:author: 7F7064D9-75D0-41AD-B798-870EF2E72280)

Abstract.—Analysis of morphology and sequences of the 658bp barcoding region of COI support four distinct species in the genus Frumenta Busck, 1939, two of which are described new: Frumenta davidi McCarty, new species,andFrumenta dianeae McCarty, new species. Frumenta solanophaga Adamski and Brown, 2002, is a junior synonym of F. nephelomicta (Meyrick, 1930). Frumenta nundinella (Zeller, 1873) and F. nephelomicta are redescribed. Sequence data for F. dianeae were unavailable, but the morphology of this species is distinct. We found three morphological features that may deﬁne Frumenta and provide support for the placement of Frumenta within Gnor- imoschemini Povolny, 1964, including a culcitula revealed by scanning electron micrographs. Images of adult habitus and male and female genitalia are provided as well as a map showing species distributions and a key to the species. Key Words: biocontrol, COI barcodes, haplotype network, morphology, Solanum, systematics, taxonomy DOI: 10.4289/0013-8797.122.2.415

Prior to this work, the genus Frumenta Frumenta solanophaga Adamski and Busck, 1939 was comprised of three spe- Brown, 2002 was the most recently de- cies known only from the Nearctic region. scribed taxon to date. Tribal placement of Busck (1939) implicitly diagnosed the Frumenta, however, has been enigmatic. genus for the single species Gelechia Throughout his professional career, nundinella Zeller (1873) stating that it Povolny presented an ever-changing land- was probably related to Gnorimoschema scape of classiﬁcation in the tribe Gnor- Busck, 1900 based on similarities he imoschemini Povolny, 1964. He created noted (i.e., “the character of the palpi, dozens of new taxa and notably switched scaling, and aedeagus”). The genus was his own generic combinations. In regards monotypic until Hodges (1983) included to Frumenta, Povolny (1967) hypothesized Asapharca nephelomicta Meyrick, 1930. its inclusion in the tribe Gnorimoschemini

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without examining either of the then in- culcitula in the males. We also document cluded species let alone F. nundinella.He the presence of diagnostic pleural pouches explicitly stated his decision was based in females of all species, which could be a only on the figures and description of putative synapomorphy for the genus. Busck (1939). The classification of Gnor- imoschemini and the placement of Fru- MATERIALS AND METHODS menta among Gelechiinae Stainton, 1854 A total of 172 pinned adult specimens are untested hypotheses. More importantly, and64genitaliapreparations were exam- Povolny’s varied classifications have been ined from the U.S. National Museum problematic in relation to pest diagnostics (USNM), Smithsonian Institution, Wash- as many species in the tribe feed on Sola- ington, DC, and seven specimens and naceous crops and are potential invasives three genitalia preparations from the Ca- in certain World regions. Indeed, species nadian National Collection (CNC) in Ot- of Frumenta are specialist herbivores on tawa, Canada. Additional locality and plants in the genus Solanum Linnaeus. BOLD data were included from the fol- Two Nearctic species of Solanum are lowing specimens: one specimen from known hosts of Frumenta. Frumenta the Mississippi Entomological Museum nundinella has been recorded numerous (MEM), Mississippi State University, times feeding on Carolina horsenettle (S. Starkville, Mississippi; and one specimen carolinense Linnaeus) where larvae feed from the Biodiversity Institute of Ontario on either leaves or fruits in different gen- (BIO), Guelph, Ontario, Canada. Holo- erations (Bailey 1978). Frumenta neph- types of F. nundinella and F. nephelomicta elomicta and F. solanophaga have both were examined at the Natural History been reared from silverleaf nightshade (S. Museum (NHM), London, United King- elaeagnifolium Cavanilles) (Adamski and dom. All USNM and CNC specimens Brown 2002). These two species of So- were labeled with a unique identifier and lanum, like numerous other North Amer- corresponding barcode and databased. ican species, have invaded other regions Additional information about the speci- (i.e., Europe and Africa), where they have mens can be found in the online database become noxious weeds. Because of this USNM Department of Entomology Col- impact, species of Frumenta have been lection (http://collections.nmnh.si.edu/search/ studied as candidates for biocontrol (Bailey ento/, indexed by the unique identifier 1978; Neser et al. 1989; Olckers 1995; USNMENTnnnnnnn). The holotypes of Wells et al. 1986). Olckers (1995) recog- new taxa are deposited in the USNM. nized at least one new species of Fru- Label data are given verbatim within menta in the course of his biological study quotations for the holotypes. but did not describe it. A distribution map for all species was An opportunity arose to examine the created using R (R Development Core specimens mentioned in Olckers (1995) at Team 2019) in the RStudio (RStudio the National Insect Collection of the U.S. Team 2016) IDE implementing the pack- National Museum (USNM). While re- ages ‘openstreetmap’ (Fellows 2016), visiting these specimens, we confirmed ‘mapdata’ (Brownrigg 2018), and ‘ggmap’ Olckers’ new species and discovered an and ‘ggplot’ (Kahle and Wickham 2013). additional undescribed species in the ge- Neighbor-joining trees were constructed nus. In this work we describe these two in Mesquite (Maddison and Maddison new species and review the taxonomy of 2019) implementing the package Ninja Frumenta. We report the discovery of the (Wheeler 2009) under the Kimura

2-parameter model of evolution (Kimura exception of F. dianeae, for which we 1980). A haplotype parsimony network only had one female specimen. We was created using R (R Development placed the wings in 90% EtOH, then Core Team 2019) in the RStudio (RStudio subsequently into a solution of 5.25% Team 2016) IDE. The haplotype com- bleach for several minutes, until the mand in ‘pegas’ (Paradis 2010) was used coloring of the scales faded and the veins to create the haplotype network based on could be easily seen. They were rinsed a distance matrix created using the Ki- again in 90% EtOH and placed on a glass mura 2-parameter model of evolution slide. Once the wings dried, we placed a (Kimura 1980). Details of analyzed se- glass coverslip over them using a water quences and voucher specimens are avail- soluble glue as an adhesive. able on the Barcode of Life Data System Preparation of moth genitalia fol- (http://www.boldsystems.org) in the public lowed Clarke (1941), except we used dataset ‘DS-FRUMENTA’ (http://dx.doi.org/ MercurochromeTN or orange G and 10.5883/DS-FRUMENTA) and have been chlorazol black as stains, and we exam- submitted to GenBank (see BOLD dataset ined the slide preparations with both for GenBank accessions). dissecting and compound microscopes. A Visionary Digital Imaging SystemTM We followed Adamski and Brown (2002) with a Canon 30D camera and a 65mm for slide-mounting male genitalia; the lens was used for photographing adult dorsal part was separated from the ven- specimens. We stacked images using tral part on one side, and the genitalia Helicon FocusTM stacking software. were unrolled and mounted ﬂat on a An Environmental Scanning Electron slide. For each species, one male spec- Microscope (SEM) was used to obtain imen was treated in a solution of 20% detailed images of the male genital KOH at room temperature overnight, structure. For this process, we brushed cleaned, and then placed in glycerine. off the scales on the posterior abdominal This was done to study and illustrate segments of an intact male abdomen, the morphology of the genitalic struc- making the genitalia more visible. We tures in situ, as slide-mounting causes then removed the abdomen and posi- distortion and provides only one view. tioned it on an aluminum stub so that the This method allowed us to make better posterior end faced dorsally (straight comparisons between species, as we up). The abdomen was glued in place found the relative positions of certain using PELCOÒ Colloidal Graphite (Iso- parts (such as the positioning of the propanol Base) glue, and it was not coated. valves relative to the uncus and the po- We used EDX analysis (H), charge-up re- sitioning of the phallus within the vin- duction mode (L), and mix image mode culum) to be important in separating (M) for each image at a working distance species. In the descriptions, we refer to of 6.5 mm. After taking the images, we left characters being found in either rolled the abdomen on the aluminum stub, la- (undissected, glycerine) or unrolled (dis- beled it, and placed it next to the specimen sected, slide-mounted) specimens. Be- in the collection. cause of the limited amount of material, Wing slide preparations were modi- only one specimen from each species fied from Borror et al. (1981). We re- was studied in this manner. We also moved the right forewing and hindwing examined female genitalia using this from a male of each species, with the method, but we found it provided no

