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ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS IN SCIENCE
June 28, 1965
___
C3.L.US
TABLE OF CONTENTS and
AUTHOR INDEX 1963-1964 Nos. 64-85
Los Angeles County Museum • Exposition Park
Los Angeles, Calif. 90007
2
Contributions in Science
TABLE OF CONTENTS
No. 64. A definition and classification of the tribe Stiriini (Lepidoptera: Noctuidae), by Charles L. Hogue. 129 pp., 32 figs. April 10, 1963.
No. 65. A new Costa Rican salamander (genus Oedipina) with a re-exami- nation of O. collaris and O. serpens, by Arden H. Brame, Jr. 12 pp., 3 figs. March 18, 1963.
No. 66. A new toadfish of the genus Poriehthys from Caribbean Panama, by David K. Caldwell and Melba C. Caldwell. 8 pp., 3 figs. June 14, 1963.
No. 67. Marine shore fishes from near Puerto Limon, Caribbean Costa Rica, by David K. Caldwell. 11 pp., 2 figs. June 14, 1963.
No. 68. A collection of reptiles and amphibians from the highland faunal assemblage of western Mexico, by Roy W. McDiarmid. 15 pp., 5 figs. June 14, 1963.
No. 69. The salamanders of South America, by Arden H. Brame, Jr. and David B. Wake. 72 pp., 26 figs. Oct. 11, 1963.
No. 70. Intergeneric behavior by a captive Pacific pilot whale, by Melba C. Caldwell, David H. Brown and David K. Caldwell. 12 pp., 6 figs. Oct. 4, 1963.
No. 71. Studies on the lizard family Xantusiidae. IV. The genera, by Jay M. Savage. 38 pp., 24 figs. Oct. 11, 1963.
No. 72. The Machris Brazilian Expedition. Botany: Lythraceae, by A. Lour- teig. 10 pp., 4 figs. Oct. 11, 1963.
No. 73. Fossil birds from the Anza-Borrego Desert, by Hildegarde Howard. 33 pp., 3 pis., 1 fig. Dec. 30, 1963.
No. 74. A new chaetodont fish, Holacanthus limbanghi, from the eastern Pacific, by Wayne J. Baldwin. 8 pp., 2 figs. Dec. 30, 1963.
No. 75. A new species of Acmaea (Archaeogastropoda) from the Pleistocene of San Nicolas Island, California, by Jere H. Lipps. 15 pp., 6 figs. Dec. 30, 1963.
No. 76. A review of the fishes of the genus Pleuronichthys, by John E. Fitch. 33 pp., 7 figs. Dec. 30, 1963.
No. 77. A new poeciliid fish, Phallichthys tico, from Costa Rica, by William A. Bussing. 13 pp., 5 figs. Dec. 30, 1963.
No. 78. The chronologic and geographic range of desmostylians, by Edw. D.
Mitchell, Jr. and Charles A. Repenning. 20 pp., 4 figs. Dec. 30, 1963.
1965
Table of Contents
3
No. 79. Morphological data on two sibling species of small honey-guides, by Herbert Friedmann. 5 pp., 1 fig. Dec. 30, 1963.
No. 80. Evidence of aestivating lungfish from the Sangre de Christo forma- tion, lower Permian of northern New Mexico, by Peter Paul Vaughn. 8 pp., 1 fig. May 8, 1964.
No. 81. The role of olfaction in food location by the turkey vulture. Cathar- tes aura), by Kenneth E. Stager. 63 pp., 19 figs. June 30, 1964.
No. 82. The fish fauna of the Playa del Rey locality, a southern California marine Pleistocene deposit, by John E. Fitch. 35 pp., 49 figs. June 30, 1964.
No. 83. A new swift from Mt. Moroto, Uganda, by Herbert Friedmann. 4 pp. Dec. 8, 1964.
No. 84. Results of the 1964 Cheney Tanganyikan Expedition. Ornithology, by Herbert Friedmann and Kenneth E. Stager. 50 pp., 6 figs. Dec, 8, 1964.
No. 85. Mammals collected by the Los Angeles County Museum expedition to northeastern Venezuela, 1958, by Bernard B. Butterworth and Andrew Starrett. 8 pp. Dec. 8, 1964.
4
Contributions in Science
AUTHOR INDEX
|
Baldwin, Wayne J. |
No. 74 |
|
|
Brame, Arden H., Jr. |
Nos. 65, |
69 |
|
Brown, David H. |
No. 70 |
|
|
Bussing, William A. |
No. 77 |
|
|
Butterworth, Bernard B. |
No. 85 |
|
|
Caldwell, David K. |
Nos. 66, |
67, 70 |
|
Caldwell, Melba C. |
Nos. 66, |
70 |
|
Fitch, John E. |
Nos. 76, |
82 |
|
Friedmann, Herbert |
Nos. 79, |
83, 84 |
|
Hogue, Charles L. |
No. 64 |
|
|
Howard, Hildegarde |
No. 73 |
|
|
Lipps, Jere H. |
No. 75 |
|
|
Lourteig, A. |
No. 72 |
|
|
McDiarmid, Roy W. |
No. 68 |
|
|
Mitchell, Edw. D., Jr. |
No. 78 |
|
|
Repenning, Charles A. |
No. 78 |
|
|
Savage, Jay M. |
No. 71 |
|
|
Stager, Kenneth E. |
Nos. 81, |
84 |
|
Starrett, Andrew |
No. 85 |
|
|
Vaughn, Peter Paul |
No. 80 |
|
|
Wake, David B. |
No. 69 |
1 507,73
|
Q LOS ANGELES |
CONTRIBUTIONS |
|
COUNTY MUSEUM |
IN SCIENCE |
|
tfBER 64 |
April 10, 1963 |
|
A TRIBE |
DEFINITION AND CLASSIFICATION OF THE STIRIINI (LEPIDOPTERA: NOCTUIDAE) |
|
By Charles L. Hogue |
Los Angeles County Museum
Exposition Park
Los Angeles 7, Calif.
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum. Issues are numbered separately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientists and scientific institutions on an exchange basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information and the form in which it is presented. Priority will be given to manuscripts by staff members, or to papers dealing with specimens in the Museum’s collections. Manuscripts must conform to CONTRIBUTIONS style and will be examined for suitability by an Editorial Committee. They may also be subject to critical review by competent specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological Journals is highly recommended as a guide. (2) Typewrite material, using double spacing throughout and leaving ample margins, on only one side of 8*4 X 11 inch standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be avoided if possible. (5) Legends for figures and unavoidable footnotes should be typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method of literature citation must conform to CONTRIBUTIONS style — see number 50 and later issues. Spell out in full the title of non-English serials and places of publication. (7) A factual summary is recommended for longer papers. (8) A brief abstract should be included for all papers. This will be published at the head of each paper.
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PROOF. — Authors will be sent galley proof which should be corrected and returned promptly. Changes after the paper is in galley will be billed to the author. Unless specially requested, page proof will not be sent to the author. 100 copies of each paper will be given free to a single author or divided equally among multiple authors. Orders for additional copies should be sent to the Editor at the time corrected galley proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell Editor
CONTENTS
PAGE
Introduction 5
Acknowledgments 6
Taxonomic history 6
Geographical distribution 8
Ecology 9
A Representative Species: Basilodes Rugifrons 11
Analysis of Relationships 17
Definition of the Tribe 30
Classification 38
Appendix: Descriptions of New Species 83
Summary 88
Literature Cited 88
Illustrations 93
Note on the illustrations 95
List of abbreviations used in the illustrations 96
Illustrations 98
SMITHSONIAN
MSTITUTIOM
A DEFINITION AND CLASSIFICATION OF THE
TRIBE STIRIINI (LEPIDOPTERA: NOCTUIDAE) By Charles L. Hogue2
INTRODUCTION
The Stiriini are a structurally uniform but superficially diverse tribe of noctuid moths found only in the New World, chiefly in central and northern Mexico and the southwestern United States. As in most higher noctuid taxa, the tribe’s constituents are poorly known and, prior to the present work, virtu- ally unstudied with regard to phylogenetic relationships. The primary purpose of this study, therefore, has been to study the phylogenetic relationships in order to define the tribe and to bring its internal classification into order.
I have conformed in my presentation, as well as in my actual procedure of study, to the taxonomic principle that higher categories are to a large degree subjective and constructed on the basis of personal and practical reasons but are primarily and ideally based on some real phenomenon of organisms, in this case, their phylogenetic relationships (Simpson, 1961: see especially pp. 108-109). Hence, I have attempted to elucidate and infer phylogenetic rela- tionships separate from, and prior to, the assembling and dividing of the vari- ous groups into taxa and ranks and assigning names. This has the desired effects of keeping phylogenetic logic and conclusion clear (Arnett, 1961:107) and making them most available for future scrutiny and improvement as more in- formation becomes available. And it further satisfies, I hope, a need in sys- tematics expressed by Michener (1957:173) “. . . for all systematists to pub- lish with their works full statements of the type of relationships and other considerations behind their classifications!’
This study is based entirely on the morphology of the adult moths. Speci- mens of all the known species were studied and all but the rarest were per- sonally dissected. Dissections were mostly of the genitalia, but at least one specimen of almost every species was dissected in full and mounted on slides so that the entire endo- and exoskeleton could be studied in detail.
Unfortunately, several species of dubious taxonomic status are still un- known either as females, or more critically, as males. These are the females of “ Antaplaga ” prepontendyta Dyar, “Lythrodes” arivaca Barnes, Plagio- mimicus astigmatosus Dyar, “Stiria” iticys Dyar and Xanthiria primulina Druce, and the males of Plagiomimicus chalcospilans Dyar, P. raglena Dyar, P. unicum Barnes and Benjamin, Cirrhophanus hyperion Dyar and C. hoff- manni Hogue, new species.
1Work completed while the writer was a graduate student, Department of Zoology, University of California, Los Angeles, in partial fulfillment of the requirements for the degree of Doctor of Philosophy.
2Curator of Entomology, Los Angeles County Museum.
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Contributions in Science
No. 64
ACKNOWLEDGMENTS
It was J. G. Franclemont of the Department of Entomology of Cornell University who originally suggested this project to me and E. L. Todd, Ento- mology Research Division, A.R.S., U. S. Department of Agriculture (U. S. National Museum), who made most of the material available. I would like to express my special appreciation and thanks to these persons for same and for the other numerous suggestions and kindnesses they extended to me while 1 was engaged in this work. For criticising the final manuscript I also thank them and W. T. M. Forbes, Cambridge, Massachusetts and J. N. Belkin, De- partment of Zoology, University of California, Los Angeles. In this respect I should mention that I alone am responsible for the conclusions herein, with not all of which the above mentioned persons agree.
Acknowledgment is due my colleagues, W. A. Powder and S. Ramal- ingam, for numerous, miscellaneous helps, and to the following persons for putting at my disposal their personal collections or collections in their care: C. F. Harbison, San Diego Museum of Natural History; R. Leuschner, Los Angeles, California; L. M. Martin, Los Angeles County Museum; B. Mather, Jackson, Mississippi; P. T. Riherd, Mercedes, Texas; F. H. Rindge, American Museum of Natural History; F. P. Sala, Burbank, California; and Mr. and Mrs. L. A. Willahan, Los Angeles, California.
For financial assistance, I wish to thank the University of California, Los Angeles (graduate research grant), and the Southern California Academy of Sciences (AAAS grant).
The photographs are the work of R. J. Pence, of the University of Cali- fornia, Los Angeles.
TAXONOMIC HISTORY
When Grote ( 1874) originally described the genera Stiria and Stibadium he remarked on their close relationship. In his List of the Noctuidae of North America in the same publication he suggested a further relationship of these genera by placing them next to his earlier described Plagiomimicus. This treat- ment was the first trace of the tribe Stiriini. Basilodes and Cirrhophanus had already been described by other authors but Grote failed to see their affinities with the above genera and placed them in seemingly arbitrary positions quite apart from the others. His arrangement of these genera in his later Check List of the Noctuidae of America, North of Mexico, Part I (1875) was approxi- mately the same.
Between the time of the above lists and Grote’s New Check List of North American Moths (1882a), several new forms became known which Grote related to Stiria, Stibadium and Plagiomimicus. He grouped these in the “ New List” under a formal name, “Stiriinae!’ In addition to the three genera just mentioned, this group included the genera Polenta (synonymized by Grote with Plagiomimicus) , Fala and Acopa. Although this was the first formal dec- laration of the taxonomic group which is the subject of this study, Grote
1963
Definition and Classification of Stiriini
7
(1882b) implied the existence and position of the group in another paper pub- lished in the same year when he stated, “The following genera [Antaplaga, Plagiomimicus, Polenta, Stiria, and Stibadium ] seem to fall in between Helio- this and PlusiaV In the “New List ” Grote still excluded Basilodes, which he put in the “Calpinae’’ and Cirrhophanus, which he put with Antaplaga dimi- di at a under the “Heliothinae!’ These three groups and the “Plusiinae!’ how- ever, were contiguous in the list, indicating that Grote thought they were some- what related.
In the same year that Grote published his “New List',’ Smith ( 1881-1882) devoted a paragraph to the stiriine genera in his Synopsis of the North Ameri- can Genera of the Noctuidae (pp. 30, 35). He considered Stiria, Stibadium, Plagiomimicus and Basilodes generically indistinct. For the first time in the literature, Basilodes was united with the other stiriine genera but no statement of a formal grouping was made. In his Synopsis of the North American Helio- thinae, also published in the same year (Smith, 1882), Smith’s treatment of Antaplaga was identical to that of Grote in the “ New List’.’ Both placed the genus immediately between Pippona and Grotella.
The identity of the nominal species Polenta tepperi and Plagiomimicus richii proved to be a source of consternation to Grote for some time. At first, since some specimens he had, which were identified as tepperi, possessed the typical foretibial claw and facies of Plagiomimicus, he included tepperi in that genus and synonymized Polenta with it. The lack of a foretibial claw on the type of tepperi (which he had then not seen), however, puzzled him. He ques- tioned the statement of the original describer of the species, Morrison, and its curator, Smith, to that effect (Grote, 1883a:75) and wanted the character state verified (Grote, 1882b:75-76). Later, on the insistence of Smith (1883: 229) that the claw was in fact absent (it was, but because it had broken off), and a personal superficial examination of the type, Grote described the new species richii from his specimens with the fore claw (Grote, 1886). He then excluded Polenta tepperi from Plagiomimicus and the “Stiriinae” and re- placed it with his new richii (Grote, 1886; 1890:69). He followed this treat- ment until Smith (1893:263-264) stated his error of observation and cor- rected the situation by synonymizing richii with tepperi.
In his publications following the ‘New List” Grote continued to revise and expand the “Stiriinae!’ To the group, in his discussion On the North American Calpinae to Heliothinae (Grote, 1883a), were added the genera Basilodes, Chamaeclea, Cirrhophanus, and Neumoegenia which were new or formerly excluded. Cirrhophanus was only provisionally included for Grote thought that it might be distantly related to Gortyna, a non-stiriine genus. Two further genera, Hypsoropha and Plusiodonta, were added, without ex- planation, in his Introduction to a Study of the North American Noctuidae, published also in 1883 (Grote, 1883b). In both of these papers, Grote con- tinued to allude to the possible affinities of the Stiriinae with the Plusiinae be- cause of similarities in wing shape and metallic coloring.
Grote continued to follow essentially the same classification in his later
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Contributions in Science
No. 64
lists and discussions (Grote, 1890a; 1895), oscillating in his placement of Cirrhophanus with the Stiriinae and with the Heliothinae. He excluded some genera included earlier, such as Hypsoropha and Plusiodonta, and added some species to Basilodes which are now placed in the closely allied genus Chalco- pasta. He mentioned the exclusively American distribution of the group in a paper comparing the noctuid fauna of Europe and North America (Grote, 1890b: 69). It is notable that his former group names were given formal tribal status there and in his “ Revised List” (Grote, 1890). Thus, for the first time, the tribe Stiriini was formally declared.
The genera of the Stiriini have been variously treated by other workers. Smith’s (1891:52-53; 1893:261-266; 1903:48) and Dyar’s (1903:183, 196- 197) lists and catalogues generally followed Grote’s arrangement. Dyar segre- gated Antaplaga more than Smith but still put it near other heliothidine genera. This genus was then already becoming an ill-defined catch-all for unrelated forms.
Hampson (1910) treated all of the then known Stiriini in volume 9 of his Catalogue of the Lepidoptera Phalaenae in the British Museum. He com- pletely destroyed all semblance of a natural group with his arrangements, especially in the phylogeny of the Acronyctinae, the subfamily in which he placed the stiriine forms. The diagram, by his own careful explanation, was not intended as a true phylogeny, but it did attempt to show general phylo- genetic relationships. On the diagram, the Stiriini were widely dispersed among various foreign and American genera. This situation was obviously due to Hampson’s customary reliance on artificial characters. He described a number of new genera and species which clarified the taxonomy of the tribe in some respects.
Barnes and McDunnough (1917:70-73) and McDunnough (1938:99- 100) abandoned Hampson’s artificial classification in their check lists and generally grouped the stiriine genera naturally as did Grote.
Draudt’s ( 1919-1939) generic arrangement in volume 7 of Seitz’s Macro- lepidoptera of the World was identical to that of Hampson except in the addi- tion of the numerous new species described since Hampson.
The most recent treatment of the tribe by Forbes (1954:169, 251-252) was restricted to the species occurring in northeastern United States. His defi- nition of the tribe was, therefore, somewhat incomplete but accurate as far as it went. The genera he included are all good Stiriini except Stiriodes. Forbes was the first since Grote to treat the group formally as a tribe.
GEOGRAPHICAF DISTRIBUTION
The Stiriini have been only sparsely collected. The larger museums of the United States have the majority of the known material, but the overall amount is small compared to that for most other noctuid groups. While many of the common species of the southwestern United States are represented, primarily collected by O. C. Poling in the early 1900’s, the Mexican species are known
1963
Definition and Classification of Stiriini
9
from few localities and are represented by very few specimens, for some of these, uniques or only one sex being known. The locality data from these specimens reflects better the activities of collectors than the ranges of the moths. Few noctuid collectors have been active in Mexico in recent years. Most material dates back to the late 19th and early 20th centuries and is mainly from two sources: (1) the sporadic collections for the Godman-Salvin, Bio- logia Centrali- Americana project studied by Druce and (2) miscellaneous ac- cumulations sent to the United States National Museum by Roberto Muller and William Schaus and studied by Dyar.
Because of the paucity of material, only a few broad, tentative generaliza- tions about the distribution of the Stiriini can be made. The tribe appears to be entirely Central and North American. No known noctuids from the Old World or South America can be included in it. The center of distribution, where most of the species occur, seems to be central and northern Mexico and the southwestern United States. The collections from the northeastern and midwestern United States are mostly sporadic and of only a few species. There is very little material from extreme southern Mexico and other Central Ameri- can countries.
Except for a few from central Mexico and northeastern United States, all of the collections are from arid or semiarid areas. The tribe thus seems to be generally adapted to dry climates and edaphic conditions and is probably as- sociated with xerophytic plant types. This may explain the significance of certain stiriine structures such as the foretibial “claw” and frontal protuber- ance. The corneous, heavy appearance of these structures suggests that they are fossorial adaptations for digging through either a pupal cell wall or the overlying soil, through which the adult moth must pass when emerging from its underground pupation site. These materials may be expected to be quite compact and hard in arid regions. As far as I know, however, there are no recorded observations or experiments to demonstrate such a function for these structures. The frontal protuberance, and foretibial “claws” also tend to occur in many other unrelated noctuids of arid regions.
ECOLOGY
There is even less information available on the ecology of the Stiriini than about their geographical distribution. No life histories have been fully eluci- dated and other kinds of ecological data are almost completely unrecorded. According to the literature, only six per cent of the species have even been reared so that at least their food plants, larvae and pupae are known.
Superficial descriptions of larvae and pupae and listings of host plants have been recorded for the following species: Plagiomimicus expallidus (Crumb, 1956:211), P. pityochromus (Crumb, 1956:211), P. spumosus (Murtfeldt, 1894; Comstock, 1946; Crumb, 1956:210), Cirrhophanus tri- angulifer (Jones, 1937; Crumb, 1956:212), Basilodes chrysopis (Crumb, 1956:213) and B. pepita (Dyar, 1921; Crumb, 1956:213).
10
Contributions in Science
No. 64
From such incomplete information it would be dangerous to make any generalizations. However, a few consistencies are apparent from the data which at least are worthy of note. They may possibly indicate some broad tendencies, since the species listed above belong to diverse genera.
The larvae of all the species just noted feed on plants of the family Compositae, especially the flower heads of ambrosias. This constancy in food preferences, as far as demonstrated, lends support to the validity of the tribe as a phyletic unit. Furthermore, this association may explain the cryptic colors of the larvae and adults of some species. Jones (1937) reports finding a specimen of Cirrhophanus triangulifer . . sitting by day in the flower-head of Bidens involucrata, the golden yellow-brown of its wings matching closely the color of the flower!’ He noted also that the flower-feeding larvae matched the flower-heads closely in color. A similar association is known for other noctuid species, especially diurnal ones.
The larva of Plagiomimicus spumosus feeds on sunflowers. But instead of feeding on the surface of the flower-head it burrows into the interior and is not cryptically colored but pale and unmarked like borers in general (Com- stock, 1946). Likewise, the adult is not colored like the flower and probably does not rest among its petals during the day.
Similarities in the morphology of the larvae of the few stiriine species of which he had specimens led Crumb ( 1956) to group them into a single taxon, “Amphipyrinae, group 5!’ He included four other genera in this group which may be related to the Stiriini.
The larva of Cirrhophanus triangulifer and Plagiomimicus spumosus are known to form an extra hard, underground pupation cell. The foretibial “claw” and frontal protuberance of these species might be modifications for pene- trating the walls of these cells during emergence. (See remarks on these struc- tures under “Geographical distribution!’ section immediately above).
The dates on specimen labels generally indicate the seasonal occurrence of the adults. By far the majority of species have been collected in the fall or late summer, a lesser number in the spring, and a few in both, indicating bi- voltism. Mid-summer and mid-winter thus seem to be poor times for collecting Stiriini. The adult flight periods probably correspond to the flowering seasons of the host plants.
1963
Definition and Classification of Stiriini
11
A REPRESENTATIVE SPECIES: BASILODES RU GIF RONS
Most of the characteristic attributes of the Stiriini are only generally and vaguely describable in words. To convey the nature of these attributes more fully and accurately, I have illustrated many of them. The illustrations, how- ever, are restricted to genitalic and surface features since these were the only ones universally available and of the greatest taxonomic importance. There- fore, I feel it is necessary to illustrate in some detail at least one species, Basilodes rugifrons (Grote).
