Monograph |
Corresponding author: Ryan A. St Laurent ( rstlaurent@flmnh.ufl.edu ) Academic editor: Andrew Mitchell
© 2019 Ryan A. St Laurent, Akito Y. Kawahara.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
St Laurent RA, Kawahara AY (2019) Reclassification of the Sack-bearer Moths (Lepidoptera, Mimallonoidea, Mimallonidae). ZooKeys 815: 1-114. https://doi.org/10.3897/zookeys.815.27335
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A backbone molecular phylogeny of Mimallonidae, based on 47 species and 515 loci, was recently published. That study resolved some of the major relationships in the family, but taxon sampling was limited and a classification of the family was not formally presented for all species. Here morphological phylogenetic analyses in parsimony and maximum likelihood (ML) frameworks were conducted that included 192 species and 55 morphological characters. A phylogenetic analysis was also conducted on the morphological dataset with a topological constraint based on the 515 locus tree from the previous study. Results show that nearly all species can be confidently placed in a genus using morphological phylogenetics. The presence of a frenulum, a character that was historically used to distinguish major groups of Mimallonidae, varies within and among genera. Based on our phylogenetic results, the classification of Mimallonidae is revised, which now includes 291 species in 41 genera. Descriptions of three new genera are included: Fatellalla gen. n., Citralla gen. n., and Lepismalla gen. n. The following taxonomic changes were made in the present article: 43 new/revived combinations (in Aceclostria Vuillot, Arcinnus Herbin, Bedosia Schaus, Bedosiallo St Laurent & Kawahara, Cicinnus Blanchard, Citralla, Druentica Strand, Fatellalla, Lacosoma Grote, Lepismalla, Mimallo Hübner, Procinnus Herbin, Psychocampa Grote, Roelmana Schaus, and Thaelia Herbin), two new species-level synonyms (melini Bryk is synonymized with viemanda Schaus, jaruga Jones is synonymized with hamata Walker), one revived synonymy (roscida Dognin is resynonymized with externa Moore), seven new statuses (in Druentica, Macessoga Schaus, and Trogoptera Herrich-Schäffer), six revived statuses (in Aceclostria, Cicinnus, Druentica, Psychocampa, and Zaphanta Dyar), and one new designation of nomen nudum. In order to alleviate nomenclatural problems, twelve lectotypes are designated (for Tolypida amaryllis (Schaus), Trogoptera althora Schaus, Adalgisa croesa Schaus, Alheita pulloides (Dognin), Lacosoma briasiaSchaus, Lacosoma diederica Schaus, Lacosoma raydela Schaus, Psychocampa lacuna (Schaus), Cicinnus corallina Dognin, Cicinnus latris Schaus, Cicinnus solvens Schaus, Cicinnus tuisana Schaus) as well as a neotype for Mimallo despecta Walker (= Cicinnus despecta). This paper also provides apomorphies for each genus and a morphological key to genera. Annotations are given to aid researchers in understanding all changes made herein, and images of male and female and their genitalia are present for nearly all type species.
Citralla , evolution, Fatellalla , Lepismalla , morphology, Neotropical, New World, revised classification, taxonomy
The sack-bearer moths (Mimallonidae) have been poorly studied in terms of their evolution, systematics, and natural history. The family is the only representative in the Mimallonoidea, and nearly all species are Neotropical (
Mimallonidae have extraordinary morphological diversity.
The only formal phylogenetic study of Mimallonidae is the molecular analysis of
The current study aims to incorporate morphological data to assess apomorphies and revise the classification of the family. This article has four main goals:
1 Conduct morphological phylogenetic analyses of Mimallonidae.
2 Present a key to genera of Mimallonidae, based on male genitalia morphology.
3 Provide a generic classification of Mimallonidae with apomorphies based on new phylogenetic results.
4 Present a complete, annotated checklist of Mimallonidae that includes all species in the family.
In order build a comprehensive phylogeny of Mimallonidae (Goal 1), 192 species of Mimallonidae (66% of the family considering that 291 species are treated as valid here) were coded for 55 morphological adult characters (see Suppl. material
Morphological phylogenetic analyses were carried out in a framework of maximum likelihood (ML) and parsimony. ML analyses were conducted in IQ-TREE v. 1.6.1 (
For the parsimony analyses, we used TNT v. 1.5 (
Our taxonomic key (Goal 2) is based on male genitalia morphology, developed from the male genitalia characters in our morphological matrix (characters 1–25). We created this key so that researchers can readily identify genera of Mimallonidae, and place new or unrecognized taxa in the according genus. Our revised classification (Goal 3) follows the results from the topologically constrained, morphological phylogenetic analysis (Fig.
For the checklist (Goal 4), species that were not explicitly mentioned or sampled in
Lectotypes are designated when syntypes from multiple localities exist for a given species, or when lectotypification is necessary to determine the identity of a species. Lectotypes for species described from syntypes from the same location, which we deem to be conspecific, will not be designated here, but rather later in more in-depth genus-level revisions where interspecific relationships can be more robustly determined. A neotype is designated for Mimallo despecta since the original type(s) is/are almost certainly lost (see
Morphology terminology follows
The following institutional abbreviations are used throughout the checklist.
CDH Coll. of Daniel Herbin, Garidech, France
CEIOC Entomological Collection of the Oswaldo Cruz Institute, Rio de Janeiro, Rio de Janeiro, Brazil
CGCM Coll. of Carlos G. C. Mielke, Curitiba, Paraná́, Brazil
CGD Coll. of Guy Durand, France
CPAC Coleção Embrapa Cerrados, Planaltina, Distrito Federal, Brazil
CPL Coll. of Peter Landolt, Washington, USA
CRAS Coll. of Ryan St Laurent, Gainesville, Florida, USA
DZUP Coll. Pe. Jesus S. Moure, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil
HEC Hope Entomological Collections of Oxford University Museum of Natural History, Oxford, U.K.
ISEZ The Institute of Systematics and Evolution of Animals of the Polish Academy of Sciences, Kraków, Poland
MGCL McGuire Center for Lepidoptera and Biodiversity, Gainesville, Florida, USA
MJWC Coll. of Matthew J.W. Cock, Llannon, Wales, U.K.
MWM Museum Witt, Munich, Germany
Our morphological analysis which utilized a molecular topological constraint provides the most up-to-date picture on the evolutionary relationships of Mimallonidae (Fig.
Relationships between species were also well-supported, with essentially all species placed in the genera to which they are assigned in the present study as per the taxonomic sections of this article (Sections 3 and 4). Although the majority of genera were recovered as monophyletic in both the constrained and unconstrained ML trees (Fig.
Generic clade recovery in the unconstrained ML analysis (Suppl. material
Our results also demonstrate that morphological traits, such as the degree of development of the frenulum (
1 | Uncus deeply bifurcated, bifurcation deeper than uncus width (Fig. |
Tolypida Schaus |
– | Uncus not deeply bifurcated, but may be bidentate if not simple | 2 |
2 | Genitalia structurally asymmetrical, such that phallus and single long tusk are oriented to one side of genitalia (Fig. |
Aceclostria Vuillot |
– | Genitalia largely symmetrical (minor asymmetry may be apparent in size of projections from the ventral portion of the vinculum, or in characters related to the valvae) but not with major structural asymmetry as in Fig. |
3 |
3 | Uncus proximal margin continuous with tegumen (Fig. |
Macessoga Schaus |
– | Uncus proximal margin clearly differentiated from tegumen (all male genitalia figures herein, except Fig. |
4 |
4 | Vinculum apodemes present (e.g., Fig. |
5 |
_ | Vinculum apodemes absent | 11 |
5 | Vinculum tusks present (Figs |
6 |
– | Vinculum tusks absent (Fig. |
Cicinnus Blanchard, in part |
6 | Vinculum tusks present as single pair (Figs |
8 |
– | Vinculum tusks present as two pairs (Figs |
7 |
7 | Base of gnathos rectangular, relatively lightly sclerotized (Fig. |
Isoscella St Laurent and Carvalho |
– | Base of gnathos ovoid, relatively heavily sclerotized (Fig. |
Roelmana Schaus |
8 | Vinculum tusks sharp (Fig. |
9 |
– | Vinculum tusks blunt (Fig. |
Arcinnus Herbin |
9 | Juxta complicated or hood-shaped, fused to phallus, occupying most of central region of genitalia (e.g., Figs |
10 |
– | Juxta simple, fused to base of phallus, anchoring it to vinculum, without complex dorsal region (Fig. |
Aleyda Schaus |
10 | Juxta component dorsal to phallus distinctly hood-like, arching over phallus towards its terminus (Fig. |
Euphaneta Schaus |
– | Juxta component dorsal to phallus variable, but always complex and heavily sclerotized, not hood-like, usually with setae covered fan-like lobes symmetrically on either dorsal side of phallus (e.g., Fig. |
Cicinnus Blanchard, in part |
11 | Lobe-like ventral-anterior projection of the vinculum present (Fig. |
Bedosia Schaus |
– | Lobe-like ventral-anterior projection of the vinculum absent (vinculum base in most Mimallonidae truncated or flattened, not extended as mesal lobe) | 12 |
12 | Base of valvae with smooth, elongate or curled spine-like tusks (Figs |
13 |
– | Base of valvae without elongate or curled spine-like tusks (not including saccular projections/spines) | 18 |
13 | Valvae tusks curled, often shorter than half length of valva (Figs |
14 |
– | Valvae tusks not curled, though they may be curved, generally elongate, longer than half length of valva (Figs |
15 |
14 | Gnathos arms massive, heavily sclerotized, much larger than uncus, extending beyond saccular edge of valva (Fig. |
Lepismalla gen. n. |
– | Gnathos arms very small, smaller in size than uncus, not extending beyond saccular edge of valva (Fig. |
Pamea Walker |
15 | Gnathos present | 16 |
– | Gnathos absent | Ulaluma St Laurent & Kawahara |
16 | Valvae membranous ventrally, extensively narrowed distally, and clubbed distally (Fig. |
Micrallo St Laurent & C. Mielke |
– | Valvae not as above | 17 |
17 | Sternite VIII with claw-like spines (Fig. |
Procinnus Herbin |
– | Sternite VIII without claw-like spines (but pair of elongate arms and/or other modifications may be present) (e.g., Fig. |
