Cunicumara anae St Laurent, 2016.

Hoary appearance caused by interspersion of gray, pale-khaki, and dark-brown scales layered upon salmon to orange-brown, sandy tan ground color. Extremely long bipectinate antennae extending more than half length of the stout forewings, with distinctly long pectinations.

(1) Basally-fused and outwardly projected gnathos with paired distal extensions (Fig. 5a); (2) Phallus with dorsal, curled, horn-like juxtal processes with third, single process located between the curled pair (Fig. 5b).

St Laurent et al. (2018a) did not include this genus in their phylogeny and instead treated this genus as incertae sedis based on morphology, due to similarities with the other incertae sedis genera. The morphological phylogenetic results of the present article support a close relationship between Cunicumara and other incertae sedis genera, particularly with Menevia and Roelofa (Fig. 1, see also Suppl. materials 5–7).

The female of C. anae is unknown, and thus we are unable to figure it or its genitalia.

Cicinnus lantona Schaus, 1905.

Shape somewhat variable, but maculation consistent in presence of white apical dash on the forewing which forms a connection with a white, swooping “postmedial lunule” which may or may not form a continuous white band outlining the postmedial line (St Laurent and Dombroskie 2016).

(1) Paired, elongated, thin, tusk-like extensions reaching outwards from modified transtilla/juxtal complex which itself extends inward into the body cavity from attachments on either side of the inner costal apodemes of the valvae (Fig. 3a).

Perophora olivia Schaus, 1896.

Combination of the following two characters: forewing with black apical streak which forms a connection with a dark colored, straight or concave (or convex in some females) postmedial line and with pair of elongate darkly colored terminal abdominal tufts. Such tufts are seen in other genera but never in combination with the presence of a darkly colored forewing apical streak.

Valvae with basal mesal flap-like extension which may be finger-like or more triangular and spined; valva flap extends into center of genitalia and curves outwards (Fig. 6a).

Hydrias amaryllis Schaus, 1896.

Unmistakable yellow moths with thick gray ante- and postmedial bands spanning width of the wings, gray bands are outlined with cream or white on both sides.

Deeply bifid uncus such that bifurcation deeper than uncus width (Fig. 4a).

Cicinnus enthona Schaus, 1905.

Reinmara can be recognized by the following combination of characters: contrast between medial and submarginal areas due to diffuse pink or silvery-gray scaling delimited by strongly marked, relatively straight postmedial line; notched forewing tornus and anterior margin of hindwing. In all but one species (T. ignea St Laurent, Herbin, & C. Mielke) female and male antennae are strongly dimorphic, being bipectinate in males as is typical of Mimallonidae and dentate in females. Such dimorphism is only also observed in unrelated Roelofa.

Combination of the following characters: (1) Saccular edge of valva curled, with short spine of variable length (Fig. 18a); (2) Bulbous, asymmetrically sized setae-filled diaphragmal sacs (Fig. 18b); (3) Gnathos mesally fused and bifid with fingerlike tips distally (Fig. 18c).

Trogoptera erosa Herrich-Schäffer, [1856].

Similar in overall size and shape to Reinmara, but wings broader, margins more squared, ground coloration more earthen in tone, most species are light khaki brown, some darker. Males and females with bipectinate antennae.

Apomorphies 1 and 2 as for Reinmara; but gnathos not bifid, distally forming single point of various lengths (Fig. 19a).

Although the morphology of Reinmara and Trogoptera are similar, we note that in ongoing molecular phylogenetics of the family, which now includes denser sampling of both genera than were available in St Laurent et al. (2018a), the two genera are quite divergent genetically. Eventually it will be worthwhile to include R. ignea which displays characters found in both genera (Reinmara genitalia but Trogoptera antennae structure) in order to clarify the phylogeny of Trogopterini.

Adalgisa croesa Schaus, 1928.

Three distinct hyaline patches between the following vein pairs of the forewing: Rs2 and Rs3, M3 and CuA1, CuA1 and CuA2. There are smaller hyaline patches between most other wing veins and narrowly along distal margin of discal cell. Similarly located, smaller hyaline patches exist on the hindwing as well.

(1) Phallus very long (longer than length from base of vinculum to tip of valva) and sharp, covered in fine spine-like setae (Fig. 12a), Suppl. material 3: Plate 6, 24: 4.

Cicinnus anoca Schaus, 1905.

Small mimallonids with consistently short, triangular wings, always with a white postmedial lunule on the forewing, though this may be faint in species with accented veins in the broad, darker postmedial region (a character which itself is rather diagnostic of Alheita). Ventrally largely unmarked except for discal spot and significantly darker brown region of forewing delimited by the outline of the dorsal postmedial lunule. The similar genus Tarema is not so cleanly marked ventrally and has less falcate forewings.

(1) Gnathos plate typical of Alheitini, but with a thickly sclerotized mesal band that runs along it vertically (Fig. 13a); (2) Uncus rounded with paired mesal teeth (Fig. 13b).

We were unable to include a female A. anoca in our figures. Female Alheita are generally not largely distinct from males, with only the usual degree of sexual dimorphism (larger size, broader wings) which is usually observed in related genera.

Arianula haxairei Herbin, 2012.

Arianula is the only other mimallonid genus besides Adalgisa which has irregularly scattered hyaline patches on all wings. However, Arianula is easily recognized by the alternate configuration of these hyaline patches, such that there are fewer patches on the forewing than in Adalgisa, with the most apparent patch being between Rs4 and M1 (conversely in Adalgisa, the most apparent patch is between Rs2 and Rs3); and a unique trio of rectangular patches on the hindwing between M3, CuA1, and CuA2, not observed in this arrangement in Adalgisa.

(1) Bizarre claw-like valvae each similar in size to the tegumen + uncus, giving the overall genitalia a triangular appearance (Fig. 17a); (2) Phallus weakly sclerotized, tube-like (Fig. 17b), with elongated narrow juxtal process dorsal to phallus, which itself terminates in a multi-pronged, spined upward curved claw.

Cicinnus fatella Schaus, 1905: 326, by present designation.

The name for this new genus is derived from the type, and only known species belonging to Fatellalla: Fatellalla fatella comb. n. The name is feminine.