The following discussion and illustrations (Figs. 2-5) are not intended to be morphologically comprehensive since many details are not treated, such as head sutures, axillary sclerites, etc. But they will serve as a basic depiction of stiriine structure and also as an explanation of the terminology used in the remainder of this paper. As nearly as possible, the structures are described in the same sequence as they are in the formal taxonomic descriptions.
Head (Figs. 2a; 3a, b). The most unusual feature of the head and a char- acteristic stiriine feature is the protruding, heavily sclerotized frontal pro- tuberance (fp). It is composed of two parts, a peripheral ring (pr) and a ventral secondary prominence (seep). Below the frontal protuberance, the infraclypeal plate or clypeal shelf (csf) runs transversely across the head. In a cleared specimen the ocular diaphragm (od), which is perforated by an oval orifice, can be seen below the surface of the large, hemispherical eye (e). Each of the paired ocelli (oc) is situated immediately above the eye and behind the enlarged scape (sep) of the antenna (ant). On the ventral surface of the head the most conspicuous structures comprise the mouthparts and consist of the long proboscis (prob) and the three segmented, paired labial palpi (lp). Other less conspicuous elements of the mouthparts are the minute paired, 2-4 segmented maxillary palpi (mxp) and the paired pilifers (pf). The anterior tentorial arms (ata) traverse the head internally.
The cervical sclerite (cs) lies in the cervical membrane and connects the head with the propleuron.
Prothorax (Figs. 2a; 3a). The prothorax is much smaller than the other thoracic segments. Its simple pleura (p) unite dorsad with the flat, small notum (n) from which the large rectangulo-conical patagia (pt) arise. Cephal- oventrad the plueura connect broadly with the sternum (s). Most of the morphological sternum is no longer evident externally in the Lepidoptera but is believed to be represented internally by a flat, emarginate lamella, the lamella of the discrimen (Id). The latter is supposedly formed by medial mi- gration and invagination of the two lateral halves of the primitive pre-lepi- dopteran sternum (Weber, 1928). The same condition prevails in the meso- and metathorax.
The parapatagia (ppt) are weak straplike sclerotizations behind the patagia and above the large cephalically displaced mesothoracic spiracle (sp).
Mesothorax (Figs. 2a, b; 3a). The mesothorax is the largest and most complex of the three thoracic segments. Its pleuron is composed of several sclerites, the largest of which is the anterior episternum. The latter is incom-
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pletely subdivided into a dorsal anepisternum (aneps) and a ventral kate- pisternum (keps) by the anepisternal cleft (anepsc). An unsclerotized area of the anepisternum, the basalar cleft (bcl), lies just below the basalare (ba). The preepisternum (peps) lies immediately cephalad of the katepisternum.
Caudad of the large episternum is the V-shaped epimeron. The anterior arm of the V is termed the anterior epimeron (aepm) and the posterior arm, the posterior epimeron (pepm). A large subalare (sa) resides in the mem- brane between the tips of the arms of the V.
The pleural suture (pis) runs a devious course between the episternum and epimeron. It is straight and clearly visible ventrad but makes a curve caudad at the level of the anepisternal cleft and disappears beneath a strong overfold of the dorsal end of the anterior epimeron. Dorsally the pleural su- ture is lost in a complex of structures of doubtful morphological interpreta- tion. The most conspicuous of these are the pleural wing process (pwp) and the tegular arm (tega), the latter providing a base for the tegula (teg). In- ternally, the pleural suture is represented by the pleural ridge (plr).
The ventral margin of the largest sclerite of the thorax, the mesoscutum (sc), is complexly modified. The modifications are associated with the in- tricate articular system at the bases of the wings and include the suralare (sra) , scutal ridge (sr), scutal incision (si), adnotale (ad), and posterior notal wing process (pnp). The deep scuto-scutellar suture (sss), marked internally by the scuto-scutellar ridge (ssr), separates the scutellum (scl) from the scutum (sc). The externally reduced postnotum (pn) is hidden beneath the scutellum and is obvious only internally, laterad as the later op ostnotum (lpn) and mesad as the dorsal half of the phragma (ph) .
As in the prothorax, the sternum is mostly internal and in the form of a lamella, the lamella of the discrimen (Id), which is very large and entire, i.e., bridging completely with the furca. Another element of the sternum, the spina (spi), deeply invades the interior of the mesothorax cephalad of the lamella. Externally the spina is indicated by a deep pit in the mesosternum just anterior to a small V-shaped sclerite, the spinasternum (sst). The tip of the spina bears a cephalically projecting, flat horn, the horn of the spina (hs).
Behind the lamella and medially connected with it is a strong invagina- tion, the furca (f). The furca connects laterad with the dorsal end of the posterior epimeron by means of a f ureal bridge (fb). The phragma (ph) is very large and occupies much of the interior of the metathorax.
Metathorax (Figs. 2a; 3a). The metathorax is smaller than the meso- thorax but has most of the same elements. A large, rectangular, undivided episternum (eps) occupies the anterior third of its pleuron. The basalare (ba) is a rounded sclerite partially set off from the dorsal part of the episternum. The epimeron (epm) is partially membranous caudad and complexly modi- fied internally to form part of the tympanal complex which will be discussed separately below. The subalare (sa) resides in the membrane dorsad of the epimeron.
In contrast to the condition in the mesothorax, the pleural suture (pis)
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runs a straight path dorsoventrad between the episternum and epimeron. It runs dorsad into the pleural wing process (pwp). A small simple tegular arm (tega) resides in a notch between the pleural wing process and the basalare, but it bears no tegula.
The scutum (sc) is constricted dorsomesad by the infringement of the mesoscutellum but is well developed laterad. The ventral margin of the scutum is modified like that of the mesothorax into a suralare (sra), scutal incision (si), adnotale (ad), and posterior notal wing process, (pnp). There is no definite scutal ridge. The scutal suture (ss) marks the presence of an apodeme, the scutal phragma, which is a part of the tympanal complex to be discussed below. Behind the scutum, and separated from it by the scuto-scutellar suture (sss) , is the elongate scutellum (scl) .
Again, as in the pro- and mesothorax, the lamella of the discrimen (Id) represents most of the morphological sternum. It is similar in shape to that of the mesothorax but smaller and does not bridge completely with the complex, large furca (f) .
Legs (Figs. 2a; 3a; 4c) . The foreleg of B. rugifrons, like that of all Stiriini, is very much stouter and shorter as a whole than in most noctuids. This is especially true of the tibial segment. The tibia characteristically bears an apical “claw” composed of a seta. Some other noctuids have a similarly appearing “claw” but it is usually morphologically a spine. The tibia also bears an epiphy- sis (epi).
The mid- and hindlegs are more normal than the foreleg. Their coxae are longitudinally divided by a coxal suture (cxs) into an anterior eucoxa (euc) and posterior meron (m). The tibiae of both bear spurs : one pair subapically on the posterior face on the midleg; one pair subapically, and another pair about a third from the end, on the posterior face on the hindleg.
All three legs have five tarsal segments each (1-5) which bear plantar bristles (prbr) on their plantar surfaces. These bristles are arranged roughly in three longitudinal rows. The ungues (ung), or tarsal claws (not to be con- fused with tibial “claws”) are located at the tips of the last tarsal segments.
Of unknown function and taxonomic significance are fairly large, well- defined sensilla (sens) located subapically on the outer surfaces of the fore- tibiae. These, and other smaller sensilla, occur in fairly constant position on all the legs in the Stiriini. Their presence in noctuids has not been reported in the literature, to my knowledge.
Wings (Fig. 4b). The shape and venation of the wings are typical for most trifid noctuids. A single areole (ar), a secondary cell formed by the fusion of R3 and R,, is present. The frenulum (fr) of the hindwing is formed by very long bristles: the male has several fused ones and the female three sep- arate ones.
The wing pattern is not illustrated here (see Holland, 1903: plate XXVIII, figure 5, for a colored photograph of this species’ upper wing pat- tern) . It is somewhat modified from the primitive noctuiform pattern but some of the elements are still evident. The AM and PM lines traverse the wing
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faintly in their primitive positions. They are joined caudad by a dark area to form a roughly hexagonal spot on the inner margin. The median shade is ab- sent; the distal lines are obscured by a dark area spreading in a short distance along the outer margin. The orbicular spot and apical patch are completely absent but the reniform spot is represented by a small, round dot. For better examples of the full noctuiform pattern on the forewing in the Stiriini, see the figures of various units in the genus Plagiomimicus (e.g., Figs. 7-9).
The hindwing is patternless and evenly white above. The pale lower sur- faces of both fore- and hindwings are similar and also without pattern.
Tympanal complex (Figs. 2a; 3a, c). Elements from both the meta- thorax and the abdomen enter into the tympanal complex. The metathoracic elements are intricate derivations mainly from the metepimeron and to a lesser extent, metascutum. From the metepimeron come the four pockets (I, II, III, IV), the largest and most ventral of which is IV. Overlying the pockets is the transparent thin anterior tendon plate (atp). Between the semicircle formed by the pockets is the thin, taut tympanal membrane (t) which is separated in- distinctly from the conjunctiva (cj) by the slightly sclerotized, nodular epau- lette (ep). The alula, a broad posterior flap of the wing base bordered by the axillary cord (ax), overlies the entire complex externally. Ventrad and caudad of pocket IV is the posterior tendon plate (ptp); this structure is reduced to a narrow strip in the Stiriini; it is typically a large, convex, hat-shaped scler- otization in other trifid Noctuidae. The scutal phragma (scph), an elongate apodeme of the scutum, also enters into the tympanal complex. It is also char- acteristically reduced in the Stiriini.
The abdominal elements of the tympanal complex are the earlike hood (h), inclosing the first abdominal spiracle, and the counter tympanic cavities (etc). Where the countertympanic cavities from the two sides adjoin, the median countertympanic septum (cts) is formed. The opening of the counter- tympanic cavities to the exterior is termed the countertympanic orifice (co).
Abdomen (Fig. 4d). The abdomen has eight complete annular segments (I-VIII), each consisting of a strongly arched and well sclerotized tergite (T) and sternite (S) which are connected laterad by a broad pleural membrane. The tergite and sternite of VIII are modified slightly and associated with the genitalia in both sexes. They will be considered with those structures below. TI is quite small and partially un sclerotized. SI and SII are fused into a single plate to the sides of which are deep invaginations, the abdominal furcae (abf) . The cephalolateral corners of the other basal sternites may also be slightly invaginated as secondary abdominal furcae (sabf). A strongly sclerotized, narrow trough, the tergopleural groove ( tpg ), extends from the cephaloventral corners of TII under TI into the countertympanic cavities. The elements en- tering into the tympanal complex have already been mentioned. The first seven pleural membranes have large, central spiracles (sp).
Vestiture (Fig. 4a). The body vestiture of B. rugifrons is typical of the Stiriini. That of the head is deep, close and composed of medium, spatulate, evenly colored scales. That of the thorax is very dense, deep, spreading and
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generally composed of long spatulate scales with dentate tips. The individual scales are darker colored distad except for the extreme tips which are a sharply contrasting white. Together the scales give the thorax a rough, irrorate ap- pearance.
The thoracic vestiture is well tufted, tegular (tegt), patagial (ptt), pos- terior mesoscutal (pmesosct) and metascutal (metasct) tufts being present. The last is composed of hairlike, white scales. The abdomen is untufted except for a slight, transverse first abdominal tuft (It). The rest is generally clothed with dense, imbricated, normal, self-colored scales. The vestiture of the ventral side of the thorax and legs is typical of most noctuids, being generally scanty and composed of linear scales basad on the legs and on the body, and dense, imbricated, medium, spatulate scales distad on the legs.
Male genitalia (Figs. 5a-e). Abdominal segment VIII, because of its proximity to the genitalia and because its tergite and sternite have special pat- terns of sclerotization, is usually considered with the genitalia. In B. rugifrons the tergal sclerotization (VIIIT) is V-shaped and the sternal sclerotization (VIIIS) U-shaped.
The genitalia proper are joined to VIII by a long, loose intersegmental membrane and are composed of many varied elements of great taxonomic value. The dorsal tegumen (tg) is united laterad on each side with the ventral vinculum (vin) by a small, incompletely separated, slightly sigmoid sclerite, the pleurite (pi) (homologous with pleurite of Forbes ?) ; the whole complex forms an oval ring through the center of which passes the intromittant organ, the phallus. The phallus is composed of a strongly sclerotized, cylindrical aedeagus (ae) in which the inverted vesica (ves) normally lies. The latter is everted and inflated during copulation and bends at a right angle (dextrad) to the aedeagus. It has many long slender spines, the cornuti (cn) of the primary group (pg), on its ventral surface. A short, secondary row of small cornuti, the secondary group (sg), is situated in the angle of the vesica. The phallus is supported in the membrane between the tegumen and vinculum by a ventral, shield-shaped juxta (jx) and a dorsal pair of twin, seculate sclerites, together termed the anellus (anel) .
Articulating with the dorsal neck of the tegumen (ntg) is the uncus (un) which bears numerous short setae. The former is limited in extent cephalad generally by the furcation of the lateral portions of the tegumen and caudad by the base of the uncus. The ventral end of the vinculum is enlarged cephalad and folded inward to form the saccus (sac) .
The rectum also passes through the ring formed by the tegumen and vinculum and protrudes caudad as the tuba analis (ta). The sclerotizations in the latter, termed the scaphium (sea) (dorsal) and subscaphium (ssca) (ven- tral) are weakly developed in the Stiriini.
Articulating with the vinculum and juxta on either side are a pair of large, flat, clasping structures, the valves (val). Each valve has a compli- cated inner structure with several elements. The homologies of these elements are not yet known; therefore I use a noncommital descriptive terminology here,
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dividing the inner surface of the valve into three basic areas: (1) a narrow, dorsobasal costa (cos), a ventrobasal sacculus which is broken into a basal division (bsl) bearing a basal lobe (blbsl), and a distal division (dsl) bearing both a weak basal lobe (bldsl) and a spinelike distal lobe (dldsl) and (3) a distal, undifferentiated area bearing a row of stout setae, the corona (cor) at its margin.
Female genitalia (Figs 5f-g). Most of the components of the female genitalia of B. rugifrons are typical of only the most advanced stiriine types. The ovipositor lobes (ovlb) are pointed, well sclerotized, convex and elongate. Each of the pleura of segment VIII has a large, median, elongate, V-shaped unsclerotized area. The ventral margins of the pleura are deeply folded under (“strongly deflexed”) cephalad, arch laterad around the depressed lamella postvaginalis (lpv) and connect with the lamella antevaginalis (lav). These structures are associated with each other and with the ostium bursae (ob) or mouth of the long, narrow ductus bursae (dbu) and, when viewed ventrally, ap- pear as an integral, structural complex which, for convenience, is here termed the “ sterno-ostial concavity V This area is characteristically ovoid in B. rugi- frons.
The ductus bursae is ribbed and heavily sclerotized and expands into the large, saclike corpus bursae (crpbu). The sclerotization and ribbing of the ductus bursae continues onto the posterior third and posterior lobe of the corpus bursae. Leaving the apex of the lobe is a fine membranous duct, the ductus seminalis (dsm). The postapophysis (pap) and antapophysis (aap) are long apodemes running cephalad from the ovipositor lobes and VIII, respec- tively. They serve as places of attachment for the muscles which retract the genitalia.
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ANALYSIS OF RELATIONSHIPS
Since there are no known fossil Stiriini, I have had to infer the tribe’s in- ternal phylogenetic relationships entirely from the available attributes of the modern forms. Unfortunately, these attributes consist only of the adult in- tegumentary anatomy and scant bits of ecological and geographical informa- tion. On this basis, only a rough, general idea of the internal relationships of the tribe’s members can be formulated, but one which is sufficient to serve as a foundation for a preliminary definition and classification.
The general procedure of phylogenetic analysis which I used Is based on the traditional comparative method and will not be detailed here, being abundantly discussed and exemplified elsewhere (Cain and Harrison, 1953, in part; 1960; Simpson, 1961 : Chap. 3; Ross, 1956).
A statistical (so called “objective” or “quantitative”) treatment was at- tempted experimentally, however. The method tried was designed mostly after that employed by Hardwick (1958:20-25) and partially after that of Michener and Sokal (1957). Many such methods are now being developed for use in taxonomy (see Sokal, 1961, and papers listed by him). They may be very useful in indicating patterns of similarity and dissimilarity but cannot be re- garded as absolute measuring devices or indicators of true phylogenetic rela- tionships in any sense. I found them unsatisfactory in the present study for the following reasons (see also the criticisms of Inger, 1958:370-373) :
a. It was impossible to be truly objective in deciding what were unit characters and in breaking them into classes. As I gained experience with the group, I tended to vary in my evaluation of these factors.
b. Characters did not break down into an equal number of states3 so that their comparison directly was invalid because characters having few states would carry more grouping weight than characters having many. Furthermore, and most importantly, it seemed fundamentally inconsistent with generally accepted theory on the varying rates and processes of evolution to consider taxa equivalent and measurable in any mathematical sense.
c. In trying to be objective, I had to leave out much useful phylogenetic information such as obviously degenerate character states, primitive states, adaptive characters, etc. An attempt has been made by Cain and Harrison (1960) to integrate such data into statistical measurements of affinities (lead- ing to a natural but not necessarily phyletic classification), but it seems futile to go to such efforts in view of the dubious foundation.
d. The resulting arrangements were illogical in places, certain forms be- ing unduly approximated on the basis of a large number of adaptive or spuri- ous character states in common (e.g., size, cryptic colors, valve lobe shapes, shapes of frontal protuberances, etc.), while more important but fewer char- acters (particularly the sterno-ostial complexities of segment VIII of the
3The term “state” (= “character state”) is used throughout this paper to mean a particular condition or single variant of a varying quality of a structure, ie., a “char- acter!’ To clarify with an example : red may be a state of the character color ,
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female) received too little emphasis and allowed obviously related forms to be too greatly separated. These results may be due to the nature of the mate- rial, perhaps always coming about in compact, geologically recent groups in which adaptive traits are numerous and diverse and basic characters are few and stable (Noctuidae, Lepidoptera, Chiroptera, birds ?).
Results of the analysis
There are 100 stiriine species recognized in this study. This is such a large number from the standpoint of phylogenetic manipulation and so many are poorly known that I decided early to leave their detailed phylogeny and systematics to later studies and to work only on higher levels.
The species may be grouped on two levels. The first (lower) is com- posed of groups whose included forms are so extremely similar that they are undoubtedly very close relatives, taxonomically perhaps superspecies or spe- cies groups (as I call them in my classification), or even elements of polytypic species. My judgment is entirely empirical at this level but probably reliable in producing truly monophyletic units. Several isolated species which are not similar enough to any other to form a combined unit are treated as units of their own on the same level as the others.
Thirty-one such units were formed; they are listed with their constituent forms in Table 1. Four of these are composed of a single species, each of which is known from only one sex and so divergent from the other units or poorly known that its relationships are obscure. These are the “atypical” spe- cies mentioned in the section on the definition of the tribe. They were excluded from the phylogenetic treatment but are given summary treatment in the classification.
The above units may be further grouped into broader, roughly equivalent, monophyletic groups or phyletic units of a higher level. There are seven such units which are listed with their constituents in Table 1.
Commensurate with the general principle followed in this presentation that all classification should follow phyletic inferences, phylogenetic terminol- ogy is used exclusively in the discussion that follows. In this respect, I call the lower level, close groups of forms “primary phyletic units” (listed by Roman numerals) and the higher level groups of these, “major phyletic lines” (listed by Arabic numerals). For reference and clarity, however, I give the later ap- plied taxa names in parentheses following citation of a phyletic unit.
The immediate phylogenetic relationships of the primary phyletic units are first discussed. Then the major phyletic lines of the tribe, represented by monophyletic groups of primary phyletic units, are treated likewise after a few remarks on the value of certain characters. Both are summarized at the end in the form of phylogenetic diagram (Fig. 1 ).
Relationships of the primary phyletic units. Units I-X are only a gener- ally similar group. There is much variation in the shape of the frontal pro- tuberance, position and shape of the valve lobe, shapes of the female ovipositor lobes, bursae copulatrix, etc. Most structures of the male and female genitalia
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as well as the almost universally similar noctuiform wing pattern, however, indicate unity. With the possible exception of units IX ( Plagiomimicus psama- thochromus and argyropolius ) and X ( P . phalaenoides) , which may form a small derivative group of their own on the basis of the double-lobed, distal sacculus division, all the units form a diverse, but probably monophyletic major phyletic line ( 1 ) .
There are several character states possessed by unit I ( see character com- plexes, A-D, next section, for basis) which indicate that it is the most primi- tive of this line, indeed the most primitive stiriine unit of all. Not only the other units of this line but probably also all subsequent major phyletic lines stem from moths most like those of unit I.
Unit II ( P . laodamia ) is very similar to unit I and probably only a re- cent derivative of it. The few main different character states are obviously only slightly modified versions of their more primitive counterparts in unit I or are of little significance : ( 1 ) the pointed, upcurved ovipositor lobe still shows strong striae and is partially unsclerotized and flat like that of unit I; (2) the distally displaced valve lobe is still quite similar in shape to the central lobe of unit I and resides on an only slightly more angular and broader valve whose other features, e.g., well developed corona, small finger-like basal sacculus division lobe, are the same; (3) the single, conoidal lobe of the bursa copu- latrix, instead of two secondary lobes, is hardly significant since comparable differences occur sporadically throughout the tribe.
Units III-V (P. spumosus and allies, P. pityochromus and allies, and P. tepperi, respectively) are all independent but more extreme derivatives of I. Their morphologies are equally diverse (V is somewhat more peculiar) so it is impossible to infer the order of their derivation.
In spite of the somewhat extreme and diverse natures of units III and IV they grade together almost completely through primitive unit I. The grades are so numerous that only the best defined will be mentioned:
1. Between I and III: The single valve lobe of I shows a gradual length- ening and increasing curvature and migration basad through many of the members of III. The sequence begins with a very short lobe similar to that of ochoa (unit I) in P. jalada, proceeds through the slightly longer, more curved and basally displaced lobes of manti, corozona, astigmatosus and dimidiatus and ends with a very long, curved basal lobe in spumosus. A progressive broad- ening of the valve, reduction in the number and stoutness of the coronal setae and decreasing length of the basal sacculus division parallel the behavior of the lobe.
2. Between I and IV : The spinelike valve lobe of unit I shows a sequence of increasing spine shape and migration distad through the species of IV. The lobe of expallidus is most like that of I, being short, rounded and situated fairly near the base of the distal sacculus division. The lobe of pityochromus is more spinelike and distal, and that of concinnus is quite spinelike and lo- cated far distad.