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additional information compared to slide- present in males. This character was first mounted specimens. described by Adamski and Brown (2002) For detailed images of female pleural but was not explored in other species pouches, the pleural membrane of the of Frumenta at that time. For males, we cleaned and stained abdomen was cut propose a new diagnostic morphologi- along the left side of segments VI–VII cal feature: a laterally flat, widened and sterna VI–VII were unrolled in order to valva that broadens apically beyond the provide a clear view of otherwise over- basal arch. lapping segments. The abdomens were Placement of Frumenta within Gnor- slide-mounted in lactic acid under a cover imoschemini Povolny.—Povoln y (1967) slip. Structures were examined with a based his placement of Frumenta within Nikon Eclipse 800 compound microscope Gnorimoschemini on the figures and at magnifications of 40–100x equipped with descriptions of Busck (1939) without a Nikon DS-Fi1 digital camera. Nikon’s NIS examining the specimens himself. De- 2.3 Elements was used to assemble multi- spite this, we believe the placement of ple photos of different focal planes into Frumenta within Gnorimoschemini is single deep-focus images. For close-ups accurate. Most gnorimoschemine fe- of the pouches, differential interference males have a hook-like signum (Huemer contrast (Normarski) illumination was used and Karsholt 2010) and while it is not as to enhance depth. strongly hooked in Frumenta, it does Terms used for species descriptions come to a sharp point; in some speci- follow Huemer and Karsholt (2010), mens, this point is slightly curved, ren- with the following exception: the later- dering the signum slightly hooked. omedial projection is referred to here as The presence of a culcitula in Fru- the juxtal lobe. We also use the term menta has not been previously docu- ‘eversible scent pouch’ for the mem- mented, but we observed the presence of branous sac associated with the female a pair of membranous lobes with micro pheromone gland, and acanthus for the dentitions between the ventral mem- individual setae of the frenulum. brane of the tuba analis and the base of the gnathos using SEM imaging. This RESULTS AND DISCUSSION structure is also visible in non-dissected Review of Frumenta Busck.—Busck male specimens with exposed genitalia, (1939) wrote a fairly lengthy description but not in slide-mounted specimens. of Frumenta without explicit diagnostic This structure is likely distorted or dam- features. We conclude that one of the aged during standard dissections, as we two characters he used to identify Fru- found it to be very delicate, unlike the menta in his key to genera (Busck 1939), culcitula in other gnorimoschemines. We the “small, straight, obtusely pointed propose that this structure is a culcitula, signum” is diagnostic for the genus. In as defined by Hodges (1966). All males other gnorimoschemines, the signum is in Gnorimoschemini have a culcitula of larger and more strongly hooked. We this type and all females with a visible propose a new diagnostic morphological signum have a central finger-like projec- feature for Frumenta: the presence of a tion with a hooked tip. The combination pair of pleural invaginated pouches on of these characters in Frumenta supports the intersegmental membrane anterior the tribal placement. to the 7th pair of spiracles (Figs. 30–33) Molecular Analyses.—We failed to in the females; these pouches are not obtain sequenced data for F. dianeae and

had incomplete sequences for 13 speci- – Female ...... 4 mens (four F. nundinella,fiveF. neph- 2. Gnathos narrow and only slightly di- elomicta, and four F. davidi). To remove lated distally (Figs. 21, 24), valva length any potential impact of missing data, we beyond basal arch short (extending to created two neighbor-joining trees: one posterior margin of uncus) (Figs. 18, 21) including all taxa, regardless of se- ...... nundinella quence length (Fig. 40), and one in- – Gnathos wide and conspicuously dilated cluding only specimens with the entire distally (Figs. 20, 22), valva length be- 658bp barcode region of COI (Fig. 41). yond basal arch long (extending beyond Both neighbor-joining trees (Figs. 40– posterior margin of uncus) (Figs. 17, 19) 41) resolve three species of Frumenta (F...... 3 nundinella, F. nephelomicta, and F. da- 3. Saccus less than 1.5X length of lateral vidi). Haplotype distances among spec- arms of vinculum, phallus in situ ex- imens identified as F. solanophaga and tending posteriorly beyond juxtal lobes specimens identified as F. nephelomicta ¼ length of phallus (Figs. 19, 22, 25) are similar and cluster as a single taxon...... nephelomicta Because this corroborates with exam- – Saccus more than 2.0X length of lat- ined morphology and recorded host data, eral arms of vinculum, phallus in situ we synonymize the species. barely extending beyond juxtal lobes pos- Two specimens identified as F. nundi- teriorly (Figs. 17, 20, 23) ...... davidi nella (CCDB-31101-C11 and CNCLE 4. Width of the posterior margin of seg- P00090830) (Fig. 40) with incomplete se- ment VIII less than ½ length of seg- quences do not cluster with the other ments IX–X (Figs. 26, 28) ...... 5 specimens identified as F. nundinella.We – Width of the posterior margin of seg- presume this is artifact. Three speci- ment VIII greater than ½ length of mens identified as F. nephelomicta, segments IX–X (Figs. 27, 29) ...... 6 (CCDB-31101-C09, USNMENT00657620, 5. Apophyses posteriores 2.0X longer than and USNMENT00657619) (Fig. 40) with apophyses anteriores, apophyses poste- incomplete sequences were also distant riores extending beyond anterior margin from the other specimens identified as of segment VIII (Fig. 28); segment VIII F. nephelomicta. In this case, specimens equally long on dorsal and ventral sides, USNMENT00657620 and USNMENT overall corpus bursae cylindrical (Fig. 00657619 were from the same collecting 28) ...... nundinella event and eclosed from the same host – Apophyses posteriores 1.5X longer than as specimen CCDB-31101-C10, which apophyses anteriores, apophyses poste- clustered with the remaining specimens, riores not extending beyond anterior and are presumed to be siblings. All margin of segment VIII (Fig. 26); seg- taxa unambiguously cluster when only ment VIII longer dorsally than ventrally, specimens with complete sequences are overall corpus bursae pyriform (Fig. 26) included (Fig. 41). The parsimony ...... davidi haplotype network (Fig. 42) better il- 6. Apophyses posteriores about equal in lustrates the distances between the length to apophyses anteriores, corpus species. bursae lacking anterior lobe-like extension on anterior end (Fig. 29) ... nephelomicta Key to the Species of Frumenta – Apophyses posteriores about 1.5X lon- 1. Male ...... 2 ger than apophyses anteriores, corpus

