Athyrma Hubner 1823
Creators: name: Adams, Morton S.
name: Mccabe, Timothy L.
Publisher: name: Zenodo
Year: 2022
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Lepidoptera
Noctuidae
Athyrma

description: Athyrma Hübner, [1823] 1816
Verzeichniss bekannter Schmettlinge. Augsburg. p. 267.
Type species: Phalaena adjutrix Cramer, 1780, Uitlandsche Kapellen 3:144, pl. 272, figs. E, F, by monotypy. Type: Female. RMNH. INS.1103916. Depository: Naturalis Biodiversity Center, Leiden, Netherlands.
Athyma; Pagenstecher, 1888. Jahrbüchen des Nassauischen Vereins für Naturkunde 41:149. [Misspelling]
Original generic description: Hübner [1823] 1816, p. 267.
“Die Schwingen mit unterschiedlichen schwarzen Fleschen bezeichnet.”
Translation: The wings marked with unique black spots.
Species included by Hübner in original description:
Athyrma adjutrix Cramer. 1780
Athyrma ganglio Hübner, [1823] 1816
Athyrma interpuncta Hübner, [1823] 1816. Nomen nudum [manuscript name vide Hemming 1937, p. 160]
Re-description (based on type of genus Athyrma adjutrix Cramer, 1780).
Adult habitus. Wingspan: 35–44 mm. Females average larger. Head prominent, eyes large and naked, not lashed; antennae of both male and female filiform (Fig. B) with straight segmental bristles slightly longer than the width of the shaft, except in the ganglio complex in which the male antennae are serrulate (Fig. C); proboscis robust, labial palpi upturned to vertex or higher, third segment one half length of second, closely scaled without a brush; frons free of scales, a patch of white scales below the eyes; thoracic vestiture mouse-brown with a mixture of white and brown, mostly 3-toothed scales, no crest but a slight dorsal ridge; tegulae of broad toothed scales, concolorous with thoracic vestiture; abdomen concolorous with thorax, dorsal tufts on 1st and 2nd segments, white scale tuft at tip of male abdomen. Legs. All legs heavily scaled with a tuft of longer scales on the proximal end of the tibia; no tibial spines on any legs; foretibia without spurs; mid-tibia with a distal pair of spurs; male mid-tibia with long hairs from the base, but without the retaining groove found in Eulepidotinae; hind tibia with a pair of spurs both distally and proximally; on the proximal pair the smaller member of the pair greater than half the length of the longer; on the distal pair the smaller member approximately one third the length of the longer, the tarsi bicolored, paler at the joints, tarsal spines in a row of three, claws unmodified. Wings. (Fig. E) Appearance of the forewing distinctive. Ground color varying from dull mouse-brown to pink-tinged, some species with a fine dusting with silver scales; apex of the forewing slightly developed with a slight bulge in the middle of the forewing margin; reniform and subreniform stigmata fused into a crude "dog head" likeness, this pattern reduced particularly in the females of some species; claviform always present and delta shaped; post medial line a series of dots, often reduced but the two dots at the costa always present. In A. tuberosa Felder & Rogenhofer, 1874, and in A. brigittae sp. nov., a straight subterminal line present and conspicuous; a trace of white on inner aspect of reniform, possibly representing medial line; a row of terminal spots; fringe concolorous with forewing ground color; costa with marks at the basal line, antemedial (a.m.) line and the medial line. Hind wing a uniform mouse-brown with minimal patterning, i.e. no spots or lines.