In many of its character states, the single species of unit V is peculiar.
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The combination of states, noctuiform forewing pattern, simple VIII segment in the female genitalia, and the single median lobe of the male valve seems to place it best near the foregoing units, I, III and IV. However, most of its states are so strongly modified from those of the latter two units that the species is not detectably a derivative of either of them and even somewhat un- certainly of primitive unit I. For the present, I tentatively consider this unit equivalent with III and IV as independent derivatives of the ultimately primi- tive line represented by I.
I am even more dubious about the status of units VI (P. hutsoni and olivalis), VII (P. hachita and allies) and especially VIII (P. alesaea). The first two have many characters in common, e.g., circular unsclerotized areas in
VIII pleura in the female; similar, simple ostial regions in the female geni- talia; elongate valves with a small, broad, distal lobe in the male genitalia, etc., but their facies vaguely and inscrutably dissent throwing doubt on the homol- ogy of these characters. VIII is even more distinct: the valve lobe is set far distad, almost off the edge of the valve and is a strong spine; the wing pattern is completely absent, and the secondary group of cornuti on the vesica is absent, a few enlarged spicules on the sinistral inflation being present instead. Considering the general level of organization of the female genitalia, i.e., of the most primitive type (see next section) and the equally diverse other peculiar character states, I tentatively place these units in a roughly equivalent status as derivatives of the main ancestral stem of major phyletic line 1.
Units IX (P. argyropolius and psamathochromus ) and X (P. phalae- noides) are very similar. The latter unit has a few slightly peculiar features, e.g., ( 1 ) oblong, doubly narrowed valve instead of a broad valve with a strong- ly angled or produced distoventral corner, (2) distal lobe of the distal sacculus division rounded and with minute denticles at the tip instead of spinelike and (3) very few cornuti. These states are clearly variations of their more gen- eralized counterparts in unit IX. Because of the type of female genitalia (most primitive type; see next section), I place them near the rest of the units of major phyletic line 1, but consider them of special derivative status because of the double lobed distal sacculus division. This condition is prophetic of the bilobed condition of most of the higher Stiriini. Therefore, I consider units
IX and X as representative of the section of the ancestral stock of line 1 which gave rise to the rest of the tribe. This special position prompts me to consider these units as a second (2) major phyletic line.
XI ( Chrysoecia scira and allies), XII (C. dela and allies, subspecies ?) and XIII ( Gorgora morga) superficially appear to be very different units. In fact, they are so different at first inspection that the latest classification prior to the present one put them in three widely separated genera. The last unit, containing the single pecular species Gorgora morga was even placed in a different subfamily, the Acontiinae.
The nearly identical female genitalia of these units, however, indubitably unite them as major phyletic line 3. Other subtle but probably significant similarities are also present: (1) the zigzags in the courses of the PM lines of
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XI and XII (see especially scira and atrolinea where these lines are well de- veloped) are almost identical; (2) the orbicular spot has a tendency in all (especially XIII) to enlarge rectangularly; (3) the thoracic vestiture is “hairy”; (4) the lunules of the fore wing are usually marked; (5) the cornuti of the male vesica are diminutive centrally (not so in scira, however), etc.
Although XI may be somewhat more primitive than the other units of this line in the better developed noctuiform wing pattern, most features are equally diverse so the units are best considered equally divergent.
Unit XIV ( Chichimeca ) is distinct from all other units by its preponder- ance of degenerate character states, e.g., (1) the tiny distal sacculus division lobe (this is quite similar to a parallel degenerate state in unit XXI of major phyletic line 5, but is so withdrawn that its homology with other distal sacculus lobes is indeterminable); (2) the sclerotization and ribbing of the posterior region and lobe of the corpus bursae is obsolete and (3) the wing pattern is completely absent. Therefore, I consider this unit as an autonomous major phyletic line, 4.
Units XV-XXI ( Cirrhophanus ) are another diverse, large group like major phyletic line 1 but their unity and intragroup arrangement is much clearer because of numerous, fairly continuous intergradations. This group comprises major phyletic line 5.
The most probable primitive unit is XV (C. chryseochilus ) which still retains a fairly complete noctuiform wing pattern. This pattern is disrupted by a secondary pattern of longitudinal streaks (mainly from darkened wing veins), which is characteristic for most other members of the group, especially in the most immediate derivative, unit XVI (C. triangulifer and allies).
One species in the latter unit, C. triangulifer, shows a character state which especially relates it to units VII-XXI apart from other general similari- ties. This character is a multiple foretibial “claw!’ The claw, as it exists in this species, is a compact series of two or three, entirely separate, small setae which appear topologically to have arisen by a division of the usual, single large seta. In this condition, these setae resemble a commonly occurring anomaly (dupli- cation) in the chaetotaxy of mosquitoes (and probably in most insects) which may take part likewise in their evolution and differentiation (Belkin, 1952: 129). Further evolution appears to have been for these separate setae to fuse into a heavy unit, as it is found in the remaining units, thus producing a con- dition superfically similar to the original one. However, the fundamental, mul- tiple construction remains to reveal the mode of derivation and indicates the derivative status of the units possessing it.
The next five units, XVII (C. plesioglaucus and comstocki) , XVIII (C. papago and miaiphona) , XIX (C. hoffmanni) , XX (C. discistrigus ) and XXI (C. compositus and hyperion), are grouped mainly because of the common, multiple foretibial claw (this condition not personally confirmed for C. hy- perion) but there are other features which suggest monophyly also: (1) XVIII, XIX and XX have the streaked forewing pattern, (2) there is a se- quence of gradual loss of the primitive ovipositor lobe striae and elongation
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and curving of the lobes themselves through the units, and (3) all have lost the basal lobe of the distal sacculus division. I have not analyzed the compara- tive morphology of these units thoroughly, however (this awaits the discovery of more material, especially critically in rare species like C. hyperion and hoff- manni), and therefore consider them equally derivative for the present.
Unit XVII has a very distinctive wing pattern which hardly looks like any- thing in related units. The structure of the female genitalia and especially the thoracic vestiture, however, indubitably relate it to unit XVI. These characters have virtually identical states in the two units in all respects, even as to color of the thoracic vestiture. The very different wing colors appear to be adaptive. There is one field observation (Jones, 1937; see “Ecology”) that the yellow- orange color and streaked wing pattern typical of unit XVI has a cryptic func- tion: It camouflages the adults which have the habit of resting on similarly colored flowers. It is possible that the wing color and pattern of XVII also have a comparable, specialized cryptic function, but camouflage these moths to differently colored flowers. Even by themselves, the beautiful wings of these moths are very suggestive of a brightly colored flower. Therefore, the very different colors involved may be easily explained by the strong selective forces and simple genetic basis which have been demonstrated as involved in com- parable situations in other moths (Ford, 1955: 188).
The annectant features of the single species of unit XXII ( Cuahtemoca chalcocraspedon) are sufficient to make it a major phyletic line (6). The rea- soning for this is appropriate to the next section and is discussed there in more detail under phyletic line 6.
Major phyletic line 7 is composed of units whose great similarity suggests relatively recent divergence. The line, however, divides into two sections primarily on the basis of two types of valve lobes. In the first section, contain- ing units XXIV ( Chalcopasta territans and allies) and XXV (C. chalcotoxa and allies), the distal sacculus division bears a single, basal fingerlike lobe (secondarily absent from two species in XXV). In the second section, contain- ing units XXVI ( Basilodes chrysopis and allies) and XXVII ( B . rugifrons and allies) , the lobe is located far distad and is spinelike.
Though marked, this dichotomy is not strong considering the topology of these states. Each lobe, when present without the other, is extremely similar in shape and place of origin to a counterpart in the double lobed condition in unit XXIII (C. ellica ) and other lower Stiriini. It appears, therefore, that the latter is an inclusive condition immediately primitive to the two former mu- tually contradictory and derivative, single lobed conditions.
Though primitive in this important respect, other character states indi- cate that unit XXIII has probably diverged from its ancestors. Most conspicu- ous among these is the field of gold color (a cryptic specialization ?) covering much of the noctuiform pattern of the forewing. This state and other simi- larities relate this unit to unit XXIV. The latter is a derivative of the former as indicated by the distal sacculus lobe condition just described and the more extensive gold on the forewing.
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Unit XXV contains three species, two of which have very peculiar male genitalia. In these the valves are oddly shaped and the lobes are altogether absent (a probable degenerate condition), although a ventrally directed spur (remnant of distal lobe ?) is developed from the distal end of the sacculus. The latter is absent from one side in one species. These peculiar states and others, e.g., fusiform corpus bursae and tubular, coiled phallus vesica, separate these species from unit XXIV. However, the third species of unit XXV (C. acantha ) has normal genitalia and otherwise is similar to unit XXIV (espe- cially to C. restricta ) and connects the two units.
The two remaining stiriine units, XXVI ( Basilodes chrysopis and allies) and XXVII ( B . rugifrons and allies) are extremely similar. The latter has a somewhat peculiar wing pattern but this is obviously a simplified version of the former’s primitive noctuiform type: AM and PM lines traverse the same paths in detail (note particularly the sharp angles of the AM line basad, and of the PM line distad, at vein 1A). The area between these lines is brown in B. chrysopis of unit XXVI and remains so only in the inner area in most species of unit XXVII forming a characteristic hexagonal or square macula. A tiny dark dot in the reniform of these two forewing types further reveals the relationship.
Relationships of the major phyletic lines. As a basis for the discussion of the relationships themselves, a few preliminary remarks are necessary regard- ing the value of certain stiriine characters to higher intratribal phylogeny.
Most morphological characters of the Stiriini show either such great uni- formity or extremely diverse variation that they offer no clues to an inference of major phylogenetic trends or relationships. There are relatively few con- servatively varying characters showing fairly consistent correlation and pat- terns of parallel variation throughout the tribe as a whole so that they can be taken to reflect a picture of overall evolutionary history. In the main, I have found only the following characters and character complexes (A-D) to have application in this respect.
(A) Forewing pattern. This, the most frequently used discriminating character complex in the Lepidoptera, is usually considered of dubious phyletic value because it is so subject to homoplasy and extreme variation resulting from relatively minor genetic alterations. I find, on the contrary, that the fore- wing pattern can be of great usefulness, at least in the Stiriini, in determining relationships if carefully analyzed. In the first place, although independently derived states of this character may come to resemble each other superficially, their usual great complexity never permits them to do so in every minute de- tail. Close inspection almost always reveals fundamental differences. In the second place, regardless of the degree of specialization attained by some states, primitive elements usually persist to reveal true relationships.
The generalized noctuid pattern (noctuiform) has been worked out in detail. AM and PM transverse lines, apical patch, and orbicular and reniform spots are all very stable elements of the noctuiform pattern. They show con- siderable qualitative variation but their presence together is quite constant in
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most Noctuidae. The presence of these states may therefore be considered primitive for the Stiriini.
The noctuiform forewing pattern is rather fully expressed in several units. The genus Plagiomimicus is characterized by it, even though a few primary phyletic units show considerable modification from the noctuiform condition: ( 1 ) unit VII ( hachita and allies) has lost the reniform and orbicular spots and apical patch but retains the transverse lines; (2) VIII ( alesaea ) has lost all traces of any noctuiform elements, an obvious degeneration; (3) the pattern of VI ( olivalis and hutsoni) is very much diffused and the spots are missing but the basic transverse line pattern is still manifest.
The noctuiform pattern is retained in a few units of each major phyletic line suggesting that they are relicts of the primitive stock of their respective lines: (1) XXII ( Cuahtemoca chalcocraspedon) . This unit has, in addition, a tarnished, metallic luster prophetic of the brilliant gold of still higher units, especially XXIII-XXV {Chalcopasta) . (2) XV {Cirrhophanus chryseochilus) . The pattern of this unit is similar to that of XXII but is disrupted by a sec- ondary, lined or streaked pattern which is further modified in its derivative units in the same genus. (3) XXIII-XXV, especially XXIV ( Chalcopasta el- lica ) . These still show vestiges of the noctuiform pattern in spite of an extensive influx of brilliant gold. (4) XVI-XVII. The well developed noctuiform pat- tern of unit XXVI ( Basilodes chrysopis and allies) is probably closest to that of its own ancestors and those of its closest relatives, units XXIII-XXVII ( Chalcopasta and Basilodes) . Some variants ( B . pepita, chrysopis ) show some- what bronzy lusters or limited, diffuse, golden areas slightly more brilliant than the tarnished luster of unit XXII ( Cuahtemoca ) and inclined towards the obviously highly specialized condition in unit XXIV (C. territans and allies) where brilliant metallic gold is the predominant wing color masking most of the noctuiform elements.
(B) Male genitalia. The taxonomic value of this complex of structures in the Lepidoptera has been abundantly discussed elsewhere. Unfortunately, many opinions are based on speculations of the functional significance of the various elements and not on experimental evidence. For this reason, they should be taken with reservation. The recent studies of Callahan (1958) and Callahan and Chapin (1960) should help to remove some of these specula- tions. These authors have admirably demonstrated the mechanism of the phal- lus and bursa copulatrix in a few unrelated species from which some reliable conclusions can be drawn regarding variations in these structures. For ex- ample, there is obviously a very definite correlation between the shapes of the phallus vesica and the bursa copulatrix as well evidenced by their intimate in- terposition during copulation. One would expect these characters to vary rather conservatively, therefore, since major variations in one would require major concordant variations in the other— statistically unlikely— to retain reproductive success.
Like most stiriine structures, the male genitalia are either too constant ( e.g ., shape of vesica, distribution of cornuti) or sporadic (shape of uncus,
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shape of basal sacculus division lobe) in variation to be of much general phylo- genetic significance. I have found only three characters with states of value in this regard: (1) presence or absence and shape of lobes of distal sacculus di- vision, (2) length of tegumen neck and (3) presence or absence, number and shape of coronal setae.
The interpretation of the first of these is complicated by the dubious homologies of the various states. Various fingerlike and spinelike lobes occur on the distal division of the sacculus, but their positions are varied and incon- sistent within groups. Thus it is impossible to trace fully the changes that have occurred. Only in the higher Stiriini where their detailed shape and place of origin are constant, and in certain other groups where the various states grade together, are the lobes of undoubtedly homology.
Probably, a single, fingerlike lobe of the type found in unit I ( Plagiomimi - cus ochoa and resolutus ) of the first major phyletic line is the most primitive. It greatly resembles in shape and position the lobe of related Nocloa and allies and to a great extent many Heliothidini and Oncocnemedini (these two tribes may be fairly closely related to the Stiriini). The single lobe in the other units of the same line probably corresponds to it even though it varies in shape and position a great deal.
A second lobe was subsequently acquired (morphological source un- known) by higher Stiriini and persisted with the former as an intermediate primitive state from which several unilobed degenerations subsequently oc- curred.
The length of the tegumen neck shows a trend of variation generally paralleling that of other characters (to be discussed) and in this respect is prob- ably phylogenetically significant. In units I-X ( Plagiomimicus ) the neck is extremely short and erect like the condition in most related noctuids. This state therefore appears to be primitive. Increasing length and recumbence, especially well expressed in units XV ( Cirrhophanus chryseochilus) , XXII ( Cuahtemoca ) and XXVI-XXVII (Basilodes) , is a derivative trend.
A final male genitalic character showing one probable primitive state con- cerns the presence or absence, number, shape and distribution of the coronal setae of the valve. The Acronyctinae normally have a well developed, even row of stout coronal setae along the entire distal edge of the valve. On the basis of prevalence, this state would appear to be primitive for the subfamily and like- wise for the included Stiriini.
(C) Female genitalia. In inferring the relationships of the major phyletic lines I have placed the greatest reliance on characters of the female genitalia, especially those of the pleura, sterno-ostial region, i.e., ventral pleural margins and sternum of segment VIII, and the ostial lamellae. Lesser reliance was placed on characters of the ovipostior lobes. The several elements of this struc- tural complex coordinately show a sequence of increasing complexity in the lines from one to the other. This sequence is so logical topologically and com- pletely intergrading that it is likely a valid indicator of the actual evolutionary progression of the tribe.
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Basically, the sequence is as given below. A few units of each line may show parallel, secondary modifications, of course, e.g., declining ventral mar- gin of VIII pleuron, pointed ovipositor lobes, but most units, especially the more primitive ones, exemplify the sequence stages well.
In the simplest, initial condition in line 1 (Fig. 7k), the pleura of abdomi- nal segment VIII are completely and evenly sclerotized and confluent with the sternal region, their ventral margins not being defined. No special, complex sterno-ostial concavity is present, therefore. The lamella postvaginalis is not a discrete sclerite. The latter is represented only by an extended sternal area, which, with the 1. antevaginalis, forms a simple ring around the ostium bursae.
In line 2, obviously secondarily modified lamellae (forming a sclerotized chamber, Fig. 14k) obscure the arrangement to some degree, but there are essentially no changes, the separation of this line being based on male genitalic characters (see below).
Line 3 exhibits the first logical step towards greater complexity in these structures (Fig. 16k). An elongate median area in each of the pleura of VIII becomes unsclerotized (strong striae still remain to suggest an intermediate stage in a desclerotization process). Still, no special, complex sterno-ostial con- cavity is present but the ventral margins of the pleura become sharply defined though remaining undeflexed and partially contiguous. The lamella postvagi- nalis becomes a vaguely defined sclerite separate from the rest of the sternum (now largely membranous) and fills the space between the cephalically diverg- ing ventral pleural margins. The lamellae now are proper sclerites but still ring the ostium bursae simply.
Plagiomimicus resolutus of unit I of major phyletic line 1 shows an incli- nation towards the above condition, more so than any other member of the line.
The pleura of VIII themselves are essentially unchanged in line 4 (Chi- chimeca) but their ventral margins become slightly deflexed (underfolded), entirely separated, and divergent cephalad (Fig. 19k). The lamella postvagi- nalis enlarges and fills the area between the diverging margins of VIII. The lamella antevaginalis is reduced but forms a slightly bowed ring around the ostium bursae with the 1. postvaginalis. Thus, an ill-defined sterno-ostial con- cavity is formed.
In line 5 ( Cirrhophanus ) greater complexity appears (Fig. 21k). The pleura of VIII themselves are still essentially unchanged with large, elongate, unsclerotized areas containing striae (these are lost secondarily in some units of this line) but their ventral margins separate even more and become more deep- ly deflexed (or secondarily strongly declining). The sternum of VIII becomes a depressed, oval area, as viewed ventrally, with a well defined, oval lamella postvaginalis situated between the pleural margins and the lamella antevagi- nalis forming a bowed ring around the ostium bursae with the 1. postvaginalis.
The sterno-ostial concavity may be considered well developed in the pre- ceding line but not so completely as in the most advanced stiriine lines, 6 (Cuahtemoca) and 7 (Chalcopasta and Basilodes). Here, these structures reach
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their ultimate modification (Figs. 26k, 30k). The ventral margins of the pleura are well separated and greatly deflexed and curve out around a well defined, variously shaped (usually transverse, semilunar or ligulate, with an eroded posterior margin) lamella postvaginalis. The latter forms a strongly bowed ring around the ostium bursae with the lamella antevaginalis. The unsclerotized areas of VIII pleura remain but usually completely lose their striae (weakly manifest in some).
Changes in two other characters of the female genitalia generally parallel this sequence. One concerns a very minute and sometimes obscure feature, i.e., striae on the ovipositor lobes and in the unsclerotized areas of VIII. The other concerns the shape of the ovipositor lobes.
The ovipositor lobes are blunt, short and flat, and have well developed striae in line 1, which has a simple sterno-ostial region. They remain generally so in the lines with slightly more complicated ostial regions and completely sclerotized VIII pleura, but become pointed, long and convex and without striae in the most derivative units although these may be retained in some cases ( e.g ., unit XVI of line 5). There is an intermediate phase wherein the lobes are only slightly convex and the striae are reduced in number and strength (units XXII, Cuahtemoca chalcocraspedon, and XV, Cirrhophanus chryseo- chilus ). These units also show similar weak striae in the unsclerotized area of the VIII pleura, indicating the transitional phase between the condition where an unsclerotized area is present with strong striae and with blunt lobes, and the condition where an unsclerotized area and pointed lobes are present but no striae remain in the former.
The flat, blunt lobes are topologically (and functionally ?, see next para- graph) closer to the general noctuid type, i.e., soft membranous pads, than the pointed lobes, and therefore probably are more primitive.
I might speculate on the function of the heavily sclerotized, pointed lobes. Their shape and rigidity suggests that they are used by the female moth to in- sert eggs in some confined space, perhaps between flower petals (the known larvae are flower feeders) or in the soil or bark crevices for protection against rigorous desert conditions (most species range in arid regions). If such is the case, this state is to be considered a derivative specialization since most noctuids deposit their eggs in general open situations.
(D) A final obscure, but probably phylogenetically significant structure, the scutal phragma, shows a trend in the same direction as those discussed above. In line 1, this structure is somewhat keel-shaped, projecting moderately into the thoracic cavity. This is closer to the type found in most noctuids (a very strongly keeled and strongly projecting apodeme) than to the type in the more advanced Stiriini (a barely projecting low ridge); therefore, it is likely to be primitive.
In light of the foregoing considerations, the phylogenetic relationships of the major phyletic lines of the Stiriini appear to be as follows:
The most primitive major phyletic line is number 1 (most of the genus Plagiomimicus) as indicated by its preponderance of primitive states: (1) well
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developed noctuiform forewing pattern, (2) very short, erect tegumen neck, (3) well developed corona, (4) moderately keeled scutal phragma, (5) simple sterno-ostial region, (6) completely sclerotized VIII pleura which are conflu- ent with the sternum, and (7) flat, blunt ovipositor lobes with well developed striae.
Only one primary phyletic unit of this line, however, possesses all of these states: I (P. ochoa and resolutus). The remaining units are independent deriva- tives not on the main line of tribal evolution and show various special modifica- tions of all these primitive states, except those in segment VIII of the female genitalia. Subsequent major lines probably stemmed from moths most like those of unit I.
Major phyletic line number 2 is distinct from 1 only in one significant feature, a double, instead of a single lobed distal sacculus division. This feature could be an independent acquisition but in view of the great similarity in shape and position of these lobes with those of the many remaining higher Stiriini in which they are double (especially units XVI: Cirrhophanus dyari and allies and XII: Chrysoecia dela and allies), it appears that this line represents the section of line 1 which gave rise to all the more derivative major lines.
As indicated by the only slightly more complex female genitalia and simi- lar double lobed, distal sacculus division, major phyletic line 3 (Chrysoecia and Gorgora ) is the closest relative and a derivative of line 2. The noctuiform pattern and most other primitive states are strongly modified in all the mem- bers of this line. No unit therefore can be considered relictual.