This new genus can be recognized by the following combination of characters in the male (the only known sex): triangular forewings with pink antemedial and medial areas, which contrast with the orange-brown submarginal area. The submarginal area is clearly delimited by a pair of thin, straight, preapical, blackish brown postmedial lines. The rounded hindwings display the same patterning as the forewings. The coloration of the body is that of the ground color of the wings: orange-brown, with a distinct darker brown lateral line along either side of the abdomen, as well as a distinct dark brown tuft of elongated scales extending from the terminus of the abdomen. The genitalia are distinct due to the truncated, flattened uncus and the knob-like saccular process at the base of each valva, characters together not observed in any other known Mimallonidae. The gnathos plate is typical of alheitine Lacosominae; and the short, rounded valvae, narrow pair of valva apodeme arms, and thick bunch of elongate setae originating from the diaphragm are similar to those seen in Tarema and Alheita. In Fatellalla, the arms cross over each other mesally and are sharply tipped unlike in these other genera.

(1) Uncus flattened, truncated (Fig. 106I); (2) Sharply tipped valva apodeme arms which cross mesally below uniquely shaped (mesally rounded) gnathos plate (Fig. 106II).

Male.Head: Tan-brown ground color, frons dark brown, eyes very large, occupying more than two-thirds area of head, bordered posteriorly by dark scales; antenna coloration dark tan, antenna almost entirely bipectinate, distalmost 9–10 antennomers dentate; labial palpus three segmented. Thorax: Coloration pinkish orange with red hue (fading to light brown in old specimens), scales covering prothorax grayer, contrasting against lighter remainder of thorax. Legs: Coloration as for thorax, vestiture thick, long. Tibial spurs elongate, covered in scales except for tip, roughly half length of first tarsal segment. Forewing dorsum: Forewing length: 15.5–17.0 mm, avg.: 16.3 mm, wingspan: 29.0–4.5 mm, n = 2. Triangular, margin mostly straight except slightly convex below apex and slight indentation at tornus. Ground color orange brown, but antemedial and medial areas evenly tinted with pink, faint gray suffusion present submarginally. Antemedial line absent; preapical postmedial line formed by two thin, parallel dark brown lines with medial area pink coloration between them. Postmedial lines slightly wavy basally. Pink coloration of medial area does not extend beyond postmedial lines. Discal mark present as pink ovoid region slightly lighter than surrounding medial area. Fringe poorly preserved in examined specimens, but darker brown than submarginal area. Forewing ventrum: Darker brown than forewing dorsum, medial pink suffusion bleeds into submarginal area, postmedial line convex and indented at intersections with veins. Hindwing dorsum: Coloration, patterning as for forewing dorsum, but discal mark absent, submarginal area comparatively wider than medial area. Hindwing ventrum: Following same pattern as forewing ventrum, but lighter overall. Frenulum present as single bristle. Venation: Typical of Mimallonidae. Abdomen: Dorsal coloration as for thorax, laterally very dark brown, nearly black, dark coloration continues on to elongated paintbrush-like tuft of dark-brown scales extending from terminus of abdomen. Vestiture thick, long. Genitalia: (Fig. 106) n= 4. Vinculum ovoid, ventrally rounded. Paired sclerotized arms extend outward from sclerotized base of costal valva apodemes. Uncus simple, truncated distally, rectangular. Gnathos well-sclerotized, forming mesally lobed plate below uncus, gnathos spanning tegumen. Valvae small relative to tegumen + uncus, valvae rounded, hardly extending beyond vinculum-tegumen juncture. Saccular edge of valvae with heavily sclerotized knobs near valval base. Diaphragm with dense setae extending outward above phallus. Juxta fused to phallus, encircling it, lateral margins of juxta extend one quarter length of phallus as triangle directed toward apex of phallus, ventral lip of juxta connects to vinculum. Base of phallus weakly sclerotized with undefined coecum phallus. Phallus cylindrical, downwardly angled distally, vesica spiculate. Female. Unknown

We describe Fatellalla for F. fatella, an Amazonian species widely distributed in Ecuador, the Brazilian Amazon, and French Guiana. This species is exceptionally rare in collections, with less than ten specimens known to us in global collections. Although St Laurent et al. (2018a) did not sample F. fatella, this species clearly belongs to Lacosominae: Alheitini due to the presence of the fused gnathos “plate” which is also observed in alheitine genera Thaelia, Tarema, and Alheita. The genitalia of Fatellalla are most similar to those of Tarema and Alheita (see St Laurent et al. (2017b)), the presence of elongated valvae apodeme extensions basally not unlike those of Tarema, support a close relationship between these genera. Our morphological phylogenetic analyses also support the placement of Fatellalla within Alheitini due to close placement with several other alheitine genera, namely Alheita (Fig. 1, see also Suppl. materials 5–7). Although Fatellalla is nested within a poorly supported Alheita clade, we maintain this genus as valid pending molecular phylogenetic analyses due to the extremely divergent uncus shape and external appearance which is inconsistent with any observed Alheita. We note also a particularly long branch length (relative to other Alheita) in our ML analyses, emphasizing the divergent morphological characters of this odd genus.

Schaus’ (1928) placement of this species in Druentica was unjustified and erroneous.

Cicinnus caudina Schaus, 1905.

Antemedial/ medial areas strongly contrast against the darker, chestnut brown postmedial/submarginal areas, these two distinct regions of color are divided by a pure white, sinuate postmedial line. Trogoptera mana Schaus displays a similar coloration scheme but the forewing has a distinct tornal notch and an anterior hindwing notch typical of Trogopterini, which is absent in Herbinalla. The particularly acute hindwing anal angle, which is accentuated by darker scales at the tips, is largely unique to this genus.

(1) Seemingly floating sclerotized plate apparently homologous with plate-like gnathos of Alheitini, covered in fine setae, situated centrally within the diaphragm, plate not connected to remainder of the genitalia by any sclerotization (Fig. 15b); (2) The only alheitine genus with narrow and downwardly curved uncus (Fig. 15a) (this itself is not an apomorphy due to elongated/narrow uncus in other mimallonid genera, however its presence coupled with other typical alheitine traits such as valva shape, are unique to this genus).

Tarema rivara Schaus, 1896.

Hoary in appearance due to generous amounts of light gray scales present over the entirety of the wings and body. Tarema are similar to Alheita, particularly by the small size of the moths of both genera, and the postmedial lunule. However, the ventral surface of Tarema is not as plainly maculated due to the presence of postmedial lines and the light gray scales.

(1) Uniquely shaped, often spiked projections emanating from the costal valva apodeme (Fig. 14a); (2) Robust gnathos plate not affixed to tegumen along its length as in related genera (Fig. 14b; Suppl. material 3: Plate 3, 11: A); (3) Pair of elongated spines on phallus, one shorter spine present dorsally, the other more elongated extending laterally nearly length of phallus (Fig. 14c).