Venation. (Fig. A) Venation of the fore and hind wing typical of Erebidae, except for distortion of the male forewing by a unique sexual modification (alar fovea) below the costa (C) about midway along its length; ventral surface of fovea with a conspicuous patch of pale-colored scales (Fig. D); forewing costa convex above fovea while subcosta (Sc) similarly distorted below fovea into a concave curve; radial vein (R1) originating from middle of the discal cell and gently curving beneath the fovea; areole present below fovea; R2 arising very near the end of the upper border of the areole; R3 and R4 stalked from the end of the areole; R3 terminating at the costa before the apex of the forewing; R4 terminating at the apex; R5 originating from end of the areole; first medial vein (M1) arising from middle of the lower border of the areole; M2 arising very close to M3 from the cell, distant from M1; M1 appearing quadrifid with the cubital veins (Cu1 and Cu2); middle discocellular cross-vein absent; one anal vein (A2) present. Hind wing: Sc and Rs fused near the base; areole absent; cubitus vein appearing quadrifid with Rs and M1 branching from the apex of the discal cell; no middle discocellular cross-vein; two anal veins but no trace of vestigial R1 at the base. Male frenulum with one stout bristle; female frenulum with two bristles (three bristles commonly encountered in the Erebidae).
Male genitalia. (Fig. F). Expanded base of uncus with a simple, dorsal, round-tipped spur; tip of the uncus tridentate, a second simple spur approximately midway on its ventral aspect; anal tube (scaphium plus subscaphium (Pierce, 1909)) membranous. The scaphium sclerotized in most Erebidae (Fibiger and Lafontaine, 2004; Lafontaine and Schmidt, 2010). Valve: cucullus rounded with a patch of nondeciduous spinules on its costa, these not in a single marginal row; clasper "thumb-like" apparently to fit into "sockets" of the female subgenital sternite; digitus, ampulla, and editum absent; juxta elongate, appearing in two distinct halves; sacculus and vinculum unmodified. Phallus a broad, straight tube proportionately large for the size of the genital capsule; vesica with four diverticula, all but one with an apical patch of non-deciduous spines; patch of spines on each lobe approximately the same size, the lobes extending at oblique angles from the central core. Eighth abdominal segment modified as a supra-genital plate with a bicuspid area of sclerotization.
Female genitalia. (Fig. G). Intersegmental membrane between last (8th) sternite and subgenital plate conspicuously modified with a pair of heavily sclerotized hemispheric sockets, which do not appear sensory. These intersegmental sockets developed from anterior edge of eighth sternite, a plate-like portion separated from the body of the sternite. Bursa copulatrix functionally divided into recognizable ostium bursae, ductus bursae, corpora bursae, and appendix bursae; ostium bursae and ductus bursae unmodified; corpora bursae membranous except where variously modified by an area of sclerotization; appendix bursae not developed in most species but defined by the point of attachment with ductus seminalis.
Larva. (Fig. H). Ultimate instar a typical Erebidae semi-looper, with abdominal prolegs reduced in size on segments three and four; general habitus similar to species of Zale Hübner, 1818; head and body with middorsal broad stripe with undulating margins typical of many Zale species. When "looping" or provoked, 1st and 2nd abdominal segments form a hump, with a conspicuous black or yellow and red middorsal transverse band (eyespot); 5th and 6th abdominal prolegs and terminal (anal) proleg unmodified.
Pupa. Enclosed within a leaf; with surface flocculence often seen in Erebinae.
Food plants. Known larval food plant records are in the Dilleniaceae and Vitaceae (Janzen & Hallwachs, 2019).
Etymology. Athyrma is Greek for “without an entrance”. However, there is a familiar Greek toy, popular in Hübner’s time, called an Athyrma. It is a figurine in the shape of an animal. Perhaps Hübner’s choice of the name was a subtle reference to the dog head pattern on the forewing of most members of the eponymous genus.
Behavioral observations. Three species of flying Athyrma (A. tuberosa, A. adjutrix, and A. dormitrix) were observed at a mercury vapor lamp in French Guiana. No wing-produced sound was detected from these three species despite numerous observations (TLM). Heliocheilus paradoxus (Grote, 1865), with a similar forewing fovea, but this fovea devoid of scales, was heard making ticking sounds in flight around collecting lights in Arizona (TLM).
In Athyrma the fovea is crowded with specialized scales that persist even in worn individuals. These sex scales are paler colored than ordinary wing scales. The fovea is densely packed with long-stemmed scales, which are broadly rounded at the tips. These scales are sheltered by the foveal pocket. A hook-like process of unknown function, buried completely by the sex scales, hangs from the middle of the leading edge of the fovea (Fig. A). We propose this alar fovea is a character shared by the common ancestor of the genus Athyrma and exclusively by its descendants.