As mentioned in the preceding section, major line 4 (unit XIV: Chichi- meca) is distinctive and degenerate in most character states. The female geni- talia, however, are of a type most similar to that of line 3 but slightly more complex. The line, therefore, appears to be the closest derivative of that line.
Major phyletic line 5 is large and diversified, most units being rather spe- cially modified. However, there are two units (XV and XVI) exhibiting several other primitive states whose level of female genitalic complexity (see above) situates them as immediate derivatives of line 4 and primitive to the remaining Stiriini.
The wing pattern and shape of the single species of line 6 (Cuahtemoca chalcocraspedon ) is strikingly similar (even to scattered blue scales) to that of one species (ochoa) of the most primitive stiriine unit, unit I. The female and male genitalia of this line, however, are quite advanced and of the type found in line 7 (Chalcopasta and Basilodes). Furthermore, the forewing is per- fused with tarnished gold of a shade similar to one species in unit XXVII (Basilodes pepita). The gold resides in the basal and distal sectors and reni- form and orbicular spots almost exactly like the brilliant gold of unit XXIII (Chalcopasta ellica).
The line, therefore, may be considered an annectant relict representing closely the ancestral stock derived very early from line 5 which gave rise to the remaining, most advanced line 7.
Units XXIII-XXVII (Chalcopasta and Basilodes ) of line 7 are without
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question the most highly advanced Stiriini. This is indicated mainly by the complex female genitalia which represent the derivative extreme of the se- quence of variation in these structures. In addition, the tegumen neck is quite long (not so extreme in Chalcopasta except in primitive ellica) or very long and arched and the ovipositor lobes are completely without any trace of striae (except in ellica ). The noctuiform forewing pattern and corona are still fairly well developed in most species, however. The former shows at least partially from beneath the secondarily acquired gold masking colors.
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DEFINITION OF THE TRIBE
Description of Adults
General. Facies robust. Size small to medium large. Colors predominant- ly brown, grey-brown, orange-yellow and brownish-yellow, a few forms white and metallic gold. Integument moderately sclerotized in most, darkly pig- mented in some. Sexes similar, female slightly larger than male.
Head. Size medium. Frons with large, heavily sclerotized protuberance of varied but basically characteristic shape and composition, i.e., raised outer, circular or ovate ring with a secondary central, subcentral or ventral conical or broad elevation; a few atypical but easily derivable types, e.g., deep smooth crater without any central elevation, rectangular, rough plateau without cen- tral depression, or transverse trough with lower external node. Clypeal shelf strong, prominent. Proboscis fully developed, apex unmodified except for nor- mal sense organs. Labial palpus moderately long, approaching or moderately exceeding frons and slightly upcurved; apical segment short to very short (nearly spherical), penultimate segment and basal segments usually subequal, former rarely somewhat longer or shorter than latter. Eye large, globoid and naked, without lashes. Ocelli present. Antenna simple, filiform or slight ser- rate, sparingly ciliate. Cervical sclerite with propleural arm very short.
Thorax. Large, cuboid. Lamella of prodiscrimen shallowly to moderately emarginate; parapatagia weakly sclerotized. Mesoscutum and mesoscutellum strongly convex; lateropostnotum moderately projecting internally; horn of spina well developed. Other sclerites and sutures typical of most Acronyctinae, very constant in shape and position.
Legs. Foreleg short; tibia very short and stout, shorter than or subequal to first tarsal segment, bearing strong, heavily sclerotized apical “claw” com- posed of a spinelike or shovel-shaped, single, stout seta or several apically fused setae; “claw” bearing apex of tibia often produced and strongly angled. Rest of foreleg and mid- and hindlegs moderate in length; tarsi with numerous sharp plantar bristles (rarely blunt tipped), arranged roughly in three rows, lateral tarsal bristles usually short; ungues small. No “spines” on mid- and hindtibiae; spurs well developed, normal.
Wings. Forewing elongate, triangular; apex acute to nearly a right angle, tornus obtuse, latter sometimes with small fringe tooth, both sharp or smoothly rounded; margins straight or slightly convex, outer margin entire, crenulate in some. Pattern and colors varied, basic noctuiform elements primitively present but suppressed or totally obscured in cryptic and degenerate, derived forms; when former, PM line sharply angled distad beneath apical patch, projecting through ST line; AM line oblique, straight or slightly angled over cell Cu; dashes absent; orbicular and reniform usually fairly well developed; subreniform and claviform absent; median shade always reduced and displaced distad. Hind- wing normal in shape; pattern lacking except for faint exterior line, varied in color, usually white, brown or fuscous. Frenulum of female with three free
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setae, of male with several fused setae. Venation of both wings of normal trifid type.
Tympanal complex. Generally simple. Epaulette weak, nodular. Pockets all present: I and II nearly vertical, strong, with broad orifices; III weak in some, varying in position from close to IV to close to II; IV large, with small to large orifice. Biigel absent. Anterior tendon plate large. Posterior tendon plate greatly reduced, a narrow strip. Scutal phragma also greatly reduced, a long, low or slightly lobed, hanging flap. Countertympanic cavities moderate in size, ovoid. Hood small, undivided, inclosing spiracle I. Alula small.
Abdomen. Short, stout at base, strongly tapering caudad. Abdominal furca moderately deep, slightly hooked at tip. Small accessory abdominal fur- cae developed in other basal segments. Sternites and tergites well sclerotized and arched. Brushes absent.
Vestiture. Generally rough. No special hair pencils, scale patches, etc. (except in male, a short, transverse patch of linear scales in base of U-shaped sternal sclerotization of abdominal segment VIII) . Head, including palpi, with dense, deep, close and pure vestiture, composed of small to medium, spatu- late, dentate, evenly-colored scales. Dorsum of thorax with dense, deep, spread- ing and pure or mixed vestiture, composed of elongate spatulate or linear, self- colored or distally dark scales (except for white tips); with following tufts: rounded or transverse patagial; spreading and often strongly erect tegular; slight anterior mesoscutal in some; strong posterior mesoscutal, sometimes lat- eral mesoscutal and other small extra mesoscutal tufts present; loose spreading meta scutal: all tufts poorly developed in many. Venter of thorax with scanty, shallow, loose and mixed vestiture, composed of medium spatulate and very long linear scales and hairlike linear scales. Legs with dense, deep, compact and mixed vestiture composed of linear, elongate and medium spatulate scales basad and dense, shallow, imbricated scales distad. Abdomen with shallow, rich, imbricated and pure vestiture, composed of evenly colored normal scales; f untufted except for slight, transverse tuft on TI.
Male genitalia. Tegumen broad or somewhat narrow, unlobed, with short erect to long curved neck. Uncus simple, cylindrical or medially swollen, straight or slightly undulate, sharply pointed apically. Vinculum equal to or shorter than tegumen, attached to latter laterad by small sigmoidally-shaped flaps (pleurites ? of Forbes); saccus short, stout or elongate, sometimes with small, anterior nipple. Valves symmetrical, simple, generally oblong in shape, parallel-sided, some with convex lower margin. Corona well developed and simple to weak or almost obsolete. Costa often broadly expanded. Sacculus ; divided into basal and distal divisions; basal division bearing a basal lobe, latter a low mound to a long fingerlike lobe; distal division bearing basal setose ver- rucae or a weak to well defined, fingerlike lobe and/or a distal spinelike or fingerlike lobe. Phallus with aedeagus well sclerotized; caecum small; vesica simple, ovoid to pear-shaped, moderately inflated and bent acutely dextrad on inflation, slight sinistral inflation in some as well; vesica armed with numer- 1 ous (few in some) distal, basally-directed, long to short cornuti, usually a sec-
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ondary row or patch of spinelets or denticles in crotch of angled vesica; vesica densely spiculate on side opposite cornuti. Anellus divided, with triangular, crescent-shaped or ovoid, paired, fused, approximated or moderately separated sclerites. Juxta subquadrate, kite- or shield-shaped with elongate, median rounded ridge. Tuba analis long; scaphium and subscaphium weak. VIII tergal sclerotization V- or U-shaped, arms sometimes fused and swollen caudad; sternal sclerotization U-shaped, inclosing transverse patch of long, linear scales.
Female genitalia. Ovipositor lobe partially or completely sclerotized, of two intergrading types; one thin, short, flat, and blunt, the other rigid, elongate and convex with pointed, straight or slightly upcurved tips, longitudinal striae well developed in former type, reduced or absent in latter. Postapophysis and antapophysis long, usually slender, stocky in some, subequal in length, former slightly longer than latter. Segment VIII varied, of three main intergrading types: ( 1 ) pleuron evenly sclerotized and confluent with sternum, ventral mar- gin not defined; no special, complex sterno-ostial concavity, to (2) pleuron with large, elongate, curved, median unsclerotized area containing distinct striae, ventral margin defined evenly along entire length, not deflected, con- tiguous mesad or caudad with opposite margin; no special, complex sterno- ostial concavity, to (3) pleuron with a large, variously shaped, median un- sclerotized area with or without striae, ventral margin defined along entire length, moderately to strongly deflexed cephalad and (with opposite margin) inclosing a depressed lamella postvaginalis and associated with it and 1. ante- vaginalis to form a generally oval, special, complex sterno-ostial concavity. Lamella postvaginalis not defined to well defined, ligulate to circular in shape; 1. antevaginalis ligulate; both lamellae usually forming a simple or bowed ring around ostium or rarely specially sclerotized and fused into a chamber. Ductus bursae moderately long, broad, usually well sclerotized and ribbed cephalad, membranous caudad, often with a quadrangular sclerotized plate in crotch of corpus bursae lobe. Corpus bursae large, a simple, ellipsoid or cudgel-shaped sac except for simple or lobulate ventral or lateral (sinistrad) posterior lobe; lobe and posterior third usually strongly ribbed and sclerotized like the ductus, remainder membranous; no signa; ductus seminalis leaving apex or outer corner of lobe.
Diagnosis
The Stiriini are defined and diagnosed in full, of course, only by the com- bination of all the character states listed above. There are, however, a few fair- ly unique and constant features which could be used as “key characters!’ These are (1) the extremely short, stout foretibia, (2) the single, strong, apical tibial “claw” composed of a single stout seta or several apically fused setae, (3) the frontal protuberance composed of a peripheral ring around a subcentral or ventral secondary prominence, (4) the sharply right-angled phallus, the vesica of which has a distal group of cornuti on one side and small, secondary basal group in the angle, (5) the simple bursa copulatrix with posterior third ribbed
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and sclerotized like the ductus, (6) the generally rigid, well sclerotized oviposi- tor lobes, and (7) the rough vestiture and prominent and often erect tegular and posterior dorsal thoracic tufts. Usually several of these states occur in all species of the Stiriini and all are combined in most species.
Integrity and affinities
It is highly probable that the species included in the Stiriini (Table I), as here defined, form a natural monophyletic unit. The large number of similar and completely intergrading character states shown by its components is alone strong evidence of monophyly. The unit is so compact that it should perhaps be considered a lower taxonomic category than a tribe, possibly a supergenus or subtribe. However, the tribal status is best retained until the Noctuidae are better studied as a whole, and such categories become more meaningful taxo- nomically. Tribes as such are at least handy working units for comprehensive studies though not always equivalent.
The phylogenetic position of the Stiriini within the trifid noctuids (Pan- theinae, Acronyctinae, Amphipyrinae, Agrotinae, Hadeninae, Cuculliinae) is uncertain. The various subfamilies in this group are presently defined on rather artificial characters, their comparative anatomy and basic relationships remain- ing uninvestigated. The subfamilies to which the stiriine genera are presently assigned by Forbes (1954) , Acronyctinae, and McDunnough ( 1938) , Amphi- pyrinae, particularly are of dubious integrity since they are largely defined by negative, superficial character states (e.g., lack of mid- and hind-tibial spines, hairless and unlashed eyes). It appears certain, at least, that the tribe is a rela- tively specialized (fossorial? foretibial “claws” and frontal protuberance) and secondarily simplified (reduced scutal phragma, reduced posterior tendon plate and simple tympanum) phyletic unit but I can give only an opinion as to its primitive relatives based on only a few indications and by no means a compre- hensive survey of the trifids.
The Stiriini seem to be derived from a generalized agrotine stock under- going loss of mid- and hindtibial spines, reduction of foretibial “claws’’ scutal phragma and posterior tendon plate, and acquisition of a frontal protuber- ance and other specific, specialized features (wing pattern, valve lobes, ovi- positor lobes, etc.). The derivation was either direct or, more likely, through intermediate phases possibly represented by the generally similar Heliothidini or Oncocnemedini which have less simplified (usually), spined mid- and hind- tibiae, multiple foretibial “claws’’ fully developed scutal phragma and posterior tendon plate. Both of these groups have multiple foretibial “claws” which could be a primitive condition giving rise to the single stiriine “claw” through loss of all but the functionally most important terminal one and the male genitalia are simple like those of the Stiriini (valve usually with a single, simple median lobe). Furthermore, many of the Oncocnemedini have a phallus vesica which is angled and has cornuti much like the Stiriini.
In spite of the compactness of the tribe, there are a few atypical forms which are impossible to classify comfortably within it. Only the males of these
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are known. They may be remotely primitive or derivative Stiriini or homo- plastically similar species belonging to other phyletic units. 1 have considered them tentatively as the former since their few available character states and those of the tribe occur in similar combinations even though they are not in- dividually identical. These species and their most conspicuous, atypical char- acter states are the following:
1. prepontendyta Dyar, 1914 (Antaplaga) . Valve broad at base, sacculus and costa very narrow, with wide membrane between. Lateral tarsal setae heavy, with blunt tips.
2. primulina Druce, 1 8 8’9 (Metoponia) . Frontal protuberance poorly de- veloped, only a slight, smooth bulge on the frons.
3. iticys Dyar, 1914 ( Stiria ). Lobe of distal division of the sacculus (basal ?, distal ? lobe) very short, heavy and spatulate. Frontal protuberance a circu- lar plateau with higher, rough, central elevation and dorsal, semicircular ridge.
4. arivaca Barnes, 1907 {Lythrodes) . This species has stiriine character- istics as far as I can determine but it is known from only a single, holotype male which I have not been able to study in enough detail to venture anything but a guess as to its proper intratribal position (see “Classification”).
There is a moderately large array of other noctuid species which show some attributes of the Stiriini. These species may represent Stiriini still more atypical than the above or may be the tribe’s closest and possibly primitive relatives. They are here excluded from the tribe for lacking one or more of the most characteristic stiriine states, or for having some outstandingly non- stiriine state. For example, all forms were excluded which did not have a fore- tibial “claw” composed of a single, stout seta or several apically fused setae, or did have unmodified soft, padlike female ovipositor lobes. These non-stiriine character states are always associated (correlated?) with other such states which further suggests a separate integrity.
All of the species and genera mentioned in the next three paragraphs share some basic character states with the Stiriini, the most significant probably be- ing the greatly reduced posterior tendon plate and scutal phragma. Each, how- ever, is similar to the Stiriini in only a few character states: general stiriine facies; presence of a frontal protuberance, and foretibial “claw” (when pres- ent, morphologically a spine not a single seta or group of fused setae) ; simple tympanum; long, numerous cornuti on vesica of phallus, etc. The possible re- lationships of these forms to the Stiriini have been implied in the arrangements of various authors but have yet to be studied and elucidated fully. This whole complex of generally similar forms may constitute a phyletic group of many tribes of which the Stiriini are but one.
The genera Nocloa Smith, 1906, Lythrodes Smith, 1903, Oslaria Dyar, 1904, Redingtonia Barnes and McDunnough, 1912, Chalcamistis Dyar, 1916, Ruacodes Hampson, 1906, Argyrhoda Hampson, 1910, Paramiana Barnes and Benjamin, 1924, Phaioecia Dyar, 1911, and Euamiana Barnes and Benjamin, 1927 (and others?), include very similar species and undoubtedly should be lumped into one genus. Tarsal segments 2-4 of the short, “clawless” foreleg
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are very short (nearly spherical), the foretarsal ungues are very much larger and heavier than those of the mid- and hindlegs, the phallus is straight (vesica not angled), with long, slender cornuti and the valve has a single, median lobe and a diffuse corona. “ StibadiurrT murisca Schaus, 1921, also belongs with these genera.
As mentioned parenthetically above, several species have a foretibial “claw” similar to that of the Stiriini. However, this “claw” is morphologically a spine (no basal alveolus) rather than a seta or group of fused setae. There- fore, the resemblance is due to homoplasy rather than homology. Other simi- larities are present which are probably also homoplastic. These species and their features somewhat resembling those of the Stiriini are as follows: Fala ptychophora Grote, 1875, frontal protuberance, female bursa copulatrix and male valve; “Antaplaga” koebeli Riley, 1893, frontal protuberance and thoracic tufting; “Tarache” pyralina Schaus, 1904, frontal protuberance, forewing pat- tern and wing shape; Rolua monetifera Dyar, 1915, and “Xanthothrix” stag- matogon Dyar, 1921 (belongs in the genus Rolua ) , and Narthecophora pulvera Smith, 1900, male genitalia.
Other species and genera generally similar to the Stiriini and possibly not too distantly related to them are Tristyla alboplagiata Smith, 1893, Hem- inocloa mirabilis (Neumoegan, 1884), Chamaeclea Grote, 1883, Azenia Grote, 1882, Satrapodes Hampson, 1910, Neumoegenia Grote, 1882, Ogdo- conta Butler, 1891, and Stiriodes Hampson, 1910. None of these have an armed foretibia, however, and most have a frontal protuberance of a different type than that of the Stiriini. Stiriodes was included in the Stiriini by Forbes ( 1*954) but, although the genus has something of a stiriine facies, it is abundantly dis- tinct structurally.
Species which at some time have been erroneously included in stiriine genera in important publications are given below :
1. aliaga Barnes, 1905, in Stiria originally. It was transferred to Nocloa by Hampson (1910:254) where it may be left pending further study.
2. crenulosum Dyar, 1909, in Stibadium originally and by Hampson (1910:528) and Draudt (1919-1939:294). The generic status of this species is as yet undefined. It is very close to “ Stibadium ” navium (number 1 1 below) .
3. duplicatus Smith, 1891, in Cirrhophanus originally and by Smith (1893:261), Dyar (1903:196), and Hampson (1910:202, questionably). This species was placed in the new genus Phaioecia by Dyar ( 19 11). It is very close to Nocloa aliaga (number 1 above) .
4. grisescens Barnes and McDunnough, 1910, in Antaplaga originally. This species is close to the genotype of Grotella and probably belongs in that genus.
5. koebeli Riley, 1893, in Antaplaga originally and by Dyar (1903); in Chalcopasta by Hampson (1910:222), Draudt (1919-1939:308) and Mc- Dunnough (1938:101). The generic status of this species is as yet undefined.
6. melanocrypta Dyar, 1912, in Antaplaga originally and by Draudt
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(1919-1939:325). This species is very close to the genotype of Grotella and undoubtedly belongs in that genus.
7. mina Schaus, 1894, in Basilodes originally and by Druce (1889:503). It was transferred to the new genus Satrapodes by Hampson (1910:217) and should be left there pending further study.
8. mirabilis Neumoegan, 1884, in Basilodes originally and by Smith (1893:262), Dyar (1903:196) and Hampson (1910:208, questionably). It was segregated into the new genus Heminocloa by Barnes and Benjamin ( 1 924: 165-166) and is best retained there pending further study.
9. murisca Schaus, 1921, in Stibadium originally and by Draudt (1919- 1939:293). It belongs with Nocloa and related genera (see above) in an as yet undefined generic position.
10. nanata Neumoegan, 1884, in Stiria originally and by Smith ( 1893: 262 ) and Dyar ( 1 903 : 1 97 ) . It was transferred to Nocloa by Hampson (1910: 255) and should be left there pending further study.
11. navium Harvey, 1875 (Telesilla) , in Stibadium by Smith (1893:263), Dyar (1913:197), Hampson (1910:109), Draudt ( 1 9 1 9- 1 93 9 : 294 ) , and Mc- Dunnough (1938:99). It belongs with crenulosum in an as yet undefined generic position.
12. philobia Druce, 1889 {Zatrephes) , in Basilodes by Hampson (1910: 207) and Draudt (1919-1939:304). The taxonomic position of this species is undefined; it should be returned to Zatrephes pending further study.
13. pyralina Schaus, 1904 {Tar ache) , in Antaplaga by Hampson (1910: 385) and Draudt ( 1919-1939:324). The generic status of this species is as yet undefined.
14. pyronea Druce, 1895 {Grotella) , in Antaplaga by Hampson (1910: 389) and Draudt (1919-1939:325). This species is very close to the genotype of Grotella, and undoubtedly belongs in that genus where it was placed orig- inally.
15. viridescens Schaus, 1914, in Stibadium originally. It was transferred to Oslaria by Hampson (1910:257) and should be left there pending further study.
16. viridifera Grote, 1882 {Zotheca), in Plagiomimicus by Smith ( 1891 : 53; 1893:264) and Dyar (1903: 197). It belongs with viridescens Schaus (see above) in the genus Oslaria.
The affinities of albigutta, fastigiata and tristriga, described near the genus Basilodes by Herrich-Schaeffer (1868, Zoologisch-mineralogisher verein, Regensburg, Correspondenz-blatt 22:181) from Cuba and known only from the types until recently were in doubt. I am informed by E. L. Todd ( personal communication) that they are erebines.
In the original description of her new plusiine genus Plusiophaes, Prout ( 1921 : 123-124) stated that the genus is probably distantly related to Chalco- pasta. Her opinion undoubtedly derived from the fact that both genera possess similarly shaped, partly metallic gold forewings. Probably this same similarity
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caused several true species of Chalcopasta to be originally described in the genus Plusia, a genus related to Plusiophaes. Grote’s inclination to relate the Stiriini to the Plusiinae was undoubtedly prompted by these same similarities. Neither Plusia, Plusiophaes, nor any of the Plusiinae are actually closely re- lated to Chalcopasta, the similarities being due without question to converg- ence. The convergence is strikingly great between some species, e.g., Phyto- metra stenochrysis Warren of Japan (Plusiinae) and Chalcopasta sinuata (Stiriini).
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CLASSIFICATION
Introduction
Based on the phylogenetic inferences developed in the preceding section, which I believe are in best accord with present information, I am proposing the following classification of the Stiriini. The classification should be considered preliminary in view of the incompleteness of the phylogenetic inferences neces- sitated by the restricted data. It awaits further improvement when more at- tributes than only the adult skeletal morphology become available, e.g., early stages, life histories, geographical distribution, etc.