Druentica Strand |
18 | Uncus broadly rectangular (Fig. |
Fatellalla gen. n. |
– | Uncus variable, usually triangular, may be somewhat flattened or finger-like apically, but basally triangular | 19 |
19 | Pronounced coecum phallus absent (Figs |
39 |
– | Pronounced coecum phallus present (e.g., Figs |
20 |
20 | Coecum phallus simple (not bifurcated) (Fig. |
22 |
– | Coecum phallus bifurcated (Fig. |
21 |
21 | Transtilla/valvae apodemes forming complex sclerotization situated inward into body cavity from which elongated spine-like tusks project outward through center of genitalia (Fig. |
Menevia Schaus |
– | Transtilla/valvae apodemes not developed as above, elongated spine-like tusks absent (but saccular spines present) | Cunicumara St Laurent |
22 | Phallus snake-like (narrow, curved, variable in length) (Fig. |
Auroriana St Laurent & C. Mielke |
– | Phallus and gnathos (if present) not as above | 23 |
23 | Distal tip of valvae extremely narrowed, forming spine (Fig. |
Arianula Herbin |
– | Distal margin of valvae variable, but not as above | 24 |
24 | Gnathos absent (note, valvae apodeme extensions may be confused with gnathos) | 25 |
– | Gnathos present (note gnathos may be typical with paired projections (e.g., Figs |
29 |
25 | Valvae apodemes lobe-like, covered in setae (Fig. |
Lurama Schaus |
– | Valvae apodemes not as above | 26 |
26 | Valva apodeme/transtilla extensions columnar, broad, spanning near entire dorsal-ventral height of vinculum (Fig. |
Tostallo St Laurent & C. Mielke |
– | Valvae apodemes not as above | 27 |
27 | Valvae apodemes filamentous, elongate (Fig. |
Mimallo Hübner |
– | Valvae apodemes not as above | 28 |
28 | Valvae apodemes wrinkled triangular projections (Fig. |
Eadmuna Schaus |
– | Valvae apodemes not as above | Thaelia Herbin, in part |
29 | Diaphragm with four discrete setae-filled sacks (upper pair of sacks may be significantly smaller and/or thicker than lower pair) (Figs |
30 |
– | Diaphragm without four discrete setae-filled sacks | 31 |
30 | Gnathos distally separated as pair of finger-like projections, or slight bifurcation (Fig. |
Reinmara Schaus |
– | Gnathos distally a single finger-like projection, not separated distally (Fig. |
Trogoptera Herrich-Schäffer |
31 | Gnathos with paired distal projections (e.g., Figs |
35 |
– | Gnathos with single projection, or not projected (often plate-like) (Figs |
32 |
32 | Uncus apex and gnathos both significantly more heavily sclerotized than surrounding tegumen and situated very near each other, appearing pincer-like from the lateral aspect (Fig. |
Zaphanta Dyar |
– | Uncus and gnathos (if present) variable, not as above | 33 |
33 | Gnathos as single mesal plate, not synscleritous with surrounding genitalia (Fig. |
Herbinalla St Laurent & Kawahara |
– | Gnathos not as above | 34 |
34 | Gnathos projected mesally, heavily sclerotized, forming closed region below uncus (Fig. |
Thaelia Herbin, in part |
– | Gnathos unfused mesally or fused, but not heavily sclerotized/mesally projected (Fig. |
Ulmara Schaus |
35 | Phallus sword-like, sharp, covered in minute spines (Fig. |
Adalgisa Schaus |
– | Phallus not as above: narrow, tubular (Fig. |
36 |
36 | Either side of phallus flanked by knob-like sclerotization of juxta, these knobs usually only attached to phallus by membrane, not strongly fused to it by sclerotization (Figs |
37 |
– | Either side of phallus not flanked by knob-like sclerotization of juxta | 38 |
37 | Gnathos arms with minute teeth ventrally, or with short protuberance near apex (Fig. |
Biterolfa Schaus |
– | Gnathos arms either wrinkled, or elongated, finely tapering distally, never toothed or with additional dorsal protuberances (Fig. |
Psychocampa Grote |
38 | Mesal base of valvae with upturned projection, this projection usually spined (Fig. |
Roelofa Schaus |
– | Mesal base of valvae simple | Bedosiallo St Laurent & Kawahara |
39 | Gnathos replaced by smooth, flattened plate (Figs |
42 |
– | Gnathos either apparently absent, or reduced to pair of narrow sclerotizations extending below uncus, from which various projections may emanate (Figs |
40 |
40 | Uncus broadly triangular (Fig. |
Citralla gen. n. |
– | Uncus very narrow, sharp, or more acutely triangular | 41 |
41 | Gnathos synscleritous with juxtal complex and/or connected to diaphragm by patches of setae (Fig. |
Lacosoma Grote |
– | Gnathos reduced to finely tapering sclerotizations extending downward below subuncal projections, therefore uncus/gnathos complex not synscleritous with juxta | Vanenga Schaus |
42 | Phallus with elongated lateral projections ending in sharp tooth-like points (Fig. |
Tarema Schaus |
– | Phallus variable, but not as above, never with elongated, sharp-tipped lateral projections (Fig. |
Alheita Schaus |
For the generic classification of Mimallonidae, we follow the phylogeny in Fig.
Zaphanta infantilis Dyar, 1910.
Very small moths, among the smallest in body size within Mimallonidae, with forewing length ranging from 9–11 mm. Coloration also diagnostic: wing ground color yellow with purplish-pink antemedial regions. Ventrum of both fore- and hindwings with apparent antemedial as well as postmedial lines.
(1) Uncus and a subuncal sclerotization fused, heavily sclerotized apically forming a conical structure with tegumen (Fig.
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.
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 (
(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.
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.
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.
Auroriana colombiana St Laurent & C. Mielke, 2016.
Orange-brown ground color with diffuse pink coloration on all regions of wings, with a tornal notch on the forewing only, anterior margin of the hindwing smooth. Similarly colored brown and pink Mimallonidae have a notch on the anterior margin of the hindwing (though this may be weak) as well as a usually deeper notch on the forewing tornus, and/or the pink coloration is not suffused across the whole wing in these species, but rather clearly delimited by the postmedial line. For example, see Fatellalla gen. n. below, a similarly colored genus with more clearly distinct medial and submarginal pink coloration.
Distally downward curved, snake-like phallus (Fig.
Cicinnus esperans Schaus, 1905.
Combination of the following characters: silvery-gray or brown ground color, forewing with hyaline or sub-hyaline patch bisected by the M2 vein; dorsal postmedial line incomplete, formed by brown crescents between veins; the presence of smooth wing margins without a sharply falcate forewing apex.
Combination of the following characters: (1) Uncus clearly differentiated from tegumen as separate smaller structure (Fig.
Perophora fabia Druce, 1887.
Similar to Eadmuna, but ground coloration more yellow to yellow-brown, with continuous dorsal postmedial line, which is mostly straight (except for a sharp angle toward costa), not incomplete and crenulate as in Eadmuna.
Combination of the following characters: (1) Uncus undifferentiated from tegumen, forming large (relative to remainder of genitalia) rounded triangle (Fig.
Bombyx amilia Cramer, 1780.
Combination of the following characters: wing margins irregular, postmedial lines always with distinct maculation of variable thickness spanning from postmedial line to wing margin, postmedial maculation wider on hindwing; forewing always with hyaline patch bisected by M2.
Pair of twisted, wrinkled, tendril-like elongations originating from vinculum near inner costal base of valvae (Fig.
Perophora albescens Jones, 1912.
Unmistakable, bird dropping-like white and brown coloration, white ground color combined with rounded forewings is unique to this genus.
Transtilla/valva apodeme formed by two columnar structures with multiple invaginations and internal wrinkles (Fig.
Tostallo was not included in
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.
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.
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
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.
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.
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.
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.
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.
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
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.
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.
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
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
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.
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.
According to
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.
(1) Subuncus projections extend from ventral margins of tegumen, extended outward most usually as a simply triangular protrusion (Fig.
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.
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.
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.
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.
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 (
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.
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.
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.
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.
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.
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.
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.
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
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.
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.
Bedosiallo was found to be a strongly supported clade in
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.
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.
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.
We were unable to figure female genitalia of Arcinnus, but direct the reader to figs 54 and 55 in
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
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.
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.
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
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.
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.
