Agriphila bistrialis: type lost. Given the difficulty of identifying species of Doryodes from an illustration, and the lack of a type locality for Agriphila bistrialis, we designate a neotype in order to preserve the long-standing identity of the inland species of Doryodes. Neotype ♂, USA, North Carolina, Carteret Co., Croatan National Forest, Sam Hatcher Road, 23 April 2006, J. Bolling Sullivan. BOLD barcode Sample ID: 06-NCCC-932 [USNM].

Ligia acutaria is nominally described from southern Russia, although already in 1852 Herrich-Schäffer suspected it was mislabeled. The type is lost, so to ensure that the current synonymy is maintained, we designate the neotype of Agriphila bistrialis as Neotype of Ligia acutaria also.

We have examined material from North Carolina east of the Piedmont, from most of Florida except for the Keys and Panhandle and a single male from a power cut near Grand Bay National Wildlife Refuge, Jackson County, Mississippi. Specimens from Florida, Mississippi and North Carolina are closely similar in appearance, genitalia and barcodes.

Superficially, adults usually can be identified by the narrower dark stripe on the forewing and their relatively small size (forewing length: ♂ 13.0–15.5 mm, ♀ 14.5–16.0 mm). Compared to D. spadaria the medial longitudinal stripe on the forewing of D. bistrialis is much narrower and the hindwings are whitish not buff through June emergences. Later in the year, D. spadaria hindwings become more whitish, so wing length and the width of the longitudinal line must be relied upon to distinguish the species, or genital characters must be examined for positive identification. Females of D. bistrialis do not overlap those of D. spadaria in size, females of D. spadaria having a forewing length of 18.0–21.0 mm. The male vesica is also diagnostic in D. bistrialis in that there is a line of smaller cornuti extending along the trunk usually in three patches, whereas in D. spadaria there are two side-by-side cornuti on the basal trunk of the vesica. In D. bistrialis, diverticulum 1 is 2–3 × as long as wide and has a deeply-serrated rooster-comb-like cornutus at the apex; D. spadaria has no diverticulum in this position and the cornuti in the vesica are triangular, sometimes minutely serrated on one side. D. bistrialis can be distinguished from D. fusselli by the shape of diverticulum 1, which is rounded in D. fusselli and about as long as wide; also the apex of the vesica in D. bistrialis is symmetrical with a triangular cornutus on a pouch on each side (diverticula 4 and 5). D. bistrialis can be distinguished from D. reineckei and D. latistriga by the wing pattern and by size and from D. broui by the characters of the vesica. Along the Gulf Coast D. bistrialis could be confused with D. broui and D. tenuistriga, but characters of the vesica (D. broui, Fig. 40 and D. tenuistriga, Fig. 42, versus D. bistrialis, Fig. 33) readily distinguish these three species and D. bistrialis occurs farther inland. The female genitalia of D. bistrialis are elongated with a more compact, less differentiated, appendix bursae than in D. spadaria.

Doryodes bistrialis, unlike all other species in the genus, occurs mainly inland away from coastal salt marshes. It occurs in pine savannas where wiregrass (Aristida stricta), the presumed food plant, is abundant. It has only been recorded in North Carolina, Mississippi and Florida, and it follows the distribution of the presumed foodplant, Aristida stricta. The species is on the wing from April through October in North Carolina, and throughout the year in Florida. The species appears to be associated with wiregrass, but we were unable to successfully rear it on cut grasses. Eggs should be placed on potted Aristida and larvae monitored to determine their preference for the grass or detritus in the base of the grass clump. In North Carolina the savannas are usually a half mile or more inland from coastal marshes and extend westward into the Sandhills adjacent to the piedmont. It is possible that the salt marsh species and the wiregrass species could occur in the same or very close areas where coastal marshes penetrate inland but we did not find such areas.

Holotype ♂. Florida, Pinellas Co., Ft. Desoto Park nr St. Petersburg, 4 Jan. 1968, J. D. Lafontaine. CNC. Paratypes: 2 ♂. Florida. Gulf Co., Rd. to Cape San Blas, nr Port St Joe, 29.8°N, 85.3°W, 31 July 1998, Jeff Slotten, genitalia slide FLMNH-MGCL 02944 (1 ♂). Florida, Sarasota Co., Siesta Key, 3 Feb. 1952, C.P. Kimball, genitalia slide FLMNH-MGCL 02948 (1 ♂). FSCA.

This species is named after Ft. De Soto Park, Florida.

This species is superficially indistinguishable from D. spadaria, D. fusselli, and D. broui, although its range on the Gulf Coast of Florida is north and west of the range of D. spadaria and D. fusselli, and east of the range of D. broui. Forewing length is 16 mm, on the small size for D. spadaria, but within the normal range of the other similar species. The species can be definitively identified only by the male vesica, particularly by the long, narrow diverticulum 1, which is about 4–5 × as long as its basal width and with an apical, deeply serrated rooster-comb-like cornutus; diverticulum 2 is on the left side of the vesica, not posterior as in D. bistrialis; diverticulum 4 is preapical on the right, without a cornutus, and diverticulum 5 forms a projecting lobe with a cone-shaped diverticulum on its left side.