I have made the classification particularly conservative by expanding the limits of genera both to emphasize relationships and to preserve nomencla- torial usage as much as possible without greatly affecting the approximate equivalence of taxa. Accordingly, I have recognized very few genera, proposed only two new genera (these were absolutely necessary) and refrained from applying formal names to the 31 species groups. The genera roughly corre- spond to the major phyletic lines. The species groups all equal the primary phyletic units.
As has been mentioned repeatedly in this paper, the Stiriini are basically a very compact and uniform series of intergrading forms. What variation there is, is very diverse and inconsistent, there being but few moderately conserva- tive characters and many annectant situations. The latter strongly suggests re- cency of origin of the tribe and, possibly, hybridization.
Partly as a result of these characteristics and partly as a result of my extreme conservatism in recognizing genera, I have found it exceedingly dif- ficult to characterize the tribe’s higher categories (genera) though relatively easy to characterize its lower categories (species groups). This is a common situation with natural classifications, and a difficulty which I believe has re- tarded the natural classification of the Noctuidae in general. This family is outstandingly a basically uniform, superficially diverse group also. Unfortu- nately, superficial characters form the basis of most classifications to date.
Diagnoses and key
For the sake of expediency and ease of reference, I have expanded the following phylogenetic key to include the diagnoses of the genera and species groups. As far as possible, easily observed features of both males and females are given, the most diagnostic being first. Exceptions are common, but if the whole set of character states is utilized fairly good discrimination is ensured.
1 . Female genitalia: Pleuron of abdominal segment VIII completely and evenly sclerotized or with a large indistinct but definitely oval unsclerotized area; ventral margin not defined (at least not cephalad) so that pleuron and sternum confluent. Male genitalia : Tegumen neck usually very short and erect. General: Pattern noctuiform, at least transverse lines present, never any metallic or irridescent colors, wing veins never darkened to produce a streaked pattern Genus 1: PLAGIOMIMICUS 2
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Female genitalia: Pleuron of abdominal segment VIII with a median, elongate unsclerotized area; ventral margin entirely defined, separating pleuron from sternum. Male genitalia: Tegumen neck usually moderate or very long. General: Pattern varied, if noctuiform, then generally brassy or brilliant metallic gold or with areas of such color, wing veins some- times darkened to produce a streaked pattern 10
2. (1) Female genitalia: Pleuron of abdominal segment VIII completely and
evenly sclerotized. Male genitalia: Distal sacculus division with two lobes or a single long, or moderately long, spikelike, or straplike basal lobe or a single strong, narrow, spinelike distal lobe 3
Female genitalia: Pleuron of abdominal segment VIII with a large, oval unsclerotized area. Male genitalia: Distal sacculus division with a single, short, poorly developed, broad spinelike lobe set far distad 8
3. (2) Male genitalia: Distal sacculus division with a single lobe. Female
genitalia: Ostial lamellae not specially modified into a large, purse- shaped chamber or if so modified, then mouth of chamber wider than width of chamber. General: Forewing color varied, never with fine, brownish-gold AM and PM lines, reniform and orbicular usually strong. 4
Male genitalia: Distal sacculus division with two lobes. Female genitalia: Ostial lamellae modified into a large, purse-shaped chamber with mouth narrower than width of chamber (one exception, phalaenoides, not so modified but with forewing as follows). General: Forewing color light, even, silvery-grey with fine, brownish-gold transverse lines, reniform and orbicular obsolescent 9
4. (3) Female genitalia: Ovipositor lobe with well defined striae. Male geni-
talia: Corona strong, along entire distal valve margin 5
Female genitalia: Ovipositor lobe without striae. Male genitalia: Corona weak, never along entire distal valve margin, usually restricted to dis- todorsal tip, often curving obliquely onto inner valve surface 6
5. (4) Female genitalia: Ovipositor lobe straight, flat and blunt. Lobe of corpus
bursae with two secondary, smaller lobes. Male genitalia: Distal sacculus division with lobe erect, set basad. Phallus vesica slightly irregular, inflated on anterior side distad. General: Angulation of PM line beneath apical
patch not extreme, reaching only to middle of patch
Species group I: P. ochoa & resolutus
Female genitalia: Ovipositor lobe curving dorsad, slightly convex and pointed. Lobe of corpus bursae simple, conoidal. Male genitalia: Distal sacculus division with lobe reclinate, set far distad. Phallus vesica smooth-
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ly pear-shaped, not inflated secondarily. General: Angulation of PM line beneath apical patch extreme, reaching to, or past, far edge of patch, almost touching outer margin Species group II: P. laodamia
6. (4) General: Forewing multicolored, markings distinct and well developed.
7
General: Forewing white, self-colored, markings entirely absent .......
Species group VIII: P. alesaea
7. (6) (Note three alternatives.) General: Frontal protuberance an oval ring
with a broad lower elevation. Wing color usually predominantly brown. Orbicular spot not markedly dark. Male genitalia: Distal sacculus division with a basal, fingerlike, or straplike lobe, lobe sometimes quite long and curving mesad. Uncus gradually curving. Costa well expanded. Basal sac- culus division very short, rectangular. Saccus slender, tapered. Female genitalia : Ductus bursae entirely membranous except for distinct ostial lamellae which may be modified into a chamber. Setae of ovipositor lobe
long. Postapophysis and antapophysis subequal
Species group III: P. spumosus & allies
General: Frontal protuberance a circular, smooth-bottomed cup with no secondary prominence. Wing color predominantly brown. Orbicular spot markedly dark. Male genitalia: Distal sacculus division with a heavy spine- like lobe usually set toward distal edge of valve. Uncus undulate. Costa barely expanded. Basal sacculus division short, rectangular. Saccus slen- der, tapered. Female genitalia: Ductus bursae entirely membranous, no sclerotized chambers. Setae of ovipositor lobe long. Postapophysis and antapophysis subequal Species group IV : P. pityochromus & allies
General: Frontal protuberance an inverted, heart-shaped ring with lower lip produced into a conical prominence. Wing color light, with olive-green markings. Orbicular spot obsolete. Male genitalia: Distal sacculus di- vision with a thin, spikelike lobe set near edge of valve. Uncus gradually curved. Costa expanded obliquely. Basal sacculus division very broad, triangular. Saccus stout, cylindrical. Female genitalia: Ductus bursae very heavily sclerotized and confluent with lamellae to form a bowl- shaped chamber. Setae of ovipositor lobe minute. Postapophysis almost one-third longer than antapophysis Species group V: P. tepperi
8. (2) General: Forewing with broad, diffuse, fuliginous green AM and PM
lines. ST line an indistinct, continuous shade. Integument darkly pig- mented. Female genitalia: Ovipositor lobe elongate, bluntly pointed
Species group VI : P. olivalis & hutsoni
General: Forewing with well defined, thin, black AM and PM lines. ST line broken into a row of dots or absent. Integument normally pigmented.
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Definition and Classification of Stiriini
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Female genitalia: Ovipositor lobe triangular, sharply pointed
Species group VII: P. hachita & allies
9. (3) Male genitalia: Distoventral corner of valve produced or strongly angled. Distal lobe of distal sacculus division pointed, without minute denticles. Costa expanded obliquely. Cornuti of phallus vesica numerous (at least more than twenty). Female genitalia: Lamellae modified into a
large, purse-shaped chamber with a narrowed mouth
Species group IX : P. argyropolius & psamathochromus
Male genitalia: Distoventral corner of valve not produced nor strongly angled. Distal lobe of distal sacculus division with minute denticles on rounded tip. Costa moderately expanded, not oblique. Cornuti of phallus vesica very few (less than five). Female genitalia: Lamellae not modified into a narrow-mouthed, purse-shaped chamber, instead, an indistinct,
small chamber with a wide mouth present
Species group X: P. phalaenoides
10. (1) Female genitalia: Ventral margins of VIII pleura entirely and evenly defined, convex and contiguous mesad or caudad, not deflexed. Pleural unsclerotized area curved and with strong striae. General: A low, semi- circular ridge below frontal protuberance (above clypeal shelf). Male genitalia: Cornuti of phallus vesica diminutive in center of primary group (except C.scira) 11
Female genitalia: Ventral margins of VIII pleuron unevenly defined cephalad, approximate caudad, not contiguous. Pleural unsclerotized area curved or V-shaped, usually without striae. General: Never a semicircular ridge below frontal protuberance. Male genitalia: Cornuti of phallus vesica subequal or varying in size longitudinally, not toward center of group 13
11. (10) General: Forewing predominantly orange or multicolored, discoidal macula white when present. Transverse lines usually strong, at least PM weakly evident. Plantar bristles sharply pointed. Male genitalia: Valve oblong, tip broad. Distal sacculus division with two separate full lobes. Corona of several slender setae .... Genus 2: CHRYSOECIA .... 12
General: Forewing predominantly black with a large, quadrate, orange discoidal macula. Transverse lines not at all evident, masked by black color. Plantar bristles with rounded tips. Male genitalia: Valve triangular, tip pointed. Distal sacculus division with a single, bifurcated lobe. Corona
of two very stout setae
Genus 3: GORGORA \ Species group XIII: G. morga
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12. (11) General: Forewing multicolored, white in distal and basal sectors,
median sector predominantly brown with scattered blue scales; transverse lines somewhat indistinct. Cup of frontal protuberance generally round. Male genitalia: Lobe of basal sacculus division a short, thin, irregular finger or a broad, short peak. Basal lobe of distal sacculus division weak, arising some distance from distal lobe. Secondary group of cornuti of
phallus vesica a row of many, short, subequal denticles
Species group XI : C. scira & allies
General: Forewing bicolored, ground color orange, zigzag transverse lines usually strong, rarely obsolescent; no blue scales. Cup of frontal pro- tuberance a transverse trough. Male genitalia: Lobe of basal sacculus di- vision a long, curved, broad finger. Basal lobe of distal sacculus division strong, arising close to distal lobe. Secondary group of cornuti of phallus
vesica reduced to a few unequal, fairly long spines
Species group XII: C. dela & allies
13. ( 10) General: Forewing pattern entirely absent, self-colored, usually sooty
white, if orange, fringe concolorous. Forewing elongate. Male genitalia: Costa greatly expanded. Distal sacculus division with a single, tiny, spike- like lobe. Female genitalia: Greatly attenuated. Sclerotization and ribbing
of ductus and corpus bursae obsolescent
.... Genus 4: CHICHIMECA; Species group XV: C. thoracica & allies
General: Forewing pattern present, if self-colored, fringe contrasting dark. Forewing shape varied, not elongate. Male genitalia: Costa not greatly expanded, usually hardly expanded at all. Distal sacculus division almost always with a strong, spinelike lobe. Female genitalia: Not at- tenuated, usually quite compact. Sclerotization and ribbing of ductus and corpus bursae strong 14
14. (13) General: Forewing never with brilliant metallic or brassy gold col-
ors; pattern usually streaked, i.e., with darkened veins and streaks between veins. Female genitalia: Ventral margin of VIII abdominal pleuron mod- erately deflexed or declining. Striae in unsclerotized area of VIII pleuron well defined, weak or absent. Striae of ovipositor lobe almost always present Genus 5: C1RRHOPHANUS . 15
General: Forewing with brilliant metallic or brassy gold areas or if dull colored then streaks not present. Female genitalia: Ventral margin of VIII abdominal pleuron greatly deflexed or declining. Striae in unsclero- tized area of VIII pleuron and on ovipositor lobe usually absent (rarely weak) 21
15. (14) General: Foretibial “claw” a simple seta or group of three entirely
separate setae. Forewing pattern streaked; streaks in cell Cu forming a
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trident (base distad). Male genitalia: Distal sacculus division with two lobes, or with one, weakly sclerotized, fingerlike lobe 16
General: Foretibial “claw” composed of several, apically-fused setae. Forewing streaked or not, but streaks in cell Cu never forming a trident. Male genitalia: Distal sacculus division with a single spinelike, heavily sclerotized lobe 17
16. (15) Male genitalia: Distal sacculus division with a single, long, basal
fingerlike lobe. Tegumen neck very long. Phallus vesica with an appendic- ular lobe. General: Forewing ground color dark purplish basad, marginal sector lighter, brownish yellow. Female genitalia: Ovipositor lobe convex, well sclerotized, sagittate Species group XV : C. chryseochilus
Male genitalia: Distal sacculus division with two lobes. Tegumen neck short. Phallus vesica without any appendicular lobes. General: Forewing ground color orange, marginal sector lighter, but not brownish-yellow. Female genitalia: Ovipositor lobe flat, partially unsclerotized, broad and blunt Species group XVI: C. triangulifer & allies
17. (15) General: Forewing with a purplish iridescent cast. Male genitalia:
Distal sacculus division with lobe very heavy, long, and set at far distal edge of valve. Female genitalia: Sterno-ostial concavity triangular. Ovi- positor lobe short, pointed, curved upward, with strong striae
Species group XVII : C. plesioglaucus & comstocki
General: Forewing generally dull orange-yellow or brownish-white. Male genitalia: Distal sacculus division with small lobe set in from distal edge of valve. Female genitalia: Sterno-ostial concavity subquadrate or flask- shaped. Ovipositor lobe not with above combination of character states. 18
18. (17) General: Forewing ground color dull orange, streaked with definite,
though diffuse, transverse lines. Fringe concolorous with wing. Female genitalia: Ovipositor lobe straight, flat, blunt, without striae. Male geni- talia: Secondary group of cornuti of phallus vesica present, a long curved
row of sharp, equal spinelets
Species group XVIII: C. papago & miaiphona
General: Forewing ground color pale white or yellowish, with streaks but no definite transverse lines, instead a diffuse stain in discoidal area or forewing ground color orange, self-colored. Fringe marked, not entirely concolorous with wing. Female genitalia: Ovipositor lobe curved, pointed, striae weak or absent. Male genitalia: Discrete secondary group of cornuti of phallus vesica absent although a separate group of scattered, curved- tipped spicules may be present 19
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19. (18) General: Forewing with terminal line distinct. Female genitalia: Ovi-
positor lobe elongate. Ductus bursae sclerotized and ribbed for less than one-half its length. Species group XIX: C. hoffmanni
General: Forewing with terminal line absent. Female genitalia: Ovipositor lobe short, triangular. Ductus bursae sclerotized and ribbed well over one-half its length 20
20. (19) General: Forewing streaked, fringe light, interrupted by extensions of streaks. Female genitalia: Ductus bursae short, much shorter than corpus. Corpus bursae ellipsoid, with sclerotization and ribbing normal; lobe and ductus seminalis posterior. Dorsal edge of ovipositor lobe curved, striae absent or very weak. Male genitalia: Distal sacculus division lobe
reclinate. Cornuti of phallus vesica in one continuous group
Species group XX: C. discistrigus
General: Forewing not streaked, fringe evenly dark. Female genitalia: Ductus bursae very long, about as long as corpus. Corpus bursae ovate, transverse, without sclerotization and ribbing; lobe and ductus seminalis directed at right angle to ductus bursae. Dorsal edge of ovipositor lobe nearly straight; striae distinct. Male genitalia: Distal sacculus division lobe
erect. Cornuti of phallus vesica in two widely separated groups
Species group XXI: C. compositus
21. (14) General: Forewing with brassy-gold restricted to marginal and basal
sectors and to reniform and orbicular; scattered blue scales along costal area. Male genitalia: Distal sacculus division with two lobes of about equal
length, set close together
. . Genus 6: CUAHTEMOCA -, Species group XXII: C. chalcocraspedon
General : Forewing with brassy gold color general or brilliant metallic gold in distal and basal sectors and in reniform and orbicular or generally dull colored. Male genitalia: Distal sacculus division with no lobes, one lobe or two lobes, if latter then set far apart or unequal in length 22
22. (21) General: Forewing with brilliant metallic gold over much of wing in
well defined fields. No black dot in reniform. Outer margin entire. Male genitalia: Distal sacculus division with no lobes or with well developed, basal, fingerlike lobe; an erect, distal, spinelike lobe absent altogether or reduced and set well on valve close to basal lobe. Corona weak or absent. Female genitalia : Ventral margin of VIII pleuron strongly declining or de- flexed Genus 7: CHALCOPASTA 23
General : Forewing without brilliant metallic gold; brassy or yellow gold may be present but if so then generally distributed over entire wing or in diffuse areas. A black dot evident in reniform. Outer margin crenulate. Male genitalia: Distal sacculus division with well developed, distal, spine-
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Definition and Classification of Stiriini
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like lobe; basal lobe usually absent or represented at most by small, setose verrucae, if well developed then set far from distal lobe and the latter then set far to edge of valve. Female genitalia: Ventral margins of VIII pleuron always strongly deflexed Genus 8. BASILODES 25
23. (22) General: Median sector of forewing brown, only basal and distal
sectors gold, both reniform and orbicular filled with gold. Male genitalia : Distal sacculus division with two lobes. Tegumen neck quite long, arched. Female genitalia: Ovipositor lobe straight, saggitate, with blunt tips. Un-
sclerotized area of VIII pleuron with weak striae
Species group XXIII: C. ellica
General: Median sector of forewing always gold; gold may extend over most of wing but not in reniform. Male genitalia: Distal sacculus division with single, erect, fingerlike lobe or without erect lobe. Tegumen neck short. Female genitalia: Ovipositor lobe angled dorsad, acuminate, with sharp tips. Unsclerotized area of VIII pleuron without striae 24
24. (23) General: Gold masking most of forewing pattern, always entirely
covering distal third of wing. Male genitalia: Distal sacculus division with single, basal, erect, fingerlike lobe. Phallus vesica pear-shaped, swollen end basad. Female genitalia: Corpus bursae ellipsoid-oblong, lobe nor- mal Species group XXIV : C. territans & allies
General: Gold restricted to oblique median area of forewing, weak or absent in distal third. Male genitalia: Distal sacculus division with a basal, fingerlike, lobe or without lobe, if former then phallus vesica swollen distad, if latter then a distal spur pointing ventrad on valve and phallus vesica tubular and coiled. Female genitalia: Corpus bursae ellipsoid-ob- long or fusiform, if former then lobe displaced cephalad
Species group XXV : C. chalcotoxa & allies
25. (22) General: Frontal protuberance an oval to heart-shaped hollow cup
with ventral portion slightly convex. Forewing coloring bronzy or with some yellow gold, no fringe tooth at tornus. Male genitalia: Phallus vesica elongate, secondary group of cornuti absent. Female genitalia: Ventral margins of VIII pleura close together, distance about equal to the width
of the ventral sclerotized half of the neighboring pleuron
Species group XXVI: B. chrysopis & allies
General: Frontal protuberance a heart-shaped ring with a subcentral truncate, conical secondary prominence. Forewing coloring never metal- lic gold; usually yellow and with a large hexagonal or quadrate macula mesad in inner area, small fringe tooth at tornus. Male genitalia: Phallus vesica pear-shaped, secondary group of cornuti present. Female genitalia: Ventral margins of VIII pleura widely separated, distance about twice the
width of the ventral sclerotized half of the neighboring pleuron
Species group XXVII: B. rugifrons & allies
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Genus 1: PLAGIOMIMICUS
1873, Plagiomimicus Grote. Buffalo Soc. Nat. Sci., B. 1:182. Type species:
P. pityochromus Grote, 1873; Albany, Alabama; monobasic.
1874, Stibadium Grote. Buffalo Soc. Nat. Sci., B. 2:74. Type species: S. spu- mosum Grote, 1874; Kansas; monobasic. Synonymy of Smith, 1882. Brooklyn Ent. Soc., B. 5:30, 35.
1875, Polenta Morrison. Boston Soc. Nat. Hist., Proc. 18:124. Type species: Schinia tepperi Morrison, 1875; Texas; monobasic. Synonymy of Grote, 1880. Brooklyn Ent. Soc., B. 3:38.
1877 , Antaplaga Grote. Canad. Ent. 9:70-71. Type species: A. dimidiata Grote, 1877; Colorado; monobasic. NEW SYNONYMY.
191 8, Neophaeus Dyar. U. S. Natl. Mus., Proc. 54:350. Type species: N. chalcospilans Dyar, 1918; Mexico; monobasic. NEW SYNONYMY.
Remarks
This large and perhaps unwieldy genus includes two major phyletic lines,
1 and 2. Possibly some of its diverse species groups are worthy of generic separation, most likely those of line 2, but in keeping with my policy of ex- treme conservatism in this preliminary study, I prefer to recognize a single broad genus. This also obviates the necessity of proposing several new generic names which might suffer synonymy when the group is studied more exten- ( sively.
The genus is very difficult to characterize as a whole because of its diverse constituents. The description below is only very generally applicable except for the female genitalic structures, primarily those of segment VIII which are the most constant and diagnostic. The single lobe of the distal sacculus division of the valve is extremely variable in position and shape throughout the genus but fairly constant in species groups. A double lobe is the only defining char- acter state separating the species groups on the second major phyletic line.
This state may have special significance, however, in representing a major evo- lutionary step.
Forbes (1954:171), in his key to the acronyctine genera, erroneously ascribes two “spines” (setal “claws”) to the foretibia of the genus “ Plagio - j mimicus,> (including only two species, pityochromus and concinnus) .
Description
General. A diverse genus. Typically medium in size and robustness, a few fairly large and small forms. Dull colored, usually brown. Frontal pro- tuberance quite varied in shape, usually an oval ring with a broad, rough ventral elevation, a few odd types. Foretibial “claw” a single massive seta, often flattened and curved. Forewing pattern typically noctuiform, dull col- ored, brown, some lighter colors: yellow, white. Thoracic vestiture of hair- like to spatulate, dentate scales; stiriine tufts present in most, poorly developed in some.
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Definition and Classification of Stiriini
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Male genitalia. Tegumen usually broad, narrow in some, neck short to very short, erect. Uncus short to moderately long, gradually curving, straight or gradually undulate. Saccus elongate. Valve variously shaped, usually oblong, ventral margin strongly convex in some; costa usually well expanded; basal sacculus division short, with broad, short lobe; distal division with single lobe, varied in shape and position; corona usually well developed, sometimes dif- fuse. Phallus vesica swollen, often expanded somewhat sinistrad as well as dextrad; cornuti well developed, numerous, moderately long, secondary group often with fairly long spinelets. Anellus sclerites crescent-shaped to triangular. Juxta subquadrate to shield-shaped. VIII tergal and sternal sclerotizations varied, former usually V-shaped, latter U-shaped.
Female genitalia. Ovipositor lobe varied in shape and degree of sclero- tization, from flat, partially sclerotized with striae to elongate, convex, heavily sclerotized and sharply pointed types, without striae. VIII pleuron evenly sclerotized (sometimes with small, irregular, unsclerotized areas), confluent with sternum, ventral margin not defined; no special, complex sterno-ostial concavity present. Lamella postvaginalis confluent with sternum, not a dis- crete sclerite, rarely fused with 1. antevaginalis (and sometimes ductus bursae also) to form a specially shaped, heavily sclerotized chamber. Corpus bursae fusiform to cudgel-shaped; posterior lobe usually well sclerotized and ribbed; sclerotized plate in crotch of lobe usually well developed and defined.