In French Guiana an adult A. dormitrix was observed feeding on rotting fruit three meters from the ground (TLM). The fruit of this unidentified tree was persistent. Athyrma adults have not been observed at fermented banana bait in the canopy or subcanopy (TLM).
KEYS TO THE SPECIES OF ATHYRMA
Prior to using the keys, the specimens should be sorted to sex and male antennal modification.
Dichotomous keys are as follows: Key A, male habitus; Key B, female habitus; Key C, male genitalia of species with serrulate antennae; Key D, male genitalia of species with filiform antennae; Key E, female genitalia.
Key A: Male Athyrma based on habitus but including known distributions and characters of the vesica where necessary. The forewing pattern has been illustrated (Fig. E) and all the salient features are labelled. The large mark near the wing base, distally bordered by the antemedial [a.m.] line, will hereafter be called the “delta mark”. The irregular mark that encompasses the region of the reniform and often extending to the subreniform will hereafter be called the “dog head”. When fully developed, it takes the shape of a dog’s head and neck.
1 Subterminal line straight and distinct, from near forewing apex to outer two thirds of hind margin..................... 2
- Subterminal line absent or indistinct, never a solid straight line, may be represented by a series of spots................. 3
2 Costa mark at wing base present or absent; delta mark with less pronounced “shelf” (dissection required; Fig. 19); vesica with the two spine-bearing diverticula straight and directed distally (away from phallus); Central America............. brigittae
- Costa mark at wing base present; delta mark with more pronounced “shelf” (dissection required; Fig. 18) vesica with two spinebearing diverticula twisted in an incomplete spiral and directed proximally (towards the phallus); South America.. tuberosa
3 Antennae serrulate (Fig. C) with bristle tips reflexed forward so that the bristles are so proximate as to nearly touch each other; dog head usually reduced.............................................................................. 4
- Antennae filiform (Fig. B) with bristles that have their tips nearly straight or with entire bristle very gently curved throughout length, well-separated; dog head variable.................................................................. 8
4 Dog head with a well-developed neck; delta mark with distal edge bulging; Peru .......................... tapichensis
- Dog head lacks a neck or has a reduced neck, but if present then delta mark with distal edge straight – not bulging; widespread........................................................................................ ... 5
5 Dog head typically lacks a neck; portion of forewing, adjacent and beneath dog head, with darker brown scales (dissection required); vesica with only one patch of cornuti (in the form of short, non-deciduous spines); Florida, Cuba, Central America....................................................................................... ciboney (in part)
- Dog head often has start of a neck; region around and below reniform mark is pale (dissection required); vesica with two separate patches of cornuti; Central and South America....................................................... 6
6 Central America (dissection required); vesica with five diverticula (Fig. 66), the three non-spine bearing, basal diverticula conjoined at base and all lack apical spinules (use high magnification).............................. ciboney (in part)
- South America (dissection required); vesica with four or five diverticula, one of the three basal diverticula not conjoined with the others (one very distant) and two (of the three most basal diverticula) terminate in spinules (use high magnification).... 7
7 Northern South America; (dissection required, Fig. 65) non-spine bearing diverticulum longer than spine bearing diverticula.............................................................................................. ganglio
- Northern and Central South America; (dissection required, Fig. 69) non-spine bearing diverticula are shorter than spine bearing diverticula................................................................................... ... orbana
8 Costa at base without black mark or this mark is faint (this basal mark nearly touches thorax); black dog head, when present, with comparatively thin neck; French Guiana and Brazil ................................................. nodosa
- Costa at base with distinct dark mark (this basal mark nearly touches thorax); dog head fully expressed with a thick neck.. 9
9 Slightly elongate white spot at base of skull of dog head appearing indented into margin............................ 10
- Slightly elongate white spot at base of skull of dog head appearing confluent with margin........................... 11
10 A.m. costal mark more than double the width of the basal mark; northern coast of South America ............... dormitrix
- A.m. costal mark usually only slightly larger than basal mark; Panama ....................................... cryani
11 Distal margin (a.m. line) with delta mark bulging at middle (Figs. 1–2) .......................................... 12
- Distal margin (a.m. line) of delta mark straight (Fig. 10) or slightly bulging...................................... 15
12 Delta mark bulging to form a square (Fig. 5); Peru ..................................................... .. yasuni
- Delta mark slightly bulging at middle or straight........................................................... 13
13 Delta mark slightly bulging at middle; widespread in South America............................................ 14
- Delta mark straight; coastal Brazil ............................................................ itatiaia (in part)