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bursae with lobe-like extension on an- Midtibia as above, but with wider pale terior end (Fig. 27) ...... dianeae yellowish-white bands. Hindlegs pale yellowish white, hindtibia with piliform pale yellowish-white scales and two small Frumenta davidi McCarty, patches of dark-brown scales, one basal, new species one medial. Tarsomeres of all legs with dark-brown scales tipped pale yellowish http://zoobank.org/B3B5AB0D- white, each with pale yellowish-white api- 55CA-4B46-9244-CAD674BCC7AD (Figs. 1–3, 11–12, 17, 20, 23, 26) cal bands. Forewing length 6.0–10.0 mm (mean = 8.6 mm, n = 25); dorsal surface Etymology.—The species epithet da- pale yellowish white. Base of wing with vidi is a patronym in honor of the first one or two dark-brown spots, absent in author’s father. This honor is bestowed on some specimens. Discal cell with two ir- David for nurturing and supporting the regularly shaped dark-brown spots on the entomological passion of his daughter and apical portion which are diagonal to one traveling with her on countless lepidop- another. Basal third with one dark-brown terological expeditions across the United spot posterior to CuP. Costa with 3–4 States. dark-brown spots; 1–2 basal, one medial, Diagnosis.—Frumenta davidi can be one apical. Marginal spots dark brown, recognized by having the forewing with separate or confluent, forming an uneven more white scales in a yellowish-white band; fringe pale yellowish white. Ventral wing pattern; the termen delineated by a surface dark brownish gray with distal row of marginal spots at the base of the costa and outer margin pale yellowish yellowish-white fringe; a saccus that is white with dark-brown spots. Venation more than 2X longer than the lateral arms (Fig. 11) with R4 and R5 branched sub- 2 of the vinculum; a phallus that barely ex- apically near / 3 from discal cell; discal tends beyond the juxtal lobes in intact cell closed diagonally with rounded apex; specimens; narrow posterior abdominal apical part of M1 parallel with R5;M2 segments in the female; apophyses poste- closer to M3 than to M1;CuA1 and CuA2 riores that do not extend beyond the an- parallel. Hindwing dorsal surface and terior portion of segment VIII; segment ventral surfaces pale brown or dark brown, VIII of the female that is longer dorsally with pale yellowish-white fringe. Vena- than ventrally; and a signum with the tion (Fig. 12) with Sc+R1, RS, and M1 length the width of the ductus bursae. curved divergently from M2 and M3; Description.—Head: Vertex, frons, and Sc+R1 and RS connected by base of R1 1 haustellum pale yellowish white. Scape of near / 3 length of discal cell; CuA1 and antenna pale yellowish white, flagellum CuA2 nearly parallel. Frenulum with one pale yellowish white with some brown acanthus in male, three acanthi in female. scales. Labial palpus pale yellowish white; Abdomen: Pale yellowish white on dorsal second segment longer than third, scales and ventral surfaces. Female abdomen on ventral surface divergent from base, with pair of small, invaginated pleural forming brush-like array; third segment pouches on intersegmental membrane with appressed scales. Thorax:Tegulae anterior to 7th spiracle (see Figs. 30–33 of and mesonotum pale yellowish white. F. nephelomicta and F. nundinella). Male Forelegs dark brown, except foretibia with genitalia (Figs. 17, 20, 23) with tegumen two pale yellowish-white bands, one basal, sub-triangular (unrolled). Uncus trapezoi- one medial. Midlegs pale yellowish white. dal, equally long as it is wide, with lateral

Figs. 1–10. Dorsal habitus of Frumenta spp. 1–3, F. davidi, n. sp., paratypes, ♂ USN- MENT01196881, ♂ USNMENT01480131, ♀ USNMENT01480129. 4, F. dianeae, n. sp., holotype, ♀ USNMENT01196883. 5–7, F. nundinella, ♂ USNMENT01480061, ♂ USNMENT01480063, ♂ USN- MENT01480038. 8–10, F. nephelomicta ♀ USNMENT01480082, ♀ USNMENT01480120 ♀ USN- MENT01196864.

margins setose (rolled); lateral margins of F. nephelomicta). Gnathos curved, may appear inwardly curved on slide- tongue-shaped, widening medially, taper- mounted specimens as an artifact of ﬂat- ing into narrowly rounded apex (rolled). tening. Culcitula with short, linear arms, Anterior diaphragma above phallus and arms abruptly dilated distally, forming ventrad tegumen expanded on median line sub-rectangular, medially concave, open as convex, darkened, pedunculate process; extensions; densely covered with micro lateral edges almost contiguous with in- dentitions (not illustrated; see Figs. 34–38 ner medial margin of tegumen (unrolled).

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Figs. 11–16. Wing venation of male Frumenta spp. 11–12, F. davidi, n. sp., paratype, slide USNM84264. 13–14, F. nundinella, slide USNM84268. 15–16, F. nephelomicta, slide USNM84283.

Valva long, extending beyond posterior (unrolled). Female genitalia (Fig. 26) with margin of uncus; basally narrow, wider posterior abdominal segments long and beyond basal arch; valva beyond basal narrow, with width of posterior margin of arch long, setose (rolled). Vinculum with segment VIII less than ½ length of seg- elongate saccus, more than 2.0X longer ments IX–X. Eversible scent pouch pres- than lateral arms of vinculum (rolled). ent dorsally between segment VIII and Juxtal lobes short and pointed posteriorly, papillae anales (not shown; see Fig. 28 of setose (unrolled). Phallus long and narrow F. nundinella); papillae anales with long with bulbous base, extending about 1/10 and short setae interspersed. Apophyses length beyond juxtal lobes (rolled); ductus posteriores long and narrow, not extending ejaculatorius entering dorsally into base beyond anterior margin of segment VIII.

Figs. 17–19. Male genitalia of Frumenta spp., rolled lateral view. 17, F. davidi, n. sp., paratype, USNM84263, gel capsule. 18, F. nundinella, USNM84217, gel capsule. 19, F. nephelomicta, USNM84261, gel capsule.

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Figs. 20–22. Male genitalia of Frumenta spp., rolled ventral view. 20, F. davidi, n. sp., paratype, USNM84263, gel capsule. 21, F. nundinella, USNM84217, gel capsule. 22, F. nephelomicta, USNM84261, gel capsule.

Figs. 23–25. Male genitalia of Frumenta spp., unrolled view. 23, F. davidi n. sp., paratype, USNM82139. 24, F. nundinella, USNM82137. 25, F. nephelomicta, USNM82135.

Apophyses posteriores about 1.5X longer outwardly beyond curvature of corpus than apophyses anteriores. Anterior mar- bursae. gin of segment VIII scalloped shallowly. Type material.—Holotype:♀ “Texas, Dorsal side of segment VIII longer than Brownsville, coll. IX–X [Sep.–Oct.] 1963, ventral side. Ductus bursae elongate, nar- Iss[ued] XI 1963, Brown & Jackson,” row, and slightly dilated at anterior end “stem gall on Solanum eleagnifolium [sic],” near inception of ductus seminalis; in- “♀genitalia slide by D. A[damski], USNM ception point with lateral arms of a 82140” [green label], “Specimen ID rhomboid plate wrapping around ductus USNMENT01480118,” “HOLOTYPE Fru- bursae. Corpus bursae elongate, broad- menta davidi ♀ McCarty, 2020” [red label], ened anteriorly. Signum small with slightly “USNMENT01480118”, [USNM]. Para- pointed apex; base of signum protruded types: 13 ♂♂,12♀♀: UNITED STATES:

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Figs. 26–29. Female genitalia of Frumenta spp. 26, F. davidi, n. sp., holotype, USNM82140. 27, F. dianeae, n. sp., holotype, USNM84254. 28, F. nundinella, USNM82138. 29, F. nephelomicta, USNM82136.