St Laurent et al. (2018a) did not include this genus in their phylogeny and instead placed this genus in Lacosominae: Alheitini based on morphology, due to strong similarities with the genus Alheita that was included. Our ongoing molecular work, which includes broadened taxon coverage, and Tarema, fully supports this placement (St Laurent et al. in prep.). Furthermore, the morphological analyses carried out in the present study place Tarema nested within Alheitini, either sister to Herbinalla (constrained ML analysis, Fig. 1) or sister to the remainder of Alheitini (unconstrained ML and parsimony analyses, Suppl. materials 5, 7 respectively).

Thaelia linamariae Herbin, 2016.

The coloration of Thaelia species is variable, but wing patterning is consistent. Externally, Thaelia have an apical postmedial line and elongated, falcate forewings, thereby making them appear very distinct from other Alheitini (all other Alheitini genera have preapical postmedial lines and less falcate, shorter, more triangular wings).

See diagnosis and remarks below.

The definition of Thaelia here follows St Laurent et al. (2018a), based on phylogenetic placement of T. linamariae Herbin (type species of Thaelia), T. subrubiginosa (Dognin), and T. anysia (Schaus) within a robustly supported clade (fig. 1 in St Laurent et al. (2018a)). All Thaelia species, sensu this work, were examined morphologically, and included in our morphological phylogenetic analyses, and all form a well-supported clade in all analyses (see Fig. 1, but also Suppl. materials 5 and 7). There are three distinct male genitalia patterns within this genus: (1) gnathos present, valva apodeme extensions absent, and cornuti present (T. linamariae, T. beniensis Herbin); (2) gnathos present, valva apodeme extensions present, cornuti absent (T. anysia); 3) gnathos absent, valva apodeme extensions present, cornuti absent (T. subrubiginosa, T. inornata (Druce)). We do not consider it warranted to further break Thaelia into separate genera at this time considering the consistent external morphology among these species. Ongoing molecular phylogenetics which include denser sampling of Thaelia further support a single genus concept for the species that we place in Thaelia.

Trogoptera rumina Druce, 1894: 355, by present designation.

The name for this new genus is derived from citrus (Latin) referring to the lemon-yellow coloration of the type species of rumina, the only recognized species in the genus. The name is feminine.

This new genus can be recognized by the following combination of characters: bright yellow coloration with gray and pink shading on the tornal region of the forewing and anal angle of the hindwing. The postmedial line of the forewing is faint, crenulate, and incomplete, existing only apically, as a small splotch halfway across the wing, and along the tornal shading. The hindwing displays similar maculation. Ventrally, the antemedial area of the forewing is shaded gray and pink, making Citralla and Zaphanta the only Mimallonidae genera with completely shaded antemedial regions of the ventral surface of the forewings. Citralla, however, lacks the ventral antemedial line present in Zaphanta. The prothoracic tibia has a prominent tuft of pink scales that is seen nowhere else in Mimallonidae. The male genitalia are simple, but unique in the absence of gnathos and transtilla projections, and by the simple triangular uncus and narrow valvae. The phallus is nondescript and largely similar to that of Lacosoma. The female genitalia are most similar to the related Vanenga, but display narrower papillae anales, ostium bursae, and ductus bursae. In Vanenga the confluence of the ostium bursae and ductus bursae is almost as wide as segment VIII, but in Citralla this part of the ductus bursae is only about one quarter the width of VIII, compare Figs 57, 119.

Combination of the following characters: (1) Gnathos reduced to narrow bars below uncus which lack both mesal extensions and subuncus projections typical of related genera (Lacosoma and Vanenga); (2) Simple, smooth, triangular uncus and narrow valvae (relative to sharply triangular or extremely narrow uncus of Lacosoma and Vanenga).

Male.Head: Gray-brown, eyes very large, occupying more than two-thirds area of head, bordered posteriorly by dark scales; antenna coloration light tan, antenna bipectinate to tip, distalmost 10–12 pectinations significantly shorter; labial palpus three segmented, but segments difficult to discern due to compact scaling. Thorax: Coloration light yellow with scales along posterior prothoracic margin and junction with abdomen very faint pink and gray, ventrum pale gray. Legs: Coloration variable, prothoracic leg: femur light purple-gray, tibia yellow with light gray scales before juncture with tarsus, prominent tuft of pink scales present on inner margin of tibia apex, tarsus yellow. Mesothoracic leg: femur and tibia light gray, tarsus yellow with some gray scaling apically. Metathoracic leg: all segments predominantly yellow with some gray scaling at terminus of tibia and apex of tarsus. Tibial spurs elongate, narrow, dorsally covered in scales, ventral surface and tip naked, length roughly half length of first tarsal segment. Forewing dorsum: Forewing length: 9–14 mm, avg: 11 mm, wingspan: 19–27 mm, n = 16. Triangular, margin nearly straight. Ground color light yellow. Antemedial faint, pink, irregular, antemedial area may be slightly suffused with pink; preapical postmedial line irregular, incomplete, existing only near tornus, apex, and halfway across length of wing as single splotch. Postmedial line outwardly shaded with gray and pink, particularly along tornus where pink suffusion reaches wing margin. Discal mark present as light gray ovoid splotch. Fringe checkered off-white and orange-brown, slightly crenulate. Forewing ventrum: Nearly identical to forewing dorsum, but antemedial line absent and antemedial area completely shaded by pink and gray, discal spot more pronounced. Hindwing dorsum: Coloration, patterning as for forewing dorsum, but discal mark faint or absent, antemedial line absent. Hindwing ventrum: Following same pattern as forewing ventrum. Frenulum present as single bristle. Venation: Typical of Mimallonidae. Abdomen: Dorsal coloration as for thorax, but slightly darker. Genitalia: (Fig. 118) n = 3. Vinculum ovoid, ventrally inwardly notched. Uncus simple, triangular, ventrally membranous. Gnathos and transtilla absent, but sclerotized bars extend downward from uncus/tegumen junction. Valvae narrow, triangular, simple; base of valvae extend centrally above vinculum base such that valvae cannot be fully spread. Juxta fused to phallus, encircling it, extending dorsally above phallus as flattened process. Phallus cylindrical, basally truncated. Female.Head: As for male, but antennae smaller overall. Thorax, Legs: As for male. Forewing dorsum: Forewing length: 11.5–15.0 mm, avg: 13.8, wingspan: 26–30 mm, n = 7. As for male, but slightly broader overall. Forewing ventrum: As for male, but slightly broader overall. Hindwing dorsum: As for male, but slightly broader overall. Hindwing ventrum: Following same pattern as forewing ventrum. Frenulum as multiple bristles. Abdomen: As for male, but more robust. Genitalia: (Fig. 119) n = 1. Tergite VIII forms smooth, thickened posteriorly directed arch, mesally with cup-like indentation at dorsal base of papillae anales. VIII weakly sclerotized laterally. Apophyses anteriores thick, truncated distally, roughly half length of apophyses posteriores which are outwardly bent halfway along length. Lamella ante- and postvaginalis poorly preserved, but weakly sclerotized without distinguishing features. Ductus bursae long, narrow, about three times the length of VIII-X, ductus widest at convergence with ostium bursae, but remaining very narrow along remainder of length. Corpus bursae small in length in comparison to elongated ductus bursae and large papillae anales, shape balloon-like. Papillae anales narrow, elongated, ventrally angled such that apical ridges of papillae anales and opening between them situated ventrally in nearly same plane as ostium bursae.