Species group I: P . ochoa and resolutus Figure 6
Description
General. Size medium, expanse about 30mm. Frontal protuberance sim- ple, an elongate to oval, rough surfaced plateau (slightly hollowed in ochoa). Foretibial “claw” broad and heavy, arising from moderately projecting, straight extension of tibial apex. Forewing apex acute, sharp, tornus moderately sharp; pattern noctuiform, ground color brown, markings darker brown, angulation of PM line beneath apical patch sharply acute, scattered blue scales in costal and inner areas. Hindwing brown, exterior line distinct. Scutal phragma mod- erately projecting. Thoracic vestiture of mixed linear and spatulate scales; tufts normal.
Male genitalia. Tegumen broad, slightly bulbous dorsad in ochoa, neck very short and erect. Uncus moderately long, cylindrical, curving sharply basad, more gradually distad; setae especially dense and erect dorsad. Saccus elongate, rectangular, rapidly narrowed cephalad. Valve oblong, distal half slightly narrowed; costa moderately expanded; basal sacculus division mod- erately long, ending rather abruptly, lobe a narrow, short finger; distal division with single, fingerlike lobe arising mesad; corona a single row of several, bristlelike, equal setae running along entire distal edge of valve. Phallus vesica slightly irregular, inflated cephalodistad; cornuti numerous, moderately long and heavy distad, shorter and thinner basad, secondary group a long row of
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short, cuspidate denticles, an irregular sclerotized area between patches; aedeagus somewhat angled. Anellus sclerites triangular. Juxta subquadrate. VIII tergal sclerotization V-shaped, arms and base narrow, former weak cau- dad; sternal sclerotization U-shaped.
Female genitalia. Ovipositor lobe of primitive, broad, flat, blunt type, striae distinct. Postapophysis almost twice as long as antapophysis, both slender. VIII pleuron with slight unsclerotized area centrally (striae faintly showing here in ochoa), ventral margin vaguely defined in resolutus. Sternum slightly unsclerotized caudad, jutting cephalad. Corpus bursae ellipsoid; pos- terior lobe with two short secondary lobes, sclerotization and ribbing mod- erately strong; sclerotized plate in crotch of first secondary lobe, a second plate also in crotch of second lobe.
Remarks
This group is quite distinct from the others in the genus in having the ovipositor lobes of the primitive type. Several other character states are also primitive and set the group apart (see “Phylogenetic analysis”). It may be advisable therefore to recognize the group as a separate genus but since there are only two species and there is general resemblance to the other Plagiomimi- cus, I choose not to follow this course
Included species
1. resolutus Dyar, 1909. Ent. Soc. Wash., Proc. 11:26 (Plagiomimicus) .
2. ochoa Barnes, 1904. Canad. Ent. 36:241-242 (Stibadium) .
=dollii Smith, 1908. N. Y. Acad. Sci., Ann. 18:118-119 (Plagio- mimicus). Synonymy of Barnes and McDunnough, 1917. Check List Lepid. Boreal Amer. p. 70.
Species group II : P. laodamia Figure 7
Description
General. Size medium large, expanse about 40mm. Frontal protuberance oval to heart-shaped, depressed dorsad and with broad lower elevation. Fore- tibial “claw” broad, flat, arising from moderately produced, straight tibial apex. Forewing apex acute, slightly sharp, tornus moderately sharp; pattern noctui- form, ground color medium brown, markings darker brown, medium shade not distinct, PM angulation beneath apical patch extreme, nearly reaching outer margin. Hindwing brown, exterior line distinct, offset distad. Thoracic vestiture mainly of linear dentate scales; anterior mesoscutal tuft developed in addition to other normal tufts.
Male genitalia. Tegumen broad, neck very short, erect. Uncus very short, stout, narrowed apically, curving sharply basad, setae long and dense dorsad. Saccus elongate, narrowed gradually cephalad. Valve elongate, rectangular, distal half slightly narrowed; costa moderately expanded; basal sacculus di- vision broad, moderately long, lobe narrow, prominent; distal division with
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Definition and Classification of Stiriini
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single spinelike lobe arising far distad, near edge of valve; corona a row of several stout, equal setae running along entire distal edge of valve. Phallus vesica normal, cornuti numerous, long, stout, closely bunched; secondary group a row of subequal, short denticles (longer sinistrad). Anellus sclerites crescent-shaped, fused cephalad. Juxta kite-shaped. VIII tergal sclerotization U-shaped, nearly circular, arms and base broad; sternal sclerotization U- shaped, arms not expanded.
Female genitalia. Ovipositor lobe slightly convex, pointed, and curved dorsad; striae distinct. Postapophysis about one-third longer than antapophy- sis, latter slightly sinuate. VIII pleuron completely sclerotized and confluent with sternum. Lamellae jutting slightly cephalad, forming a simple ring around ostium bursae. Corpus bursae cudgel-shaped, somewhat truncate cephalad; posterior lobe conoidal, sclerotization and ribbing strong; plate in crotch of lobe well developed.
Included species
3. laodamia Druce, 1890. Zool. Soc. London, Proc. p. 520 (Plusiodes) . The figure of this species is incorrectly labeled as laverna by Draudt (1919- 1939: pi. 43, row a, fourth from left).
Species group III. P. spumosus and allies Figure 8
Description
General. Size medium to medium-small, expanse about 23-35mm. Frontal protuberance an oval ring with a broad, moderately prominent lower eleva- tion. Foretibial “claw” flat, arising from strongly produced, straight tibial apex. Forewing apex strongly acute, to nearly a right angle, sharp, costal and outer margins moderately to slightly convex, tornus rounded to sharp; pattern varied but basically noctuiform, ground color brown, markings of darker shade (some special patterns and colors: yellow submarginal sector in aureolus, nearly all white in dimidiatus, crescent-shaped and oval, silver maculae in olvello, powdery suffusion in hilli), apical patch prominent in most, reniform and orbicular obsolescent in some, PM angulation beneath apical patch barely obtuse to sharply acute. Hindwing white or brown, markings obsolescent. Thoracic vestiture varied, usually of long, spatulate, dentate scales, some with mixed types; tufts normal.
Male genitalia. Tegumen broad, neck very short, erect. Uncus moderately long to long, cylindrical, slightly swollen subapically in most, gradually curv- ing. Saccus quite elongate, narrowed gradually cephalad, with rounded point. Valve oblong and slightly narrowed distad or broad, with straight dorsal and convex ventral margins; costa greatly expanded; basal sacculus division broad, swollen and very short, ending abruptly, lobe very broad and short; distal di- vision with single lobe, short and spikelike to very long and curving mesad, always arising about the center, sometimes a tiny secondary lobe present basad in addition; corona of slender setae at dorsal tip of valve, sometimes running
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obliquely onto inner surface. Phallus vesica normal, expanded sinistrad in some; cornuti numerous, varied in shape: moderately long and slender or stout and subequal in some, long, stout distad and shorter basad in others, secondary group a row of spinelets, latter shorter dextrad, quite long and angled sinistrad. Anellus sclerites oval, widely separated or triangular or crescent-shaped and approximated. Juxta subquadrate to shield-shaped. VIII tergal sclerotization narrowly V-shaped in most, arms weak or fused into a heavy convexity in hilli; sternal sclerotization U-shaped.
Female genitalia. Ovipositor lobe convex, varied in shape: apex pointed, usually abruptly narrowed subapically, evenly and heavily sclerotized, without striae, often with long setae (nearly reaching tip of lobe). Postapophysis slightly longer than antapophysis and often much heavier. VIII pleuron com- pletely sclerotized (a very slight medial unsclerotized area in some, never con- taining striae) and confluent with sternum. Sternum membranous caudad producing an indefinite lamella postvaginalis, latter jutting cephalad and con- tinuous laterad with 1. antevaginalis to form a wide, simple ring around os- tium, sometimes slightly enlarged to form an indefinite, squarish chamber or sclerotized and fused into a purse-shaped, wide-mouthed chamber. Corpus bursae cudgel-shaped or oblong, slightly curved in some; posterior lobe broad, sclerotization and ribbing strong in most, almost absent in some (those with oblong corpora) ; plate in crotch of lobe well developed.
Remarks
This is the largest species group in the genus and the tribe. It contains climidiatus, the genotype of Antaplaga, which formerly formed the nucleus for this conglomerate genus in spite of its obvious distinctive appearance (men- tioned by various authors, e.g., Barnes and Benjamin, 1924:170). Should anyone feel that this species group should have generic status of its own, the nominal genus Stibadium is available for it and carries a meaningful usage established by the McDunnough check list ( 1938) .
This group has been related to the next in the phylogeny section and could be combined with it to form a separate genus. Species group V ( P . tepperi ) may be included also to form a genus Plagiomimicus which would be more compact than the present and perhaps more useful.
One new species, bajae, is described herein (see appendix).
Included species
4. jalada Schaus, 1898. N. Y. Ent. Soc., J. 6: 144 (Stibadium).
5. aureolus Edwards, 1882. Papilio 2: 126-127 (Stibadium).
6. bajae Hogue, new species. (See appendix) .
7. olvello Barnes, 1907. Canad. Ent. 39:94-95 (Stibadium) .
8. raglena Dyar, 1912. U. S. Natl. Mus., Proc. 42:68 (Stibadium) .
9. unicus Barnes and Benjamin, 1926. Insec. Incit. Mens. 14:4-5 (Sti- badium).
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Definition and Classification of Stiriini
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10. chalcospilans Dyar, 1918. U. S. Natl. Mus., Proc. 54:350 ( Neo - phaeus ).
11. laverna Druce, 1898. Biol. Cent. Amer., Meter. 2:496-497; 3: pi. 95, fig. 22 ( Thalpochares ).
12. manti Barnes, 1904. Canad. Ent. 36:243 (Stibadium).
13. dimidiatus Grote, 1877. Canad. Ent. 9:71 (Antaplaga).
14. corozona Schaus, 1898. N. Y. Ent. Soc., J. 6: 144 ( Stibadium ).
15. astigmatosus Dyar, 1921. Insec. Incit. Mens. 9:42 {Stibadium) .
16. curiosus Neumoegan, 1883. Papilio 3: 141-142 ( Stibadium ).
17. spumosus Grote, 1874. Buffalo Soc. Nat. Sci., B. 2:74 {Stibadium) .
18. mavina Barnes and McDunnough, 1910. Canad. Ent. 42:250-251 {Sti- badium). I suspect that this species was based on a faded or aberrant specimen of the preceding and should be synonymized with it. I have not examined the types, however, so leave a formal determination of synonymy to future students.
19. hilli Barnes and Benjamin, 1923. Contrib. Nat. Hist. Lepid. N. Amer. 5:83-84 (Stiria).
=caliente Hill, 1923. So. Calif. Acad. Sci., B. 22: 17; fig. {Antaplaga) . Synonymy of Hill, 1923. Ibid. p. 19.
Species group IV : P. pityochromus and allies Figure 9
Description
General. Size medium to medium-small, expanse about 25-33 mm. Frontal protuberance a circular, smooth-bottomed prominent cup with no secondary prominence. Foretibial “claw” shovel-shaped, arising from strongly produced, strongly angled tibial apex. Forewing apex acute, moderately sharp, tornus rounded; pattern noctuiform, ground color medium to light brown, transverse lines light, apical patch prominent, orbicular markedly dark (absent in expal- lidus). Hindwing white to brown. Thoracic vestiture of spatulate scales; tufts normal.
Male genitalia. Tegumen moderately broad, neck very short and erect. Uncus moderately long, swollen mesad, undulate; setae fairly numerous, longer ventrad. Saccus elongate, narrowed cephalad with a rounded point. Valve ob- long, apex rounded; costa little expanded; basal sacculus division moderately broad, ending somewhat abruptly, lobe short; distal division with a single, mod- erately long, curved, spinelike lobe set far distad near edge of valve (except in concinnus where it is more basal as in preceding unit) ; corona of few, slender setae, restricted to dorsal apex of valve or normal. Phallus vesica normal; cor- nuti numerous, subequal, moderately long and stout, secondary group a patch of fairly long, stout spinelets. Anellus sclerites widely separated, crescent- shaped. Juxta broadly kite-shaped. VIII tergal sclerotization triangular, heavily sclerotized and convex caudad; sternal sclerotization U-shaped, arms broad- ened caudad.
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Female genitalia. Ovipositor lobe convex, evenly and strongly sclerotized, apex pointed, usually abruptly narrowed subapically, without striae, with long setae (about reaching tip of lobe). Postapophysis slightly longer and heavier than antapophysis, both usually slender (stout in expallidus) . VIII pleuron completely sclerotized (small irregular unsclerotized areas in some, contain- ing no striae) and confluent with sternum. Sternum unsclerotized caudad, pro- ducing an indefinite lamella postvaginalis, latter jutting cephalad and connect- ing with 1. antevaginalis to form a wide, simple ring around ostium or enlarged to form a squarish, indefinite chamber. Corpus bursae cudgel-shaped, curved; posterior lobe with two secondary lobes, sclerotization and ribbing strong; sclerotized plate in crotch of first lobe well developed.
Remarks
As mentioned above, this group is not sharply differentiated from the preceding. At present it corresponds approximately ( concinnus added) to the genus Plagiomimicus in the restricted sense of previous authors (Draudt, 1919- 1939:313; McDunnough, 1939:99-100).
Included species
20. expallidus Grote, 1882. Papilio 2:185 (Plagiomimicus).
21. triplagiatus Smith, 1890. Ent. Amer. 6: 139 (Plagiomimicus).
22. pityochromus Grote, 1873. Buffalo Soc. Nat. Sci., B. 1:182 (Plagio- mimicus).
—medius Morrison, 1875. Boston Soc. Nat. Hist., Proc. 18:123-124 (Schinia). Synonymy of Grote, 1882. New Check List N. Amer. Moths p. 35. This name applies to specimens with a split reniform. There may be at least a subspecific difference involved but it awaits determination.
23. concinnus Dyar, 1909. Ent. Soc. Wash., Proc. 11:23 (Stibadium).
Species group V : P. tepperi Figure 10
Description
General. Size somewhat varied, generally small, expanse about 23-28 mm. Frontal protuberance an inverted, heart-shaped ring with lower lip produced into a conical prominence. Foretibial “claw” tiny, arising from unproduced tibial apex. Forewing apex acute, sharp, outer margin shallowly undulate, tornus rounded; pattern noctuiform, ground color olive green, lines white, api- cal patch prominent, reniform rudimentary, orbicular absent. Hindwing white to brown, markings obsolescent. Thoracic vestiture of spatulate scales; tufts normal.
Male genitalia. Tegumen very broad, neck very short, erect. Uncus short, stout, gradually curving; setae short, erect and longer dorsad. Saccus stout, elongate, cylindrical, rounded cephalad. Valve very short, almost rhomboid; costa expanded obliquely; basal sacculus division very broad, triangular, end-
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Definition and Classification of Stiriini
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ing abruptly, lobe a short, straight finger; distal division with a single, long, thin, spinelike lobe, set near edge of valve; no discrete corona. Phallus vesica small; cornuti stout, shorter basad, secondary group absent. Anellus sclerites crescent- shaped. Juxta rectangular. VIII tergal sclerotization bracket-shaped; sternal sclerotization of two parallel, separated strips, weakly connected cephalad.
Female genitalia. Ovipositor lobe convex, evenly and heavily sclerotized, acuminate, setae short. Postapophysis almost a third longer than antapophysis and much heavier. VIII pleuron completely sclerotized and confluent with sternum. Sternum slightly unsclerotized caudad. Ductus bursae very heavily sclerotized and confluent with lamellae to form a bowl-shaped chamber. Corpus bursae very narrow caudad, gradually swollen cephalad; posterior lobe with weak secondary lobes; sclerotization and ribbing strong.
Remarks
As mentioned in the phylogeny section, this species group (one species, tepperi ) deviates somewhat from the two preceding groups. It might, therefore, be considered as an autonomous genus to which the name Polenta may be ap- plied (as has been done by most recent authors).
Included species
24. tepperi Morrison, 1875. Acad. Nat. Sci. Philadelphia, Proc. pp. 68-69 (Schinia).
—richii Grote, 1886. Canad. Ent. 18:99-100 {Plagiomimicus) . This synonymy is discussed in detail in the section on taxonomic history.
Species group VI: P. hutsoni and olivalis Figure 1 1
Description
General. Size medium small or small, expanse about 20 or 30mm. Frontal protuberance an inverted heart-shaped ring with lower lip produced into a conical prominence. Legs slender, foretibial “claw” moderately strong, arising from strongly projecting, straight tibial apex. Forewing apex acute, rounded, tornus rounded; pattern as follows: median and submarginal sectors white, suffuse, fuliginous, brownish-green transverse lines, reniform and orbicular absent, apical patch obsolete. Hindwing brownish-black, markings obsolete. Thoracic vestiture hairy {olivalis), or of short spatulate scales {hutsoni) ; tufts poorly developed in both.
Male genitalia. Tegumen moderately broad, neck short. Uncus moder- ately long, cylindrical, tip slightly narrowed, straight; setae short and sparse. Saccus elongate or stout. Valve oblong; costa small; basal sacculus division short, ending diffusely, lobe a broad, low mound; distal division with a single, short, stout spine, set far distad; corona diffuse, weak. Phallus vesica slightly inflated sinistrad; cornuti moderately long {olivalis) or very short {hutsoni) and subequal, secondary group a short row of spinelets on a convex, sclerotized
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plate. Anellus sclerites separated, U-shaped. Juxta shield-shaped. VIII tergal sclerotization of parallel, entirely separate strips; sternal sclerotization a nearly complete circle, broad caudad, narrow cephalad.
Female genitalia. Ovipositor lobe convex, evenly and heavily sclerotized, elongate, bluntly pointed. Postapophysis and antapophysis slender, subequal. VIII segment very short, pleuron with large, central, oval, unsclerotized area, confluent with sternum. Sternum unsclerotized caudad to form an ill-defined, chevron-shaped lamella postvaginalis; 1. antevaginalis ligulate, lamellae not jutting cephalad, flush with segment. Ductus bursae completely membranous. Corpus bursae cudgel-shaped; posterior lobe simple, sclerotization and ribbing very weak, essentially absent in hutsoni.
Included species
25. hutsoni Smith, 1907. Amer. Ent. Soc., Trans. 33:140 ( Stibadium ). =fuliginosus Smith, 1907. Amer. Ent. Soc., Trans. 33.140-141 (Sti- badium). Synonymy of Barnes and McDunnough, 1916. Contrib. Nat. Hist. Lepid. N. Amer. 3:168. I have seen the types of these nominal species and agree with this synonymy.
26. olivalis Barnes and McDunnough, 1916. Contrib. Nat. Hist. Lepid. N. Amer. 3: 12-13, pi. 3, fig. 8 ( Stiria ).
Species group VII: P. hachita and allies Figure 12
Description
General. Size medium-small, expanse about 25-28mm. Frontal protuber- ance a moderately deep, circular, smooth cup with prominent lower lip. Fore- tibial “claw” shovel-shaped, arising from a strongly projecting, slightly angled tibial apex. Forewing apex acute, rounded, tornus rounded; pattern as follows: broad white median sector between well defined, black AM and PM lines, outer and basal sectors white also or contrasting yellow, ST line broken into a row of small dots (absent in biundulalis) . Thoracic vestiture of linear scales; tufts poorly developed.
Male genitalia. Tegumen moderately broad, neck short. Uncus moder- ately long, slightly swollen mesad, straight to undulate; setae short. Saccus elongate, pointed cephalad. Valve oblong, costa small; basal sacculus division short, ending diffusely, lobe only a broad, flat mound; distal division with a single, short, stout spine, set far distad; corona diffuse, weak. Phallus vesica slightly inflated sinistrad; cornuti numerous, moderately long and subequal, secondary group a row of long spinelets. Anellus sclerites crescent-shaped, fused or separated. Juxta shield-shaped. VIII tergal sclerotizations two entirely separate, parallel strips; sternal sclerotization U-shaped, arms very broad caudad.
Female genitalia. Ovipositor lobe convex, evenly and heavily sclerotized, triangular, sharply pointed. Postapophysis and antapophysis subequal, stout.
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Definition and Classification of Stiriini
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VIII very short, pleuron with large, central, oval, unsclerotized area, confluent with sternum. Lamellae as in preceding species group. Ductus bursae com- pletely membranous. Corpus bursae cudgel-shaped; posterior lobe simple; sclerotization and ribbing very weak.
Included species
27. hachita Barnes, 1904. Canad. Ent. 36:241 (Antaplaga) .
28. sexseriatus Grote, 1881. Papilio 1:155 ( Grotella ).
29. biundulalis Zeller, 1872. Zool. Bot. Ges. Wien, Verhandl. 22:502-503, taf. Ill, figs. 1 4a- 1 4c ( Sedenia ).
Species group VIII : P. alesaea Figure 13
Description
General. Size small, expanse about 23mm. Frontal protuberance an oval or heart-shaped ring inclosing a low, subcentral, truncate, secondary promi- nence. Foretibial “claw” quite heavy and long, arising from strongly project- ing, straight tibial apex. Forewing short, shape nearly deltoid; pattern com- pletely absent, self-colored, white. Hindwing patternless and white also. Thor- acic vestiture of spatulate scales; tufts poorly developed (? my specimens rubbed).
Male genitalia. Tegumen moderately broad, neck short. Uncus short, stout, swollen mesad; setae dense, longer and erect dorsad. Saccus elongate, pointed cephalad. Valve oblong; costa small; basal sacculus division moder- ately broad, ending diffusely, lobe a short pommel; distal division with a single, long, sharp, stout spine set very far distad and extending off the valve margin; corona weak. Phallus vesica slightly inflated sinistrad; cornuti few, distal ones very short and stout, set on a curved, sclerotized plate, basal ones slightly longer and finer, free from plate, secondary group absent, a few enlarged spicules in a row on sinistral inflation instead. Anellus sclerites crescent- shaped, separated. Juxta broadly shield-shaped.
Female genitalia. Ovipositor lobe convex, acuminate, strongly and evenly sclerotized. Postapophysis about one-quarter longer than antapophysis, both slender. VIII pleuron completely sclerotized, confluent with sternum. Sternum unsclerotized caudad and jutting cephalad as a semicircular plate (lamella postvaginalis) to form a flat, simple chamber with the 1. antevaginalis. Corpus bursae oblong; lobe simple, sclerotization and ribbing strong and extending over one-third of the entire corpus.
