Argyria gonogramma Dyar, 1915

Landry, Bernard, Bilat, Julia, Hayden, James, Solis, M. Alma, Lees, David C., Alvarez, Nadir, Leger, Theo & Gauthier, Jeremy, 2023, The identity of Argyria lacteella (Fabricius, 1794) (Lepidoptera, Pyraloidea, Crambinae), synonyms, and related species revealed by morphology and DNA capture in type specimens, ZooKeys 1146, pp. 1-42 : 1

Argyria gonogramma Dyar, 1915

Figs 8 , 12 , 13 , 15 , 16 , 18 , 28 , 30 , 34

Argyria gonogramma Dyar, 1915: 87-88. Type locality: Bermuda. Błeszyński and Collins (1962: 213).

= Argyria pusillalis Hübner, 1818: 30, [36], [38], figs 167, 168. Type locality: [USA, Maryland] Baltimore. Nomen dubium.

Argyria pusillalis = Argyria pussillalis [sic] Hübner, 1818: 28; original misspelling.

Argyria lacteella (Fabricius, 1794): Fernald 1896: 72, plate V fig. 5; Grossbeck 1917: 126; Kimball 1965: 233; Tan 1984: 96 et seq.; Ferguson et al. 1991: 40; Munroe 1995: 35 (in part); Martinez and Brown 2007: 81, fig. 9.

Type material examined.

Holotype ♂ (Figs 8 , 15 , 16 ), with label data as follows: 1- "Bermuda, | 11.3.BWI | F.M. Jones", 2- "V-3 | D", 3- "Type No. | 18244 | U.S.N.M.", 4- "Argyria | gonogramma | Type Dyar", 5- "♂ genitalia | slide, 29Apr[il].'32 | C.H. #27", 6- "Genitalia Slide | By 107,454 | USNM"; deposited in the NMNH.

Other specimens examined.

411 specimens (see Suppl. material 2).

Morphological diagnosis.

In this small satiny-white moth measuring between 10.5 and 13.5 mm in wingspan, the median markings of the forewing (Figs 8 , 12 ) usually include a well-marked blackish-brown spot on the discal cell that is connected by curved lines to an oblique bar on the costa and a thin triangle on the dorsal margin. On the forewing costa, subapically, a thin curving bar is not followed by a triangle, but usually by 1-3 horizontal lines reaching the terminal margin below the apex. Relatively dark brown forms, with less contrasting markings (Fig. 13 ) have been collected in Alabama, Florida, and Louisiana (CUIC, FSCA, MHNG, NMNH) from December to April. In forewing markings this species is closest to A. lacteella (Fig. 11 ) in which there usually is a clear subapical triangle on the costa and for which the median spot, if present, is paler brown and usually smaller than the costal and dorsal triangles. In the absence of a subapical triangle on the forewing costa A. gonogramma is also similar to A. diplomochalis (Figs 9 , 14 ), which, however, doesn’t have any indication of a median spot or of any line between the median spot of the dorsal margin and the costa. In the male genitalia Argyria gonogramma (Figs 15 , 16 , 18 ) has the basal projection of the valva shorter than that of A. lacteella (Fig. 17 ) and just barely curving (Fig. 16 ); the cornuti on the vesica are also longer and thicker than those of A. lacteella . In the female genitalia (Fig. 28 ) only one wide, sclerotised pocket can be found anterior of the ostium bursae, whereas A. lacteella (Figs 24 , 27 ) has two pockets in the same area.

Molecular results.

Phylogenetic inference reveals that Argyria gonogramma constitutes a homogeneous clade. The monophyletic clade is identified in both GMYC and PTP species delimitation approaches, but it is not found in the ABGD approach and is grouped with A. lacteella (Fig. 1 ). This clade shows very low genetic variability within the COI barcode with an average intraspecific divergence of only 0.49% (Fig. 2 ). This low genetic diversity may be the result of different evolutionary processes, including recent colonisation. This species is mainly present in the US where it overlaps with A. lacteella in Florida (Figs 33 , 34 ).

Distribution.