Citralla is here described for the unique species C. rumina comb. n., which is distributed from Guatemala to southeastern Brazil. Further taxonomic investigations into the various populations of C. rumina will undoubtedly reveal several cryptic species, as we have observed slight external morphological distinctions in populations in Southeastern Brazil and the Amazon, in comparison with topotypical material from Panama and nearby Costa Rica.

Ongoing molecular phylogenetic work which includes Citralla consistently places this genus sister to Vanenga, which together form a clade sister to Lacosoma (St Laurent in prep.). Morphological analyses are less consistent in placement, with our morphological analyses recovering Citralla sister to (unconstrained ML analysis) or nested within Lacosoma (constrained ML and MP analyses). Regardless, tribal placement of the new genus is confidently in Lacosomini. The substantial reduction of the gnathos and juxtal configuration are most similar to those of both Lacosoma and Vanenga than to any other know Mimallonidae genus. But the uncus shape and pink scale tufts on the forelegs are unique to C. rumina. The larvae of C. rumina (Fig. 120) have been reared and photographed by Janzen and Hallwachs (2017), revealing morphology remarkably similar to various species of Lacosoma, two species of which have been reared by the first author (St Laurent and Carvalho 2017b, St Laurent et al. 2017). In particular, the striated appearance of the head and prothoracic shield, as well as the less rugose anal plate are largely consistent with Lacosoma as opposed to the more uniform coloration and generally more robust anal shield of other groups such as Mimalloninae and Cicinninae. We therefore, consider these characters as additional information supporting our decision to place Citralla within the Lacosominae: Lacosomini.

According to Janzen and Hallwachs (2017), the host plant of C. rumina in Costa Rica is Eugenia salamensis Donn. Smith (Myrtaceae). Myrtaceae is a host plant family that is frequently a larval resource for Mimallonidae (St Laurent et al. 2018).

Lacosoma chiridota Grote, 1864.

Lacosoma is one of the most speciose genera of Mimallonidae, and displays a great deal of variation. They can easily be recognized by their genitalia (see Apomorphies below), but are externally more variable. Externally, Lacosoma are some of the smallest Mimallonidae in overall size, most species have crenulated wing margins and pink, salmon, and gray coloration. The general shape and size of Lacosoma species is more diagnostic than any one other external character. The combination of the general characters of small size, falcate forewings, and crenulated margins allow for the recognition of most species in the genus. Few Central American species display straight wing margins (such as L. elassa (Franclemont) and L. morgani Herbin), but are of the usual size and coloration for Lacosoma in general. The more uniquely colored species display the typical falcate wing shape and crenulate wing margins. Examples of uniquely colored species include those that are all pink: L. maldera Schaus; nearly all black: L. syrinx (Druce) and L. briasia Schaus (Fig. 163); and pink and yellow: L. valera Schaus and L. valeroides Herbin and C. Mielke.

(1) Subuncus projections extend from ventral margins of tegumen, extended outward most usually as a simply triangular protrusion (Fig. 20a), but in some Central American species (for example L. elassa, L. morgani) these projections are more elongated and bent; (2) Gnathos projected ventrally below uncus, connecting to complex dorsal projections of juxta, together forming intricate sclerotizations with variable projections (Fig. 20b; Suppl. material 3: Plate 1, 6: 0, 6: A).

Perophora mera Dognin, 1924.

Small mimallonids with short triangular forewings with distinct straight postmedial line, ground coloration pale tan-orange with pink coloration throughout the antemedial and medial areas.

(1) Uncus as long (roughly length of valva), narrow spine-like in shape, not obviously triangular as in essentially all other mimallonid genera (Fig. 21a); (2) Subuncus projections similar in shape to those of Lacosoma, but originating from downward lateral projections of tegumen extending well below costal margin of valva (Fig. 21b), as opposed to extending directly outward from ventral margin of tegumen above or very near costal valva margin as in Lacosoma (se Fig. 20a).

Cicinnus partha Schaus, 1905.

Most Druentica are silvery gray medium sized mimallonids with a distinct, usually straight postmedial line, and a preapical black dot along the costa where the postmedial line angles towards the costa, meeting it. Most species have straight forewing edges, but a few (such as D. rotundula (Dognin), D. muta (Dognin), D. mutara (Schaus), and D. brosica (Schaus)) have crenulated wing margins not unlike some Lacosoma and Mimallo, hence erroneous placement of several species of Druentica in these unrelated genera. Druentica is another of the larger mimallonid genera, and thus there is more interspecific variation in this genus than in other genera previously treated above. However, male genitalia of Druentica are immediately recognizable as such.

Combination of the following characters: (1) Slightly curved tusk-like arms extending from valva base (Fig. 22a); (2) Paired, mesally separated, elongated gnathos arms with distal flattening extend from immediately below uncus, gnathos arms roughly equal to length of uncus (Fig. 22b).

Cicinnus montagnaniae Herbin, 2012: 14, by present designation.

The name for this new genus is derived from the small, silvery coloration of the sole Lepismalla species, reminiscent of silvery Zygentoma in the genus Lepisma Linneaus. The ending –alla/-allo has been commonly applied in Mimallonidae. The genus name is feminine.