External structural characters as described for genus. Forewing length 16 mm; forewing whitish buff with slightly darker-buff and pale-gray streaks; a prominent blackish-brown stripe along the middle of wing, curving upward and tapered at about ¾ from base; stripe narrower than for D. spadaria and D. okaloosa, but wider than for D. bistrialis; stripe bordered by narrow white line above extending to ¾ from base, and with similar white line below stripe extending from above forewing tornus almost to wing apex. Hind wing white with very faint buffy tone. Male genitalia mainly as described for genus. Dorsal heavily-sclerotized margin of valve extending beyond middle membranous part, then tapered abruptly into a sharp spine; ventral sclerotized margin of valve not evenly tapered, but widens slightly preapically then tapered to blunt point free from inner membranous part of valve. Aedeagus cylindrical, 8–9 × as long as mesial width. Vesica with swollen area distal to end of aedeagus, 0.4–0.5 × as long as aedeagus and about 2 × as long as wide, with two spinule-covered sclerotized plates, these partially or completely divided into as many as four plates each with less spinules; diverticulum 1 finger-like, 4.5–5.5 × as long as mesial width, with apical serrated cornutus; diverticulum 2 rounded, on left side at base of diverticulum 1, with shark-fin-like cornutus; diverticulum 3 quadrate, on right side and without a cornutus; diverticulum 4 preapical, on right side without a cornutus; diverticulum 5 forming a bulbous apical lobe with a rounded spine-tipped cornutus on left side.

This species is known only from three male specimens, all from the Gulf Coast of Florida between Sarasota County and Gulf County. Collecting dates are in January, February, and July. Nothing is known of its biology except it is associated with coastal salt marshes.

Holotype ♂, Florida, Okaloosa Co., Shalimar, black light trap, 3461, H. G. Hilton, genitalia slide FLMNH-MGCL 02951. FSCA.

The species name is in honor of the Okaloosa, a tribe of the Creek Nation and longtime inhabitants of the area.

This species probably occurs with D. desoto and D. reineckei in the salt marshes and tidal creeks throughout the coastal panhandle area of Florida. The species is slightly larger than D. bistrialis, which may occur nearby but inland. The washed out appearance of D. reineickei immediately distinguishes it from D. okaloosa. Doryodes desoto is very similar and at present dissection of the male genitalia is the only reliable way to distinguish these two species, although based on the single specimen of D. okaloosa, it appears that D. okaloosa is broader winged than D. desoto, but not as broad winged as D. reineckei. The abundance of large cornuti in the vesica, as well as the spatulate lower process of the valve and the short, broad aedeagus distinguishes D. okaloosa from all other Doryodes species.

Forewing length 16.5 mm; forewing stripe dark brown, narrower than for D. spadaria but wider than for D. desoto; forewing wider and browner than in D. spadaria and D. desoto, but not as wide as in D. reineckei, and longitudinal stripe sharply defined, unlike that of D. reineckei. Antennae missing but presumed to be bipectinate as in other species in genus. Hind wing white with buff tinge. Male genitalia mainly as described for genus. Dorsal heavily-sclerotized margin of valve wider and less well defined than in other species except for tapered spine-like apex; apex not rounded as in D. tenuistriga, but wider than in other species; ventral sclerotized margin of valve slightly tapered to a broadly spatulate apex, much as in D. tenuistriga. Aedeagus cylindrical, about 5 × as long as mesial width. Vesica with swollen area distal to end of aedeagus about 0.5 × as long as aedeagus and about 2 × as long as wide, with four large spine-covered sclerotized plates; diverticulum 1 slightly longer than wide with large deeply-serrated cornutus near apex; three preapical diverticula, each with large shark-fin-like cornutus; arrangement of large cornuti not seen in any other species of Doryodes.

At present this species is known from the holotype collected in Okaloosa County, Florida. It likely ranges south in the coastal brackish marshes toward the St. Petersburg/Tampa area and to the west along the Florida coast but little collecting has been done in salt marshes of the Florida Panhandle.

Doryodes spadaria: type lost. It is impossible to identify this species from the description given by Guenée, but the type, listed as being from Florida, would most likely be from the East Coast, which would mean it represented the species now known either as D. bistrialis, D. fusselli or D. spadaria. In order to maintain the longstanding identity of this species as the widespread species in salt marshes of the Atlantic Coast, we designate the lectotype of Themma divisa Walker as neotype of Doryodes spadaria, an action that will ensure the current identity and synonymy. The data are given under Themma divisa. Themma divisa: lectotype ♂, here designated, BMNH, examined; male in good condition except right antenna and tip of left antenna missing; forewing length 18.1 mm; data “E. Doubleday, St John’s Bluff, E Florida/ Themma divisa/ syn-type [blue circle]/ 40-1-14-84/ S.O. 153.” Tunza promptella: holotype ♂, no locality, BMNH, examined. Doryodes spadaria race grandipennis: syntypes ♂, ♀, USNM, examined.

