Corresponding author: M. Alma Solis (
Academic editor: D. Lafontaine
The Neotropical genus
With respect to their actual biology,
In the present work, we treat newly assembled historical and recent material from the Western Hemisphere. Our purpose is to refine the circumscription and composition of
Pinned specimens were examined with an incandescent light source (reflected light). Male and female genitalic preparations varied, some of those pre-dating this study having accumulated from several sources. Most were prepared following
This work drew in part from an effort to treat taxa with taxonomic and nomenclatural problems identified during preliminary surveys of pyraloids in the extensive Costa Rican collection of D. Janzen and W. Hallwachs. Because the genitalic characters of
The following abbreviations refer to collections from which specimen material forms the basis of this work:
Following the identification of a suite of male secondary sexual characters suspected of diagnosing multiple species pairs, we undertook a preliminary DNA barcoding exploration of two putative species for which relatively recent (~20 year old) material existed. Sequencing was done using standard protocols at the Biodiversity Institute of Ontario (
All characters were equally weighted and coded as unordered. Preference was given to combinations of binary and inapplicable coding schemes over multistate characters. Conspecificity of male and female specimens was inferred by locality when other biological information was unavailable. Preliminary phylogenetic analysis involved parsimony searches via the ratchet routine (island-hopper, 1000 iterations per rep) in Windada/Winclada (1000 iterations per rep;
The scope of our treatment of described
Initial examination of specimens tentatively identified as
A total of 64 characters (56 binary, 8 multistate; 5 head, 13 thoracic, 13 abdominal and tympanal, and 25 male genitalic, and 8 female genitalic) were adduced and coded (below; see Appendix I for the full matrix). Inapplicable and missing data were coded with “-” and “?”, respectively.
0. Ocelli: (0) absent; (1) present
1. Proboscis: (0) reduced, inconspicuous; (1) conspicuous (
2. Frons: (0) of normal contour, evenly rounded; (1) conical or expressed as a small hump; (2) carinate or otherwise modified (
3. Length of labial palpus: (0) extending beyond clypeus; (1) not extending beyond clypeus
4. Maxillary palpi: (0) extending anteriorly beyond frons; (1) not reaching anterior margin of frons
5. Forewing (Rs3, Rs4): (0) bases separate; (1) stalked (
6. Forewing (M1, M2): (0) bases separate; (1) stalked (
7. Medial area: (0) contrast with basal and postmedial areas subtle, almost unicolorous except for lines, spot; (1) contrast between medial area and basal and postmedial areas sharp
8. Forewing coloration: (0) compound, ground color not uniform in any given area (antemedial, medial, postmedial;
9. Concentration of white scales apical to antemedial line: (0) absent, or if present then only diffusely; (1) present (
10. Hindwing (HW) postmedial line: (0) not conspicuous or nearing inner margin; (1) distinct, approaching or reaching inner margin (
11. Distance between postmedial line and wing terminus: (0) narrow (
12. Wing lines: (0) dark on light ground; (1) light on dark ground (
13. HW (M2M3+CuA1): (0) bases separate; (1) stalked (
14. Male hind leg secondary sexual complex consisting of a flattened hind tibial spur with flattened scales and basal tarsus with concave spoon-like modification: (0) absent; (1) present (
15. Dark patch amidst hind tibial scales: (0) absent; (1) present (
16. Tufts of epipleural setae: (0) absent; (1) present
17. Female medial hind tibial spurs: (0) one pair (medial pair absent); (1) two pair (medial pair present)
18. Bullae tympani invaginated in Sternum 2: (0) not (all ingroup taxa;
19. Saccus tympani invagination: (0) short, not beyond puteolus; (1) deep, with posterior ridge, but not prominently invaginated posteriad (
20. Saccus: (0) not prominent (
21. Mesal extent of saccus (@ pons): (0) short (