The single species of Lepismalla, L. montagnaniae comb. n., is recognizable by the almost complete lack of markings (except for faint irregular postmedial lines and heavy black discal marking) on the dorsal surface of the wings, which combined with the small size and falcate wing shape is unique in the family Mimallonidae. Genitalia are robust structures, with prominent gnathos arms that extend distally below the saccular margin of the valvae. In this way the gnathos arms are somewhat similar to those of related genera Procinnus Herbin and Micrallo St Laurent and C. Mielke, but extend much farther ventrally relative to the valvae. The basal valva arms typical of Druenticini are present, but highly reduced, and are flanked by setae covered regions of the sacculus. This genus lacks the claw-like sternite VIII extensions in the male genitalia which are typical of the related genus Procinnus, and has much shorter valvae tusks than either Procinnus or Micrallo.

(1) Large gnathos arms extend ventrally below saccular edge of valvae such that valvae appear dorsally to gnathos (Fig. 131I); (2) Base of valvae with tiny tusk-like arms (smaller than in any other Druenticini) (Fig. 131II).

Male.Head: Light brown, eyes very large, occupying more than two-thirds area of head; antenna coloration light tan, antenna entirely bipectinate, distalmost quarter length of antennae with pectinations markedly shorter; labial palpus apparently two segmented, though a third segment may be present but small. Thorax: Light brown, lightly speckled with darker petiolate scales. Legs: Coloration as for thorax, vestiture thick, long. Tibial spurs elongate. Forewing dorsum: Forewing length: 14.5–16.0 mm, avg.: 14.6 mm, wingspan: 28–35 mm, n = 6. Triangular, margin concave mesally forming falcate apex. Ground color silvery gray and light brown, overall lightly flecked with dark brown petiolate scales. Antemedial line absent; preapical postmedial line faint, outwardly convex, consisting of numerous barely distinguishable individual dark brown petiolate scales. Discal mark present as irregular black splotch heavily contrasting against light ground color and otherwise largely unmarked surface. Fringe light brown, lightening to cream near tornus. Forewing ventrum: As for forewing dorsum, but more brown than gray, anal region light tan, apex lighter gray than surrounding brown area; postmedial line slightly more well-defined; discal mark fainter than for dorsum. Hindwing dorsum: Coloration, patterning as for forewing dorsum, but discal mark much fainter to nearly absent. Fringe nearly white along most of wing margin. Hindwing ventrum: Following same pattern as forewing ventrum, but lighter overall, more of a continuation of light tan of anal area of forewing ventrum. Frenulum as single bristle, though difficult to see. Venation: Typical of Mimallonidae. Abdomen: Dorsal coloration as for thorax, ventrally lighter gray. Vestiture appearing thinner in comparison with thorax. Sternite of VIII with narrow pair of sclerotizations. Genitalia: (Fig. 131) n = 2. Vinculum ovoid, ventrally slightly pointed. Uncus simple, broad, triangular, distally narrowed to thin point. Gnathos massive, the most distinct aspect of the genitalia, gnathos heavily sclerotized with paired mesal arms which extend beyond and below saccular edge of valvae, gnathos arms distally and inwardly membranous, small tooth present at apex of each arm, gnathos arms fused mesally at base by narrow sclerotized band. Valvae small relative to gnathos, somewhat rectangular in shape, terminally with slight saccular lobe. Pair of small (length less than one third length of valvae), tusk-like curled sclerotized arms extend outward from saccular edge of valvae, tusk-like arms inwardly flanked by protruding setae covered sacculus. Juxta as thin sclerotization ventral to phallus forming connection with vinculum. Phallus curved when viewed laterally, viewed dorsally/ventrally phallus flattened distally with membranous opening such that phallus appears spade-like with sclerotized ring forming margins of spade shape, basal half of phallus as membranous sack. Female. Unknown

Lepismalla is described for the unique Amazonian taxon L. montagnaniae, which is so far outwardly and by male genitalia, unlike any other Mimallonidae, though is most similar to related genera Procinnus and Micrallo. In the original description, it was suggested that “Cicinnus” montagnaniae might be more properly placed in a new genus (Herbin 2012). After examining genitalia of all genera in prior to and in preparation of the present work, it is clear the genitalia of L. montagnaniae do not conform to any other generic concept of Mimallonidae, and certainly this species is unique externally as well. However, the subfamily and tribal apomorphies provided by St Laurent et al. (2018a) for Druenticini are consistent with the general male genitalia characters seen in L. montagnaniae. Our morphological and ongoing molecular phylogenetic analyses also consistently place this taxon as sister to the druenticine genus Procinnus, within a robustly supported Druenticini clade (St Laurent et al. in prep.).

The shape of the phallus of Lepismalla is very similar to that of other druenticine genera Micrallo and Procinnus. The phallus in all three genera is flattened with a unsclerotized central region in the distal half, which is clearly visible from the dorsal or ventral aspect. These genera also display a membranous sack-like region along the basal dorsal half of the phallus, though the size of this structure differs between the genera. Paired sclerotizations of the VIII sternite typical of Druenticinae, including Micrallo and Procinnus, are present in Lepismalla, but are reduced to narrow sclerotized strips in the otherwise membranous intersegmental region. The silvery gray coloration of Lepismalla is a coloration scheme seen in almost all Druenticini genera (except the darkly colored Ulaluma and some Procinnus), and the gnathos configuration is not unlike that of Procinnus and Micrallo, but simply more robust and ventrally farther reaching, such that in Lepismalla the gnathos arms actually reach below the saccular edge of the valvae. The basal valvae arms typical of Druenticini are present in Lepismalla, albeit greatly reduced.

Lepismalla montagnaniae is rare in collections, with only a handful of specimens known to us in global collections. Almost all examined material comes from the Amazon Rainforest, although one specimen, unfortunately destroyed in the fire at the Museu Nacional, Rio de Janeiro, Brazil, was from Mato Grosso do Sul, Brazil, in quite a different habitat on the border of Cerrado and Pantanal. An additional specimen in CPAC is from Distrito Federal in the Brazilian Cerrado. Future, finer scale, examinations of all known Lepismalla specimens may eventually reveal that this is not a monotypic genus, though it is certainly not diverse.

Micrallo minutus St Laurent & C. Mielke, 2016.

Tiny to small (forewing length: 11.5–15 mm) silver gray mimallonids, with elongate triangular forewings, and well-defined, straight forewing postmedial lines that are perpendicularly angled toward the costa, immediately separates this genus from others in the family.

(1) Deeply curled, concave tegumen + vinculum shaped like a broad cesta of Jai alai in the lateral aspect; (2) Basally membranous, distally narrowed, terminally clubbed valvae (Fig. 23a); (3) Membranous pockets encase valvae, contain thick, elongate deciduous setae (Fig. 23b).