Mimallonoidea Burmeister, 1878
Mimallonidae Burmeister, 1878
Perophoridae Plötz, 1885; (
Ptochopsychidae Grote, 1896
Lacosomatidae Brues & Melander, 1915
Zaphantinae St Laurent & Kawahara, 2018: 739
Zaphanta Dyar, 1910: 85
infantilis Dyar, 1910: 85* (Guyana,
fraterna Schaus, 1912: 48, stat. rev. (Costa Rica: Limón,
Incertae Sedis
Cunicumara St Laurent, 2016: 86
anae St Laurent, 2016: 88* (Bolivia: Santa Cruz,
Menevia Schaus, 1928: 665
lantona species-group
lantona (Schaus, 1905: 327)* (Cicinnus) (French Guiana,
magna St Laurent & Dombroskie, 2016: 49 (Brazil: Santa Catarina, DZUP)
rosea St Laurent & Dombroskie, 2016: 44 (Ecuador: Napo,
torvamessoria St Laurent & Dombroskie, 2016: 47 (Peru: Puno,
lucara species-group
lucara (Schaus, 1905: 328) (Cicinnus) (French Guiana,
menapia St Laurent & Dombroskie, 2016: 57 (Guatemala: Izabal,
mielkei St Laurent & Dombroskie, 2016: 59 (Brazil: Minas Gerais, DZUP)
ostia species-group
ostia (Druce, 1898: 447) (Perophora) (Panama: Chiriquí,
pallida Herbin & C. Mielke, 2014: 147 (Brazil: Maranhão, DZUP)
parostia Schaus, 1928: 667 (Unknown,
perostia‡ in
plagiata species-group: plagiata subgroup
alurca Herbin & C. Mielke, 2014: 146 (Brazil: Maranhão, DZUP)
ulcara‡ in Herbin & C. Mielke (2014), misspelling
australis St Laurent & Dombroskie, 2016: 86 (Brazil: Santa Catarina,
elegans‡ Franclemont, unavailable manuscript name5
plagiata (Walker, 1855: 1341) (Mimallo) (Brazil: Rio de Janeiro,
superba‡ Jones, unavailable manuscript name5
plagiata species-group: vulgaris subgroup
cordillera St Laurent & Dombroskie, 2016: 107 (Peru: Puno,
delphinus St Laurent & Dombroskie, 2016: 109 (Brazil: Distrito Federal, CPAC)
franclemonti St Laurent & Dombroskie, 2016: 97 (Brazil: Santa Catarina,
falco‡ Franclemont, unavailable manuscript name5
vulgaris St Laurent & Dombroskie, 2016: 89 (French Guiana,
vulgaricula St Laurent & Dombroskie, 2016: 103 (Brazil: Amazonas,
Roelofa Schaus, 1928: 640
elyanae Herbin & C. Mielke, 2014: 142 (Brazil: Maranhão, DZUP)
hegewischi (Druce, 1887: 227) (Perophora) (Mexico,
hegewishi‡ Schaus, 1928: 640, misspelling
maricia Schaus, 1928: 640 (Brazil: Rio de Janeiro,
narga (Schaus, 1905: 329) (Cicinnus) (Surinam,
maera (Schaus, 1913: 5) (Cicinnus) (Brazil: Santa Catarina,
olivia (Schaus, 1896b: 52)* (Perophora) (Colombia,
Tolypida Schaus, 1928: 663
amaryllis (Schaus, 1896a: 143)* (Hydrias) (Brazil: São Paulo,
alboflava (Dognin, 1917: 17) (Cicinnus) (Brazil: Santa Catarina,
spitzi Pearson, 1984: 463 (Brazil: Goiás,
Aurorianinae St Laurent & Kawahara, 2018: 741
Auroriana St Laurent & C. Mielke, 2016: 123
colombiana St Laurent & C. Mielke, 2016: 127* (Colombia: Meta,
florianensis (Herbin, 2012: 23) (Cicinnus) (French Guiana,
gemma St Laurent & C. Mielke, 2016: 134 (Brazil: Santa Catarina, DZUP)
Mimalloninae Burmeister, 1878
Eadmuna Schaus, 1928: 663
esperans (Schaus, 1905: 327)* (Cicinnus) (Brazil: Espírito Santo,
guianensis St Laurent & Dombroskie, 2015: 59 (French Guiana,
paloa Schaus, 1933: 487 (Brazil: São Paulo,
pulverula (Schaus, 1896b: 52) (Perophora) (Brazil: São Paulo,
Macessoga Schaus, 1928: 664
aelfrida Schaus, 1928: 664 (Brazil: Minas Gerais,
fabia (Druce, 1887: 227)* (Perophora) (Panama,
hoppia Schaus, 1928: 665, stat. n. (Brazil: Rio de Janeiro,
hyginia Schaus, 1928: 665, stat. n. (Brazil: Minas Gerais,
laxa (Dognin, 1912: 173) (Cicinnus) (Argentina: Misiones,
lucero‡ in Piñas (2004), nomen nudum13
Mimallo Hübner, 1820: 190
Mamillo‡ in
almeidai Pearson, 1951: 325 (Brazil: Rio de Janeiro, CEIOC)
amilia (Cramer, 1780: 130)* (Bombyx) (Surinam,
vorax (Sepp, [1832]: 47) (Bombyx) (Surinam, Unknown)
aemilia‡ in Herrich-Schäffer (1865), misspelling
grisea (Schaus, 1896b: 52), comb. n. (Perophora) (Brazil: Paraná,
hector Dognin, 1924: 30 (Brazil: Santa Catarina,
neoamilia Pearson, 1951: 322 (Brazil: Rio de Janeiro, CEIOC)
saturata Walker, 1855: 1340, nomen dubium (Brazil: Rio de Janeiro, Unknown)15
Tostallo St Laurent & C. Mielke, 2016: 119
albescens (Jones, 1912: 435)* (Perophora) (Brazil: São Paulo,
Lacosominae Dyar, 1893
Trogopterini St Laurent & Kawahara, 2018: 745
Reinmara Schaus, 1928: 654
andensis St Laurent, Herbin & C. Mielke, 2017c: 110 (Bolivia: “N. Yungas”,
atlantica St Laurent, Herbin & C. Mielke, 2017c: 108 (Brazil: Espírito Santo, DZUP)
enthona (Schaus, 1905: 325)* (Cicinnus) (French Guiana,
ignea St Laurent, Herbin & C. Mielke, 2017c: 123 (Brazil: Santa Catarina, DZUP)
minasa Schaus, 1928: 655 (Brazil: Minas Gerais,
occidentalis St Laurent, Herbin & C. Mielke, 2017c: 112 (Ecuador: El Oro, MWM)16
wolfei Herbin & C. Mielke, 2014: 144 (Brazil: Maranhão, DZUP)
Trogoptera Herrich-Schäffer, [1856]: 60
Tomoptera‡ in Burmeister, 1878, misspelling
althora Schaus, 1928: 652 (Guatemala: Izabal,
belilia Schaus, 1928: 653 (Brazil: Pará,
callinica Schaus, 1928: 653 (Brazil: Rio Grande do Sul,
dietricha Schaus, 1934: 94 (Brazil: Rio de Janeiro,
erosa Herrich-Schäffer, [1856]: 60* (Brazil: Rio de Janeiro, Unknown)18
excavata (Walker, 1855: 1154) (Pamea) (Brazil, probably Rio de Janeiro, HEC)18
guianaca Schaus, 1928: 652 (French Guiana,
jonica Schaus, 1928: 653, stat. n. (Paraguay: Guairá,
mahala Schaus, 1928: 652, stat. n. (Ecuador: Loja,
mana Schaus, 1928: 654 (French Guiana,
maroniensis Dyar, 1910: 86 (Pamea) (French Guiana,
maroniensis‡ Dognin, 1910: 39 (Pamea) (French Guiana,
micalha Schaus, 1928: 653, stat. n. (Mexico: Veracruz,
noaha Schaus, 1928: 652, stat. n. (Mexico: Veracruz,
notata (Walker, 1855: 1155) (Pamea) (Brazil, probably Rio de Janeiro, HEC)25
salvita Schaus, 1928: 653 (Brazil: São Paulo,
sao Druce, 1894: 355 (Costa Rica: Limón,
semililacea (Dognin, 1916: 33) (Pamea) (French Guiana,
tirzaha Schaus, 1928: 653 (Panama: Chiriquí,
Alheitini St Laurent & Kawahara, 2018: 743
Adalgisa Schaus, 1928: 670
croesa Schaus, 1928: 671* (Brazil: Santa Catarina,
stellifera Schaus, 1928: 671 (Paraguay: “Molinas”,
Alheita Schaus, 1928: 668
adelitae Herbin, 2016: 202 (French Guiana,
anoca (Schaus, 1905: 327)* (Cicinnus) (French Guiana,
beroalda Schaus, 1928: 668 (Guatemala: Izabal,
counamama Herbin, 2015: 86 (French Guiana,
cymbelina Schaus, 1928: 668 (Brazil: Pará,
hermieri Herbin, 2015: 85 (French Guiana,
obscura Herbin, 2016: 200 (French Guiana,
pulla (Dognin, 1912: 173) (Cicinnus) (Colombia,
pulloides (Dognin, 1921: 19) (Perophora) (Colombia: Cundinamarca,
rionica Schaus, 1928: 669 (Brazil: Amazonas,
subnotata (Dognin, 1921: 18) (Perophora) (Peru: Mariscal Ramón Castilla,
Arianula Herbin, 2012: 25
feiranovensis Herbin & C. Mielke, 2014: 139 (Brazil: Maranhão, DZUP)
haxairei Herbin, 2012: 27* (French Guiana,
Fatellalla St Laurent & Kawahara, gen. n. 30
fatella (Schaus, 1905: 326)*, comb. n. (Cicinnus) (French Guiana,
Herbinalla St Laurent & Kawahara, 2018: 75431
caudina (Schaus, 1905: 326)* (Cicinnus) (French Guiana,
Tarema Schaus, 1896b: 55
bruna St Laurent, Herbin, & C. Mielke, 2017b: 134 (Brazil: São Paulo,
fuscosa Jones, 1908: 173 (Brazil: Paraná,
rivara Schaus, 1896b: 55* (Brazil: São Paulo,
macarina Schaus, 1928: 670 (Brazil: São Paulo,
Thaelia Herbin, 2016: 178
anysia (Schaus, 1928: 651) (Cicinnus) (French Guiana,
beniensis Herbin, 2016: 182 (Bolivia: Beni,
inornata (Druce, 1905: 90), comb. n. (Bombyx) (Peru: Puno,
linamariae Herbin, 2016: 179* (French Guiana,
subrubiginosa (Dognin, 1916: 19) (Cicinnus) (Colombia: Cundinamarca,
Lacosomini Dyar, 1893