22. Puteoli: (0) absent or indistinguishable from saccus tympani (all ingroup taxa;
23. Processus tympani: (0) inconspicuous; (1) approximately semi-circular (
24. Fornix, protrusion over venula prima: (0) protruded over slightly, flat; (1) far removed from edge (all ingroup taxa;
25. Fornical ulna: (0) > 90 degrees or low arc (all ingroup taxa;
26. Sclerotization of fornix: (0) light to moderate; (1) heavy (all ingroup taxa;
27. Fornix: (0) robust, broad (
28. Venulae secundae: (0) wide, gently tapered (
29. Tergo-sternal sclerite: (0) present, not elongate (
30. Coremata on 4th abdominal segment: (0) absent; (1) present (
31. Gnathos-ventrotergal rods complex: (0) absent; (1) present (all ingroup taxa;
32. Gnathos, middle process: (0) absent; (1) present (all ingroup taxa;
33. Dorsal ridges of tegumen: (0) absent, split to uncus, or inverted U; (1) cruciate, crossing near uncus (
34. Teguminal sulcus: (0) absent; (1) present (all ingroup taxa;
35. Uncus tip trefoil shaped: (0) absent (
36. Shape of trefoil, if present: (0) reduced, rhomboid (
37. Uncus edges: (0) simple, undifferentiated (
38. Uncus, interior (under-surface: (0) clear, without relief (
39. Valva - outer margin: (0) entire or emarginate, but continuous, such that trajectory of valval membrane continues apically (
40. Glabrous central area of valva: (0) Absent [valva elongate, setose]; (1) truncate - squared or subquadrate (emarginate) (
41. Intra-saccular process: (0) absent; (1) slightly raised bump, flange, or paddle centrally located on inner face of valva (
42. Intra-saccular process, adornment: (0) denticled or rugose (
43. Saccular bend: (0) absent (
44. Saccular margin: (0) angled close to vinculum (
45. Saccular bend angled versus rounded: (0) angled, 90 degrees (
46. Ventro-medial setal comb: (0) absent; (1) present (
47. Localized patch or cluster of ventral, saccular spine-like setae: (0) absent; (1) present (
48. Ventro-marginal setae: (0) absent or rudimentary (
49. Isolation of costal bar: (0) <75% along length of costa (
50. Secondary lobe of valva: (0) absent (
51. Recurved or decumbent setal plume associated with end of costa: (0) absent (
52. Setae arranged in recurved hook-shaped cluster: (0) absent; (1) present (
53. Scales arranged in terminal black dots on male abdomen: (0) absent; (1) present, conspicuous (all ingroup taxa)
54. Phallus - cornuti: (0) absent (
55. Number of cornuti: (0) one (
56. Antrum: (0) present, chalice-like (
57. Colliculum: (0) absent (
58. Narrow, differentially sclerotized band around center of colliculum: (0) absent (
59. Ductus bursae: (0) effectively absent or inconspicuously short (
60. Sclerotization on floor of ductus bursae: (0) absent, indiscernable (
61. Corpus bursae: (0) elongate (
62. Attachment of ductus bursae to corpus bursae: (0) basal (
63. Modification of Sternum 8: (0) absent; (1) present (all ingroup taxa)
After surveying all the described
Habitus of adults.
Head, lateral view.
Wings.
Thoracic and leg structures in
Tympanal organs. Collection and/or dissection numbers follow country of origin; when label data presented elsewhere, annotated as such.
Tympanal organs and male abdominal segments.
Male, female genitalia.
Male, female genitalia.
Male, female genitalia.
Male, female genitalia.
We were unable to discern consistently different characters among
From cladistic analysis eight most parsimonious trees obtain (L=102, CI=71, RI=84), the strict consensus of which (L=108, CI=67, RI=81) is presented (
Strict consensus (L=108, CI=67, RI=81) of eight most parsimonious trees (L=102, CI=71, RI=84) obtained from a cladistic analysis of morphological data with unambiguous character state transformations indicated. Numbered hatch marks on nodes refer to characters undergoing forward changes (solid=unreversed; hollow=reversed). Bremer values are indicated below relevant branches.
The monophyly of
The second major intrageneric grouping, the
Morphologically, this second, perhaps more enigmatic species-group, is less homogeneous than that surrounding the type species of
Brazil.