Figs. 30–33. Pleural pouches on 7th intersegmental membrane of female Frumenta spp. 30, 32, F. nephelomicta, gel capsule MIC8223. 31, 33 F. nundinella, gel capsule MIC8222. 30, 31, Abdominal segments 6–7, unrolled, magnification 40x. 32, Outer aspect of pouch, magnification 100x. 33, Inner aspect of pouch, magnification 100x.

Texas: 7 ♂♂ (USNMENT01196881, USN 01480125, USNMENT01480126, USN- MENT01480123, USNMENT01480124, MENT01196879, USNMENT01196880, USNMENT01480115, USNMENT01480116, USNMENT01480127, USNMENT01480129, USNMENT01480119, USNMENT01480117, USNMENT01480128, USNMENT01480130, USNM82139, USNM84264, USNM USNM84253, USNM84239, USNM84213, 84263, USNM84212), 8 ♀♀ (USNMENT USNM84257), Brownsville, Sep.–Oct.,

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Figs. 34–38. Scanning electron micrographs of culcitula in F. nephelomicta, USNMENT01196861. 34, Ventral view of male genitalia, with uncus, culcitula, gnathos, and valvae present. 35, Closeup of uncus and culcitula. 36, Closeup of left side of culcitula. 37–38, Closeup of left side of culcitula showing micro dentitions.

1963; 4 ♂♂ (USNMENT01196873, USNM84266, USNM84256), Mission, USNMENT01196874, USNMENT01196875, Aug. 28–Sep. 4, 1989, Lot AcP9231; 2 ♂♂ USNMENT01196877, USNM84255), 2 ♀♀ (USNMENT01480131, USNMENT01196882, (USNMENT01196876, USNMENT01196878, USNM9851, USNM9850), Llano Co.,

Fig. 39. Distribution map of Frumenta spp.

May 10, 1959; 1 ♀ (USNMENT01480132), on S. carolinense (Carolina horsenettle). Mercedes, [no date]; 1 ♀ (USNMENT- Olckers (1995) reports that F. davidi 01480133), San Benito, Jul. 16-23 [no (designated “Frumenta species A” in the year]. paper) formed both fruit and stem galls, Distribution.—All of the specimens the latter being an adaptation to the ab- were collected in Texas with the most sence of ﬂower buds on the host. To date, common locality being Brownsville. F. davidi is the only Frumenta to have Twenty-four of the 26 specimens were been reared from two different host spe- collected near the U.S./Mexican border, cies. It is also the only member of the while the remaining two were found in genus to form both fruit and stem galls as Llano County in central Texas (Fig. 39). opposed to forming only fruit galls. Six Despite the distance between these two specimens from lot AcP9231 are vouchers localities, the genitalia of the specimens from a study mentioned by Olckers (1995), from Llano County match those of the who documented attempts to introduce specimens collected in southern Texas. F. davidi into South Africa to control S. Biology.—Twenty-two of the 26 speci- elaeagnifolium. This plant is native to the mens in the USNM collection were reared southwestern U.S. and northern Mexico. from stem galls on Solanum elaeagnifo- However, it has been introduced to South lium (silverleaf nightshade), while two Africa, where it is a noxious weed (Neser others were reared from oval stem galls et al. 1989; Wells et al. 1986). While the

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Fig. 40. Neighbor-joining tree of Frumenta spp. including all specimens sequenced from the barcoding region of COI in reference to the type species of the tribe Gnorimoschemini, Gnorimoschema gallaesolidaginis. Specimens with partial sequences indicated by dashed lines at terminals. Specimens are referenced to unique identiﬁers used in the text and partial locality data (*female holotype of Fru- menta solanophaga).

attempt to introduce F. davidi was more author’s mother. This honor is bestowed successful than attempts to introduce F. on Diane for nurturing and supporting the nephelomicta for the same purpose, F. entomological passion of her daughter davidi failed to become established after and, through homeschooling, giving her three years. Despite the conditions being daughter more time and freedom outdoors more favorable in South Africa than in to chase butterflies. its native range, native hymenopterous Diagnosis.—Frumenta dianeae can be parasitoids decimated the moth population recognized by having a yellowish-white (Olckers 1995, Olckers et al. 1999). forewing with two dark-brown bands on Therefore, it is unlikely that F. davidi will the outer margin that are separated by a become established in the region and be a band of pale yellowish white; a dark-brown useful biocontrol agent for S. elaeagnifo- forewing fringe; wide posterior abdominal lium in South Africa. There have been no segments; segments between segment VIII subsequent attempts to introduce this spe- and the papillae anales that are very short; cies into South Africa (Olckers 1995). and a corpus bursae with a small lobe-like extension on the anterior side. Frumenta dianeae McCarty, Description.—Head: Vertex and haus- new species tellum pale yellowish white; scales on frons worn off. Scape of antenna pale http://zoobank.org/F975DA08-0099- yellowish white, flagellum pale yellow- 4EB4-85BB-3FF7DD85270B ish white with some brown scales. Labial (Figs. 4, 27) palpus pale yellowish white; second seg- Etymology.—The species epithet dia- ment longer than third. Thorax: Tegulae neae is a patronym in honor of the first pale yellowish white, mesonotum brown.

Fig. 41. Neighbor-joining tree of Frumenta spp. including only specimens with 658bp sequenced from the barcoding region of COI in reference to the type species of the tribe Gnorimoschemini, Gnorimoschema gallaesolidaginis. Specimens are referenced to unique identiﬁers used in the text and partial locality data (*female holotype of Frumenta solanophaga).

Forelegs missing on specimen. Midlegs Abdomen: Pale yellowish white on dorsal pale yellowish white with two dark-brown and ventral surfaces. Female abdomen bands, one medial, one apical. Tarsomeres with pair of small, invaginated pleural with dark-brown scales tipped with pale pouches on intersegmental membrane yellowish white, each with pale yellowish- anterior to 7th spiracle (see Figs. 30–33 of white apical bands. One hindleg missing, F. nephelomicta and F. nundinella). Fe- other with descaled femur remaining. male genitalia (Fig. 27) with posterior Forewing length 10.0 mm; dorsal surface abdominal segments long and wide, with pale yellowish white. Discal cell with width of posterior margin of segment VIII three irregularly shaped dark-brown spots, greater than ½ lengths of segments IX–X. two on basal third, one on distal third. Eversible scent pouch present dorsally Costa with three dark-brown spots; two between segment VIII and papillae anales basal, one medially. Margin with two (not illustrated; see Fig. 28 of F. nundi- dark-brown bands, separated by a pale nella); papillae anales short and wide with yellowish-white band. Fringe dark brown. long and short setae. Apophyses posteri- Ventral surface dark brownish gray, with ores long and narrow, extending beyond distal costa and outer margin pale yel- anterior margin of segment VIII. Apoph- lowish white. Hindwing dorsal surface yses posteriores about 1.5X longer than pale brownish gray, fringe pale yellow- apophyses anteriores. Anterior margin of ish white. Ventral surface pale yellowish segment VIII scalloped deeply. Dorsal and white, with dark-brown scales at apex. ventral sides of segment VIII equally long. Frenulum with three acanthi in female. Ductus bursae elongate, relatively narrow,