Citralla St Laurent & Kawahara, gen. n. 35
rumina (Druce, 1894: 355)*, comb. n. (Trogoptera) (Panama: Chiriquí,
Lacosoma Grote, 1864: 77
Naniteta Franclemont, 1973: 1036
arantium Herbin, 2016: 191 (French Guiana,
arizonicum Dyar, 1898: 44 (USA: Arizona,
asea Schaus, 1928: 662 (Colombia: Cundinamarca,
aurora Dognin, 1916: 17 (Colombia: Boyacá,
bigodia Schaus, 1928: 662 (Brazil: Amazonas,
briasia Schaus, 1928: 659 (Colombia: Quindío,
canens Herbin, 2016: 198 (French Guiana,
cantia Schaus, 1928: 661 (Brazil: Amazonas,
chiridota Grote, 1864: 78* (USA: Pennsylvania,
diederica Schaus, 1928: 660 (Bolivia: La Paz,
elassa (Franclemont, 1973: 11) (Naniteta) (USA: Texas,
horii St Laurent & C. Mielke, 2018: 20 (Brazil: Paraná, DZUP)
julietta Dyar, 1913: 316 (Mexico: Veracruz,
ladema Dognin, 1920: 11 (Colombia: Boyacá,
ludolpha Schaus, 1928: 660 (Venezuela: Carabobo,
streckeri‡, unavailable manuscript name39
lygia (Schaus, 1912: 56) (Cicinnus) (Costa Rica: Limón,
maldera Schaus, 1934: 94 (Brazil: Rio de Janeiro,
medalla Dyar, 1913: 316 (Mexico: Morelos,
miradorensis Herbin & Monzón, 2015: 178 (Guatemala: Petén,
morgani Herbin & Monzón, 2015: 180 (Guatemala: Petén,
ostrinum Herbin, 2016: 196 (French Guiana,
otalla Schaus, 1905: 330 (French Guiana,
oyapoca Schaus, 1928: 662 (French Guiana,
perplexa Schaus, 1920: 151 (Guatemala: Izabal,
philastria Schaus, 1928: 660 (Brazil: Amazonas,
puniceum Herbin, 2016: 193 (French Guiana,
raydela Schaus, 1928: 660 (Guatemala: Izabal,
rosea (Dognin, 1905: 120) (Perophora) (Colombia: Cauca,
schausi Dognin, 1923: 17 (French Guiana,
rosea‡ Schaus, 1905: 328, preoccupied
subrufescens (Dognin, 1916: 32) (Cicinnus) (Colombia: Tolima,
syrinx (Druce, 1898: 447) (Mimallo) (Panama: Chiriquí,
turnina Schaus, 1928: 660 (Brazil: Amazonas,
ursmara (Schaus, 1928: 656), comb. n. (Druentia) (Bolivia: Cochabamba,
valera Schaus, 1928: 662 (Venezuela: Trujillo,
valeroides Herbin & C. Mielke, 2014: 145 (Brazil: Maranhão, DZUP)
violacea (Sepp, [1848]: 67), (Bombyx) (Surinam, Unknown)45
vulfreda Schaus, 1928: 659 (Colombia: Chocó,
zonoma Schaus, 1928: 662 (Mexico: Veracruz,
Vanenga Schaus, 1928: 664
mediorosea St Laurent & Herbin, 2017: 95 (Brazil: Santa Catarina,
flavirosa‡ Jones, unavailable manuscript name
meroides‡/meroidea‡ Schaus, unavailable manuscript name
roseatincta‡ Schaus, unavailable manuscript name
mera (Dognin, 1924: 31)* (Perophora) (Brazil: Pará,
Druenticinae St Laurent & Kawahara, 2018: 745
Druenticini St Laurent & Kawahara, 2018: 746
Druentica Strand, 1932: 145 (replacement name)47
Druentia‡ Schaus, 1928: 655, preoccupied
alsa (Schaus, 1910: 422) (Cicinnus) (Costa Rica: Cartago,
brosica (Schaus, 1928: 636) (Mimallo) (French Guiana,
caquetensis (Schaus, 1928: 655) (Druentia) (Colombia: Caquetá,
coralie Herbin, 2016: 167 (French Guiana,
corana (Schaus, 1928: 657) (Druentia) (Colombia: Quindío,
derrica (Schaus, 1928: 657) (Druentia) (Peru: Puno,
differenciata (Bryk, 1953: 185), comb. n. et stat. n. (Cicinnus) (Peru: San Martín,
fanoveira Herbin & C. Mielke, 2014: 141 (Brazil: Maranhão,
garretti (Schaus, 1934: 92), comb. n. (Mimallo) (Brazil: Rio de Janeiro,
imperita (Dognin, 1905: 120) (Perophora) (Peru: Puno,
inscita (Schaus, 1890: 46) (Perophora) (Mexico: Veracruz,
inscitoides (Dognin, 1923: 24), stat. rev. (Perophora) (Brazil: Amazonas,
lola (Schaus, 1905: 328), comb. n. (Cicinnus) (French Guiana,
macallia (Schaus, 1928: 655) (Druentia) (Colombia: Putomayo,
melastoma Herbin, 2016: 171 (French Guiana,
muta (Dognin, 1912: 172) (Cicinnus) (French Guiana,
mutara (Schaus, 1933: 486) (Druentia) (Brazil: Rio de Janeiro,
narita (Dognin, 1912: 172) (Cicinnus) (Colombia: Valle del Cauca,
putidula (Dognin, 1912: 171), comb. n. (Cicinnus) (Colombia: Valle del Cauca,
partha (Schaus, 1905: 325)* (Cicinnus) (French Guiana,
patawa Herbin, 2016: 169 (French Guiana,
rotundula (Dognin, 1916: 21) (Cicinnus) (Colombia: Tolima,
scissa (Herrich-Schäffer, [1856]: 60) (Euclea) (Brazil,
zikana (Schaus, 1928: 655) (Druentia) (Brazil: Rio de Janeiro,
Lepismalla St Laurent & Kawahara, gen. n.54
montagnaniae (Herbin, 2012: 14)*, comb. n. (Cicinnus) (French Guiana,
Micrallo St Laurent & C. Mielke, 2016: 13655
minutus St Laurent & C. Mielke, 2016: 139* (Brazil: Piauí, DZUP)
Pamea Walker, 1855: 115356
Pomea‡ in Burmeister, 1878, misspelling
albistriga Walker, 1855: 1154* (Brazil, probably Rio de Janeiro,
dotta Schaus, 1928: 667 (Brazil: Paraná,
nana (Herrich-Schäffer, [1856]: 60) (Euclea) (Brazil: Rio de Janeiro, Unknown)
Procinnus Herbin, 2016: 175
acuta (Schaus, 1892: 327) (Perophora) (Brazil: Rio de Janeiro,
cahureli Herbin, 2016: 176* (French Guiana,
plana (Walker, 1855: 1338) (Mimallo) (Brazil, Unknown, probably lost)
diagonalis (Herrich-Schäffer, [1856]: 60) (Euclea) (Brazil,
producta (Dognin, 1901: 176), comb. n. (Perophora) (Colombia: Cauca,
Ulaluma St Laurent & Kawahara, 2018: 755
valva (Schaus, 1905: 329)* (Cicinnus) (French Guiana,
Luramini St Laurent & Kawahara, 2018: 748
Lurama Schaus, 1928: 667
Luramana Strand, 1932: 147 (unnecessary replacement name)
penia (Dognin, 1919: 6)* (Perophora) (Colombia: Distrito Capital,
quindiuna Schaus, 1928: 668 (Colombia: Quindío,
Ulmara Schaus, 1928: 666
azurula St Laurent, 2016: 83 (Peru: Huánuco,
conjuncta St Laurent, 2016: 79 (Ecuador: Loja,
dombroskiei St Laurent, 2016: 84 (Peru: Puno,
rotunda (Dognin, 1916: 20)* (Cicinnus) (Colombia: Tolima,
Cicinninae Schaus, 1912
Psychocampini St Laurent & Kawahara, 2018: 753
Biterolfa Schaus, 1928: 666
althea (Schaus, 1905: 326)* (Cicinnus) (French Guiana,
rana St Laurent, Giusti, & Herbin, 2017: 89 (Ecuador: Napo, MGCL)
yupanqui‡ in Piñas (2004), nomen nudum61
tinalandia St Laurent, Giusti & Herbin, 2017: 93 (Ecuador: Pichincha, MGCL)
Psychocampa Grote & Robinson, 1867: 37462
alcuna (Dognin, 1918: 26), stat. rev. (Cicinnus) (French Guiana,
bactriana (Butler, 1878: 77), comb. n. (Perophora) (Brazil: Amazonas/Pará,
batesii (Newman, 1854: 5), comb. n. (Perophora) (Brazil: Pará,
bilinea (Schaus, 1904: 141), comb. n. (Perophora) (Brazil: Paraná,
callipius (Schaus, 1928: 650), comb. n. (Cicinnus) (French Guiana,
camarinus (Schaus, 1928: 648), comb. n. (Cicinnus) (Peru: Loreto,
candacus (Schaus, 1928: 649), comb. n. (Cicinnus) (French Guiana,
concolor Grote & Robinson, 1867: 375* (Brazil: Pará, Unknown)
cunona (Schaus, 1905: 330) (Cicinnus) (French Guiana,
stenia (Dyar, 1910: 86) (Cicinnus) (Brazil: Pará,
funebris (Schaus, 1896b: 51), comb. n. (Perophora) (Brazil: Paraná,
gaujoni (Dognin, 1922: 26), comb. n. (Perophora) (Ecuador: Loja,
hamata (Walker, 1855: 975) (Perophora) (Brazil: Rio de Janeiro,
jaruga (Jones, 1912: 436), syn. n. (Perophora) (Brazil: São Paulo,
joanna (Schaus, 1905: 321) (Cicinnus) (French Guiana,
kohlli Herbin, 2012: 9 (French Guiana,
lacuna (Schaus, 1910: 418), stat. rev. (Cicinnus) (Costa Rica: Cartago,
madenus (Schaus, 1928: 648), comb. n. (Cicinnus) (Brazil: Amazonas,
magnapuncta (Kaye, 1901: 157), comb. n. (Perophora) (Trinidad,
manalca (Schaus, 1928: 648), comb. n. (Cicinnus) (French Guiana,
marona (Schaus, 1905: 323) (Cicinnus) (French Guiana,
mulatro (Schaus, 1920: 151), comb. n. (Cicinnus) (Guatemala: Izabal,
narseres (Schaus, 1928: 647), comb. n. (Cicinnus) (Bolivia: La Paz,
unalca (Schaus, 1905: 325) (Cicinnus) (Guyana,
undiscata (Dognin, 1923: 33), stat. rev. (Perophora) (Brazil: São Paulo,