Canada: Quebec, Nova Scotia, Prince Edward Island. USA: Maine, New York, New Jersey, Maryland, North Carolina, South Carolina, Georgia, Florida.

External structural characters as described for genus. Doryodes spadaria is the most widespread and common species in the genus, and except for Doryodes fusselli in coastal North Carolina, all specimens of Doryodes from salt marshes along the Atlantic Coast of Canada and the United States that we have seen are Doryodes spadaria. Adults are relatively larger (forewing length: ♂ 13–20 mm, most commonly 16–18 mm; ♀ 18–21 mm, most commonly 19 mm) than those of Doryodes bistrialis (forewing length: ♂, 13.5–15.5 mm, ♀ 14.5–16.0 mm), the species most likely to be confused with it in southeastern United States outside of North Carolina where the smaller D. fusselli also occurs in salt marshes. The forewing ground color in males varies from whitish buff to yellow buff with gray streaks; the longitudinal dark stripe is dark brown and conspicuously wider than in D. bistrialis; the ground color in females averages paler than that of males and the wings and dark longitudinal stripe are narrower. In the male genitalia, most structural characters are as described for the genus; the sclerotized costal margin of the valve is more heavily sclerotized apically and extends farther beyond the central membranous part of the valve than in most other species; the sclerotized ventral margin of the valve ends in a blunt point before the end of the central part of the valve. Two examples of the genitalia are illustrated, one to show the typical orientation of the vesica in comparison with other species (Fig. 37), and a ventral orientation of the vesica (Fig. 38) to illustrate the shape and positions of the two preapical diverticula and their cornuti. The vesica is slightly shorter than the aedeagus; the basal, swollen part of the vesica is armed with two thorn-like cornuti on sclerotized plates; the left preapical diverticulum (# 5) is tapered to a large triangular cornutus, whereas the right preapical diverticulum (# 4) has a bulge in the middle and a smaller cornutus. In the female genitalia the corpus bursae is elongated with the part anterior to the opening of the ductus bursae swollen laterally and with longitudinal “ribbons” of sclerotization. The appendix bursae is lightly sclerotized and slightly bilobed posteriorly. The ductus bursae is almost as long as the corpus bursae, and is heavily sclerotized dorsally and ventrally with the plates slightly narrower mesially and expanded laterally and more heavily sclerotized at their junction with the corpus bursae; the ventral plate at the end of the ductus bursae is extended posteriorly as a quadrangular plate that projects over the opening to the ductus with the posterior margin of the plate concave. The anterior and posterior apophyses are similar in length (posterior slightly longer), about half the length of the ductus. The anal papillae are lightly sclerotized, produced ventrally anteriorly, rounded posteriorly, with the surface setose.

Doryodes spadaria is widely distributed in coastal salt marshes on the Atlantic Coast of Canada and the United States from eastern Quebec to southern Florida. A few inland records in southern Florida (e.g., Kissimmee Prairie) are anomalous, unless there is brackish water in these areas. Adults occur in Canada and northeastern United States from June to August. In southeastern United States there are spring and summer generations and at least three generations in Florida.

Holotype ♂, North Carolina, New Hanover Co., Fort Fisher Maritime Forest, A trail, 775504 W 335833 N, 15 Watt UV trap, Coastal fringe Evergreen Forest, June 3, 1995, J. Bolling Sullivan, Richard Broadwell & Brad Smith. USNM. Paratypes: 8 ♂, 4 ♀. North Carolina. Same data as for holotype (2 ♂, 2 ♀); same locality and collector as holotype, June 27, 1995 (1 ♂, 1 ♀), April 19, 1995 (1 ♂), November 8, 1994 (1 ♂), October 5, 1994 (1 ♀). North Carolina, Carteret Co., Ft Macon State Park, 15 July 1999 (1 ♂); Carteret Co., maritime scrub, N34.697°, W-76.683°, 29 Aug. 2005, J. Bolling Sullivan, barcodes 05-NCCC-523, & 05-NCCC-524 (2 ♂). BIO, CNC, JBSC, USNM.

The species name is in honor of John Fussell from Morehead City, North Carolina, who has worked tirelessly for decades to describe and protect the unique flora and fauna of the North Carolina coastal plain, particularly the Croatan National Forest. All of our lives are richer for his efforts.

This species occurs with D. spadaria in the salt marshes and tidal creeks throughout coastal North Carolina Adults are slightly larger than D. bistrialis, but noticeably smaller (especially females) than D. spadaria. The medial chocolate stripe on the forewing is broader than in D. bistrialis, but narrower than that of D. spadaria. Spring males are larger than those of the summer and fall generations, so they are more easily confused with D. spadaria. The hind wing is white, without the buff coloring of D. spadaria; in late summer some males of D. spadaria can have white hind wings, but size ranges for the two species do not overlap in this generation. Similar changes in size with season are seen for many other species (Sullivan and Miller 2007). The vesica differs in having a diffuse line of cornuti along the basal trunk and by the symmetrical pair of cornuti-tipped diverticula at the apex. Doryodes spadaria is described in full, so only differences from it are given in the description.