“
In the species most readily identifiable as
All
Larvae are internal feeders that may induce galls, and pupate within the host. The only known host plant records are in
Key to species of
1 | Forewing generally silvery gray or gray brown with white shading in vicinity of antemedial and postmedial lines, particularly in medial area and at outermost edge of postmedial line; or dark brown with poorly contrasted markings except for postmedial line. Hindwing postmedial line conspicuous, nearing inner margin. Frons conical. Valva simple or reduced, sub-quadrate or emarginate/mildly bilobed; lacking a straight, prominent coastal arm; medial projection or flange arising from within sacculus; apex of costa lacking a tuft or plume, or a fleshy, setose subcostal lobe. Uncus mucronate, hoodlike. Tegumen dividedwherein the two tablet-shaped, bubble-like sections meet centrally for some or most of their length. Juxta more or less horseshoe shaped. Tympanal apparatus with saccus indistinct, posterior ridge lightly if at all sclerotized, grading into second sternite; venulae secundae not sharply tapering inward caudally; fornix broad, robust | 2 |
1’ | Forewing ground color straw or light gray, uniform or with a contrasting gray medial area suffused with white; or with outer margin and basal areas rust colored ( |
4 |
2(1) | Forewing medial area variously but diffusely shaded, generally without sharp contrast or orbicular spot; basal area not traversed by a white band. Uncus tapering to a distinct, ventrally directed squared tip. Tegumen invaginate such that sulcus joining two teguminal hemispheres extends less than 40% of length of tegumen. Intra-saccular process smooth, not conspicuously denticled | 3 |
2’ | Forewing medial area variously shaded, but often with contrast, an orbicular spot varyingly distinct or inconspicuous, if present; the basal area usually traversed by a white band. Uncus broadly tapered with a simple rounded nipple. Teguminal sulcus extends most of length of tegumen. Intra-saccular process rugose or denticled |
|
3(2) | Forewing color variously shaded with white scaling; lines or variously shaded regions conspicuous. Orbicular spot faint, if present. Hind wing slate gray. Valva truncate, rounded, entire |
|
3’ | Forewing shaded chocolate brown, markings not obvious. Hind wings dark gray. Valva slightly emarginate |
|
4(1’) | FW medial area suffused with white basad; postmedial line with broad, gentle costal bulge. Uncus dorsoventrally flattened, edges nearly carinate; uncus tip broad, neither acutely sharp nor sculpted with trefoil shape. Valva lacking a trigger-like process below costa; costa with mane-like tuft of elongate setae, recurved medially. Phallus simple, cornuti absent. Corpus bursae elongate, without signa | 5 |
4’ | FW shading either unicolorous or with medial area more darkly shaded than both basal and postmedial areas. Uncus tip swollen, either obovate or squared, in latter case with lateral edges thickened. Valva with a trigger-like process arising from within sacculus along ventral edge, and a conspicuous fleshy subcostal lobe and setose plume; costa lacking tuft of elongate setae medially recurved. Phallus with two prominent cornuti. Corpus bursae more or less globular, rarely with signa | 6 |
5(4) | Basal area of FW with a brown ovoid spot, delineated by white bands crossing from wing base to antemedial line. Uncus tapered towards blunt squarish tip at a roughly 60 degree angle. Valva entire, not emarginate, without distinct upper and lower extensions; center of valva unadorned; intra-saccular structures indistinct. Phallus simple, naked, without cornuti |
|
5’ | Basal area of FW rust colored, mottled. Uncus broad, lateral edges parallel, tapering to a wide, gently rounded tip at a roughly 45 degree angle. Valva with upper and lower extensions, the lower sclerotized dorsad; intrasaccular flange conspicuous, adorned with both surficial and adjacent setal clusters. Phallus with a single cornutus. |
|
6(4’) | FW uniform mouse gray or mottled, in latter case with medial area more darkly shaded. Uncus rounded, obovate with distinct, rhomboid nipple; uncal edges not reinforced or swollen. Valva gently rounded ventrally with moderate to elongate lateral process, ventro-medial edge with a distinct comb of elongate setae | 7 |
6’ | FW ground color gray or straw colored, contrasting gray medial area in some specimens. Uncus squared or scooplike in appearance with lateral edges swollen, sometimes conspicuously so, with or without a pronounced central ridge, tip hastate or trefoil-like; valva either elbowed or sharply angled towards mid-point, but not gently rounded, lacking an elongate process distally, ventro-medial edge without a distinct comb of elongate setae | 8 |
7(6) | FW mottled in appearance, medial area slightly darker than basal and postmedial areas; orbicular spot pronounced. Epipleural setae absent. Ventral trigger-like process on valva rudimentary, if present; subcostal lobe robust, squat, <=3x longer than wide |
|
7’ | FW gray, unicolorous; orbicular spot faint. Epipleural setae present. Ventral trigger-like process pronounced; subcostal lobe elongate and narrow, ~5x longer than wide |
|
8 (6’) | Uncus with conspicuous, prominent central ridge. Elongate lateral lobe of valva absent; subcostal lobe not elongate; ventral edge of valva not conspicuously elbowed close to vinculum; central membranous area of valva conspicuously longer than wide | 9 |
8’ | Uncus with a uniformly smooth contour. Subcostal lobe pronounced, finger-like; ventral edge of valva angled or elbowed sharply (not rounded) approximately mid-way between vinculum and lateral edge of valva; central membranous area of valva not conspicuously longer than wide | 10 |
9(8) | Female with two pairs of hind tibial spurs; male with coremata present on 4th abdominal segment, flattened hind tibial spur, specialized hind tibial scales with embedded dark patch, cuplike metatarsal modification, epipleural setae |
|
9’ | Female with single pair of hind tibial spurs; male secondary sexual features above absent |
|
10(8’) | Male secondary sexual characters (including coremata on 4th abdominal segment, cf. 10) all present |
|
10’ | Male secondary sexual characters (cf. 10) absent |
|
(19♂, 10♀, 1 sex undet.).