viemanda (Schaus, 1928: 648), comb. n. (Cicinnus) (Brazil: Amazonas,
Bedosiini St Laurent & Kawahara, 2018: 752
Bedosia Schaus, 1928: 657
balca (Schaus, 1905: 323) (Cicinnus) (Guyana,
dulcis (Schaus, 1910: 420), comb. n. (Cicinnus) (Costa Rica: Cartago,
euthymius (Schaus, 1928: 649), comb. n. (Cicinnus) (Colombia: Chocó,
fraterna (Schaus, 1905: 330)* (Cicinnus) (French Guiana,
gilia (Schaus, 1905: 322) (Cicinnus) (French Guiana,
itamaraty (Foetterle, 1902: 641) (Mimallo) (Brazil: Rio de Janeiro,
ligina (Schaus, 1910: 419) (Cicinnus) (Costa Rica: Cartago,
olasis (Schaus, 1928: 645), comb. n. (Cicinnus) (“Central America”,
strigifera (C. & R. Felder, 1874: 8) (Perophora) (Brazil,
trailii (Butler, 1878: 77) (Perophora) (Brazil: Amazonas,
venata (Dognin, 1916: 31) (Cicinnus) (French Guiana,
turgida (Schaus, 1910: 420) (Cicinnus) (Costa Rica: Limón,
yenuga Herbin, 2016: 188 (French Guiana,
Bedosiallo St Laurent & Kawahara, 2018: 756
forbesi species-group
eugenia (Schaus, 1905: 324) (Cicinnus) (French Guiana,
forbesi (Schaus, 1928: 644)* (Cicinnus) (French Guiana,
sylvia (Schaus, 1920: 152) (Cicinnus) (Guatemala: Izabal,
volucris (Schaus, 1910: 421) (Cicinnus) (Costa Rica: Cartago,
gentilis species-group89
gentilis (Schaus, 1910: 419), comb. n. (Cicinnus) (Costa Rica: Cartago,
minimalis (Herbin & C. Mielke, 2014: 137), comb. n. (Cicinnus) (Brazil: Maranhão, DZUP)
moengus species-group
moengus (Schaus, 1928: 649) (Cicinnus) (Surinam,
styx (Herbin, 2012: 6) (Psychocampa) (French Guiana,
Cicinnini Schaus, 1912
Aceclostria Vuillot, 1893: CLXXXII
Aceclostria‡ Schaus, 1928: 670, preoccupied
cordubensis (Berg, 1882: 279), comb. n. (Mimallo) (Argentina: Córdoba, Unknown)90
schulzii (Weyenbergh, 1882: 141) (Mimallo) (Argentina: Córdoba,
mus Vuillot, 1893: CLXXXII* (Brazil: São Paulo, Unknown)
callinica‡ Schaus, unavailable manuscript name91
deprava (Schaus, 1896b: 52) (Perophora) (Brazil: São Paulo,
villaricensis (Schaus, 1933: 487) (Eadmuna) (Paraguay: Guairá,
nigrescens (Schaus, 1896b: 51), stat. rev. et comb. n. (Perophora) (Brazil: Paraná,
Aleyda Schaus, 1928: 641
accipiter (Dognin, 1916: 20)* (Cicinnus) (Panama: Chiriquí,
heppneri St Laurent, McCabe, & Malm, 2018: 162 (Peru: San Martín, MGCL)94
Arcinnus Herbin, 2016: 183
hoedli Herbin, 2016: 185* (French Guiana,
xingua (Dognin, 1923: 23), comb. n. (Perophora) (Brazil: Pará,
Cicinnus Blanchard, 1852: 6696
Saccophora‡ Harris in Doubleday, 1841: 101, preoccupied
Perophora‡ Harris, 1841: 299, preoccupied
Gonogramma Boisduval, 1872: 9697
Ptochopsyche Grote, 1896: [4] (replacement name for Perophora Harris)
Group 1 (Cicinnussensu stricto)98
chaubaudi Dyar, 1914b: 391 (Mexico: Estado de México,
incerta (Möschler, 1878: 676) (Mimallo) (Surinam,
latris Schaus, 1910: 418 (Costa Rica: Cartago,
melgibsoni Herbin & Monzón, 2015: 184 (Guatemala: Petén,
melsheimeri (Harris in Doubleday, 1841: 101) (Saccophora) (USA: Pennsylvania,
melsheimerii (Harris, 1841: 299) (Perophora) (USA: Pennsylvania,
egenaria (Walker, 1866: 1575) (Arhodia) (No locality given, but probably USA,
primolus Schaus, 1928: 647 (No locality given, but probably USA,
mexicana (Druce, 1898: 446) (Perophora) (Mexico: Veracruz,
orthane Blanchard, 1852: 66* (Chile: Concepción [probably erroneous, likely Brazil],
orthana‡ Berg, 1876: 170, misspelling
pudens Schaus, 1911: 191 (Costa Rica: Cartago,
solvens Dyar, 1914a: 453 (Panama: Panama,
tuisana Schaus, 1910: 417 (Costa Rica: Cartago,
Group 2 (Cicinnussensu lato)106
bahamensis St Laurent & McCabe, 2016: 561 (Bahamas: Great Exuma,
belaria (Schaus, 1928: 637), comb. n. (Psychocampa) (Brazil: Minas Gerais,
cerradensis Herbin & C. Mielke, 2014: 133 (Brazil: Maranhão, DZUP)
conlani Herbin & C. Mielke, 2014: 135 (Brazil: Maranhão, DZUP)
corallina Dognin, 1918: 27 (Colombia: Cundinamarca,
corcovada (Schaus, 1892: 326) (Perophora) (Brazil: Rio de Janeiro,
despecta (Walker, 1855) (Mimallo) (Brazil: Rio de Janeiro,
curtisea (Weyenbergh, 1874: 220) (Mimallo) (Argentina, probably Buenos Aires, Unknown)109, 110
sanguinolenta (C. & R. Felder, 1874: 8) (Perophora) (Brazil,
sachinius Schaus, 1934: 93 (Brazil: Minas Gerais,
eminens (Dognin, 1923: 16) (Perophora) (Colombia: Distrito Capitol,
externa (Moore, 1882: 358) (Perophora) (Brazil: São Paulo, Unknown)
roscida Dognin, 1910: 38, syn. rev. (Brazil: São Paulo,
falcoargenteus St Laurent & McCabe, 2016: 566 (Venezuela: Carabobo,
felderia Schaus, 1928: 643 (Mexico: Guerrero,
floris St Laurent, 2017: 23 (Panama: Panamá [Taboga Island],
fogia Schaus, 1905: 321 (French Guiana,
giustii St Laurent, 2017: 25 (Colombia: Valle del Cauca,
hanseni Herbin & Monzón, 2015: 182 (Guatemala: Petén,
malca Schaus, 1905: 322 (French Guiana,
manicora (Schaus, 1928: 637), comb. n. (Psychocampa) (Brazil: Amazonas,
maranhensis Herbin & C. Mielke, 2014: 132 (Brazil: Maranhão, DZUP)
motagus Schaus, 1920: 151 (Guatemala: Izabal,
musa (Schaus, 1896b: 51) (Perophora) (Brazil: Paraná,
packardii (Grote, 1865: 251) (Perophora) (Cuba,
packardi‡ misspelling, various authors e.g.,
patawaensis Herbin, 2012: 11 (French Guiana,
submarcata Schaus, 1905: 320 (French Guiana,
thermesia (Jones, 1921: 353) (Perophora) (Brazil: São Paulo,
Group 3 (Cicinnussensu lato)117
beta Schaus, 1910: 421 (Costa Rica: Cartago,
gaia Herbin, 2015: 84 (French Guiana,
kawensis Herbin, 2012: 16 (French Guiana,
mamorensis Herbin, 2012: 21 (Bolivia: Beni,
trini St Laurent & Cock, 2017: 55 (Trinidad,
veigli (Schaus, 1934: 92) (Psychocampa) (Brazil: Minas Gerais,
Group undetermined (Cicinnussensu lato)
bibula (Dognin, 1924: 31), comb. n. (Perophora) (Brazil: Pará,
haromana Herbin & C. Mielke, 2014: 134 (Brazil: Maranhão, DZUP)
lemoulti Schaus, 1905: 329, comb. rev. (Cicinnus) (French Guiana,
mawaja (Dognin, 1922: 27), comb. n. (Perophora) (Brazil: Pará,
Euphaneta Schaus, 1928: 637 (replacement name)
Phaneta‡ Walker, 1855: 1382, preoccupied
divisa (Walker, 1855: 1383)* (Phaneta) (Brazil: Pará,
romani Bryk, 1953: 186 (Peru: Ucayali,
Isoscella St Laurent & Carvalho, 2017b: 93
andina St Laurent & Carvalho, 2017b: 106 (Peru: Junín, MGCL)
ecuadoriana St Laurent & Carvalho, 2017b: 97 (Ecuador: Napo, MWM)
leva St Laurent & Carvalho, 2017b: 101 (Peru: Puno,
peigleri St Laurent & Carvalho, 2017b: 104 (Ecuador: Carchi, MWM)
ventana (Dognin, 1897: 243)* (Perophora) (Venezuela: Mérida,
Roelmana Schaus, 1928: 671125
beneluzi (Herbin, 2012: 19), comb. n. (Cicinnus) (French Guiana,
brasiliensis (Herbin & C. Mielke, 2014: 138), comb. n. (Cicinnus) (Brazil: Maranhão, DZUP)127
doralica (Schaus, 1928: 637), comb. n. (Psychocampa) (Colombia: Boyacá,
fenestrata (Jones, 1912: 436), comb. n. (Perophora) (Brazil: Paraná,
laguerrei (Herbin, 2012: 2) (Psychocampa) (French Guiana,
maloba (Schaus, 1905: 324)* (Cicinnus) (French Guiana,
pluridiscata (Dognin, 1916: 18) (Cicinnus) (Peru: Puno,
prominens (Schaus, 1910: 417), comb. n. (Cicinnus) (Costa Rica: Cartago,
vitreata (Schaus, 1905: 324), comb. n. (Cicinnus) (French Guiana,
1 The type locality of Z. infantilis has remained unclear since its original description.
2 As part of an ongoing revision of Zaphanta, which includes dissections of all Zaphanta species and populations, it is apparent that the Central American Z. fraterna stat. rev. is markedly distinct from the Amazonian Z. infantilis (St Laurent and Giusti in prep.). We therefore revalidate Z. fraterna in the present checklist.