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Fig. 42. Parsimony haplotype network of Frumenta spp. in reference to the type species of the tribe Gnorimoschemini, Gnorimoschema gallaesolidaginis, superimposed on geography. Included specimens have complete sequences (658bp) from the barcoding region of COI. Circles represent different haplotypes and slices of circles represent individual specimens with the same haplotype. Haplotypes are positioned geographically on the map. Ticks on connecting lines indicate the number of base pair dif- ferences between haplotype pairs. (HT = the female holotype of Frumenta solanophaga is included in this haplotype).

and slightly dilated at anterior end near “HOLOTYPE Frumenta dianeae ♀ inception of ductus seminalis; inception McCarty, 2020” [red label], “USNMENT point with lateral arms of a rhomboid plate 01196883,” [USNM]. wrapping around ductus bursae. Corpus Distribution.—The only locality from bursae elongate, broadened anteriorly; which this species has been collected is anterior side with a small lobe-like exten- Madera Canyon, Arizona (Fig. 39). sion. Signum large with pointed apex; base Biology.—Unknown. of signum protruded outwardly beyond curvature of corpus bursae. Frumenta nundinella (Zeller) Type material.—Holotype ♀ “Madera (Figs. 5–7, 13–14, 18, 21, 24, 28, 31, 33) Canyon 4400’, Santa Rita Mts., Pima Co. Arizona, 31 August 1960, J. G. Gelechia nundinella Zeller 1873: 256. Franclemont,” “♀ genitalia slide by M. Frumenta nundinella: Busck 1939: 577. McCarty, USNM 84254” [green label], Gelechia beneﬁcentella Murtfeldt 1881: “Specimen ID USNMENT 01196883,” 245. Synonymized by Dyar 1903: 515.

Diagnosis.—Frumenta nundinella can nally with acuminate apex; base of R4-R5 be recognized by having an orange-brown and M1 basally divergent. M2 basally forewing mottled with dark-brown spots; a closer to M3, distally closer to M1;CuA1- gnathos that widens medially and ends CuA2 slightly closer near base, nearly par- with a blunt apex; a short valva with a allel from slightly beyond base. Hindwing truncated apical margin that does not ex- dorsal surface brown to grayish brown. tend beyond the posterior margin of the Fringe grayish orange. Ventral surface uncus; large juxtal lobes; narrow posterior similar, except costa darker. Venation abdominal segments in the female; (Fig. 14) with Sc+R1 and RS divergent apophyses posteriores that are 2.0X longer from anterodistal margin of discal cell; than the apophyses anteriores. Sc+R1 and RS connected by base of R1 Description.—Head: Vertex, frons, and near ½ length of discal cell. M1 straight, haustellum grayish orange, with some M2-M3 near parallel. M3 and CuA1 diver- scales tipped dark brown. Scape of antenna gent from slightly beyond discal cell. CuA1 dark brown, ﬂagellum grayish orange in- shorter than CuA2. Frenulum with one termixed with dark-brown scales. Labial acanthus in male, three acanthi in female. palpus grayish orange; second segment Abdomen: Dorsally grayish orange with longer than third, with two dark-brown median slightly darker in some specimens, bands on outer surface, one basally, one giving appearance of longitudinal stripe apically; ventrally, scales divergent from (Fig. 5). Ventrally grayish orange, inter- base, forming brush-like array; third seg- mixed with dark-brown scales. Female ment with appressed scales, banded basally abdomen with pair of small, invaginated and apically dark brown. Thorax:Tegulae pleural pouches on intersegmental mem- grayish orange; basal scales tipped dark brane anterior to 7th spiracle (Figs. 31, 33). brown, in some specimens all scales tipped Male genitalia (Figs. 18, 21, 24) with dark brown. Mesonotum grayish orange tegumen sub-triangular (unrolled). Uncus with three longitudinal dark-brown bands, trapezoidal, equally long as it is wide, one medially, two lateral. Forelegs grayish with lateral edges setose (rolled); lateral orange intermixed with dark-brown scales. margins may appear inwardly curved on Foretibia dark brown with three grayish- slide-mounted specimens as a product of orange bands, one basal, one medially, one ﬂattening. Culcitula with short, linear arms, apical. Midlegs and hindlegs patterned like arms abruptly dilated distally, forming forelegs, except hindtibia with long, pili- sub-rectangular, medially concave, open form, grayish-orange scales and two pairs extensions; densely covered with micro of spurs. Tarsomeres of all legs dark dentitions (not shown; see Figs. 34–38 of brown, with grayish orange apical bands. F. nephelomicta). Gnathos narrow, curved, Forewing length 6.0–9.0 mm (mean = tongue-shaped, widening medially, ending 8.1 mm, n = 25): dorsal surface grayish with blunt apex (rolled). Anterior dia- orange with dark-brown mottled pattern; phragma above phallus and ventrad tegu- mottling thickest along costa and on basal men expanded on median line as convex, two thirds. Outer margin lined with one darkened, pedunculate process; lateral row of dark-brown spots. Fringe dark edges almost contiguous with inner medial brown. Ventral surface dark brown. Costa margin of tegumen (unrolled). Valva short, grayish orange. Apical margin with single extending to posterior margin of uncus; row of grayish-orange spots. Venation (Fig. basally narrow, gradually widening from 13) with R4 and R5 divergent near ½ length basalarchwithtruncateapicalmargin; from discal cell; discal cell closed diago- apical part of valva beyond basal arch short,