Holotype (♀, USNM): Castro, Parana; Collection Wm Schaus; [red type label] type 10575; Acontia? medalbi [sic] sp. Schs; Pyralie Schoenobiana gen. nov.; USNM Genitalia Slide by DA ♀ 107,899.
Specimens of
(
Unknown.
Unknown.
Brazil, Peru.
Below we summarize material examined for
Holotype (♀, BMNH): Holotype [round white label w/ red border]; El Tumbador, Guatemala, Champion; Godman-Salvin, Coll. 1904-1., B.C.A. Lep.-Het., Pionea chanesalis Druce; Pionea chanesalis Druce, type [hand written]; Genitalia Slide by DA, ♀. [Holotype of
Specimens of
(
Larvae have been reared from
Mexico, Guatemala; Costa Rica; Venezuela.
Unknown.
In size, with Mexican specimens appearing smaller in wingspan (FW length 4.5–7.0 mm) than Central American specimens.
It is with some trepidation that we synonymize both
(
Unknown.
Markings may be obscured in some specimens, rendering them more or less uniformly gray brown.
The specific epithet refers to the dark wing shading of this species and is treated as a noun in apposition.
Unknown.
Central Ecuador.
Male (
Unknown.
The specific epithet is from the Latin for showy or handsome.
Unknown. Adults active in October.
Southeastern Brazil (Bahia, Rio de Janeiro).
(16♂, 16♀, 1 sex undet.).
Male (
Unknown.
Unremarkable.
Unknown. Recorded adult activity mid-June (Mexico), January–June (Costa Rica), 8 March–23 April (Nicaragua), 12 April–25 May (Venezuela), December (Bolivia).
Mexico, Costa Rica, Nicaragua, Venezuela, Bolivia.
Species diagnoses for the
Thorax | Abdomen | Genitalia | ||||||||
---|---|---|---|---|---|---|---|---|---|---|
Flattened hind tibial spur | Elongatehind tibial scales | Dark patch embedded within tibial scales | Epipleural setal tufts | Concavemeta-tarsal structure | Coremata | Saccular bend or ulna | Ventro-medial setal comb | Distribution of ventro-marginal setae | Hook-shaped decumbent costal setal cluster | |
themis | + | 0/+ | 0/+ | + | + | Short | NA | 0 | 0 | 0 |
rasa | 0 | 0 | NA | 0 | 0 | NA | NA | 0 | 0 | 0 |
nyx | + | + | 0 | 0 | + | Short | Round | + | Localized | 0 |
clotho | + | + | + | + | + | Long | Round | + | Localized | + |
lachesis | + | + | 0/+ | + | + | Short | Angled | 0 | Even | + |
atropos | 0 | 0 | NA | 0 | 0 | NA | Angled | 0 | Even | + |
Male. (
Unknown.
This species varies most obviously in size (5.3 mm–10.0 mm in male forewing length), and based on the examination of several anomalous specimens from the British Virgin Islands, the presence of male secondary sexual characteristics (tibial hair pencils and abdominal coremata) do not perfectly covary: Hair-penciled males with and without coremata are noted from Guana Island, examples annotated and/or segregated in “Material examined” section above; see also Discussion.
The specific epithet refers to the Greek Titaness and embodiment of divine order and is treated as a noun in apposition.
Possibly associated with
Brazil, Cayman Islands, Costa Rica, Cuba, Dominican Republic, Florida (USA), Jamaica, Mexico, Panama, Puerto Rico, Venezuela. The Sanibel Island, Florida record represents the only known United States occurrence of any
(22♂, 32♀).