3 The monotypic Cunicumara was considered “incertae sedis” along with Menevia, Roelofa, and Tolypida by
4 Unfortunately, the type locality of M. parostia is unknown. It has been suggested that M. pallida may be a synonym of M. parostia (
5 The taxonomy of M. plagiata and related species was confusing until the revision of Menevia by
6 The original primary type of M. plagiata is lost, along with many types described by Walker originating from the Fry collection (
7
8 Confusion has surrounded the type locality and original combination of Tolypida amaryllis, the type species of Tolypida. Motada amaryllis Schaus, 1896a: 142 and Hydrias amaryllis Schaus, 1896a: 143, both bearing the specific name amaryllis, were described by Schaus in the same publication, and both had the type locality of Castro, Paraná, Brazil (
9 Eadmuna paloa was treated by
10 Eadmuna pulverula was transferred from Cicinnus to Eadmuna by
11 Macessoga hoppia stat. n. and M. hyginia stat. n. were both originally described as “forms” of M. aelfrida by
12 Macessoga laxa was placed in Druentica by
13 As mentioned in
14 Mimallo grisea comb. n. was placed in Trogoptera since
15 Mimallo saturata‡ was treated as nomen dubium by
16
17 Trogoptera althora was described from an indeterminate number of specimens from Guatemala and Costa Rica. Considering that this may lead to some confusion in identifying “true” T. althora, we designate a lectotype for the specimen labeled as the “type” in the
18 The identity of Trogoptera erosa has not been clear due to the lack of any type material, besides the figure in Herrich-Schäffer, [1856]. Trogoptera erosa is darker, with more sharply angled forewing apices than in the other common southeastern Brazilian species Trogoptera notata/excavata which may or may not prove to be synonymous names since the type of T. notata is male and that of T. excavata is female.
19
20 Trogoptera jonica stat. n. was originally treated as a subspecies of T. althora by
21 Trogoptera mahlaha stat. n. was originally treated as a “form” of T. althora by
22 Trogoptera maroniensis (Dyar) is a senior homonym of T. maroniensis (Dognin).
23 Trogoptera micalha stat. n. was originally treated as a subspecies of T. althora by
24 Trogoptera noaha stat. n. was originally treated as a “form” of T. althora by
25 Trogoptera notata is a problematic species because its holotype specimen lacks specific locality data within Brazil, though which was probably Rio de Janeiro since most of the Walker-era Mimallonidae were collected in that Brazilian state.
26 Trogoptera salvita has been often confused with T. jonica. However, the two species appear to be distinct, with the wing margins of female T. salvita being more convex. Trogoptera salvita is found in the Brazilian Cerrado, whereas T. jonica is so far known from Paraguay and neighboring Argentina, Brazil, and Bolivia. The type locality given by
27 Adalgisa croesa was described from an indeterminate number of specimens from Santa Catarina, Brazil, and French Guiana. Considering the likelihood that Adalgisa from these two localities may eventually prove to be distinct species, and in order to stabilize the identity of A. croesa, we designate a lectotype for the specimen labeled as the “type” in the
28 Alheita pulloides was described from four male specimens from Guatemala, Colombia, French Guiana, and Peru. Although
29
30 We describe a new genus, Fatellalla gen. n., for F. fatella comb. n. Although this genus was not part of the phylogenetic analyses of
31 The monotypic genus Herbinalla was described by St Laurent and Kawahara in
32 Tarema macarina was treated as synonym of T. rivara by
33 Thaelia subrubiginosa and T. anysia were erroneously placed in Alheita and Cicinnus (respectively) by Schaus. Both species were transferred to Thaelia by St Laurent et al. (2018a) based on their phylogenetic results. External morphology also supports the new combinations of these two species. However, genitalia of T. subrubiginosa and T. anysia are rather distinct from the type species of Thaelia figured in
34 Thaelia inornata comb. n. was omitted from
35 We describe a new genus, Citralla gen. n., for C. rumina comb. n. Based on male genitalia, larval characteristics, and our morphological phylogenetic analyses, we can unambiguously place this new genus in Lacosominae: Lacosomini and ongoing molecular phylogenetics supports the novelty of this new genus (St Laurent in prep.). See genus description above for diagnosis, apomorphies, description, and figures for this genus. The type locality for “Trogoptera” rumina in
36 Naniteta was described by
37 Lacosoma briasia was described from an indeterminate number of specimens from far reaching localities in Brazil, Colombia, and French Guiana. Considering that this may lead to some confusion in identifying “true” L. briasia, we designate a lectotype for the specimen labeled as the “type” in the
38 Lacosoma diederica was described from an indeterminate number of specimens from Brazil, Bolivia, and Paraguay. Considering that this may lead to some confusion in identifying “true” L. diederica, we designate a lectotype for the specimen labeled as the “type” in the
39 There is a specimen labeled “Type” in
40 Lacosoma morgani may be a synonym of L. elassa; both species are nearly identical in male genitalia and external morphology, but the two species are described from distant localities (Guatemala for L. morgani, and southern Texas, USA for L. elassa). More material of L. elassa, especially from farther south, would be needed to compare directly with L. morgani.
41 Lacosoma raydela was described from an indeterminate number of specimens from Guatemala and Costa Rica. Considering that this may lead to some confusion in identifying “true” L. raydela, we designate a lectotype for the specimen labeled as the “type” in the
42 Lacosoma schausi was proposed as a replacement name by
43 Lacosoma subrufescens was transferred to Lacosoma from Trogoptera by
44 Lacosoma ursmara comb. n. is transferred from Druentica based on the discovery of female L. asea, which is remarkably similar to the female type of L. ursmara. The two species are from far separated type localities (L. asea in Colombia, L. ursmara in Bolivia), so we do not treat them as synonyms pending an inclusive revision of Lacosoma. Both species are Andean endemics and are relatively rare in collections. The morphological phylogenetic analyses consistently recover L. ursmara nested within Lacosoma.
45 Lacosoma violacea is a taxon of uncertain identity. No specimens have been found attributed to this name, and the only information pertaining to this species are the original figures and text in Sepp [1818]. Externally, the specimen figured in Sepp is unlike any other known Mimallonidae, particularly the blue coloration and arrangement of the postmedial line on the ventrum of the forewings. In fact, the color blue is not known from any Mimallonidae. It is possible that Sepp’s illustration is a misrepresentation of a more characteristically colored mimallonid species, and thus we consider Sepp’s illustration unreliable for accurately assigning the name violacea.
46 Vanenga mediorosea was described for the Vanenga populations inhabiting the Brazilian Atlantic Forest, the Pampa, Paraguay, and Argentina. This species has had at least three manuscript names variously applied to it in the
47 Druentica was proposed by
48 Druentica brosica was transferred to Druentica from Mimallo in
49 Druentica differenciata comb. n. et stat. n. was described by
50 Druentica garretti comb. n. displays external and genital morphology (MWM genitalia preparation 35.544) similar to D. brosica, and the former is unlike any described Mimallo.
51 Druentica inscitoides stat. rev. was treated as a synonym of D. scissa by
52 Druentica lola comb. n. is hereby transferred to Druentica from Lacosoma based on external and genital morphology. Druentica lola is more similar to D. mutara than any Lacosoma species, though a genitalia dissection (St Laurent diss.: CPC2 in CRAS) of D. lola from French Guiana reveals distinct basal valvae arms, valvae, and phallus structure consistent with more typical Druentica species. The morphological phylogenetic analyses consistently recover D. lola nested within Druentica.
53 Druentica putidula comb. n. is hereby transferred to Druentica from Cicinnus based on external and genital morphology. All morphological phylogenetic analyses consistently recover D. putidula nested within Druentica.
54 We describe a new genus, Lepismalla gen. n., for the unique species L. montagnaniae comb. n. Based on male genitalia and our morphological phylogenetic analyses, we can unequivocally place this new genus in Druenticini, refuting placement in Cicinnus as the original combination had proposed. See genus description above for diagnosis, apomorphies, description, and figures for this genus. Ongoing molecular phylogenetics which includes Lepismalla concurs with the novelty of Lepismalla as a valid genus, and its placement in Druenticini (St Laurent in prep.).
55 Micrallo was preliminarily placed in Druenticini by
56 The identity of the three species of Pamea is uncertain. We have not been able to locate the type(s) of P. nana, but the figures in Herrich-Schäffer (1856) display a male specimen not unlike the male types of P. albistriga and P. dotta. Pending a revision of Pamea, the three species are currently treated as valid.
57 We transfer Procinnus producta comb. n. from Cicinnus to Procinnus based on external and genitalia morphology. The genitalia of a male were examined by brushing, and are unambiguous, matching very closely the described Procinnus species. Due to the rarity of this species we did not fully dissect P. producta at this time, and thus did not code it for morphology. The genitalia were readily visible due to brushing, however, and thus we are confident of its generic placement. In the original description,
58 As explained in
59
60 Subsequent to the description of U. azurula, an additional specimen from Peru: Junín: 38 km SW of Mazamari, 2540 m (deposited in the MGCL) has been examined. This specimen was collected roughly halfway between the type localities of U. azurula and U. dombroskiei, and did not show any intermediate characters between these two species. This specimen displays external and male genitalia characteristics typical of U. azurula, supporting the validity of this rarely collected species.
61 As mentioned in
62 See
63 Psychocampa alcuna stat. rev. was previously treated as synonym of P. hamata (
64 Psychocampa bactriana comb. n. belongs to a relatively large group of similar species including P. marona, a species included in the phylogenetic study of
65 Psychocampa batesii comb. n. is very similar to P. bactriana. Upon direct comparison of the holotype of P. bactriana (
66 Although outwardly similar to Cicinnus s. s., a dissection of a male P. bilinea comb. n. surprisingly revealed genitalia diagnostic of Psychocampa, and therefore we move this species to Psychocampa from Cicinnus. All of our morphological phylogenetic analyses support this transfer as well.
67 Psychocampa callipius comb. n., P. candacus comb. n., P. gaujoni comb. n., and P. narseres comb. n. are hereby transferred to Psychocampa from Cicinnus based on phylogenetic and morphological data which places P. marona within the Psychocampa clade sensu
68 Since the original description of “Cicinnus” camarinus in
69 Psychocampa funebris comb. n. is hereby transferred to Psychocampa based on examination of genitalia which revealed characters completely in line with those of Psychocampasensu
70 As previously discussed in Annotation 63, Psychocampa hamata was transferred to Psychocampa based on morphology and the molecular results of
71 Yet another taxonomic issue concerning the Psychocampa hamata/alcuna complex arises from the name jaruga, proposed by
72 Psychocampa joanna and P. unalca were transferred to Psychocampa from Cicinnus by
73 Psychocampa lacuna stat. rev. was treated as synonym by
74 Psychocampa madenus comb. n. possesses the genitalia typical of Psychocampa (sensu
75 Psychocampa magnapuncta comb. n. belongs to a complex of similar species in this genus. As per
76 Psychocampa manalca comb. n. possesses the genitalia typical of Psychocampa (sensu
77 Psychocampa marona was transferred to Psychocampa from Cicinnus by
78 Psychocampa mulatro comb. n. was first noted to be related to P. undiscata by
79 Psychocampa unalca was transferred to Psychocampa from Cicinnus by
80 Psychocampa undiscata stat. rev. was erroneously synonymized with P. joanna in
81 Psychocampa viemanda comb. n. is externally difficult to distinguish from P. unalca, but P. viemanda is larger with darker maculation and has wider submarginal areas. Considering the close relationship of P. unalca and P. viemanda, and the incontrovertible evidence placing P. unalca in Psychocampa based on external morphology and molecular data in
82 The name melini syn. n. was proposed as a subspecies of P. joanna by
83 Bedosia dulcis comb. n. is hereby transferred to Cicinnus based on a genitalia examination of a Costa Rican specimen (genitalia vial NHMUK010402311 in
84 Bedosia euthymius comb. n. is most similar in external morphology to B. dulcis, more so than any Cicinnus and is therefore transferred to Bedosia. We have also examined the genitalia, confirming placement of euthymius in Bedosia. Ongoing molecular phylogenomics which include B. euthymius furthermore supports this transfer (St Laurent et al. in prep.).