Remarks
The single species in this group was undoubtedly placed originally in the genus Antaplaga because of the lack of any wing pattern in common with sev- eral species formerly included in that genus but segregated in the present study as the genus Chichimeca. There is definitely no close relationship between
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alesaea and species of Chichimeca, the similarity in wing ornamentation having arisen undoubtedly by parallel degeneration and loss of pattern frequently seen in noctuids.
Included species
30. alesaea Dyar, 1918. U. S. Natl. Mus., Proc. 54:350 (Antaplaga) .
Species group IX: P. psamathochromus and argyropolius Figure 14
Description
General. Size medium, expanse about 30-3 3mm. Frontal protuberance an oval ring with a broad, low, ventral elevation. Foretibial “claw” strong and flat, arising from strongly produced, straight tibial apex. Forewing apex acute, sharp, tornus moderately sharp; pattern noctuiform but simplified: color light silvery-grey or powdery grey-brown, when former, AM and PM lines fine, brownish-gold ( argyropolius ), when latter, generally like most species in group III (psamathochromus) , orbicular and reniform obsolescent in both. Hindwing pale, markings obsolete. Thoracic vestiture primarily of linear scales; tufts poorly developed.
Male genitalia. Tegumen unusually narrow, ventral arms long, neck short. Uncus short, fusiform; setae numerous, longer and erect dorsad. Saccus very long, rounded cephalad. Valve rhomboid-oblong, distoventral corner produced or strongly angled; costa expanded moderately obliquely; basal sacculus di- vision short, swollen, deltoid, nearly confluent with distal division, lobe a short, broad or narrow finger; distal division with two lobes: basal irregular, finger- like, distal a strong, curved spine; corona weak. Phallus vesica normal; cor- nuti numerous, moderately long distad, shorter basad (latter extending far basad over ventral surface in psamathochromus ) , secondary group a short row of several spinelets, these longer and curved sinistrad in argyropolius, subequal in psamathochromus. Anellus sclerites long, approximated cephalad. Juxta subquadrate, shield-shaped. VIII tergal sclerotization U-shaped, base very broad; sternal sclerotization U-shaped, arms broad.
Female genitalia. Ovipositor lobe convex, evenly and heavily sclerotized, sagittate. Postapophysis about one-fifth longer than antapophysis, both very slender. VIII pleuron completely sclerotized, confluent with sternum (ventral margin faintly defined caudad in psamathochromus) . Lamellae specially sclero- tized and fused into a purse-shaped, narrow-mouthed chamber. Corpus bursae oblong or cudgel-shaped; lobe simple, sclerotized plate in crotch of lobe ex- tensive, extending onto lobe considerably; sclerotization and ribbing moder- ately strong.
Remarks
This species group and the next are related and distinct from the others in the genus Plagiomimicus in possessing a double lobe on the distal sacculus
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division of the male valve. The special phyletic significance of this state has already been discussed and may be sufficient to warrant the separation of this group as a separate genus.
Included species
31. psamathochromus Dyar, 1909. Ent. Soc. Wash., Proc. 1 1 :22-23 (57/- badium).
32. argyropolius Dyar, 1914. U. S. Natl. Mus., Proc. 47:376 ( Stiria ).
Species group X: P. phalaenoides Figure 15
Description
General. Size medium, expanse about 28mm. Frontal protuberance an ovoid ring with a subcentral, broad, truncate lower elevation. Foretibial “claw” moderately heavy and shovel-shaped, arising from strongly projecting, straight tibial apex. Forewing almost identical in all respects to that of argyropolius in preceding group but markings a little darker. Hindwing pale, markings obso- lete. Thoracic vestiture of mixed linear and spatulate scales; tufts poorly de- veloped.
Male genitalia. Tegumen moderately broad, ventral arms long, neck mod- erately long. Uncus short and stout; setae numerous, longer and erect dorsad. Saccus elongate, rounded cephalad. Valve oblong, angular, twice narrowed, once mesad, once again distad, apex rounded, acuminate; costa moderately expanded; basal sacculus division inflated, deltoid, somewhat confluent with distal division, lobe a short, broad finger; distal division with two lobes, both fingerlike, basal broader, distal with minute denticles apically; corona diffuse. Phallus vesica cylindrical, slightly expanded sinistrad; cornuti very few (2-5) and short, secondary group a patch of several unequal denticles arising from sclerotized plate, latter extending distad nearly to primary group. Anellus sclerites ovoid, separated. Juxta shield-shaped, VIII tergal sclerotization nearly circular, broad cephalad; sternal sclerotization broadly U-shaped, arms ex- panded.
Female genitalia. Ovipositor lobe convex, evenly and heavily sclerotized, short, acuminate. Postapophysis nearly twice as long as antapophysis. VIII pleuron completely sclerotized, confluent with sternum. Sternum unsclerotized caudad, lamellae not modified into a large, narrow-mouthed, purse-shaped chamber, instead, an indistinct, small chamber with a wide mouth present. Corpus bursae elongate, cudgel-shaped; lobe simple, sclerotized plate in crotch well developed; sclerotization and ribbing moderately strong but somewhat re- stricted in area.
Included species
33. phalaenoides Dyar, 1918. Insec. Incit. Mens. 6:133-134 (Stiria).
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Genus 2: CHRYSOEC1A
1910, Chrysoecia Hampson. Catalogue Lepid. Phalaenae Brit. Mus. 9:182. Type species: Anthoecia scira Druce, 1889; Ruicon, Guerrero, Mexico; monobasic.
Remarks
Hampson originally erected this genus to include the single species scira. I have enlarged the genus to include two of the three primary phyletic units of major phyletic line 3, excluding the primary phyletic unit represented by one species, Gorgora morga (unit XIII) which I treat as generically distinct (see below). The two species groups (units XI and XII) of this genus are super- ficially very different but have female genitalia of the same type.
Description
General. Medium-small in size. Frontal protuberance of two types: (1) a shallow, round cup with slightly produced ventral lip or (2) a transverse trough with a ventral, external, median node, a low, semicircular ridge below both types on the frons. Foretibial “claw” a single seta. Forewing pattern of two types (see descriptions below), PM line always strongly zigzagged, reni- form and orbicular (especially latter) with a tendency to enlarge and square out, lunules usually marked. Thoracic vestiture usually somewhat hairy with poorly developed tufts.
Male genitalia. Tegumen normal in breadth or somewhat narrowed, neck short and erect or moderately long. Uncus usually moderately long and cylin- drical. Valve oblong, usually slightly curved; costa moderately expanded; basal sacculus division broad, often with a fairly long, fingerlike lobe (may be re- duced); distal division always with two lobes: basal lobe fingerlike and often weak, distal lobe spinelike; coronal setae slender. Phallus vesica normal, slight- ly angular; cornuti often diminutive in center of primary group, secondary group a row of denticles or a very few (2-5), unequal spines.
Female genitalia. Uniform. Ovipositor lobe of primitive, blunt flat type, striae distinct. VIII pleuron with large, elongate, median, unsclerotized area containing distinct striae, ventral margin defined evenly along entire length, contiguous or approximate with opposite margin. Sternum membranous or sclerotized. Lamellae ill-defined, much as in preceding genus, forming a sim- ple ring around the ostium bursae. No special, complex sterno-ostial concavity present. Ductus bursae membranous or heavily sclerotized and confluent with lamellae. Corpus bursae ellipsoid or elongate; lobe simple; sclerotization and ribbing strong.
Species group XI: C. scira and allies Figure 16
Description
General. Size medium-small, expanse about 25-30mm. Frontal protuber- ance a shallow, small round cup, with ventral lip slightly produced externally,
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a low, semicircular ridge below this on frons. Foretibial “claw” moderately strong, arising from slightly produced tibial apex. Forewing apex rounded, tornus rounded and margins moderately convex; pattern and coloring com- plex: AM and PM lines zigzagged and dark, orbicular and reniform obsoles- cent, center of former greatly expanded in some, scattered blue scales through median sector and costal area, outer and basal areas white. Hindwing fuscous, lighter basad, markings obsolescent. Thoracic vestiture of linear to moderately spatulate scales; tufting usually poorly developed, small, lateral and anterior mesoscutal tufts developed in scira.
Male genitalia. Tegumen slender, ventral arms somewhat long, neck mod- erately long. Uncus moderately long to long, cylindrical. Saccus triangular to oblong. Valve oblong, straight or slightly upcurved; costa moderately ex- panded; basal sacculus division broad, ending diffusely, lobe a short, thin finger or broad, short peak; distal division with weak basal lobe, distal lobe a sharp, moderately long spine arising near edge of valve; coronal setae slender. Phallus vesica slightly angular; cornuti several, moderately long and stout in scira, more numerous, moderately long and stout only peripherad (diminutive cen- trad) in all others, secondary group a row of short, subequal denticles. Anellus sclerites crescent-shaped to ovate. Juxta shield-shaped. VIII tergal sclerotiza- tion V-shaped; sternal sclerotization U-shaped, arms slightly expanded.
Female genitalia. Postapophysis slightly longer than antapophysis. Ven- tral margins of VIII pleura contiguous mesad. Lamella postvaginalis vaguely triangular, jutting cephalad slightly with ligulate 1. antevaginalis. Ductus bursae membranous caudad, becoming abruptly sclerotized and ribbed about halfway. Corpus bursae ellipsoid; lobe simple; sclerotization and ribbing strong.
Included species
34. scira Druce, 1889. Biol. Cent. Amer., Heter. 1:301; 3: pi. 28, fig. 5 (Anthoecia) .
=benjamini Hill, 1924. So. Calif. Acad. Sci., B. 23:158 ( Chamoclea , sic) [ Chamaeclea ]. NEW SYNONYMY. The Arizona specimens of this spe- cies given this name by Hill may be subspecifically distinct from scira but I do not regard them worthy of such recognition.
35. gladiola Barnes, 1907. Canad. Ent. 39:67 ( Chamaclea , sic) [ Chamae- clea].
3 6. requies Dyar, 1909. Ent. Soc. Wash., Proc. 11:21-22 ( Centrartha , sic) [Centrarthra].
Species group XII: C. atrolinea and allies Figure 17
Description
General. Size medium-small, expanse about 28mm. Frontal protuberance a transverse, shallow trough with a ventral, external node, a low, semi-circular ridge below this on frons. Foretibial “claw” moderately strong, flattened, aris- ing from slightly produced tibial apex. Forewing apex rounded, tornus mod-
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erately sharp; pattern varied with regard to presence or absence of elements but extremely constant in position and color: ground color orange-yellow, markings black, thin zigzagging lines, median shade complete, lunules marked. Hindwing fuscous. Thoracic vestiture hairy; tufts poorly developed.
Male genitalia. Tegumen moderately broad, neck short, erect. Uncus moderately long, sharply curved basad, swollen distad. Saccus elongate, rounded cephalad. Valve oblong, slightly upcurved; costa expanded somewhat obliquely; basal sacculus division broad, somewhat inflated, ending diffusely, lobe a long, curved finger; distal division lobes about equal, in length, directed at right angles, basal lobe fingerlike, distal lobe spinelike; coronal setae slender, few. Phallus vesica normal; cornuti moderately long, stout peripherad, short and thin centrad, secondary group of a few, unequal, fairly long spines. Anellus sclerites triangular, approximate cephalad. Juxta subquadrate. VIII tergal sclerotization V-shaped; sternal sclerotization U-shaped.
Female genitalia. Postapophysis almost one-third longer than antapophy- sis. Ventral margins of VIII approximate caudad. Ill-defined lamella post- vaginalis filling entire space between ventral VIII margins. Ductus bursae heav- ily sclerotized and fused with lamellae. Corpus bursae elongate; lobe simple; sclerotization and ribbing strong.
Included species
37. atrolinea Barnes and McDunnough, 1912. Contrib. Nat. Hist. Lepid. N. Amer. 1:25, pi. l,fig. 3 (Antaplaga) .
38. dela Druce, 1894. Ann. Mag. Nat. Hist., Ser. 6, 13:361 (Acontia) .
39. stigmatosa Dyar, 1912. U. S. Natl. Mus., Proc. 42:67 (Antaplaga) .
40. hemicrocea Dyar, 1912. U. S. Natl. Mus., Proc. 42:67-68 (Anta- plaga).
These four species are considered by Draudt (1919-1939:324) as two species, dela and atrolinea with stigmatosa and hemicrocea as forms of dela. He mentions, however, that even atrolinea may be nothing more than a north- ern form of dela. I tend to concur with the latter view although there is some possibility that hemicrocea, which is the most distinctive in wing pattern, may actually constitute a separate species. These forms need further study to de- termine their proper status.
Genus 3: GORGORA
1914, Gorgora Dyar. U. S. Natl. Mus., Proc. 47:377. Type species: G. morga Dyar, 1914; Mexico City, Mexico; monobasic.
Remarks
There is only a single species group containing one species in this genus but it is so distinct from the other members of major phyletic line 3 on the basis of the male genitalia that I choose to recognize its original generic status even though the female genitalia do not deviate from the type typical of the line. Dyar described this genus originally in the subfamily Erastriinae (Aeon-
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tiinae) possibly by reason of its gaudy colors. It has now been shifted to an entirely different subfamily removing an enigma for the acontiine specialists.
Description
See description of species group below.
Species group XIII: G. morga Figure 18
Description
General. Size medium, expanse about 32mm. Frontal protuberance a sim- ple, rough oblong plateau slightly hollowed dorsad, a low semicircular ridge below this on frons. Foretibial “claw” moderately stout, arising from slightly produced tibial apex; plantar bristles long with rounded tips. Forewing shape somewhat elongate, margins somewhat convex, apex and tornus rounded; pat- tern unique: ground color shining black with a large quadrangular, orange discal spot, marginal area orange, lunules black. Hindwing fuscous. Thoracic vestiture hairy, orange; tufts poorly developed.
Male genitalia. Tegumen broad, ventral arms moderately long, neck very short and erect. Uncus moderately long, gradually curving, slightly swollen subapically; setae numerous, moderately long. Valve triangular-oblong; costa moderately expanded; basal sacculus division very broad, ending diffusely, lobe a short pommel; distal division with a single, unequally forked, heavily sclerotized lobe, arising mesad; corona represented by two, very short, stout apical setae. Phallus vesica normal; cornuti moderately long, stout peripherad, diminutive centrad, secondary patch a curved row of short spinelets. Anellus sclerites crescent-shaped. Juxta subquadrate. VIII tergal sclerotization V- shaped; sternal sclerotization square.
Female genitalia. (Described from photograph of holotype.) Ovipositor lobe of primitive, flat, blunt type, striae distinct. Postapophysis almost one- fourth longer than antapophysis. VIII pleuron with an elongate, median un- sclerotized area containing distinct striae, ventral margin defined but not de- flected, contiguous mesad with opposite margin. No special, complex sterno- ostial concavity present. Lamellae not determinable from photograph. Corpus bursae elongate; lobe simple; sclerotization and ribbing strong.
Included species
41. morga Dyar, 1914. U. S. Natl. Mus., Proc. 47:378 ( Gorgora ).
Genus 4: CHICHIMECA Hogue, new genus Type species: Eulithosia thoracica Edwards, 1884; Arizona.
Remarks
The peculiar degenerate nature and fairly numerous species of the single primary phyletic unit XIV of the fourth major phyletic line make it desirable
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to recognize it as a distinct genus. No other name being available, I propose the new genus Chichimeca.
The genus is made up of several species formerly placed in the nominal genus Antaplaga. The general appearance of these was responsible largely for a few Grotella species and Plagiomimicus alesaea being lumped into this genus. All have similar facies and self-colored, white (rarely yellow-orange) wings but, of course, are distinct structurally.
Description
See description of species group below.
Species group XIV : C. thoracica and allies Figure 19
Description
General. Size medium-small to small, expanse about 22-32mm. Frontal protuberance a circular to oval ring inclosing a subcentral, truncate secondary prominence. Foretibial “claw” quite stout, shovel-shaped, arising from mod- erately produced, straight tibial apex. Forewing shape elongate, all margins moderately convex, apex acute, tornus broadly rounded; pattern absent, self- colored, sooty white to yellow-orange. Hindwing varied in color, usually like forewing, pattern obsolete. Thoracic vestiture somewhat hairy, tufts poorly developed.
Male genitalia. Tegumen moderately broad, neck short and erect. Uncus moderately long, cylindrical, slightly swollen mesad or subapically, slightly undulate; setae sparse and short. Saccus oblong, nippled cephalad. Valve ob- long, slightly curved; costa greatly expanded; basal sacculus division short, lobe pommel-shaped; distal division with a single, small spikelike lobe arising mesad; corona weak, usually running obliquely onto inner surface of valve. Phallus vesica normal; cornuti numerous, short, spiculate, secondary group a short row of spinelets. Anellus sclerites crescent-shaped, fused. Juxta shield- shaped. VIII tergal sclerotization narrowly U-shaped; sternal sclerotization U- shaped, arms broad.
Female genitalia. Attenuated. Ovipositor lobe of primitive flat, blunt type, striae distinct. Postapophysis about one-fifth longer than antapophysis, both very slender and attenuated. VIII pleuron with an elongate, median, unsclero- tized area containing distinct striae, ventral margin defined and slightly de- flected, approximate with opposite margin caudad, diverging out around lamella postvaginalis and associated with it and 1. antevaginalis to form an ill-defined sterno-ostial concavity. Lamella postvaginalis elongate, oval; I. antevaginalis weak, ligulate, bowed cephalad. Ductus bursae entirely membranous. Corpus bursae elongate, constricted gradually mesad; lobe simple, pointed; sclerotiza- tion and ribbing obsolescent.
Remarks
The species in this group are quite uniform. Several nominal species
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Definition and Classification of Stiriini
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among the smaller white forms (numbers 44-51) may be synonymized upon further study.
Included species
42. thoracica Edwards, 1884. Papilio 4:44 (Eulithosia) .
43 . spreta Draudt, 1927 in Seitz. Macrolepid. World 7:325, pi. 47, row a ( Antaplaga ).
44. subfumosa Dyar, 1909. Ent. Soc. Wash., Proc. 1 1:25-26 (Antaplaga) .
45. anaemica Draudt, 1927 in Seitz. Macrolepid. World 7:324, pi. 46, row k ( Antaplaga ) . Originally described as a new form of subfumosa. Its exact status in this difficult group awaits further study.
46. salacon Druce, 1895. Ann. Mag. Nat. Hist., Ser. 6, 16:39 (Grotella) .
47. simplicia Dyar, 1926. Insec. Incit. Mens. 14:185 (Antaplaga) .
48. dulcita Schaus, 1898. N. Y. Ent. Soc., J. 6:144 (Grotella).
49. pseudovarra Dyar, 1926. Insec. Incit. Mens. 14: 184 (Antaplaga) .
50. varrara Dyar, 1918. U. S. Natl. Mus., Proc. 54:350 (Antaplaga) .
51. muricolor Dyar, 1926. Insec. Incit. Mens. 14:184 (Antaplaga) .
Genus 5. CIRRHOPHANUS
1812, Cirrhophanus Grote. Canad. Ent. 4:187. Type species: C. triangulifer Grote, 1872; Missouri ?; monobasic.
1884, Eulithosia Edwards. Papilio 4:43. Type species: E. composita Edwards, 1884; Arizona; selection of Hampson, 1910. Catalogue Lepid. Phalaenae Brit. Mus. 9:384. NEW SYNONYMY.
1910, Hoplolythra Hampson. Catalogue Lepid. Phalaenae Brit. Mus. 9:214. Type species: Lythrodes discistriga Smith, 1903; syntypes: Walter’s Station, California, Southern Arizona; monobasic. NEW SYNONYMY. 1918, Pumora Dyar. U. S. Natl. Mus., Proc. 54:348. Type species: P. hyperion Dyar, 1918: Cuernavaca, Mexico; monobasic. NEW SYNONYMY.
Remarks
This genus, comprising major phyletic line 5, is, like Plagiomimicus, an- other large and somewhat unwieldy assemblage of primary phyletic units. There is greater uniformity of structure here, however. The female genitalia are consistently similar, the special sterno-ostial concavity always being pres- ent as a rounded depression between well defined and moderately deflexed or declining ventral margins of the VIII abdominal pleura. The latter have a large V-shaped unsclerotized median area sometimes containing striae. The ovi- positor lobe is generally of the typical primitive, flat, blunt type but a few secondary, relatively minor modifications of this occur. The wing pattern is characteristically streaked and orange is a frequent ground color.
Most of the atypical members of this genus have in common a multiple foretibial “claw’’ which immediately classes them as derivative. Simplified wing
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pattern and small size are other more conspicuous, derivative features. Some of these atypical forms have had separate generic status formerly, e.g., Pumora (unit XXI) and Hoplolythra (unit XX). These may prove to be valid with future study. For the present, however, I choose to recognize a large, encom- passing genus to emphasize similarities rather than dissimilarities.
Description
General. A large and varied genus. Size small to medium-large. Frontal protuberance usually an oval to reniform ring with lower lip mesally produced into a conical, secondary prominence or an inverted, heart-shaped ring with a central, truncate, secondary prominence ( chryseochilus only). Foretibial “claw” a single seta, a group of two or three separate setae or a multiple group of fused setae. Forewing shape fairly constant, angles slightly rounded, tornus often sharply angled; pattern varied but usually with streaks due to darkened veins and lines between veins (streaks in cell Cu often forming a trident, three pronged figure, with base distad) ; colors varied but usually orange or yellowish- orange with darker orange or brown markings. Hindwing varied in color (may be bicolored: black and orange), markings obsolete. Thoracic vestiture quite varied, almost always of spatulate or broad scales; tufts strong, the normal stiriine tufts present plus additional mesoscutal tufts in some.
Male genitalia. Tegumen broad, neck short to very long. Uncus usually fairly long and curving. Valve usually rhomboid-oblong or simply oblong; costa expanded variously; basal sacculus division moderately long, lobe usu- ally short but may be quite long; distal division with at least one lobe which is usually distal and spinelike but sometimes two lobes present which are close together (even on common plate) ; corona always weak. Phallus vesica normal, simple (with small appendicular lobe in chryseochilus) ; cornuti varied, primary group usually of numerous, long spines, secondary group usually present and composed of short spinelets.
Female genitalia. Ovipositor lobe varied, usually pointed and with striae, not markedly modified from primitive stiriine flat, blunt type. VIII pleuron with an elongate, median unsclerotized area (containing weak striae in some), ventral margin defined and moderately deflexed or declining cephalad, approxi- mate caudad with opposite margin and curving or diverging cephalad around lamella postvaginalis and associated with it and 1. antevaginalis to form a spe- cial, complex sterno-ostial concavity, latter usually circular or subquadrate in shape. Lamella postvaginalis round or somewhat reduced and ill-defined; 1. antevaginalis ligulate, bowed. Corpus bursae very varied in shape (see descrip- tions of species groups); sclerotization and ribbing almost always strong.