St Laurent et al. (2018a) did not include this genus in their phylogeny and instead placed this genus in Druenticinae: Druenticini based on morphology, due to strong genitalia similarities with the sampled genus Procinnus, particularly the flattened, spade-like phallus, which we now also observe in the newly described Lepismalla. All morphological phylogenetic analyses carried out here also place Micrallo within Druenticini, either sister to Ulaluma (parsimony analysis) or to Procinnus + Lepismalla (both ML analyses), thus confirming the tribal placement of St Laurent et al. (2018a).

An undescribed species that likely belongs to this genus is larger and browner than M. minutus, but otherwise bears the same arrangements of wing markings and genital configuration.

Pamea albistriga Walker, 1855.

Quite similar to some of the smaller Andean Druentica species, but instead consistently have white outlined postmedial lines and white suffusions submarginally which contrast against a pale gray ground color finely stippled with darker petiolate scales. Pamea lack the black costal dot of Druentica.

In general, similar to Druentica but (1) Gnathos arms smaller, flatter distally, and very weakly sclerotized, almost membranous at their termini (Fig. 25a); (2) Basal valvae arms coiled (forming loop) (Fig. 25b), arms never simply curved as in Druentica (e.g., Fig. 22a).

St Laurent et al. (2018a) did not include this genus in their phylogeny and instead placed this genus in Druenticinae: Druenticini based on morphology, due to clear similarities to the genus Druentica (particularly the gnathos shape), Druentica was included by these authors. As was the case with the two previously treated Druenticini genera not formally included in any molecular phylogenetic analyses, the male genitalia morphology of Pamea allow confident placement of this genus within Druenticini. Our morphological phylogenetic analyses confirm this placement, consistently placing Pamea sister to the remainder of Druenticini (both ML analyses). Our ongoing molecular work which includes Pamea, also support this placement (St Laurent et al. in prep.).

Procinnus cahureli Herbin, 2016.

Narrow, elongate, sharply acutely triangular wings; coloration varies from silver-gray to brown, but wing shape along with lack of hyaline patches on all wings, and presence of well-defined discal marking unify this small genus.

(1) Pair of elongated, sharp projections extend from either side of VIII sternite, which together with the basal valva arms typical of Druenticini and low-set gnathos arms, give the lower half of the genitalia an overall appearance of possessing a massive multi-fingered claw (Fig. 26a).

We note that one undescribed species is known to the first author to belong to this genus based on clear male genitalia characters, but has significantly stouter wings than other Procinnus and a preapical postmedial line. The coloration and discal markings of this undescribed taxon are however, typical of this genus. Also in this and another undescribed Procinnus species known to the first author, there are fewer sternite VIII arms, but those that are present are the same shape and size as in other described taxa. This combined with the valvae and gnathos shapes are not unlike those of more typical Procinnus.

Cicinnus valva Schaus, 1905.

Small (average forewing length = 14.1 mm) blackish purple ground color, with yellow postmedial lines on all wings. The forewing margin is sharply pointed mesally at the tip of CuA1.

The combined absence of the gnathos and the presence of valvae tusks (Fig. 24).

Perophora penia Dognin, 1919.

Yellow-brown, or in fresh specimens, more golden tan, with distinctly brown antemedial and postmedial lines, and veins accented by the same brown coloration, all of which strongly contrast against the lighter ground color.

Combination of the following characters: (1) Rectangular valvae, distally concave along margin (Fig. 27a), (2) Parallel setae-covered lobes projecting from the valva apodeme (Fig. 27b) (considered “transtilla” in St Laurent (2016)); (3) Gnathos absent; (4) Thin, fishhook-shaped phallus (Fig. 27c).

Cicinnus rotunda Dognin, 1916.

Unique, blueish (even somewhat iridescent in fresh specimens) black mimallonids usually with crenulated wing margins, broad wings; comparatively large antennae with some of the longest pectinations in Mimallonidae (this trait is similar to Cunicumara). The toothed postmedial lines are inwardly lined with pale brown which contrasts against the dark ground color.

Similar to Lurama, but the combination of the following characters distinguishes Ulmara: (1) gnathos present (Fig. 28a) and (2) sternite VIII with bilobed feet-like sclerotizations covered in elongate setae (Fig. 28b).

Cicinnus althea Schaus, 1905.

All three species of Biterolfa are similar, primarily differentiated by genitalia characters. Externally Biterolfa are brown to red-brown with a dark gray-brown postmedial line outwardly margined with a thin gray submarginal region which narrows from the tornus to the falcate apex. Distinctive, parallel, white outlined dark streaks span across the middle of the forewing towards the costa.

Combination of following characters: (1) Broad, stout, simple valvae without any spines, protrusions, or arms (Fig. 29a); (2) Juxta with paired knob-like lobes situated laterally on either side of phallus (Fig. 29b); (3) Prominent gnathos, mesally unsclerotized, with pair of robust, thickly sclerotized arms (Fig. 29c); ventrum of arms may be toothed, terminus of gnathos arms distinctly narrowed with finger-like process at tip of each arm, often with secondary dorsal apical protrusion from tip of arm.

See remarks of Psychocampa below.

Psychocampa concolor Grote & Robinson, 1867.

Externally Psychocampa display typical cicinnine shape and large size, and always lack hyaline patches, however it is difficult to generalize about this relatively large genus beyond that. Despite external variability across the genus, Psychocampa genitalia are quite homogenous, and therefore species belonging to this genus are easily recognizable by their genitalia, which are dramatically distinct and simple in structure in comparison with all other Cicinninae genera (except for the sister genus Biterolfa, see below).

Largely as for Biterolfa but gnathos arms more gradually narrowed, never with teeth or dorsal apical protrusions (in addition to the tapering tip).