85
86 Bedosia olasis comb. n. is similar to B. dulcis, as originally mentioned by
87 Authorship of B. strigifera (and other Mimallonidae described by C. and R. Felder (1874)) was determined with the aid of Nässig and Speidel (2007), hence it is in disagreement with the authorship listed in
88 In the original description,
89 The species Bedosiallo gentilis comb. n. and B. minimalis comb. n., previously both placed in Cicinnus, display genitalia characteristics very close to Bedosiallo. At the time of the Bedosiallo description,
90 Aceclostria cordubensis comb. n. displays the complex asymmetrical genitalia typical of A. mus, type species of Aceclostria. In A. cordubensis the complicated juxtal arrangement has more robust and more distinctly curved dorsal processes, a shorter spine that protrudes over the phallus, and the gnathos arms are smaller than in A. mus; but otherwise these features are extremely similar to all examined Aceclostria dissections examined thus far. Externally A. cordubensis is earthen in coloration, with indistinct suffuse maculation, and postmedial lines arranged in much the same way as in A. mus. Furthermore, in A. cordubensis and A. mus, the discal cell varies from fully scaled to completely hyaline in specimens collected from the same series. Such pronounced intraspecific variation in hyaline patches appears to be so far unique to Aceclostria. Distribution in dryer regions of central South America and usage of Schinus Linnaeus (Anacardiaceae) as host plants also supports the inclusion of A. cordubensis in Aceclostria (
91 A specimen of Aceclostria mus labeled as “Cicinnus callinica‡ Schaus” is present in the
92 Upon dissecting A. mus specimens from the type localities of both mus (São Paulo, Brazil, genitalia preparation NHMUK010402316 in
93 Aceclostria nigrescens stat. rev. et comb. n. has been treated as a synonym of A. cordubensis since
94 Authorship of Aleyda heppneri is St Laurent, McCabe, and Malm, 2018 as per
95 We hereby transfer Arcinnus xingua comb. n. from Cicinnus upon examination of the genitalia of the type specimen in the
96 Cicinnus is the most species-rich mimallonid genus. A great deal of taxonomic confusion has been associated with Cicinnus, and the genus has largely existed as a wastebasket taxon since its inception. Much of the confusion of the identification of “true” Cicinnus can be traced back to the type species, C. orthane. This species, described from Chile, is one of the only mimallonids recorded from that country, and therefore has been speculated to be erroneously labeled (
97 Gonogramma was named by
98 Group 1, which we treat here as Cicinnussensu stricto, includes species lacking hyaline patches on both the fore and hindwings, diffuse coloration with heavy speckling of dark brown to black petiolate scales, and short mesal arms of the gnathos. The gnathos originates from near the dorsal juncture between the vinculum and the base of the uncus. Vincular arms may be present or absent (the only Cicinnini group to have this character vary).
99 The year of description for C. incerta was incorrectly given as 1877 by
100 Cicinnus latris was described from an indeterminate number of specimens from two localities in Costa Rica. Considering that syntypes and different localities are involved in the type series representing C. latris, we designate a lectotype for the male specimen labeled as the “type” in the
101 The name melsheimeri was published twice by Harris in the same year (
102
103 As previously mentioned in Annotation 96, the type locality of C. orthane may be erroneous. Mimallonidae are exceedingly rarely collected in Chile, with only one other record of the family reported by
104 Cicinnus solvens was described from three syntypes, two male and one female, from two different localities in Panama. Considering that syntypes and different localities are involved in the type series representing C. solvens, we designate a lectotype for the male specimen labeled as the “type” in the
105 Cicinnus tuisana was described from an indeterminate number of specimens from two localities in Costa Rica. Considering that syntypes and different localities are involved in the type series representing C. tuisana, we designate a lectotype for the male specimen labeled as the “type” in the
106 Group 2 (Cicinnus s. l., in part) contains species recognizable by hyaline patches on the forewings or on both fore and hindwings, as well as by the absence gnathos arms and the presence of vincular arms in the male genitalia. These taxa will likely eventually be placed in a revalidated Gonogramma once molecular phylogenetics can more robustly determine the relationships within Cicinnini.
107 The first author dissected a topotypical male specimen of C. belaria comb. n. (St Laurent diss.: 3-15-16:1 in
108 Cicinnus corallina was described from three syntypes from Colombia and Panama. Because syntypes and specimens from different localities are part of the type series of C. corallina, we designate a lectotype for the male specimen labeled as the “type” in the
109 As for Menevia plagiata (Annotation 5), the original primary type of C. despecta is lost, along with many types described by Walker originating from the Fry collection (
110 Cicinnus curtisea potentially represents a valid species considering the geographic separation of the type localities of C. curtisea and C. despecta (see also Annotation 109). The type specimen(s) of C. curtisea is/are not known to us, however, other specimens from Buenos Aires are distinct from those from southeastern Brazil.
111 Cicinnus sanguinolenta is a clear synonym of C. despecta, see also Annotation 109.
112 Cicinnus sachinius is potentially a valid species considering the geographic range and morphological differences between the type of C. sachinius and C. despecta, see also Annotation 109.
113 Cicinnus eminens was transferred to Cicinnus from Psychocampa by
114 Cicinnus roscida syn. rev. was synonymized with C. externa in
115 Although C. manicora comb. n. displays valva and phallus shapes rather unique to this species, the presence of vincular arms, absence of gnathos arms, and presence of hyaline patches on the forewings all are clear indicators of Cicinnus s. l. Group 2, and we therefore transfer this species accordingly from Psychocampa, where it was placed apparently due to overly narrowed wings and sphingiform shape, characters that are superficially similar to some Psychocampa. All of our morphological phylogenetic analyses, as well as ongoing molecular phylogenomics that include this species (St Laurent et al. in prep.), support this transfer, consistently placing C. manicora within a broader Cicinnus s. l. clade.
116
117 Group 3 (Cicinnus s. l., in part) contains species recognizable by hyaline patches on both the forewings and hindwings, as well as by the presence of gnathos and vincular arms in the male genitalia. Most species in this group were either already described in Cicinnus, or were transferred to Cicinnus from Psychocampa by
118
119 Cicinnus gaia displays bizarre, atrophied valvae, seemingly making this species appear not to be part of Cicinnus s. l. Group 3. However, all characters unique to Group 3 are present in C. gaia, including vincular and gnathos arms, and hyaline patches on the fore and hindwings. Tornal diffusions that are present in the above species are also present in C. gaia, but are less discernable due to the lack of contrast in this more darkly colored moth.
120 Cicinnus bibula comb. n. is a very rare species in natural history collections. We are aware of only seven specimens in collections, and all appear to be female, although one in the
121 The revived placement of Cicinnus lemoulti comb. rev. in Cicinnus remains uncertain. Externally, C. lemoulti is similar to Group 2 of Cicinnus s. l., particularly C. externa, and displays the typical forewing hyaline patch of this group. However, genitalia are unusual in this species, as it has a broadly triangular uncus, irregularly shaped valvae (Supplementary File 3: Plate 5, left ¾ of genitalia shown), and a phallus with a broad, ridged hammerhead-like dorsal projection (Supplementary File 3: Plate 4). The gnathos, however, is typical of Cicinnus s. l. Regardless, C. lemoulti most certainly does not belong in Psychocampa based on hyaline patch presence and genital morphology. Furthermore, all of our morphological phylogenetic analyses support this transfer, consistently placing C. lemoulti within a broader Cicinnus s. l. clade.
122 Cicinnus mawaja comb. n. is a particularly rare mimallonid, about which we know very little. We have been able to examine the genitalia of two male specimens, genitalia vial NHMUK010402304 and via Daniel Herbin (pers. comm.). The gnathos morphology, absence of vincular arms, and presence of unique saccular extensions (Supplementary File 3: Plate 4) are confounding structures that do not, together, fit any of our groups of Cicinnus s. l., and in some ways actually appear more similar to the species belonging to the Roelmana + Isoscella clade. Based on morphology, this species does not belong to Psychocampa where it has long been placed. Due to the lack of typical genitalia characters of Psychocampini, and presence of vincular apodemes (an apomorphy of Cicinnini) we preliminarily transfer this species to Cicinnus s. l. to rectify the tribal placement of this enigmatic taxon. All of our morphological phylogenetic analyses support this preliminary transfer, consistently placing C. mawaja within a broader Cicinnus s. l. clade. Ongoing molecular phylogenomics of Mimallonidae, which includes C. mawaja, support the inclusion of this taxon within the genus that will likely eventually be defined as Gonogramma, see Annotation 97. This is surprising considering the absence of hyaline patches on the wings, a character observed in all other Cicinnus s. l.