Smaller than D. spadaria; spring males (forewing length: 16–17 mm [16–20 mm for D. spadaria]), slightly larger than summer males (forewing length: 14–15 mm); some small individuals (forewing length: 12–13 mm) late in season; females of D. fusselli (forewing length: 16 mm) smaller than those of D. spadaria (forewing length: 18–21 mm) but larger than those of D. bistrialis. Hindwing pearly white. Male genitalia with valves similar to those of D. spadaria but slightly smaller (valve length: 4.45 mm versus 4.75 mm for D. spadaria). Vesica basal trunk with single band of cornuti, usually separated into two elongated sections; diverticulum 1 rounded with toothed, rooster-comb-like cornutus on dorsal side; diverticulum 2 rounded, about 1.0–1.5 × as long as wide, with triangular cornutus on ventral surface near apex; terminus of vesica with two similar cornuti-tipped diverticula projecting distally [in D. spadaria two terminal diverticula are more elongated, different in shape from each other, and project laterally]. Female genitalia differ from those of D. spadaria in more even width of ductus bursae, and appendix bursae more elongated with posterior margin even, not bilobed as in D. spadaria.

At present this species is known only from North Carolina, occurring from Dare County in the north to Brunswick and New Hanover counties in the south. It is likely that it occurs farther south but may have been overlooked as D. spadaria. Specimens have been collected from April through October and the species appears to be on the wing continuously. Eggs were obtained from a female and fed cut Spartina alterniflora Loisel. leaves and fresh and wilted Bermuda grass. The larvae survived to the second instar and were similar to those of D. spadaria (Wagner et al. 2011). Larvae have not been located in the field.

Holotype ♂, Alabama. Baldwin Co., Camp Beckwith, UV trap, Spartina-Juncus marsh, N 30.39538°; W -87.84657°; 5 Aug. 2009. J. Bolling Sullivan. USNM. Paratypes: 44 ♂, 9 ♀. Alabama. Same data as holotype (2 ♂); same locality and collector as holotype, 7, 9 & 14 August 2009 (3 ♂); Baldwin Co., Weeks Bay Preserve, N 30.414°, W -87.833°; 4 Aug. 2009. J. Bolling Sullivan, barcodes 09-MISC-046, 047, 048, 049 (4 ♂). Louisiana. St. Tammany Parish, 4.2 mi NE Abita Springs, sec 24, T6S R12E, N 30°30.986’, W 89°57.276’, 25 Feb. 1997, V.A. Brou Jr. CNC slide ♂ 16049 (1 ♂); same locality and collector, 20 May & 27 Nov. 1983 (2 ♂), 23 May 1984 (1 ♂), 26 March 1985 (1 ♂), 27 March 1987 (1 ♂), 14 Aug. 1988 (1 ♂), 13 Jan. & 20 March 1989 (2 ♂), 13 Sept. 1998 (1 ♂), 28 May 2000 (1 ♂), 12 Oct. 2001 (1 ♂), 27 Jan. & 16 March 2002 (2 ♂), 3 April 2008, Barcode CNCLEP00113508 (1 ♂), 11 April 2008 (1 ♂), 12 April & 28 Oct. 2009 (2 ♂, 1 ♀), 21 March 2010 (1 ♂), 7 Feb. 2011, CNC slide 16058 (1 ♀), 22 April 2012, Barcode CNCLEP00113566) (1 ♂). Cameron Parish, Johnson’s Bayou, 16 March 2002, V.A Brou Jr. (1 ♀). Lafourche Parish, near Golden Meadow, 5 March 2006, V.A. Brou Jr., CNC slide ♂ 16054 (1 ♂); same locality and collector, 2 July 2005 (1 ♀), 5 March 2006 (6 ♂). Mississippi. Harrison County, Long Beach, 17 June 1992, V.A. Brou Jr. (1 ♀); same locality, 2 June 1996, CNC slide ♂ 16679, R. Kergosien (1 ♂), same locality and collector, 2 May 1997, CNC slide ♂ 16681 (1 ♂), 20 Nov. 1992, CNC slide ♂ 16706 (1 ♂), 17 May 1996, CNC slide ♀ 16440, (1 ♀). Jackson County, Gulf Coast Islands National Seashore, 19–20 April 1985, R.L. Brown (1 ♀). Jackson Co., Ocean Springs, 12, 15, 17 June 1992 & 1 Aug. 1992 (5 ♂, 2 ♀), V.A. Brou Jr. CNC, JBSC, MEM, USNM, VABC.

The name of this species refers to the width of the longitudinal dark stripe on the forewing in both sexes.

We have examined material from southern Alabama, Mississippi, and Louisiana. Specimens in Bold database exhibit considerable heterogeneity but all sequences are within 0.8% of each other.