(3♂, 13♀), USNM. Holotype ♂ (
(
Male (
Unknown.
The specific epithet refers to the absence of male hind tibial and metatarsal structures and epipleural setal tufts (presumably secondary sexual characters) present in other
Unknown. Adults active June (Mexico), July (Cuba), and July, September (Dominican Republic).
Mexico, Cuba, Dominican Republic (essentially, vic. Gulf of Mexico).
It was suggested by V.O. Becker (pers. comm., following the submission of this work) that the name
Male (
Unknown
Unknown. Adults March–May.
Northern Venezuela.
Male. (
Unknown.
The specific epithet refers to the youngest of the three fates in Greek mythology, responsible for spinning the thread of human life, and is treated as a noun in apposition.
Unknown. Adults December.
Southern Ecuador.
(
Unknown.
In Greek mythology,
Unknown. Adults in Brazil active January, April, November, December; adults in Bolivia active October–December.
Central Brazil (Rondonia east to Bahia, Ceara and Rio de Janeiro), Bolivia (Santa Cruz).
(
Unknown.
The specific epithet refers to the third of the three fates. Treated as a noun in apposition.
Unknown.
Northern Venezuela.
Given the phenomenon described by
The moths treated in this paper compise a range of geographically widespread and potentially localized cryptic species united by a range of synapomorphies. It is not possible to infer an unambiguous center of origin for
There likely exist undiscovered
By far the richest source of phylogenetic signal in our matrix are the male genitalia, accounting for almost half the characters included in our analysis. This is not unusual for species-level studies of
In contrast, it is of particular relevance to the taxonomy of
Apropos of species diagnosis, we recall that as compelling as are the raw diversity of secondary sexual characters and the demonstrations of their phylogenetic lability, there have been suggestions that the expression of such structures may be underlain by rather simple genetic systems. Following
We wish to thank J. A. Lewis (JAL) for numerous dissections, D. Adamski (DA) for dissections and preliminary character interpretation, S. Escher for illustrations, T. Litwak for assistance with figures, E. Munroe (deceased) for making his then in-progress work available to M.A. Solis, and Axel Hausmann, The Natural History Museum, Munich, Germany for his hospitality. John Rawlins of the Carnegie Museum of Natural History provided a significant loan of specimens from Bolivia and the Dominican Republic. James Hayden eagerly shared suggestions for outgroups and character codings relevant to the diagnosis of
Matrix
0 10 | 20 | 30 | 40 | 50 | 60 | ||
medalba | 0011011-01 | 10110-0001 | 0001101000 | 011210-000 | 1110--010- | -0-10-00-0 | -001 |
chanesalis | 0011011-01 | 10110-0001 | 0001101000 | 011210-000 | 1100--010- | -0-10-0100 | -001 |
umbra | 0011011-01 | 10110-0-01 | 0001101000 | 011210-000 | 1110--010- | -0-?0----- | ---1 |
speciosa | 0022011-01 | 10110-0002 | 1102101000 | 011210-000 | 2100--000- | -1010-1101 | 0001 |
ysticalis | 1122011-11 | 00110-0102 | 1102101111 | 011110-001 | 21010-0120 | 0101101101 | 0011 |
themis | 1102011010 | 0101111102 | 1102101111 | 1111111111 | 22010-0120 | 0101111111 | 1111 |
rasa | 1102011010 | 01010-0002 | 1102101111 | 0111111111 | 22010-0120 | 0101111101 | 1111 |
nyx | 1102011110 | 0101100102 | 1102101111 | 1111110001 | 3201111121 | 1101111111 | 1111 |
clotho | 1102011010 | 0101111102 | 1102101111 | 1111110001 | 3201111121 | 1111111101 | 1111 |
lachesis | 1102011110 | 01011*1102 | 1102101111 | 1111110101 | 3201100111 | 1111111111 | 1111 |
atropos | 1102011110 | 01010-0-02 | 1102101111 | 0111110001 | 3201100111 | 111111---- | ---1 |
Glaphyria | 1100100010 | 10000-0111 | 0010010110 | 000000-000 | 00?0--000- | -0-0111?-1 | 1010 |
Hellula | 1100100-01 | 00000-0110 | 0010010110 | 000000-000 | 00?0--000- | ?0-00-10-1 | 1010 |
Eustixia | 1111000--1 | 00-00-0111 | 0101010110 | 000000-000 | 00?0--000- | -0-0101101 | 1000 |