14 Dog head with a thick neck, delta mark bulge is extreme [dissection required] three spine patches on vesica...... .. adjutrix
- Dog head with neck normal, delta mark smoothly and slightly bulging [dissection required]; one spine patch on vesica ................................................................................................... urbanae
15 “Bridge” of dog head “nose” usually without adjacent contrasting pinkish or white area unless ground color mottled; South America ............................................................................................ 16
- “Bridge” of dog head “nose” usually with an adjacent pale pink to white mark; ground colors variable; Caribbean, Central America, Florida ..................................................................................... 17
16 Coastal Brazil [dissection required],with two patches of spines on vesica............................. itatiaia (in part)
- Ecuador [dissection required], with one patch of spines on vesica....................................... .. svensoni
17 A.m. line slightly bulging; ground color mottled brown and lacking pale white or pink “bridge” of dog head “nose”; Central America nearly to Mexico ........................................................................ romacki
- A.m. line straight; ground color violet gray and typically with pale white or pink area above “bridge” of dog head “nose”; Central America including Mexico, Florida, and northern Caribbean.................................... fakahatchee
Key B- Female Athyrma based on forewing markings and distribution. The female’s dog head is often so reduced that it resembles a large, squat comma.
1 Subterminal line solid, straight, and distinct, originating from close to apex and extending to outer two thirds of anal margin.................................................................................................. ... 2
- Subterminal line a series of broken dots or absent ............................................................ 3
2 South America; costal mark present or absent; delta mark with more pronounced “shelf” (Fig. 35).............. tuberosa
- Central America; costal mark present at base; delta mark with less pronounced “shelf” (Fig. 36)................ brigittae
3 No black dog head (rarely a faded brown dog head); typically two tiny black circles marking edge of p.m. line........... 4
- With large black dog head often reduced to the shape of a fat comma but sometimes complete........................ 5
4 Costal mark occurring at approximate position of origin of p.m. line (Fig. 31); forewing an even, smooth, pale brown.................................................................................................... orbana
- No costal mark near origin of p.m. line origin (Figs. 33–34); forewing heavily dusted with blue-gray scales...................................................................................................... nodosa (in part)
5 Dog head well-developed and includes a “neck” ............................................................. 6
- Dog head reduced to a circular comma-shaped mark or absent – rarely with weak extension (a partial “neck”) ........... 12
6 Elongate white spot at base of skull of dog head present and indented; forewings mottled in appearance................ 7
- Elongate white spot at base of skull of dog head present or absent, but when present not appearing indented, but confluent with margin of dog head; forewings smooth in appearance......................................................... 8
7 South America; phenotypically similar (dissection required)............................................ dormitrix
- Central America; phenotypically similar (dissection required).............................................. cryani
8 Delta mark bulging at middle on outside edge (a.m. line); black markings present on costa; South America... adjutrix in part
- Delta mark straight or bulging on distal edge, if bulging then from Central America, otherwise widespread.............. 9
9 Thickened black costal markings absent (may be a trace or a thin line at the a.m., p.m., and basal lines; French Guiana ............................................................................................ nodosa (in part)
- Thickened black costal markings present at basal and a.m. lines............................................. ... 10
10 Delta mark bulging at middle on distal edge (a.m. line).................................................. adjutrix
- Delta mark straight on distal edge....................................................................... 11
11 “Bridge” of “nose” of dog head same color as ground; “neck” of dog head present or absent, but quite thin if present..... 12
- “Bridge” of “nose” of dog head typically pale pink or white-filled; but if not pink or white filled then “neck” of dog head well developed........................................................................... fakahatchee (in part)
12 South America...................................................................................... 13
- Central America/Florida/ Jamaica ....................................................................... 15
13 Dog head “neck” fully developed.................................................................... urbanae
- Dog head “neck” thin or absent .......................................................................... 14
14 Dog head “neck”, if present, tapering................................................................. ganglio
- Dog head “neck” may be thin, but continues to “breast”..................................................... 16
15 Subterminal line usually a complete series of dots; reddish color below dog head extends to inner margin of forewing; [dissection required] (Fig. 76); Central America................................................................ romacki