Bermuda, Bahamas, widespread in the Eastern USA, from North Carolina in the North to the south of Florida, west to eastern Texas (Fig. 34 ).

Remarks.

The specimen of A. gonogramma labelled ‘Type’ in the NMNH is considered the unique holotype; the species’ description ( Dyar 1915) doesn’t indicate multiple specimens.

Argyria pusillalis Hübner is associated here with A. gonogramma and not with A. lacteella as in Munroe (1995) because at the latitude of Baltimore, Maryland, U.S.A., the type locality of A. pusillalis , only the superficially similar A. gonogramma or A. rufisignella (Zeller, 1872) could occur. Argyria nummulalis Hübner, 1818 is also known to occur in the eastern USA at the latitude of Baltimore, but this species lacks any median markings across the forewing, unlike the illustration of A. pusillalis . The name A. pusillalis is considered a nomen dubium because the original description and illustration associated with it do not allow a conclusive determination. Hübner’s collection was deposited in the Naturhistorisches Museum Wien, Vienna, Austria, which was destroyed by fire in 1835 ( Horn et al. 1990). However, although type specimens of some of Hübner’s Noctuidae species have recently been discovered in this museum (Gabor and László Ronkay, pers. comm. to BL, 11 April 2022), a search for a type specimen of A. pusillalis was not successful (S. Gaal, pers. comm. to BL, 10 August 2022). This issue could be settled by the designation of a neotype, but we refrain from doing that in order to avoid more instability in the nomenclature of this group. Also, we believe that it should be done in conjunction with a taxonomic revision of A. rufisignella , at the least.

Argyria pusillalis was originally named " pussillalis " ( Hübner 1818: 28), then mentioned as " Argyria pusillalis " on page 30 and on two indices (pages [36] and [38]), and finally as " pussillalis " again on the plate with the illustrations. Given that " pusillus " is Latin for small, it seems reasonable to believe that the original spelling " pussillalis " was in error.

Argyria gonogramma is a North American native species that was previously misidentified as A. lacteella and that has been collected in the United States since the late 1800's. The earliest specimens in the NMNH were collected by C.V. Riley from Ar[t]elier, FL, 1882 and N.[orth]C.[arolina] (undated). Another specimen collected by Boll in Texas (collection date unknown) was identified by "Rag[onot] \[18]86", and then by "CVR[iley]at the B. Mus. \[18]87".

That this species is native to the Southeastern U.S., or at least was established long before A. lacteella , is shown by earlier collecting dates for specimens in the FSCA and MGCL. For example: Florida, Sarasota Co.: 1951, Alachua Co.: 1960, Volusia Co.: 1962, Okaloosa Co.: 1963, Texas: 1978, Louisiana: 1979.

The earliest record of A. lacteella in 1979 in the USA (Florida) supports the conclusion that Tan (1984), although referring to A. lacteella , was in fact dealing with A. gonogramma . Tan’s (1984) unpublished MSc thesis provided a description of larvae, with setal maps, which were reared from egg to adult on St. Augustine grass ( Stenotaphrum secundatum (Walt.) Kuntze; Poaceae ). Tan (1984) further mentions that the early instar larvae eat the upper epidermis only and when not feeding, larvae hide in shelters made of leaves attached with silk, wherein moulting occurs. Tan (1984) also records the construction by the mature larva of a "small, compact silken case covered with frass and tiny pieces of chewed grass for pupation."

Based on collected series of specimens both Argyria gonogramma and A. lacteella now occur in sympatry and fly on the same dates in Florida, for example at Archbold Biological Station in Highlands County or in Pinellas County.

Fernald (1896) treated this species under A. lacteella (pl. V fig. 5), whereas his other illustrations on the same plate (Figs 4 , 6 ) represent the true A. lacteella . Grossbeck (1917), Kimball (most or all records, 1965), and Martinez and Brown (2007) all treated this species under A. lacteella . Melanic specimens collected in winter months account for the specimens of " A. diplomochalis " cited by Kimball (1965). This colouration variant may represent an adaptation for hiding in dry grass during the winter months and/or to obtain extra calories from the sun to allow biological activity.