This species occurs with D. broui, D. reineckei, and D. tenuistriga. It is the largest species and with the most distinctive forewing pattern. It should not be confused with D. tenuistriga, which has a narrow forewing stripe, or with D. broui, which is smaller (forewing length: 13.0–15.5 mm) and also has a narrower forewing stripe. It can be distinguished from D. reineckei by the breadth and distinctness of the longitudinal stripe; in females of D. reineckei the forewing stripe is absent or pale gray, hardly contrasting with the ground color. The most distinguishing character of the male genitalia is the elongated terminal diverticulum of the vesica that projects ventrally. Females are similar to males but have longer, more pointed forewings and whiter hindwings. The female genitalia are the most robust of any Doryodes species. The corpus bursae has a bulge on the right side that is more pointed than in the other species, and the appendix bursae is quadrate with the posterior margin almost straight and heavily sclerotized.

Forewing ground color in spring and summer specimens yellowish white to buff with gray streaking, hindwing white to whitish buff; forewing in fall and winter specimens darker with more brown shading, hindwing variably suffused with brown, especially along wing margin. Forewing length 14.5–18.0 mm (males), 17.5–20.0 mm (females), similar in length to those of D. spadaria, but wings somewhat broader. Longitudinal stripe broader than in any other species in genus. Male genitalia with cornuti in variable patches on basal trunk of vesica, larger patch dorsal and most basal, two smaller patches of cornuti lateral and more distal (but one or both can be absent). Vesica with diverticulum 1 absent, its position represented by rounded curve of vesica posterior to swollen part of vesica after aedeagus; diverticulum 2 a rounded bulge on left side of vesica with large, slightly-serrated shark-fin-like cornutus; diverticula 3 a short quadrate pouch on right side almost opposite position of diverticulum 2; diverticulum 4 cone shaped, dorsolaterally on right, with bulbous spine-tipped cornutus at base near junction with elongated ventral lobed representing diverticulum 5. Female genitalia similar to those of the other species but more robust, and appendix bursae quadrate with posterior margin almost straight and heavily sclerotized.

The adults are found in tidal creeks and salt marshes from Alabama to Louisiana. The biology is unknown, but presumed to be similar to other species of Doryodes that occur in similar habitats. Adults occur throughout the year, but concentration of collecting dates suggests a primary brood from March to May and a secondary brood in September and October (V.A. Brou Jr. pers. comm.).

Holotype ♂, Louisiana. St. Tammany Parish, 4.2 mi NE Abita Springs, sec 24, T6S R12E, N 30°30.986’, W 89°57.276’, 3 May 2010, V.A. Brou Jr.; slide CNC 16053; barcode CNCLEP 00113565. CNC. Paratypes: 98 ♂, 49 ♀. Louisiana. Same locality and collector as holotype, 25 April 1984, CNC slide ♀ 16057 (1 ♀), 24 May 1984 (1 ♀), 25 April 1985 (1 ♂), 5 & 28 May & 30 June 1986 (3 ♂), 2, 15 & 16 June & 12 Aug. 1988 (4 ♂), 5 & 20 March & 28 April 1989 (4 ♀), 16 & 22 June & 20 Dec. 1990 (2 ♂, 1 ♀), 14 April 1991 (1 ♀), 22 May & 1 Nov. 1992 (1 ♂, 1 ♀), 18 Oct. 1993 (1 ♀), 11 & 29 May, 13 June & 16 Sept. 1996 (4 ♂), 1 March & 9 April 1997 (2 ♂), 26 & 28 May 1998 (2 ♂), 5, 8 & 11 May 1999 (2 ♂, 1 ♀), 26 Jan. & 15 May 2000 (1 ♂, 1 ♀), 18 & 19 March 2002 (1 ♂, 2 ♀), 5, 11 & 31 May 2003 (3 ♂, 2 ♀), 3 June 2005 (1 ♀), 19 May & 2 & 12 June 2006 (2 ♂, 1 ♀), 20 Feb., 6 April, & 6 & 25 May 2008 (2 ♂, 2 ♀), 26 March & 12 May 2009 (2 ♂), 21 March & 13 May 2010 (1 ♂, 3 ♀), 7 & 23 April, 3, 21 & 25 May & 7 July 2011 (2 ♂, 4 ♀), 8 & 10 April & 12 May 2013 (1 ♂, 3 ♀). St. Tammany Parish, Hwy 90 at Hwy 433, 8 May 1971, E.H. Metzler, Female genitalia on slide # E.H.M. 686 (1 ♀). Cameron Parish, Johnson’s Bayou, 19 April 1985, CNC slide ♀ 16048 (1 ♂, 1 ♀), 10 & 19 Sept. 1985, (4 ♂), 14 Sept. 1990, V.A. Brou Jr. (3 ♂). Cameron Parish, Little Chenier, 14 May 1981, V.A. Brou Jr. (1 ♂). Lafourche Parish, Golden Meadow, 28 April 1975, V.A. Brou Jr. (2 ♂). Lafourche Parish, near Golden Meadow, 25 March 2007, V.A. Brou Jr. (1 ♀). St. John the Baptist Parish, Edgard, 29 March, 2 & 9 April & 9 June 1976 (4 ♂, 1 ♀), 7 April, 20 May, 5 June & 5 Aug. 1977 (4 ♂), 10 Aug., 6 May, 1 June & 7 Aug. 1978 (4 ♂, 1 ♀), 1 April, 12 & 21 May, 24 July & 17 Aug. 1979 (3 ♂, 3 ♀), 25 June, 25 July, 4 & 12 Aug., 27 Sept., 17 & 29 Oct. & 2 Dec. 1980 (7 ♂, 2 ♀), 17 April, 9 May, 4 & 6 June & 2 Oct. 1981 (6 ♂, 1 ♀), 16 & 25 March, 17 & 20 April, 12, 21 & 24 May, 4 & 26 June, 11 Aug., 6 Sept., 7 Oct. & 25 Dec. 1982 (11 ♂, 7 ♀), 14 May 1983 (1 ♂), 23 May 1984 (1 ♀), V.A. Brou Jr. St. John the Baptist Parish, Edgard, 10 May 1971, E.H. Metzler, Male genitalia on slide # E.H.M. 685 (1 ♂). Vermillion Parish, Intracoastal City, 26 July 1984, V.A. Brou Jr. (1 ♂). Mississippi. Harrison County, Long Beach, 18 May 1992, R. Kergosien (1 ♂); same collector and locality, 15 May 1997, CNC slide 16680 (1 ♂); Jackson Co., Grand Bay Nat’l Wildlife Refuge, N 30°41.3’ W 90°40.6’, coastal marsh savanna, 21 July 2014, J. Bolling Sullivan, barcodes 14-NCCC-470, 471, 472, 473, 474 & 475 (5 ♂). Texas. Jackson Co., Deutechburg, 7 Oct. 1974, A. & M.E. Blanchard, slide CNC 16682 (1 ♂); Brownsville, 6-11, Geo. Dorner (1 ♂). CNC, EHMC, JBSC, MEM, USNM, VABC.