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setose (rolled). Vinculum short with long, slide by V. Nazari, USNM 130,348” narrow saccus rounded at apex, more than [green label] “USNMENT00657958,” 2.0X longer than lateral arms of vinculum [USNM]. (rolled). Juxtal lobes large and pointed Other specimens examined.—56 ♂♂, posteriorly, setose (unrolled). Phallus long 33 ♀♀: CANADA: Ontario 1 ♂ and narrow with bulbous base, extending (CNCLEP00090829), 1 ♀ (CNCLEP- 1 about / 3 length beyond juxtal lobes (rol- 00090830), Harrow, Sep. 2–6, 1963. led); ductus ejaculatorius entering dorsally UNITED STATES: Georgia: 5 ♂♂ into base (unrolled). Female genitalia (Fig. (USNMENT01480011, USNMENT01480012, 28) with posterior abdominal segments long USNMENT01480013, USNMENT01480014, and narrow, with width of posterior margin USNMENT01480072, USNM84259, USNM- of segment VIII less than ½ length of seg- 84267), 2 ♀♀ (USNMENT01480079, USN- ments IX–X. Eversible scent pouch present MENT01480071, USNM84260), Richmond dorsally between segment VIII and papillae Hill, Bryan Co., Sep. 15–19 1944. Illinois: 9 ♂♂ anales (Fig. 28). Papillae anales long and (USNMENT01480043, USNMENT01480076, narrow, with long and short setae inter- USNMENT01480077, USNMENT01480078, spersed. Apophyses posteriores long and USNMENT01480070, USNMENT01480069, narrow, extending beyond anterior margin USNMENT01480073, USNMENT01480061, of segment VIII. Apophyses posteriores USNMENT01480067, USNM82137, USNM- about 2X longer than apophyses anteriores. 9852, USNM9854), 3 ♀♀ (USNMENT- Anterior margin of segment VIII scalloped 01480068, USNMENT01480066, shallowly. Dorsal and ventral sides of seg- USNMENT01344625, USNM9852, USNM- ment VIII equally long. Ductus bursae 82138), Decatur, Macon Co., Aug. 8–Sep. 7 elongate, relatively narrow, and slightly [no year]; 1 ♂ (USNMENT01480063), 1 dilated at anterior end near inception of ♀ (USNMENT01480060), Putnam Co., ductus seminalis; inception point with a Sep. 15, 1956 and Sep. 23, 1953; 4 ♂♂ triangular plate. Corpus bursae elongate, (USNMENT01480065, USNMENT01480074, not broadened anteriorly. Signum large with USNMENT01480064, USNMENT01480173, slightly pointed apex; base of signum pro- USNM9853, USNM9856, USNM84217), 2 truded outwardly beyond curvature of cor- ♀♀ (USNMENT01480062, USNMENT- pus bursae. 01480172, USNM9853, USNM84216), Types examined.—F. nundinella: ♀ Chicago, Cook Co., Aug. 25–Sep. 4, “Holotype” [round label with red border], 1934; 1 ♀ (USNMENT01480161), Oco- “Type” [round label with red border], nee, Shelby Co., Jul. 16-23 [no year]; 1 ♀ “B.M. ♀ Genitalia Slide No. 7116,” “Ho- (SNS10IL-00639), Manor East, 2 mi. lotype ♀ Gelechia nundinella Z., teste E. Fairfield, Wayne Co., Jun. 19, 2010. K. Sattler, 1961,” “Gelechia nundinella Indiana: 1 ♂ (USNMENT01480040), Zell., Verh Z-b GesWien. 23 p 256, TYPE Lawrence Co., Jul. 30, 1931; 1 ♀ ♀ (1873),” “Zeller Coll., Walsingham (USNMENT01480175, USNM84219), Collection, 1910–427,” “Gelechia nundi- Bedford, Lawrence Co., Aug. 28, 1931; nella Z., Texas Hg. 71” [green label], 1 ♂ (USNMENT01480041), Pike Co., “Dallas, Tex. Boll,” “368” [green label], Aug. 19, 1931. Iowa: 1 ♂ (USNMENT- “NHMUK010219656,” [NHMUK]. F. 01480042), Ames, Story Co., Jul. 26, beneficentella: ♂ “229 M., 6/15.80,” 1964. Louisiana: 5 ♂♂ (USNMENT- “Gelechia beneficentella, Murt., type” 01480044, USNMENT01480057, USN- [label with red border], “Database # MENT01480036, USNMENT01480037, USNMENT 00657958,” “♂ genitalia USNMENT01480038, USNM84214), 2

♀♀ (USNMENT01480035, USNMENT- Anderson Co., May 15, 1959. Virginia: 01480055, USNM84213), Edgard, St. 1 ♀ (USNMENT01480022), Wicomico John the Baptist Par., Jun. 21–Jul. 12, Church, Northumberland Co., Aug. 27, 1981; 1 ♂ (USNMENT01480039), 5 1914; 3 ♀♀ (USNMENT01480024, USN- ♀♀ (USNMENT01480031, USNMENT- MENT01480023, USNMENT01480045), 01480028, USNMENT01480032, USN- East Falls Church, Arlington Co., Sep. MENT01480029, USNMENT01344624, 4–19, 1929; 1 ♀ (USNMENT01480047), USNM84238), Baton Rouge, East Baton Fauquier Co., Jun. 3, 2018; 1 ♂ (USN- Rouge Par., May 22, 1970, May 25–Jun. MENT01480163), Alexandria (Rose Hill), 4, 1971, and Sep. 24, 1919; 4 ♂♂ Fairfax Co., May 7, 1976. (USNMENT01480056, USNMENT- We located twelve additional speci- 01480059, USNMENT01480058, USN- mens (7 ♂♂ [USNMENT01480046, MENT01480030, USNM84268), 2 ♀♀ USNMENT01434590, USNMENT00657963, (USNMENT01480026, USNMENT- USNMENT01480048, USNMENT01480015, 01480027), Tallulah, Madison Par., Jul. USNMENT00657961, USNMENT00657959, 27, 1910. Maryland: 1 ♂ (USNMENT- USNM9855, USNM84237]; 5 ♀♀ [USN- 01480033), Colesville, Montgomery MENT01434604, USNMENT00657960, Co., May 13, 1984; 3 ♂♂ (USNMENT- USNMENT00657962, USNMENT00657964, 01480164, USNMENT01480165, USN- USNMENT01480049, USNM9855, USNM- MENT01480166, USNM84221), McDaniel 130349]) at the USNM that were asso- (Wades Point), Talbot Co., Sep. 19–21, ciated together with the holotype of F. 1986. Missouri: 1 ♀ (USNMENT01480174, beneficentella. Of these, six are possibly USNM84218), Cooley Lake, Clay Co., Jul. in the hand of Murtfeldt and determined 18, 1968. North Carolina: 2 ♂♂ (USN- as F. beneficentella, while the remaining MENT01480034, USNMENT01480025), six are possibly in the hand of Riley. North Harlowe, Craven Co., Aug. 7–15, None of these specimens have locality 1990; 6 ♂♂ (USNMENT01480050, USN- data, and all were collected after the MENT01196862, USNMENT01480167, original description for F. beneficentella. USNMENT01480168, USNMENT01480169, The following 2 specimens were not USNMENT01480170, USNM84220), New physically examined but were taken from River State Park, Ashe Co., Oct. 8, 1999; BOLD: CANADA: Ontario: 1 (KSLEP1294), 1 ♂ (USNMENT01480171), Oliver Farm, Port Franks, Lambton Co., Aug. 7, 2018 (in New River State Park, Alleghany Co., Oct. Ken Stead’s private collection). UNITED 8–9, 1999. Ohio: 1 ♂ (USNMENT01480051), STATES: Mississippi: 1 (SL0038), Osborn, Springfield Bog Park, Summit Co., Aug. 27, Oktibbeha Co., Jun. 13, 1997 (specimen 2011. Oklahoma: 1 ♂ (USNMENT01480052), MEM13444 in MEM). Oklahoma City, Oklahoma Co., Aug. 14, 1955; Distribution.—This species has pre- 2 ♀♀ (MDOK-0566, MDOK-3127), viously been recorded in Arkansas, Geor- Bartlesville, Washington Co., Sep. 27, gia, Illinois, Indiana, Missouri, North 2008 and Aug. 15, 2009. Pennsylvania: Carolina, southwestern Ontario (Can- 1 ♀ (USNMENT01480053), Hog Island, ada), Pennsylvania, Texas, Virginia, and Delaware Co., Jun. 28, 1953. Texas: 1 ♂ the District of Columbia. (Foott 1967). (USNMENT01480054, USNM122693), We found specimens in the USNM from Laguna Park, Bosque Co., May 4, 1970; 1 Iowa, Louisiana, Maryland, Ohio, and ♀ (USNMENT01480020), West Austin, Oklahoma, and the MEM collection Travis Co., Sep. 18, 1975; 1 ♀ (USN- houses one specimen from Mississippi MENT01480021), 10 mi. SW Elkhart, (Fig. 39). This species likely occurs