The single moth at the basis of the Vermont record has been dissected and is correctly determined, but it is far outside the range since we know of no other record of A. gonogramma north of North Carolina. It was collected in sandplain habitat (M. Sabourin, pers. comm. to JH, 29 August 2022), which is consistent with the species’ habitat preference in Florida.

Figures

Figure 1. Phylogenetic inferences including 174 Argyria COI barcode sequences, i. e., 160 barcodes from BOLD, 9 COI sequences amplified by PCR (in bold), and 5 COI sequences obtained using capture from historical specimens (in red). Nodal support expressed in SH-aLRT and ultrafast bootstrap (UFBoot) is given as indicated in the caption except for nodes when SH-aLRT <80 and / or UFBoot <95. Species delimitation results including morphology, ABGD, GMYC, and PTP are indicated by different colours to represent the species proposed.

Figure 2. Mean genetic divergence (p-distance) between all samples from each species.

Figure 4. Holotype of Argyria lacteella synonym. Zebronia?? abronalis Walker, 1859 (Oxford University Museum of Natural History; OUMNH). Scale bar: 10 mm.

Figure 6. Holotype of Argyria lacteella synonym. Argyria vestalis Butler, 1878 (NHMUK Trustees of the Natural History Museum).

Figure 8. Habitus of Argyria type specimen (in NMNH) with labels underneath; holotype of Argyria gonogramma Dyar, 1915 (wingspan: 11 mm).

Figure 9. Habitus of Argyria type specimen (in NMNH) with labels underneath; lectotype of Argyria diplomochalis Dyar, 1913 (wingspan: 11 mm).

Figure 11. Habitus of additional Argyria specimen. Argyria lacteella (Florida, Putnam Co., FSCA). Scale bar: 2.5 mm.

Figure 12. Habitus of additional Argyria specimen. Argyria gonogramma (Florida, Seminole Co., MHNG). Scale bar: 2.5 mm.

Figure 13. Habitus of additional Argyria specimen. Argyria gonogramma (dark form, Louisiana, Calcasieu Co., MHNG). Scale bar: 2.5 mm.

Figure 14. Habitus of additional Argyria specimen. Argyria diplomochalis (St Croix Island, MHNG). Scale bar: 2.5 mm.

Figure 15. Male genitalia of Argyria gonogramma holotype (NMNH). Whole genitalia.

Figure 16. Male genitalia of Argyria gonogramma holotype (NMNH). Close-up of bases of valvae with tip of phallus in middle.

Figure 17. Male genitalia of Argyria species. Argyria lacteella (Florida, Alachua Co.). In FSCA. Scale bar: 500 µm.

Figure 18. Male genitalia of Argyria species. A. gonogramma (Florida, Wakulla Co.). In FSCA. Scale bar: 500 µm.

Figure 33. Distribution of Argyria lacteella (Fabricius).

Figure 34. Distribution of Argyria gonogramma Dyar.

Figures 27, 28. Female genitalia of Argyria species with ultimate segments cut dorsomedially from base to apex 27 Argyria lacteella (USA, Florida, Glades Co.) 28 A. gonogramma (USA, Florida, Orange Co.). Both in FSCA. Scale bar: 250 µm.

Figures 29 - 32. Heads of Argyria specimens 29 A. lacteella (USA, Florida, Pinellas Co.) 30 A. gonogramma (USA, Florida, Levy Co.) 31 A. diplomochalis (US Virgin Islands, St Croix) 32 A. centrifugens (Colombia, Amazonas, Leticia). All in MHNG. Scale bars: 250 µm (29 - 31), 500 µm (32).

Figures 24 - 26. Female genitalia of Argyria species 24 Argyria lacteella (holotype of A. abronalis, with spermatophore inside corpus bursae; OUMNH) 25 A. diplomochalis (Anguilla Island, BL 1889, CMNH) 26 A. centrifugens (Colombia, Amazonas, Leticia, MHNG-ENTO- 97427). Scale bars: 250 µm (24), 500 µm (25, 26).