We name this species after Vernon A. Brou, Jr. in recognition of his impressive and tireless efforts in collecting and researching the Lepidoptera of Louisiana.

This species occurs with D. latistriga, D. reineckei, and D. tenuistriga in coastal salt marsh habitats from Alabama to southern Texas. D. broui is superficially indistinguishable from D. spadaria, but differs from it in male genitalia, barcodes, and occurs far to the west of the known range of D. spadaria. It can be distinguished from D. reineckei in having a sharply-defined longitudinal dark stripe on the forewing, and from D. latistriga by the narrower forewing stripe in D. broui. It is most likely to be confused with D. tenuistriga, which typically is larger, but because of variation in both species, some specimens must be dissected or barcoded, for positive identification. The male genitalia of D. broui are most similar to those of D. latistriga, but the diverticula and cornuti are smaller, especially the terminal diverticulum (# 5), which is short and rounded, not elongated as in D. latistriga.

Forewing length: 13.0–15.5 mm (males), 13.5–17.0 mm (females), Forewing buffy brown to whitish gray with faint buffy streaks, darker forms in colder months; longitudinal stripe dark brown, similar in width to that of D. spadaria, narrower than for D. latistriga, wider than for D. tenuistriga. Male genitalia. Aedeagus 8 × as long as mesial width; vesica with dorsolateral toothed triangular cornutus on left side of basal part of vesica distal to end of aedeagus; a sclerotized plate in anterior 90° bend in vesica at position of ductus seminalis; posterior curve in vesica extended posteriorly into rounded diverticulum 1 with toothed preapical cornutus; diverticulum 2 on left side reduced to low bulge with large conical cornutus in middle; preapical posterior diverticulum (# 4) tapered with conical cornutus at distal base and also at base of short rounded apical diverticulum 5. The female genitalia are similar to those of D. latistriga, but the appendix bursae is rounded posteriorly and only lightly sclerotized.

Doryodes broui occurs from Alabama to southern Texas. Nothing is known of its biology. Adults occur throughout the year, but concentration of collecting dates suggests a primary brood between mid-March and mid-June and a secondary protracted brood between late July and mid-October (V.A. Brou Jr. pers. comm.).