- Subterminal line incomplete, often with little or no trace of dots; reddish color below dog head confined to vicinity of dog head; [dissection required] (Fig. 81); Florida, Caribbean, and Central America......................... fakahatchee (in part)
16 “Neck” of dog head complete ........................................................................... 17
- “Neck” of dog head reduced or absent, typically reduced to a squat “comma”.................................... 19
17 Dog head with “collar of neck” not constricted........................................................ .. itatiaia
- Dog head with “collar of neck” constricted................................................................ 18
18 Dog head with short “nose” (Fig. 30) ............................................................. tapichensis
- Dog head with long “nose” (Fig. 25)............................................................... svensoni
19 Costal p.m. mark indistinct [dissection required]; strap-like signa above middle of corpora bursae; northern South America................................................................................................ ganglio
- Costal p.m. mark well-defined and obvious [dissection required]; strap-like signa near middle of corpora bursae; Central America and Caribbean.......................................................................... ciboney
Key C- Male genitalia of species with serrulate antennae (largely characters of the vesica); all have claspers that are thumb-like.
1 Vesica with two or three diverticula with apical spine patches; widespread........................................ 2
- Vesica with one (rarely with two) diverticula with apical spine patch; Central America and northern Caribbean................................................................................................ … ciboney (in part)
2 Vesica with three diverticula with apical spine patches: basal one with short coarse spines, middle patch with fine spines, distal patch with large spines......................................................................... tapichensis
- Vesica with two diverticula with apical spine patches........................................................ 3
3 Vesica with only two diverticula, both with apical spine patches............................................ orbana
- Vesica with four or five diverticula, two with apical spine patches............................................... 4
4 The two diverticula with apical spine patches with a diverticulum between (Fig. 65); South America.............. ganglio
- The two diverticula with apical spine patches with no diverticulum between (Fig. 66); Central America.... ciboney (in part)
Key D- Male genitalia of species with filiform antennae (largely based on characters of the vesica).
1 Dorsal forewing with straight and prominent subterminal line ................................................... 2
- Dorsal forewing lacks distinctive subterminal line........................................................... 3
2 Clasper extends nearly to the tip of the valve; spines of both apical spine patches antrorse from phallus (Fig. 73); South America..................................................................................... tuberosa
- Clasper extends short of the tip of the valve; spines of both apical spine patches retrorse from phallus (Fig. 74); Central America.............................................................................................. brigittae
3 Vesica with 4–5 diverticula, with three apical spine patches; clasper thumb-like.................................... 4
- Vesica with 1–2 diverticula,with two or fewer spine patches; clasper variable..................................... 5
4 None of the spine patches on forked diverticulum (Fig. 57 ............................................... adjutrix
- One of the spine patches on forked diverticulum (Fig. 58)................................................ romacki
5 Vesica with two patches of spines........................................................................ 6
- Vesica with one patch of spines.......................................................................... 8
6 Vesica with two sets of spines pointing in opposite directions............................................. itatiaia
- Vesica with two sets of spines pointing in the same direction ................................................... 7
7 Apical spine patches on the same diverticulum; clasper thumb-like......................................... cryani
- Apical spine patches on separate diverticula; clasper finger-like........................................... nodosa
8 With an inconspicuous patch of a few tiny spines that are overlapping the phallus (Fig. 61).................... .. yasuni
- With a patch of large spines that are retrorse to the phallus but does not overlap the phallus ........................... 9
9 Apical spine patch on diverticulum is adjacent to the tip of the phallus.......................................... 10
- Apical spine patch is distant from the tip of the phallus...................................................... 11
10 Apical spines long (longer than phallus is wide) and numerous (typically 7 large, long spines).................. svensoni
- Apical spines short (much shorter than phallus is wide), and few (typically 4 stout and 1 or 2 delicate, short spines). urbanae
11 Spine-bearing diverticulum retrorse to phallus (Fig. 64); clasper finger-like.............................. fakahatchee
- Spine-bearing diverticulum antrorse to phallus (Fig. 59); clasper thumb-like............................... dormitrix
Key E- Female genitalia (note: several species are not reliably separated by female genitalia). The female of A. yasuni is unknown.