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throughout the eastern United States nundinella into other regions where S. and southern Canada associated with carolinense has been introduced. its host plant. Biology.—F. nundinella feeds on Frumenta nephelomicta (Meyrick) Solanum carolinense. Throughout its range, the species has two broods, one in (Figs. 8–10, 15–16, 19, 22, 25, 29, 30, May and June, and another in late July 32, 34–38) through August (Bailey and Kok 1982; Foott 1967). Adults overwinter, and fe- Asapharcha nephelomicta Meyrick 1930: males begin ovipositing on shoots of the 555 host in late May (Bailey and Kok 1982). Frumenta nephelomicta: Hodges 1983: 22. The first brood constructs leaf capsules Frumenta solanophaga Adamski and by folding two terminal leaves together Brown 2002: 1030; new synonym with silk, while the second brood forms galls in fruits of the host plant (Foott Diagnosis.—Frumenta nephelomicta 1967; Solomon 1980). Larva excavate can be recognized by having a saccus escape holes in both the leaf capsules that is less than 1.5X the length of the and fruits before pupating to facilitate lateral arms of the vinculum; a phallus adult eclosion; only a thin membrane that in situ extends posteriorly beyond remains over the exit (Solomon 1980). If the juxtal lobes ¼ length of the phallus; fruits are not present on the host during wide posterior abdominal segments in the second generation, larvae will form the female; and a ductus bursae that is leaf capsules like the first brood (Bailey more than 2X longer than the apophyses and Kok 1982). Larvae are also capable anteriores. of stimulating parthenocarpy in the ab- Description.—Head: Vertex, frons, and sence of pollination. Frumenta nundi- haustellum yellowish white. Scape and nella is one of very few insects capable flagellum of antenna yellowish white; in of inducing this phenomenon; the others some specimens, intermixed with brown are fig wasps (Solomon 1980). Bailey scales. Labial palpus yellowish white; (1978) assumed that F. nundinella would second segment longer than third, scales be a successful biocontrol agent in re- on ventral surface divergent from base, gions where its host is an introduced forming a brush-like array; third seg- species due to the moth’s host specific- ment with appressed scales. Thorax: ity, its ability to significantly inhibit Tegulae and mesonotum yellowish white. plant growth, its ability to attack multi- Forelegs yellowish white, some speci- ple plant regions, its destruction of fruit mens with foretibia with two pale brown (and seeds), and, under favorable con- bands, one basal, one medially. Midlegs ditions, the possibility for it to undergo yellowish white, some specimens with rapid population build-up, especially in brown scales intermixed. Midtibia sim- the absence of other insect competitors ilar to foretibia, but with wider pale and hymenopterous parasitoids. Surveys brown bands. Hindlegs yellowish-white, conducted in Virginia by Kok (1989) hindtibia with piliform yellowish-white showed that F. nundinella causes sub- scales; some specimens with two pale stantial but localized damage to S. car- brown bands, one medially, one distal. olinense. However, despite its potential Tarsomeres of all legs yellowish white, as a biological control agent, no trials some specimens have brown tarsomeres have been conducted on establishing F. with yellowish white apical bands.

Forewing length 9.0–12.5 mm (mean = with tegumen sub-triangular (unrolled). 11.2 mm, n = 25); dorsal surface yel- Uncus trapezoidal, longer than it is wide, lowish white, some specimens with with lateral edges setose (rolled); lateral intermixed pale brown to dark grayish- margins may appear inwardly curved on brown scales. Some specimens with slide-mounted specimens as a product of basal third of wing with 3–4 olive- ﬂattening. Culcitula with short, linear brown spots. Discal cell with 1–3 ir- arms, arms abruptly dilated distally, regularly shaped brown to olive-brown forming sub-rectangular, medially con- spots. Costa with or without 1–2 brown cave, open extensions; densely covered spots on basal third. Outer margin with with micro dentitions (Figs. 34–38). faint brown spots; in some specimens, Gnathos spoon-shaped, widening api- these spots converge, forming a band. If cally, tapering into rounded apex (rolled). present, inner edge of band lined with Anterior diaphragma above phallus and grayish yellow, outer edge lined with ventrad tegumen expanded on median either band or spots of grayish yellow. line as convex, darkened, pedunculate Fringe yellowish white to grayish yel- process; lateral edges almost contiguous low. Ventral surface brown, with distal with inner medial margin of tegumen costa and outer margin yellowish white. (unrolled). Valva long, extending beyond In some specimens, outer margin lined posterior margin of uncus; basally nar- with one row of yellowish white spots. row, gradually widening from basal arch Venation (Fig. 15) with R4 and R5 di- with broadly-rounded apical margin; vergent near ½ from discal cell; apex of valva beyond basal arch long, setose discal cell closed diagonally, apex (rolled). Vinculum with short saccus, rounded; base of R3 and base of R4-R5 about 1.5X longer than lateral arms divergent from apex of discal cell; M1 of vinculum (rolled). Juxtal lobes short distally equidistant from R5 and M2;M2 and pointed posteriorly, setose (rolled). closer to M3 than to M1;CuA1 and CuA2 Phallus long and narrow with elongate, nearly parallel basally, diverging slightly dilated base, extending about ¼ to ½ 2 from / 3 length. Hindwing dorsal sur- length beyond juxtal lobes (rolled); duc- face pale grayish brown to brown, with tus ejaculatorius entering dorsally yellowish-white fringe or grayish yellow into basal opening (unrolled). Female fringe; ventral surface similar as dorsal. genitalia (Fig. 29) with posterior ab- Venation (Fig. 16) with Sc+R1,andRS dominal segments long and wide, with curved divergently from M1;Sc+R1 and width of posterior margin of segment 1 RS connected by base of R1 near / 3 VIII greater than ½ length of segments length of discal cell; M1 and M2 con- IX–X. Eversible scent pouch present vergent near 1/3, acutely curving diver- dorsally between segment VIII and pa- gently basally and slightly distally. CuA1 pillae anales (not illustrated, see Fig. 28 straight, divergent from and CuA2 from of F. nundinella). Papillae anales short slightly beyond discal cell. Frenulum and wide, with long and short setae in- with one acanthus in male, three acanthi terspersed. Apophyses posteriores long in female. Abdomen: Yellowish white and narrow, extending beyond anterior on dorsal and ventral surfaces. Female margin of segment VIII, nearly equal in abdomen with pair of small, invagi- length to apophyses anteriores. Anterior nated pleural pouches on intersegmental margin of segment VIII scalloped deeply. membrane anterior to 7th spiracle (Figs. Ductus bursae elongate, relatively nar- 30, 32). Male genitalia (Figs. 19, 22, 25) row, and slightly dilated at anterior end