123 Originally described as a subspecies of Euphaneta divisa, E. romani was raised to full species status by
124 Isoscella ventana was treated as belonging to Psychocampa by
125 Long considered monotypic, Roelmana was found to likely include several disparate species by
126 Of all species previously and here transferred to Roelmana, R. beneluzi comb. n. displays the most divergent male genitalia, and may eventually prove to be more appropriately placed in a separate new genus. We hesitate to do so here pending ongoing molecular phylogenetic work that will likely be able to include R. beneluzi. The gnathos shape however, is consistent with other taxa placed in Roelmana here and in
127 Externally, R. brasiliensis comb. n. is nearly identical to the rare R. fenestrata comb. n., and in the original description of R. brasiliensis this similarity was unfortunately not addressed. Roelmana fenestrata is only known from the female type (erroneously reported and labeled as a male), collected in Castro, Paraná, Brazil, a locality that sits between Cerrado and Mata Atlântica. Roelmana brasiliensis is a Cerrado endemic, therefore it is possible that these two names are synonymous. However, the hyaline discal marking on the hindwing is closer to the postmedial line in R. fenestrata than in females of R. brasiliensis. Pending the availability of male specimens from near the type locality of R. fenestrata, and the loss of the type genitalia of R. fenestrata, it is not possible to reliably synonymize these names at this time. All morphological phylogenetic analyses consistently recovered R. brasiliensis as sister to R. vitreata + R. pluridiscata, thus supporting this transfer of R. fenestrata which could not be coded for morphology for the abovementioned reasons.
128 Roelmana doralica comb. n. is externally most similar to R. pluridiscata and R. vitreata, the three of which form an apparent species complex that is widely distributed throughout South America. We hereby transfer R. doralica and R. vitreata to Roelmana from Psychocampa as per
129 As discussed in the original description of R. laguerrei, R. prominens comb. n. is most similar to this species than any other in Mimallonidae, due to the purplish-blue coloration and male genitalia characteristics.
Maximum likelihood tree built using IQ-TREE with a backbone constraint (Suppl. material
Ventral view of male genitalia of type species of genera belonging to Zaphantinae, incertae sedis, and Aurorianinae. Phallus/juxtal complex is figured below the genitalia in the lateral view. 2 Zaphanta infantilis, St Laurent diss.: CPC 4 [reused with permission from
Ventral view of male genitalia of type species of genera belonging to Mimalloninae. Phallus/juxtal complex is figured below the genitalia in the lateral view, except where noted. 8 Eadmuna esperans, St Laurent diss.: 3-17-18:6 (CRAS) 9Macessoga cf fabia, St Laurent diss.: CPC 1 (CRAS) 10 Mimallo amilia, St Laurent diss.: 10-5-17:8, phallus oriented dorsally [reused with permission from
Ventral view of male genitalia of type species (except where noted) of genera belonging to Lacosominae: Alheitini. Phallus/juxtal complex is figured below the genitalia in the lateral view. 12 Adalgisa croesa, St Laurent diss.: 9-20-17:1 (CRAS) 13 Alheita rionica, St Laurent diss.: 3-27-17:2, not the type species of Alheita [reused with permission from
Ventral view of male genitalia of type species of genera belonging to Lacosominae: Trogopterini and Lacosomini. Phallus/juxtal complex is figured below the genitalia in the lateral view. 18 Reinmara enthona, St Laurent diss.: 5-18-16:1 (
Ventral view of male genitalia of type species of genera belonging to Druenticinae: Druenticini. Phallus/juxtal complex is figured below the genitalia in the lateral view. 22 Druentica partha, St Laurent diss.: 3-17-18:3 (MGCL) 23 Micrallo minutus holotype, St Laurent diss.: 10-21-15:1 [reused with permission from
Ventral view of male genitalia of type species (except where noted) of genera belonging to Druenticinae: Luramini. Phallus/juxtal complex is figured below the genitalia in the lateral view. Figures are reused with permission from
Ventral view of male genitalia of type species of genera belonging to Cicinninae: Psychocampini and Bedosiini. Phallus/juxtal complex, when excised, is figured below the genitalia in the lateral view. 29 Biterolfa althea, St Laurent diss.: 11-16:11 [reused with permission from
Ventral view of male genitalia of type species of genera belonging to Cicinninae: Cicinnini. Phallus/juxtal complex, when excised, is figured below the genitalia in the lateral view. 33 Aceclostria mus, St Laurent diss.: 10-8-14:2 (
Ventral view of female genitalia of type species (except where noted) of genera belonging to Zaphantinae, incertae sedis, and Aurorianinae. 40 Zaphanta infantilis, genitalia vial
Ventral view of female genitalia of type species (except where noted) of genera belonging to Mimalloninae. 45 Eadmuna esperans, St Laurent diss.: 2-12-16:1 (
Ventral view of female genitalia of type species (except where noted) of genera belonging to Lacosominae: Alheitini. 49 Adalgisa croesa, St Laurent diss.: 4-1-18:1 (CRAS) 50Alheita cf pulloides, St Laurent diss.: 10-10-17:3, not the type species of Alheita (MGCL) 51 Tarema rivara, St Laurent diss.: 3-14-16:8 [reused with permission from
Ventral view of female genitalia of type species (except where noted) of genera belonging to Lacosominae: Trogopterini and Lacosomini. 54 Reinmara enthona, D. Herbin genitalia diss.: H. 1103 [reused with permission from
Ventral view of female genitalia of type species (except where noted) of genera belonging to Druenticinae. 58 Druentica partha, St Laurent diss.: 3-31-18:5 (MGCL) 59Procinnus cf acuta, St Laurent diss.: LEP50136, not the type species of Procinnus (MGCL) 60 Micrallo minutus, St Laurent diss.: 10-21-15:2 [reused with permission from
Ventral view of female genitalia of type species (except where noted) of genera belonging to Cicinninae: Psychocampini and Bedosiini. 65 Biterolfa althea, D. Herbin genitalia diss.: H600 [reused with permission from
Ventral view of female genitalia of type species (except where noted) of genera belonging to Cicinninae: Cicinnini. 69 Aceclostria mus, St Laurent diss.: 3-31-18:8 (CRAS) 70 Aleyda accipiter, St Laurent diss.: 4-24-17:1 [reused with permission from
Adult specimens of type species of genera belonging to Zaphantinae, incertae sedis, and Aurorianinae. 75 Zaphanta infantilis, male [photo A Giusti] (
Adult specimens of type species of genera belonging to Mimalloninae. 85 Eadmuna esperans, male (MWM) 86 E. esperans, female (MWM) 87Macessoga cf fabia [photo S Naumann] (ISEZ) 88M. cf fabia, female (
Adult specimens of type species of genera belonging to Lacosominae: Alheitini. 93 Adalgisa croesa, male (CGCM) 94 A. croesa, female (CGCM) 95 Arianula haxairei, male [photo R Lahousse] (
Adult specimens of type species of genera belonging to Lacosominae: Trogopterini and Lacosomini. 107 Reinmara enthona, male [reused with permission from
Genitalia of Citralla rumina a ventral b lateral c phallus. Roman numerals refer to characters discussed in the text. 118 Male, Costa Rica, Heredia, Chilamate, Finca Selva Verde, St Laurent diss.: 9-20-17:4 (MGCL) 119 Female [ventral], holotype, Panama, Volcan de Chiriquí, 2000-3000 ft, genitalia vial
Adult specimens of type species of genera belonging to Druenticinae: Druenticini. 121 Druentica partha, male [reused with permission from
Male genitalia of Lepismalla montagnaniae a ventral with valvae in natural position b ventral with valvae spread c ateral d phallus. Roman numerals refer to characters discussed in the text. Note that uncus is twisted to the left in Fig.
Adult specimens of type species (except where noted) of genera belonging to Druenticinae: Luramini, note that two species of Lurama are figured to better illustrate this genus. 133 Lurama penia, male [reused with permission from
Adult specimens of type species of genera belonging to Cicinninae: Psychocampini. 138 Biterolfa althea, male [reused with permission from
Adult specimens of type species of genera belonging to Cicinninae: Bedosiini. 142 Bedosia fraterna, male (MGCL) 143 B. fraterna, female [photo P Collet] (
Adult specimens of type species of genera belonging to Cicinninae: Cicinnini. 146 Aceclostria mus, male [photo S Naumann] (
Adult specimens of type species of genera belonging to Cicinninae: Cicinnini. 153Cicinnus cf orthane, male [reused with permission
Lectotypes designated in the present work for incertae sedis and Lacosominaea dorsal b ventral. See annotations in Section 4 for complete label data for all lectotypes. All specimens are male 159 Hydrias amaryllis Schaus (= Tolypida amaryllis) 160 Trogoptera althora Schaus 161 Adalgisa croesa Schaus 162 Perophora pulloides Dognin (= Alheita pulloides) 163 Lacosoma briasia Schaus. Scale bar: 1 cm.
Lectotypes designated in the present work for Lacosominae and Cicinninaea dorsal b ventral. See annotations in Section 4 for complete label data for all lectotypes. All specimens are male. 164 Lacosoma diederica Schaus 165 Lacosoma raydela Schaus 166 Cicinnus lacuna (= Psychocampa lacuna) 167 Cicinnus corallina Dognin 168 Cicinnus latris Schaus. Scale bar: 1 cm.
Lectotypes and neotype designated in the present work for Cicinninaea dorsal b ventral. See annotations in Section 4 for complete label data for all lectotypes and the neotype. All specimens are male. 169 Cicinnus solvens Dyar, lectotype 170 Cicinnus tuisana Schaus, lectotype 171 Mimallo despecta Walker (= Cicinnus despecta), neotype. Scale bar: 1 cm.
This study would not have been possible without the kind assistance of many individuals in the various institutional and private collections in Canada, Europe, the United States, and South America. The following people permitted access to or photos from the collections utilized in this study: Courtney Richenbacher (
Table 1. Morphological character matrix for Mimallonidae and two outgroups
Data type: species data
Explanation note: Characters 1 to 25 are male genitalia, 26 to 31 are female genitalia, and 32 to 55 are external morphological characters. See Suppl. material
Explanation of the 55 characters and their states that were coded for the present study
Data type: species data
Figures denoting characters coded in Suppl. material
Data type: species data
Constraint tree in Newick format, utilized in our constrained ML analysis
Data type: phylogenetic data
Explanation note: The constraint tree was derived from the dataset2_PART2 analysis of
Unconstrained morphological ML analysis conducted in IQ-TREE
Data type: phylogenetic data
Explanation note: Support values refer to SH-aLRT/UFBoot.
Constrained morphological ML analysis conducted in IQ-TREE
Data type: phylogenetic data
Explanation note: Support values refer to SH-aLRT/UFBoot. Constraint used can be found in Suppl. material
Majority rule consensus tree of the five most parsimonious trees inferred in TNT
Data type: phylogenetic data
Explanation note: Support values refer to 100 jackknife replicates.