Holotype ♂, Alabama. Baldwin Co., Camp Beckwith, UV trap, Spartina-Juncus marsh, W 30.39538; N -87.84657; 7 Aug. 2009. J. Bolling Sullivan. USNM. Paratypes: 122 ♂, 74 ♀. Alabama. Same data as for holotype but collected 5 Aug. 2009 (one with slide CNC 16414) (5 ♂), 9 Aug. 2009, one with barcode label CNCLEP 00113509 (2 ♂); Baldwin Co., Weeks Bay Preserve, N 30.414°, W -87.833°; 4 Aug. 2009. J. Bolling Sullivan, barcodes 09-MISC-050, 051, 052, 053 (4 ♂). Louisiana. St. Tammany Parish, 4.2 mi NE Abita Springs, sec 24, T6S R12E, N 30°30.986’, W 89°57.276’, 8 April 1983, V.A. Brou Jr. (1 ♂); same locality and collector, 15 & 20 May, 2 Sept. 1983 (5 ♂, 2 ♀), 13, 26–30 April, 8 & 23 May & 8 Aug. 1984 (2 ♂, 7 ♀), 26 & 28 April 1985 (1 ♂, 1 ♀), 1 May, 10 May, CNC ♀ slide 16056, 11, 14, 16 & 22 May 1986 (5 ♂, 4 ♀), 9 & 31 May & 29 Nov. 1988 (2 ♂, 1 ♀), 8 April, 25 May & 6 June 1989 (2 ♂,1 ♀), 26 Feb., 10 March, 14 May & 2 June 1990 (3 ♂, 1 ♀), 12 April 1991 (1 ♀), 24 June 1992 (1 ♂), 14 April & 16 Nov. 1993 (2 ♂), 17 April & 4 May 1994 (2 ♂), 29 March, 23 April & 2 Aug. 1995 (3 ♂), 20 & 22 April, 20 May & 29 May 1996 (4 ♂, 2 ♀), 24 Feb. & 1 March 1997 (2 ♂), 16 June 1998 (1 ♂), 4 April 1999 (1 ♂), 19 March 2000 (2 ♀), 28 April 2001 (1 ♂), 24 Jan., 15, 16 & 22 March 2002 (3 ♂, 7 ♀), 7 & 8 April 2005 (1 ♂, 2 ♀), 27 March, 26 April & 2 May 2006 (3 ♂), 28 April, 5 May & 6 June 2007 (3 ♂), 28 March slide CNC 16410 ♂, 31 March, 11 April barcode CNCLEP 00113564, 16 April, 1 May, 19 June & 11 Aug. 2008 (4 ♂, 3 ♀), 14 Feb., 21 March, 12, 19, 21 & 28 April 2009 (7 ♂, 1 ♀), 21 March, 3, 23 & 25 April, 5, 24 & 26 May 2010 (6 ♂, 2 ♀), 21 Jan., 8, 22, 25, 26 & 27 April, 3, 8 & 28 May 2011 (8 ♂, 6 ♀), 12 Feb., 10 April 2012, barcodes CNCLEP 00113563 & 00113568, 22 April, 9 June 2012 (2 ♂, 3 ♀), 10 April 2013, barcode CNCLEP 00113511, 10 April 2013, CNC slide ♀ 16436 (2 ♀), V.A. Brou Jr. St. Tammany Parish, Hwy 90 at Hwy 433, 8 May 1971, E.H. Metzler, Female genitalia on slide # E.H.M. 683 (1 ♀). Cameron Parish, Johnson’s Bayou, 23 Oct. 1985, V.A. Brou Jr. (1 ♂). Lafourche Parish, Cut Off, 21 Feb., 11 & 19, 20, 21, & 25 April, 16 May 1975, V.A. Brou Jr. (3 ♂, 14 ♀). Lafourche Parish, near Golden Meadow, 2 July & 6 Aug. 2005 (3 ♀), 25 March 2007, V.A. Brou Jr. (1 ♀). St. John the Baptist Parish, Edgard, 23 April 1973, (1 ♂), 16 April 1975 (1 ♂), 26 March, 18 & 23 April & 1 May 1976 (5 ♂, 1 ♀), 22 April, 5, 6 & 23 May 1977 (3 ♂, 2 ♀), 1, 7, 8 & 11 May & 3 June 1978 (5 ♂), 22 March, 1, 13 & 24 April & 2 May 1979 (6 ♂, 2 ♀), 9 May 1981 (1 ♂), 6 Jan., 16 & 24 March, 2, 17, 18 & 24 April 1982 (6 ♂, 1 ♀), 14 May 1983 (2 ♂, 1♀), V.A. Brou Jr. Mississippi. Harrison County, Long Beach, 11 Feb. 1995, R. Kergosien (1 ♂); Jackson Co., G.C.R.L. Ocean Sp. [Gulf Coast Research Laboratory, Ocean Springs], 25 January 1993, R. Kergosien (1 ♂). CNC, EHMC, FSCA, JBSC, MEM, USNM, VABC.

The species is named for John P. Reinecke, a retired USDA entomologist who worked in Mississippi and developed insect organ culture techniques and detailed the anatomy of the hindgut of larval Lepidoptera.