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near inception of ductus seminalis; in- USNMENT01480081, USNMENT01480087, ception point with a rhomboid plate with USNMENT01480088, USNMENT01480086, lateral arms wrapping around ductus USNMENT01480092, USNMENT01480093, bursae. Corpus bursae elongate, narrow USNMENT01480090, USNMENT01480091, posteriorly, broadened anteriorly. Signum USNMENT01480095, USNMENT01480096, large with slightly pointed apex; base of USNMENT00657619, USNMENT01480100, signum protruded outwardly within cor- USNM12336, USNM84262, USNM84245, pus bursae. USNM82136, USNM12331, USNM130347), Types examined.—F. nephelomicta: Douglas, Cochise Co., Jul. 29, 1962 and Aug. ♀ “Holotype” [round label with red 8–Sep. 14, 1963; 1 ♂ (CMAZ-0023), 1 ♀ border], “Type” [round label with red (CMAZ-0069), 1.16 km W of Hwy 92 on E border], “Bent, New Mexico, 7000’ 10.27,” Miller Canyon Rd, Huachuca Mountains, Co- “ASAPHARCHA Meyr.,” “nephelomicta chise Co.,Sep. 30, 2009. New Mexico: 4 ♂♂ Meyr.,” “♀ genitalia on slide 9.11 1947 (USN MENT01480102, USNMENT- J.F.G.C 4794,” “Asapharcha nephelomicta 01480104, USNMENT01480105, USN- 1/1 Meyr., E. Meyrick det. in Meyrick MENT01480107, USNM84242, USNM84244, Coll.,” “Meyrick Coll., B.M. 1938–290.,” USNM82141), 3 ♀♀ (USNMENT01480101, “NHMUK010219655,” [NHMUK]. F. sol- USNMENT01480103, USNMENT01480106, anophaga: ♀ [incorrectly recorded as a USNM84241, USNM84142, USNM84243), male in original description] “MEXICO: 1 km N Animas, Hidalgo Co., Sep. 3, San Luis Potosi State, San Luis de la Paz, 1989; 3 ♀♀ (USNMENT01480108, 21.19N 100.32W, 10.ix.1999, H. G. Zim- USNMENT01480109, USNMENT01480120, mermann, AcSN 2109,” “Emerged in USNM84232, USNM84246), 2.3 mi NW Ro- Quarantine from berries of Solanum ele- deo, Hidalgo Co., Sep. 6, 1963; 1 ♂ (USN- agnifolium [sic], SOLANACEAE, Larvae MENT01480121, USNM12329), High Rolls, eat seeds and ﬂesh,” “National Coll[ection] Otero Co., Jul. Texas: 1 ♂ (USNMENT- of Insects, Pretoria, S[outh] Afr[ica],” 01480122), Palo Duro Canyon State “HOLOTYPE, Frumenta solanophagus Park, Randall/Armstrong Co., Sep. 25, Adamski + Brown” [red label], “Specimen 1968; 1 ♂ (USNMENT01196861), Shafter, ID USNMENT01196864,” “USNMENT- Presidio Co., Oct. 19, 1973; 1 ♂ (USNMENT- 01196864,” [USNM]. 01344622), Davis Mountains, Jeff Davis Other specimens examined.—28 Co., Oct. 4 1969. MEXICO: 6 ♂♂ ♂♂,26♀♀: UNITED STATES: Ari- (USNMENT01196866, USNMENT01196868, zona: 14 ♂♂ (USNMENT01480017, USNMENT01196869, USNMENT01196870, USNMENT01480114, USNMENT01480113, USNMENT01196871, USNMENT01196872, USNMENT01480111, USNMENT01480083, USNM82132, USNM81218, USNM82133, USNMENT01480084, USNMENT01480085, USNM82134), 2 ♀♀ (USNMENT01196865, USNMENT00657620, USNMENT01480094, USNMENT01196867, USNM81219, USNM- USNMENT01480089, USNMENT01480097, 81220), San Luis de la Paz, San Luis Potosi, USNMENT01480099, USNMENT01480098, Sep. 10, 1999. USNMENT01344623, USNM84231, USNM- Distribution.—Frumenta nephelomicta 84261, USNM12330, USNM84283, USNM- is found in the southwestern U.S. and 130346, USNM12332, USNM84265, central Mexico. Specimens in the USNM USNM84240, USNM82135), 18 ♀♀ collection are from Arizona, New Mexico, (USNMENT01480019, USNMENT01480016, and Texas. Specimens labeled as F. sol- USNMENT01480018, USNMENT01480110, anophaga in the collection were collected USNMENT01480082, USNMENT01480112, in San Luis Potosı´, Mexico (Fig. 39).

Additional specimens used in biological DC, for preparing the majority of the control studies were sourced from “Mex- illustrations featured in this work; Lisa ico” (without specific location) according Bartels for assisting JFL with prepara- to Neser et al. (1989). tion of Figs. 30–33; Jim Young for tak- Biology.—Frumenta nephelomicta has ing the SEM images of F. nephelomicta; been recorded on S. elaeagnifolium.Ac- Jeremy deWaard for making available cording to label data in the USNM, larvae BOLD specimen data for our analyses; are fruit feeders, consuming both seeds and Klaus Sattler and David Lees, Mi- and flesh (Adamski and Brown 2002). crolepidoptera Section, Department of Frumenta nephelomicta has been studied Entomology, Natural History Museum, as a possible biological control agent for London, United Kingdom, for making S. elaeagnifolium given its host specific- available the examination the types of ity, and it was introduced into South Af- Frumenta nundinella and F. neph- rica on three separate occasions between elomicta. Mention of trade names or 1978 and 1985 (Neser et al. 1989; commercial products in this publication Winston et al. 2014). However, it failed to is solely for the purpose of providing establish in the region; limiting factors specific information and does not imply included small release sizes during the recommendation or endorsement by the first two releases and severe drought USDA. USDA is an equal opportunity during the third release (Neser et al. provider and employer. 1989; Winston et al. 2014). Similar to F. davidi, F. nephelomicta is likely not a Literature Cited good biocontrol agent for S. elaeagnifo- Adamski, D. and J. Brown. 2002. A new species lium in South Africa, and no subsequent of Frumenta Busck (Lepidoptera: Gelechii- attempts have been done to introduce it dae: Gnorimoschemini) from Me´xico: a po- into the region (Olckers 1995). tential biocontrol agent against Solanum Remarks.—In the original description elaeagnifolium (Solanaceae). Proceedings of for F. solanophaga, Adamski and Brown the Entomological Society of Washington 104(4): 1029–1035. (2002) listed several characters that Bailey, T. E. 1978. Biology and ecology of Fru- separated this species from F. neph- menta nundinella (Zeller) (Lepidoptera: Ge- elomicta. These included a darker fore- lechiidae) and its impact on horsenettle wing pattern, a deeper basal arch of the (Solanum carolinense L.) Unpublished mas- valva, a distally wider valva, a longer ter’s thesis, Virginia Polytechnic Institute and phallus, and stouter apophyses ante- State University, Blacksburg, Virginia. Bailey, T. E. and L. T. Kok. 1982. Biology of riores. We consider these characters to Frumenta nundinella (Lepidoptera: Ge- represent variation within this species. lechiidae) on horsenettle in Virginia. The Additionally, after examining the holo- Canadian Entomologist 114: 139–144. type females of both F. nephelomicta Borror, D. J., D. M. DeLong, and C. A. Triple- and F. solanophaga as well as distribu- horn. 1981. An introduction to the study of insects, 5th ed. CBS College Publishing, tional and host data for both, we con- Philadelphia, PA. 928 pp. clude they are the same species. Brownrigg, R. (R version). 2018. Becker, R.A. and A.R. Wilks (Original S code). mapdata: Extra map databases. R package version 2.3.0. ACKNOWLEDGMENTS https://CRAN.R-project.org/package5mapdata. Busck, A. 1900. New species of moths of the We thank Young Sohn, Scientific Il- superfamily Tineina from Florida. Proceed- lustrator, Department of Entomology, ings of the United States National Museum, Smithsonian Institution, Washington, 23(1208): 225–254, pl. 1.

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