This is one of the largest Mimallonidae genera in terms of number of species, particularly after the present checklist and the work of St Laurent et al. (2018a) which cumulatively transferred many species from Cicinnus to Psychocampa based on the exceptionally homogenous and readily diagnostic genitalia of the latter genus. As stated above, it is difficult to diagnose Psychocampa based on external morphology because there are several clear species-groups within the genus which may eventually prove to be more appropriately placed in new genera that can more succinctly be unified by external morphology (though genitalia morphology is so consistent across all of these groups that additional atomization of Psychocampa into several genera may prove to be unwarranted). Genitalia morphology is largely homogenous across all Psychocampini, not only in Psychocampa, and this is clearly evident in comparing the genitalia of sister genera Psychocampa and Biterolfa, see Figs 29 and 30. However, external maculation and convincing phylogenetic results of St Laurent et al. (2018a) placing Biterolfa as sister to the larger clade of Psychocampa, which was intentionally sampled to include most of the clear species-groups of the genus, supports their valid separation as two distinct genera. Further ongoing molecular phylogenetics which continues to sample Psychocampini more densely also continues to support the separation of these two genera. Our morphological phylogenetic results here fully support the transfer of various species from Cicinnus to Psychocampa, as well as the close relationship between Psychocampa and Biterolfa, though with our increased taxon sampling, the validity of Biterolfa is called into question in a morphology-only context because it is nested within Psychocampa in unconstrained ML and parsimony morphological phylogenetic analyses. Despite this, we maintain Biterolfa and Psychocampa as valid separate genera considering the molecular results of St Laurent et al. (2018a) and the previously mentioned ongoing molecular work.

Cicinnus fraterna Schaus, 1905.

Bedosia is a rather diverse genus in coloration, shape, and size, but species belonging to this genus always have a B-shaped forewing hyaline patch (as is common in Cicinninae) and well defined postmedial lines on all wings. Most species have brown accented veins and triangular wings with straight margins or those that are slightly convex. Male genitalia, however, clearly unify this genus and distinguish Bedosia from the externally very similar Bedosiallo.

Combination of the following characters: (1) Valvae situated on dorsal half of vinculum (Fig. 31a) immediately below uncus such that ventral portion of vinculum extends downward as a well-defined projection (Fig. 31d); (2) Heavily concentrated paired set of setae emanate from diaphragm (Fig. 31b); (3) Gnathos arms massive and extremely robust, variable in shape but always arm-like and with species-specific modified tips, extended outward more so than upward (Fig. 31c).

Cicinnus forbesi Schaus, 1928.

Much like Bedosia in external shape and key features, but usually with much narrower wings; the genitalia, however, are very different from those of Bedosia, and are diagnostic of the genus (see apomorphies below).

Combination of following characters: (1) Small, heavily sclerotized gnathos situated very near uncus with mesal pair of closely parallel arms (distally fused in some species) (Fig. 32a); (2) Simple, narrow widely splayed valvae relative to widened ovoid vinculum (Fig. 32b).

Bedosiallo was found to be a strongly supported clade in St Laurent et al. (2018a), where two species of this genus that apparently represented two distinct species-groups within Bedosiallo were sampled and found sister to one another, which together were sister to Bedosia fraterna, type specie Bedosia. Although sampling of Bedosia was relatively poor in St Laurent et al. (2018a), we have since examined the genitalia of nearly all Bedosia species (including those erroneously placed in Cicinnus by Schaus (1928)), finding consistent complex male genitalia, which are readily divergent from the much simpler genitalia of Bedosiallo. In the description of Bedosiallo, all described species were dissected and their genitalia figured, displaying the consistent genitalia in that genus, as well as the consistent difference from all known Bedosia species (St Laurent et al. 2018a). Below in the checklist we transfer two additional species to Bedosiallo from Cicinnus: B. minimalis Herbin and C. Mielke, comb. n. and B. gentilis (Schaus), comb. n. These two species display genitalia much more in line with those of the described Bedosiallo species than to any Bedosia species, but do differ in some respects, particularly in the more closely fused gnathos, triangular valvae shape, juxtal configuration, and presence of cornuti (see Annotations in Section 4). Bedosiallo minimalis was included in our morphological phylogenetic analyses, and was placed within Bedosiallo in all analyses (Suppl. materials 5, 7) except the constrained ML analysis (Fig. 1). We attribute the somewhat unique genitalia of B. minimalis to result in this conflicting placement, and while B. gentilis and B. minimalis may eventually prove to belong to an entirely new genus considering such unique traits, we are confident that these small moths are more closely related to Bedosiallo than Bedosia in comparing the genitalia of all species in Bedosiini, and certainly do not belong in Cicinnus as they were originally placed.

Aceclostria mus Vuillot, 1893.

Silvery-gray to gray brown ground color, sinuate, diffuse postmedial lines, falcate forewings, and broad darker gray to dark gray brown discal marks that vary from fully scaled to fully hyaline within species (often from the same series); maculation overall is dark and diffuse giving the wings a soiled appearance.

Asymmetric, complex genitalia with phallus situated on left side (when viewed ventrally) of vinculum (Fig. 33a) with single vinculum tusk reaching out above phallus.

St Laurent et al. (2018a) did not include this genus in their phylogeny and instead placed this genus in Cicinninae: Cicinnini based on morphology, namely the presence of all apomorphies of Cicinninae and Cicinnini in what was at the time the single Aceclostria species, A. mus. Our ongoing molecular work, which includes broadened taxon coverage, and Aceclostria, fully supports this placement (St Laurent et al. in prep.). The morphological phylogenetic analyses carried out here support this placement as well, with Aceclostria always recovered within Cicinnini, sister to (unconstrained ML and parsimony analysis, Suppl. materials 5 and 7 respectively) or nested within (constrained ML analysis, Fig. 1) the broader, so far poorly resolved Cicinnussensu lato clade.

See annotations in Section 4 for information pertaining to the novel inclusion of A. cordubensis comb. n. and A. nigrescens comb. n. et stat. rev. in Aceclostria. Aceclostria was long considered a monotypic genus, but morphological examinations, including the genitalia of A. cordubensis, conclusively support these additional poorly known taxa in this genus. This genus is apparently closely associated with Anacardiaceae (see Annotations).

The Cicinnus s.l. clade is discussed in more detail below as it is the most poorly resolved set of taxa in the family.

Cicinnus accipiter Dognin, 1916.

Wings are exceptionally narrow and elongate with thin hyaline patch streaks on all wings (hyaline streak mesally kinked on hindwing, lunate on forewing). The antemedial line forms a convex semicircle, whereas the postmedial line is incomplete and mostly limited to the proximal margin of the forewing. The ante- and postmedial lines are curved toward each other or are tangent below discal cell.

Combination of the following characters: (1) Valva with saccular curl/fold which holds robust vincular tusks (Fig. 34a); (2) Phallus fused with proximally bent juxtal structure below (Fig. 34b).

Arcinnus hoedli Herbin, 2016.