This species occurs with D. latistriga, D. broui, and D. tenuistriga in coastal salt marsh habitats from Alabama to Texas. Doryodes bistrialis may occur nearby on the Gulf Coast, but inhabits inland longleaf pine savannas rather than salt marshes. Doryodes reineckei is immediately distinguished from other species of Doryodes in having a broader forewing in the male, the wing pattern appears faded and smudged, and the ventral margin of the longitudinal stripe appears to blend into the forewing ground color below it, not sharply defined as in other species. The male genitalia of D. reineckei are smaller than those of D. latistriga, but the characters of the vesica are definitive. In D. reineckei the three patches of cornuti on the basal trunk of the vesica are rotated to the left, so the basal one is more lateral than dorsal; diverticulum 1 of the vesica projects dorsally and has a rooster-comb-like cornutus, whereas in D. latistriga and D. broui there is a tapered diverticulum (or rounded lobe) projecting posteriorly with a small spined cornutus on the side, and diverticulum 2 has no cornutus in D. reineckei, whereas in the other two species there is a large cornutus at the base of the diverticulum. Females are immediately distinguishable by their whitish-gray forewing color with the longitudinal stripe is absent, or very faint, compared with the normal pattern seen in other species of Doryodes. Females have acutely pointed forewings.

Forewing length 15.0–17.0 mm (males), 17.0–20.0 mm (females). Male noticeably broader winged than other species of Doryodes. Forewing pale brown to dark gray brown, darker forms in colder months; longitudinal dark stripe paler and less sharply defined than in other species, lower margin of stripe blending into darker ground color below it; wing margin more rounded than in other species. Male genitalia with valve similar in size to that of D. spadaria but aedeagus much shorter (2.29 versus 2.97 mm), about 7.5 × as long as mesial width; basal trunk of the vesica with three patches of cornuti, two ventrolaterally on right side of aedeagus, largest one basal, dorsolaterally on left with multiple spinules on a heavily-sclerotized oval plate. Vesica above basal trunk T-shaped, elongated posteriorly-directed diverticulum 1 with a preapical rooster-comb-like cornutus on anterior surface, a small rounded diverticulum 2 on posterior right side opposite end of aedeagus, a sclerotized plate at 90° ventral angle in vesica next to ductus seminalis, and a ventral rounded apical diverticulum 5 with a conical cornutus on left side. Female genitalia disproportionally small, especially sclerotized plate in ventral wall of ductus bursae only 2.5 × as long as posterior width, plate tapered anteriorly, only 0.55–0.65 × as long as ductus bursae; ductus seminalis broad at base and gradually tapered, so appearing much wider than other species except D. tenuistriga, and, like D. tenuistriga, corpus bursae more rounded than in other species; appendix bursae a rounded lightly-sclerotized lobe with slightly bilobed posterior margin.

The species has been collected from the western panhandle of Florida along the Gulf Coast to eastern Texas. Dates are from April to August but it is likely on the wing throughout the year. Nothing is known of its biology other than its association with Spartina marshes. Adults probably occur throughout the year, but most records are from a large brood occurring between mid-March and late June, with a minor second brood in the late summer and fall (V.A. Brou Jr. pers. comm.).

Syntypes ♂, ♀, Benito, Texas, USNM, examined.

USA: Louisiana, Texas.

Doryodes tenuistriga adults usually can be distinguished from other species in the genus by the narrow forewing stripes. The forewing length varies from 16.5–18.0 mm (males) and 15.5–18.5 mm (females). Some specimens of D. broui can be similar, but the male and female genitalia are diagnostic. In the male genitalia the sclerotized areas on the dorsal and ventral margins of the valve end in broadly rounded processes well before the mainly membranous apex of the valve; in other species the process on the dorsal margin of the valve is pointed and extends to the apex of the valve, or beyond it, and the ventral process extends to, or almost to, the valve apex and is bluntly pointed or narrowly rounded. The aedeagus is short and wide, only about 5 × as long as wide. In the vesica the dorsal and ventral diverticula are similarly elongated, giving the vesica a T-shape; the vesica immediately posterior to the aedeagus has a rounded ventral lobe with a massive spine-covered sclerotized apical plate that is unique in the genus. In the female genitalia the ventral sclerotized plate on the ductus bursae is short and wide, about as long as wide and extending about ½ length of ductus; corpus bursae rounded, with protruding sclerotized lobe on posterior left opposite appendix bursae; appendix bursae rounded, lightly sclerotized.

Doryodes tenuistriga is known only from the Gulf Coast of Texas and Louisiana, occurring as far south as Brownsville, Texas. Its range overlaps those of D. broui, D. reineckei, and D. latistriga, and among these species, it is likely only to be confused with D. broui. The immature stages and larval host plants are unknown. Based on very few records, it appears that D. tenuistriga flies throughout the year but with a primary brood in April and May and a secondary protracted brood in the fall (V.A. Brou Jr. pers. comm.).

Syntype ♂, Nassau, Bahamas, BMNH, examined.

Diagnosis. Doryodes insularia is unique in the genus because of small size (forewing length: 12.5 mm), the white lines bordering the longitudinal dark stripe on the forewing are thicker than those of any other species, there is a contrasting orange-brown band below the forewing costa and another one below the white line bordering the lower margin of the black stripe, and occurrence in the Bahamas. The species is known only from the type series. The type material in the Natural History Museum, London, has not been dissected.

Bahamas. Nothing is known of the biology of D. insularia.