1 Ductus bursae with sclerotized area on appendix bursae that may or may not extend onto corpora bursae proper (Figs. 77–81); this development is to the right or behind (Fig. 77) the ductus bursae............................................ 2
- Ductus bursae without sclerotized area extending to the right................................................ ... 6
2 Sclerotized area on dorsal surface of corpora bursae................................................... dormitrix
- Sclerotized area on the right of corpora bursae, often so extreme as to develop into an appendix bursae................. 3
3 Sclerotized area lacks any extension to the right and appears as a thickening of the juncture between the ductus bursae and the corpora bursae................................................................................. urbanae
- Sclerotized area with an extension to the right.............................................................. 4
4 Appendix bursae without sclerotized area............................................................. svensoni
- Appendix bursae heavily sclerotized throughout............................................................ 5
5 Sclerotized appendix bursae adheres closely to outline of the corpora bursae (Fig. 78)........................... cryani
- Sclerotized appendix bursae a conspicuous lobe to the right (Fig. 81)................................... fakahatchee
6 Left edge of the corpora bursae with a distinct strap-like sclerotized area that wraps around left side of corpora bursae (the following species are not reliably separated by female genitalia)................................................ 7
- Left edge of the corpora bursae lacks a sclerotized area....................................................... 10
7 Corpora bursae with an uplifted bulge to the right (Figs. 84–85) (the ductus seminalis is attached so this bulge is technically an appendix bursae) ...................................................................................... 8
- Appendix bursae evenly curved to the right, the ‘uplifted bulge’ effect not apparent ................................. 9
8 Strap-like sclerotized area lightly chitinized (Fig. 85); Peru ............................................ tapichensis
- Strap-like sclerotized area heavily chitinized (Fig. 84); Central America, northern Caribbean, and Florida.......... ciboney
9 Corpora bursae shaped like a banana (Fig. 86).......................................................... orbana
- Corpora bursae only slightly curved (Fig. 82).......................................................... ganglio
10 Corpora bursae a broad irregularly shaped sac with ill-defined slightly sclerotized amorphous patches over most of the corpora bursae............................................................................................. 11
- Corpora bursae not a broad sac, and regular in shape, frequently with a pair of sclerotized areas that rarely cover most of the corpora bursae...................................................................................... 12
11 Corpora bursae almost spherical; South America...................................................... adjutrix
- Corpora bursae heavily indented on lower left (left proximal in ventral view); Central America................. romacki
12 A pair of sclerotized areas, side by side, on the ventral surface of the top half of the corpora bursae.............. brigittae
- Sclerotized area indistinct and irregular, may be up and down but never appearing as a side by side pair ................ 13
13 Sclerotized area in the bottom section of corpora bursae................................................. itatiaia
- Sclerotized area only in the top half of the corpora bursae.................................................... 14
14 Ductus bursae tapering, narrow near ostium........................................................... nodosa
- Ductus bursae broad near ostium.................................................................. tuberosa
description: Published as part of Adams, Morton S. & Mccabe, Timothy L., 2022, A Revision Of The New World Moth Genus Hübner (Lepidoptera: Erebidae), pp. 1-47 in Zootaxa 5105 (1) on pages 3-12, DOI: 10.11646/zootaxa.5105.1.1, http://zenodo.org/record/6332257