Externally not particularly distinct from several other species of Cicinnus. Arcinnus are generally smaller than most Cicinnus and always lack hyaline patches on the hindwings (similarly patterned Cicinnus have hyaline patches on both fore and hindwings). Genitalia were the primary means by which this genus was originally described, and we also consider the male genitalia to have the most robust diagnostic characters defining this genus.

(1) Uncus vestigial and not synscleritous with tegumen (Fig. 37a); (2) Most substantial component of genitalia the broad, curved vincular arms which are apparently homologous to those seen in most Cicinninae, albeit much more heavily sclerotized, thicker, and apically blunt (Fig. 37b).

St Laurent et al. (2018a) recovered Arcinnus as a valid genus sister to Euphaneta + Aleyda. In the present study, Arcinnus is recovered nested within the poorly resolved Cicinnussensu lato clade. We hesitate to synonymize this genus at this time pending broader molecular sampling of the Cicinnus s.l. clade, which is ongoing and consistently supports the validity of Arcinnus.

We were unable to figure female genitalia of Arcinnus, but direct the reader to figs 54 and 55 in Herbin (2016), where the female genitalia of the type species, A. hoedli are presented.

Cicinnus orthane Blanchard, 1852.

See below in the checklist for specific information pertaining to different species-groups within Cicinnus, but in general Cicinnus can be recognized by their falcate forewings, moderate to relatively large size compared to most other Mimallonidae genera. Small, B-shaped hyaline patches, when present, are found on all four wings or forewings only. Species without hyaline patches are usually heavily marked with irregular blotches and speckles.

These apomorphies refer to those of Cicinnussensu stricto in St Laurent et al. (2018c) and do not apply to Cicinnus s. l.: (1) Vincular arms may (e.g., C. melsheimeri (Harris) or may not (e.g., C. orthane Blanchard) be present; (2) Valvae mostly membranous, mesally more well-sclerotized, often with distinct “clasper” (Fig. 36a).

Annotations following the checklist are provided for each group of Cicinnus as we currently define them: Cicinnus s. s. (Cicinnus Group 1) and Cicinnus s. l. (which includes Cicinnus s. l. Groups 2, 3, and some species of uncertain placement). The annotations include diagnostic characters which can be treated as apomorphic of these preliminary groupings of similar species, and may eventually be used to define formal genera (e.g., the available name Gonogramma Boisduval, see Annotation 98, may eventually be resurrected to include the species treated here as Cicinnus s. l.). The morphological phylogenetic analyses do not fully resolve the relationships of species within Cicinnus, with relatively low support values for the various groups within the genus. However, Cicinnus s. l. is consistently recovered within a broader Cicinnini clade, thus tribal placement of the included species is not in question. Additional molecular sampling of Cicinnus and related genera in order to better define the clades within Cicinnus s. l. will be a focus of our future molecular work.

Phaneta divisa Walker, 1855.

This genus is similar to Aleyda, but Euphaneta are larger, have broader, more ovoid wings, lunate hyaline patches on all wings, and lack clearly distinct antemedial and postmedial lines, but rather have a dark brown antemedial area which merges with a brown costal region that extends along the length of the wing. Euphaneta are cryptically colored, apparently camouflaged resembling a woody stem.

Hood-like juxtal process (Fig. 35a) dorsal to weakly sclerotized phallus which is shorter than the juxtal hood above it; this juxtal process is a large, substantial structure comprising almost entire diaphragmal area.

We were unable to include the female of E. divisa, however sexual dimorphism in Euphaneta is not overly significant, with females only being larger and with broader wings than the males.

Perophora ventana Dognin, 1897.

Isoscella can be recognized by the following combination of characters: narrow, triangular wings with single ovoid (or more circular in some species) discal hyaline patch. The similar genus Roelmana always has broader, less elongated wings, and may lack hyaline patches entirely.

(1) Gnathos rectangular with mesal pair of thin, parallel arms equal in length to lateral bars of gnathos (Fig. 38a); (2) Vincular arms thick, not thin and tusk-like, in most species arms dorsally covered in setae (Fig. 38b); (3) Shorter, secondary pair of setae covered plate-like vincular arms located on either side of phallus (Fig. 38c).

In our morphological phylogenetic analyses Isoscella and the related Roelmana below, were poorly differentiated phylogenetically due to strong morphological similarity between these two genera. However, Roelmana, which displays more interspecific variation than within Isoscella (see below) was strongly supported as a monophyletic group in St Laurent et al. (2018a), as well as in ongoing molecular phylogenetics that continues to sample more Roelmana species. These molecular phylogenetic studies include the several morphologically distinct species-groups of Roelmana, which formed a clade sister to Isoscella. The most morphologically divergent Isoscella species, I. andina St Laurent and Carvalho, was not sequenced, but was included in our morphological analyses. This unique species is nested within Isoscella in all analyses except the constrained ML analysis (Fig. 1), rendering Isoscella paraphyletic in that analysis. We consider the placement of this taxon as sister to all Cicinnini in this analysis anomalous considering the presence of all generic Isoscella apomorphies in I. andina. The observed placement in Fig. 1 may be due to certain aspects of patterning (e.g., complete, straight ventral forewing postmedial lines in I. andina) which are unique to this species, whereas genitalia are typical of Isoscella. Therefore, based on the unconstrained analysis, we consider Isoscella to represent a monophyletic lineage, whose relationship to species of Roelmana will likely be more clearly elucidated with denser taxon sampling in our future molecular work.

Cicinnus maloba Schaus, 1905.

Wing shape of Roelmana is typical of Cicinnini, but much less variable overall within this genus than in other cicinnine genera. Forewings are sharply falcate, with the anterior margin of the forewing concave, particularly along the tornus. Two distinct wing morphological groups are found in this genus: one that has a hyaline patch on each wing (such as type species R. maloba (Schaus) and R. pluridiscata (Schaus)), whereas the other group entirely lacks hyaline patches and are instead diffusely colored with faint patterning (such as R. laguerrei (Herbin) and R. beneluzi (Herbin), comb. n.). However, genitalia are mostly homogenous in this genus and provide clear generic placement for these two otherwise outwardly distinct groups.

Similar to Isoscella but (1) Proximal component of gnathos more heavily sclerotized, more rounded than rectangular, with mesal arms more robust and thicker (Fig. 39a); (2) Primary (the longest) vincular arms sharp and tusk-like, not covered in setae (Fig. 39b).

One species, R. beneluzi, displays genitalia that are distinct from other Roelmana, and our transfer of this species from Cicinnus to Roelmana is preliminary. See Annotation 126 for further discussion.