Species group XV : C. chryseochilus Figure 20
Description
General. Size medium, expanse about 33mm. Frontal protuberance an inverted, heart-shaped ring inclosing a subcentral, truncate secondary promi-
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nence, latter barely exceeding height of ring. Foretibial “claw” a single, shovel- shaped seta arising from moderately produced tibial apex. Forewing apex acute, sharp, tornus sharp; pattern complex, basically noctuiform: ground color purplish-brown basad, veins dark producing streaked pattern, streaks in cell Cu forming a trident, marginal sector brownish-yellow, AM line obso- lescent, PM line strong. Hindwing brown, markings absent. Thoracic vestiture mixed, mostly of linear scales; tufts normal.
Male genitalia. Tegumen broad, ventral arms short, neck very long. Uncus long, cylindrical, gradually curving. Saccus oblong. Valve oblong, slightly up- curved; basal sacculus division moderately long, lobe short; distal division with a single, long, basal, fingerlike lobe; corona weak. Phallus vesica with an ap- pendicular lobe; cornuti few, moderately long, secondary group a patch of spinelets. Anellus sclerites triangular. Juxta rectangular. VIII tergal sclerotiza- tion V-shaped; sternal sclerotization U-shaped, arms broad.
Female genitalia. Ovipositor lobe convex, well sclerotized, sagittate, striae obsolescent. Postapophysis about one-fourth longer than antapophysis, both slender, latter somewhat spatulate. VIII pleuron with a large, V-shaped, median unsclerotized area containing weak striae, ventral margin moderately deflexed, curved. Sterno-ostial concavity ovoid. Lamella postvaginalis cleoid; 1. ante- vaginalis ligulate. Corpus bursae ellipsoid; lobe simple; sclerotization and rib- bing strong but restricted to extreme posterior end.
Included species
52. chryseochilus Dyar, 1909. Ent. Soc. Wash., Proc. 11:25 (Basilodes) .
Species group XVI: C. triangulifer and allies Figure 2 1
Description
General. Size medium to medium large, expanse about 34-50mm. Frontal protuberance a transverse, oval shallow cup with ventral lip mesally produced into a conical prominence. Foretibial “claw” a moderately strong, single seta or a few (2-3), small, separate setae ( triangulifer ) arising from slightly pro- duced tibial apex. Forewing apex acute, rounded, tornus rounded; pattern complex: ground color yellow-orange, generally like preceding group but noctuiform elements more modified; transverse line orange, irregular, con- fused by longitudinal streaks, etc.; streaks in cell Cu forming a trident (base distad). Hindwing fuscous to yellow, sometimes only basal area fuscous; pat- tern obsolete. Thoracic vestiture of mixed scale types; strongly and complexly tufted: in addition to normal stiriine tufts anterior, dorsolateral and lateral mesoscutal tufts developed.
Male genitalia. Tegumen varied, moderately broad to fairly narrow, ven- tral arms and neck moderately long to short. Uncus of medium length, cylin- drical and straight; setae numerous, slightly longer dorsad. Saccus elongate to stout, bluntly pointed cephalad. Valve oblong to rhomboid-oblong; costa ex-
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panded obliquely; basal sacculus division moderately long, broader basad, end- ing diffusely, lobe a short peak or a very long, straight finger; distal division with two close lobes (on single plate in triangulifer) , basal lobe fingerlike or lobulate, distal lobe spinelike with broad base; corona well developed. Phallus vesica normal; cornuti numerous, subequal, moderately long and stout, sec- ondary group a curved patch of spinelets. Anellus sclerites crescent-shaped, separate. Juxta subquadrate. VIII tergal scleroitzations V- or U-shaped, tips of arms expanded; sternal sclerotization U-shaped, arms expanded.
Female genitalia. Ovipositor lobe of primitive, flat, blunt type, striae dis- tinct. Postapophysis slightly longer than antapophysis. VIII pleuron with an elongate, median unsclerotized area (containing weak striae only in dyari ); ventral margin moderately deflexed, curved. Sterno-ostial concavity round. Lamella postvaginalis round; 1. antevaginalis ligulate. Corpus bursae varied in shape: usually tubular and curved ( dyari and closest species), bulbous ( tri- angulifer), lobe deflected laterad ( nigrifer ); sclerotization and ribbing strong.
Included species
53. dyari Cockerell, 1899. Canad. Ent. 31:288 ( Cirrophanus , sic).
54. dubifer Dyar, 1907. N. Y. Ent. Soc., J. 15:109 {Cirrhophanus) .
This species may prove to be indistinct from the preceding.
55. nigrifer Dyar, 1907. N. Y. Ent. Soc., J. 15: 108 {Cirrhophanus) .
56. magnifer Dyar, 1907. N. Y. Ent. Soc., J. 15: 108 {Cirrhophanus) .
57. triangulifer Grote, 1872. Canad. Ent. 4:187 {Cirrhophanus) . —pretiosus Morrison, 1875. Boston Soc. Nat. Hist., Proc. 18: 122-123
{Chariclea) . Synonymy of Grote, 1875. Check List Noctuidae Amer. N. of Mexico, pt. I, p. 12.
Species group XVII : C. plesioglaucus and comstocki Figure 22
Description
General. Size medium, expanse about 32mm. Frontal protuberance a reniform ring with lower lip produced mesad into a conical prominence. Fore- tibial “claw” of multiple fused setae, arising from unproduced tibial apex. Forewing shape somewhat elongate, nearly elliptical; pattern unique: ground color with iridescent purpurescent cast, dark bronzy in distal sector (a wide, diffuse, white, median sector in comstocki) , a few short, longitudinal, dark streaks. Hindwing fuscous. Thoracic vestiture exactly as in preceding group, even to coloration.
Male genitalia. Tegumen narrow, neck long and curved. Uncus short, cylindrical, tapering, straight; setae sparse and short. Saccus ellipsoid, stout. Valve rhomboid-oblong, a sharp angle dorsobasad; costa barely expanded; basal sacculus division long, nearly confluent with distal division, lobe a low peak; distal division produced distad into a strong, long, spinelike lobe; corona
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absent. Phallus vesica normal; cornuti stout, short, heavier distad and on dis- tinct sclerotized plate, weaker basad, secondary group a patch of sharp den- ticles. Anellus sclerites crescent-shaped. Juxta kite-shaped. VIII tergal sclero- tization square with open end; sternal sclerotization broadly U-shaped, arms expanded.
Female genitalia. Ovipositor lobe slightly modified from primitive flat, blunt type : slightly pointed and upcurved, striae distinct. Postapophysis slightly longer than antapophysis. Pleuron with a V-shaped unsclerotized median area (no striae), ventral margin moderately deflexed. Sterno-ostial concavity sub- triangular. Lamellae indistinct. Corpus bursae oblong; lobe simple; sclerotiza- tion and ribbing strong.
Included species
58. plesioglaucus Dyar, 1912. U. S. Natl. Mus., Proc. 42:68 (Antaplaga) .
59. comstocki Hill, 1924. So. Calif. Acad. Sci., B. 23:184, pi. 3, fig. 6 ( Chamoclea , sic) [ Chamaeclea ]. This species is very close to the preceding and considered synonymous with it or as a subspecies by some authors. I con- sider it a distinct species here and leave the determination of its proper status to future studies.
Species group XVIII: C. papago and miaiphona Figure 23
Description
General. Size medium, expanse about 28-35mm. Frontal protuberance large, an inverted heart-shaped ring with lower lip produced mesad into a conical prominence. Foretibial “claw” of multiple fused setae, arising from unproduced tibial apex. Forewing apex acute, rounded, tornus rounded; pat- tern fairly simple: ground color yellow-orange with diffuse reddish or orange transverse lines, veins streaked, streak through cell Cu simple, not like a trident. Thoracic vestiture hairy; tufts poorly developed.
Male genitalia. Tegumen moderately broad, neck short. Uncus moderately long, convex, straight; setae numerous, short, subequal. Saccus ellipsoid, pointed. Valve rhomboid-oblong; costa expanded obliquely; basal sacculus di- vision moderately long, ending diffusely, lobe a short peak; distal division with single broad, short spinelike lobe; corona weak, somewhat diffuse. Phallus vesica inflated sinistrad; cornuti subequal, short and stout, secondary group a long, curved row of sharp, equal spinelets. Anellus sclerites crescent-shaped. Juxta rectangular. VIII tergal sclerotization square, tips of arms expanded; sternal sclerotization U-shaped.
Female genitalia. Ovipositor lobe of primitive, blunt, flat type, striae ab- sent. Postapophysis slightly longer than antapophysis. Pleuron with an elongate median unsclerotized area (no striae), ventral margin moderately deflexed, nearly parallel with opposite margin. Sterno-ostial concavity subquadrate. Lamella postvaginalis weak, rectangular. Corpus bursae oblong; lobe simple; sclerotization and ribbing strong.
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Included species
60. papago Barnes, 1907. Canad. Ent. 39:’95-96 (Cirrhophanus) .
61. miaiphona Dyar, 1912. U. S. Natl. Mus., Proc. 42:69-70 ( Cirrho - phanus) .
Species group XIX. C. hoffmanni
There is only a single known specimen of this species (62.), a female. It is a new species and described in the appendix.
Species group XX: C. discistrigus Figure 24
Description
General. Size medium-small, expanse about 25-30mm. Frontal protuber- ance small, a reniform ring with lower lip produced mesad into a conical prom- inence. Foretibial “claw” small, of multiple fused setae arising from unpro- duced tibial apex. Forewing apex acute, rounded, tornus very slightly toothed; pattern vaguely similar to preceding unit: ground color brownish-yellow, streaked by darker veins, streaks intense and broadened distad especially on fringe, diffuse brownish-orange stain in discoidal and basal areas. Hindwing orange, markings obsolete. Thoracic vestiture entirely of spatulate scales; tufts normal, small anterior mesoscutal tufts developed.
Male genitalia. Tegumen medium, neck long and curved. Uncus moder- ately long, cylindrical, swollen mesad. Saccus short, pointed. Valve rhomboid- oblong, costa moderately expanded; basal sacculus division moderately long, ending in a sharp corner, lobe pommel-shaped; distal division distally produced into a short, sharp spine; corona weak. Phallus vesica normal; cornuti short and stout distad, changing to broad, uncinate spicules basad, discrete secondary group absent. Anellus sclerites crescent-shaped, fused. Juxta elongate, shield- shaped. VIII tergal sclerotization U-shaped, angles sharply produced; sternal sclerotization U-shaped, arms expanded.
Female genitalia. Ovipositor lobe short, pointed, nearly deltoid, upcurved, tip strongly sclerotized, partially unsclerotized basad, with weak striae. Posta- pophysis slightly longer than antapophysis, both stout. VIII pleuron with a V-shaped, median unsclerotized area (no striae), ventral margin moderately de flexed, nearly parallel with opposite margin. Sterno-ostial concavity ovoid. Lamella postvaginalis weakly defined, chevron-shaped; 1. antevaginalis ligu- late. Ductus bursae sclerotized and ribbed (especially on side toward corpus lobe). Corpus bursae ellipsoid, curved; lobe simple; sclerotization and ribbing strong.
Remarks
Hampson erected the genus Hoplolythra for this species and as indicated by the derivation of the name and placement of the species, he agreed with Smith, the original describer, on a relationship to Lythrodes. Both taxa do have similarly streaked forewings but are abundantly distinct structurally.
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Definition and Classification of Stiriini
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Included species
63. discistrigus Smith, 1903. Amer. Ent. Soc., Trans. 29:207 (Lythrodes) .
Species group XXI: C. compositus and hyper ion Figure 25
Description
General. Size small, expanse about 20-25mm. Frontal protuberance a reniform to heart-shaped ring with lower lip produced mesad into a conical prominence. Foretibial “claw” of multiple fused setae (? hyperion ), arising from unproduced tibial apex. Forewing apex acute, rounded; pattern of two types: (1) self-colored, yellow-orange, fringe brown ( compositus ), (2) bi- colored, median sector yellow, distal and basal sectors chocolate-brown (hy- perion). Hindwing fuscous to yellow-orange, markings obsolete. Thoracic vestiture of mixed, linear and spatulate scales; tufts poorly developed.
Male genitalia (unknown for hyperion) . Tegumen narrow, neck moder- ately long. Uncus moderately long, cylindrical, tapered, curved; setae sparse and tiny. Saccus rhomboid. Valve ellipsoid-elongate; costa moderately ex- panded; basal sacculus division moderately long, ending diffusely, lobe a broad, low mound; distal division with small, pointed, erect lobe; corona obsolete. Phallus vesica elongate; cornuti moderately long, equal, few and restricted to a small patch at far distal end, secondary group absent from normal position, a loose patch of broad, uncinate spicules located basad instead. Anellus sclerites triangular, nearly fused. Juxta rectangular. VIII tergal sclerotization V-shaped; sternal sclerotization round.
Female genitalia. Ovipositor lobe broad, pie-shaped with slightly up- curved tip, striae distinct. Postapophysis about one-third longer than anta- pophysis, both quite short. Pleuron with elongate, median unsclerotized area (no striae), ventral margin moderately deflected. Sterno-ostial concavity sub- quadrate, small. Lamella postvaginalis subquadrate; 1. antevaginalis ligulate, well defined. Ductus bursae very long and slender, well sclerotized and ribbed, especially on lobe side. Corpus bursae ovate and transverse; lobe not defined; ductus seminalis at tip directed at right angle to ductus bursae; corpus entirely membranous.
Remarks
Since hyperion is a rare species and known only from females, I have not been able to study it completely to definitely substantiate its placement in this group. The female genitalia are virtually identical to those of compositus as far as I can determine. The type of genitalia is peculiar (very long ductus bursae and transverse corpus bursae) so it would appear improbable that the similarity is spurious.
Included species
64. compositus Edwards, 1884. Papilio 4:44 (Eulithosia) .
65. hyperion Dyar, 1918. U. S. Natl. Mus., Proc. 54:348 ( Pumora ).
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Genus 6: CUAHTEMOCA Hogue, new genus Type species: Chalcopasta chalcocraspedon Dyar, 1913; Zacualpan, Mexico.
Remarks
The single species and species group of major phyletic line 6 is worthy of generic distinction on the basis of its relictual, annectant status. No other name being available, I propose the new genus Cuahtemoca.
Description
See the decription of the single species group below.
Species group XXII: C. chalcocraspedon Figure 26
Description
General. Size medium, expanse about 30-35 mm. Frontal protuberance a heart-shaped ring inclosing a ventral, conical prominence. Foretibial “claw” rounded, a single seta arising from slightly produced tibial apex. Forewing apex acute, sharp, tornus sharp, outer margin slightly concave below apex; pattern noctuiform: ground color brown in median sector, brassy gold in marginal and basal sectors and in reniform and orbicular, scattered blue scales along costal area. Hindwing grey-brown, exterior line evident. Thoracic vestiture entirely of spatulate, dentate scales; tufts normal, anterior mesoscutal tuft developed.
Male genitalia. Tegumen moderately broad, neck long. Uncus moderately long, cylindrical, straight; setae moderately dense, longer dorsad. Saccus short, pointed. Valve rhomboid-oblong, apex rounded; costa slightly expanded; basal sacculus division long, confluent with distal division, lobe a short point; distal division with two close lobes: basal long, fingerlike, distal shorter, sclerotized, spinelike. Phallus vesica normal; cornuti very numerous, long and equal, sec- ondary group absent. Anellus sclerites crescent-shaped, separated. Juxta shield- shaped. VIII tergal sclerotization U-shaped, base and tips of arms expanded; sternal sclerotization nearly circular.
Female genitalia. Ovipositor lobe sagittate, evenly sclerotized, points blunt, striae obsolescent. Postapophysis slightly longer than antapophysis, both slender. VIII pleuron with an elongate, unsclerotized median area containing obsolescent striae, ventral margin defined, moderately deflexed cephalad, arch- ing cephalad; with opposite margin inclosing a depressed lamella postvaginalis and associated with it and 1. antevaginalis to form a special, complex, oval sterno-ostial concavity. Lamellae ligulate, bowed. Ductus bursae short, mem- branous. Corpus bursae elongate, curved, slightly swollen mesad; lobe simple, ventral; sclerotization and ribbing moderately strong.
Included species
66. chalcocraspedon Dyar, 1913. U. S. Natl. Mus., Proc. 44:297 (Chalco- pasta).
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Definition and Classification of Stiriini
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Genus 7: CHALCOPASTA
1910, Chalcopasta Hampson. Catalogue Lepid. Phalaenae Brit. Mus. 9:218. Type species: Basilodes territans Edwards, 1884; Arizona; by original designation.
1912 , Rodriguesia Dyar. Ent. Soc. Wash., Proc. 14:61. Type species: Dev a ornata Ottolengui, 1898; Hot Springs, New Mexico; monobasic (= Chal- copasta howardi Edwards, 1877; Prescott, Arizona; synonymy of Barnes and McDunnough, 1916. Contrib. Lepid. N. Amer. 3:167-168).
Remarks
As discussed in the phylogeny section, major phyletic line 7 divides into two main sections primarily on the basis of the development of the lobes of the distal sacculus division: (1) a basal, fingerlike lobe or (2) a distal, spinelike lobe. The genus Chalcopasta, as here defined, stands for the first section. One unit, XXIII (C. ellica), has both lobes developed but since the distal lobe is re- duced and the wings have areas of brilliant gold and other character states re- semble more those of the first section, this unit is placed with them.
With regard to the identity of Dev a ornata Ottolengui, 1898, Barnes and McDunnough (1916: 167-168) state: “We have carefully examined the type in the National Museum and find that in maculation, squammation, and structure it is an exact match of Chalcopasta howardi with the exception of the palpi which are strongly upturned and entirely different to those of howardi. Al- though we could find no trace of glue or shellac we strongly incline to the opin- ion that the head has been neatly glued on the specimen in question, as we cannot believe that two species could resemble each other so exactly in every detail except the palpi. It would be necessary for the specimen to be relaxed to prove our contention and we must leave this to one of the curators of the museum!’ I am informed by J. G. Franglemont (personal communication), a recent curator at the National Museum, that this specimen has been relaxed and the head found in fact to be glued on. Dyar’s erection of the genus on the basis of the long palpi was, therefore, erroneous. The otherwise detailed resemblance to howardi and others of the genus Chalcopasta requires the sinking of Rod- riguesia under Chalcopasta.
Description
General. Size medium to medium-large; facies robust. Frontal protuber- ance a circular to oval ring inclosing a subcentral or central, conical promi- nence. Foretibial “claw” flattened and shovel-shaped, arising from strongly produced and sometimes slightly angled tibial apex. Forewing apex acute, sharp, tornus rounded, outer margin entire, undulate slightly in some; basic, brown noctuiform pattern overrun by discrete fields of brilliant metallic gold. Thoracic vestiture of varied scale types; tufts normal, tegular tufts often strong- ly erect.
Male genitalia. Tegumen moderately broad, neck long and arched to short and straight. Uncus moderately long to long, cylindrical and straight; setae
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numerous and short. Valve oblong, apex rounded; costa barely expanded; basal sacculus division long, ending diffusely, lobe only a low mound; distal division normally with one basal, erect, fingerlike lobe but may also have a short, distal spinelike lobe or no lobes present at all, when latter, a spur at distal end point- ing ventrad; coronal setae slender and restricted to dorsodistal margin of valve, or absent altogether. Phallus vesica usually normal in shape but may be swollen distad or tubular and coiled once; cornuti numerous, varied in shape. Anellus sclerites crescent-shaped or seculate. Juxta rectangular or shield-shaped. VIII tergal sclerotization narrowly V-shaped; sternal sclerotization U-shaped or circular.
Female genitalia. Ovipositor lobe varied, sagittate to acuminate, always heavy and well sclerotized, weak striae rarely present, usually striae absent. Postapophysis subequal to or slightly longer than antapophysis. VIII pleuron with an elongate, median, unsclerotized area without striae (weak striae in ellica), ventral margin defined, moderately deflexed or strongly declining cephalad, diverging or curving out cephalad; with opposite margin inclosing a depressed lamella postvaginalis and associated with it and 1. antevaginalis to form a special, complex sterno-ostial concavity. Lamella postvaginalis oval to ligulate; 1. antevaginalis ligulate, bowed. Corpus bursae quite varied in shape: usually ellipsoid-oblong, sometimes cudgel-shaped or fusiform; lobe normal or displaced laterad (one species: restricta)’, sclerotization and ribbing strong.
Species group XXIII: C. ellica Figure 27
Description
General. Size medium, expanse about 30mm. Frontal protuberance a rough, heart-shaped plateau with no discrete secondary prominence. Fore- tibial “claw” heavy, shovel-shaped, arising from strongly produced, straight tibial apex. Forewing apex acute, sharp, tornus rounded, costal margin very straight; pattern basically noctuiform, irrorated brown but extensively overrun in distal and basal areas by brilliant metallic gold, reniform and orbicular (and sometimes area between) also filled with this color. Hindwing brown, pattern obsolete. Thoracic vestiture of linear scales; tufts normal but somewhat poorly developed.
Male genitalia (described from photograph) . Tegumen moderately broad, neck long, arched. Uncus moderately long, cylindrical, slightly tapering, straight; setae numerous, short. Valve oblong, apex rounded; costa barely ex- panded; distal sacculus division with two lobes: basal erect, fingerlike, distal short, broad, spinelike; coronal setae slender. Juxta shield-shaped. States of other characters not determinable from photograph.
Female genitalia. Ovipositor lobe bluntly pointed, striae weakly devel- oped. Postapophysis about one-fourth longer than antapophysis. VIII pleuron with an elongate median unsclerotized area containing weak striae. Sterno- ostial concavity deltoid. Lamella postvaginalis rectangular; 1. antevaginalis
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ligulate. Ductus bursae membranous, becoming abruptly sclerotized and ribbed about halfway. Corpus bursae cudgel-shaped; lobe simple; sclerotization and ribbing strong.
Included species
67. ellica Dyar, 1915. Insec. Incit. Mens. 3:80 (Chalcopasta) . This spe- cies is incorrectly labeled as fulgens by Draudt ( 1919-1939. pi. 44, row k).
Species group XXIV : C. territans and allies Figure 28
Description
General. Size medium to medium-large, expanse about 28-40mm.; very robust facies. Frontal protuberance a circular to oval ring inclosing a sub- central, conical prominence. Foretibial “claw” strong, flattened, arising from strongly produced tibial apex. Forewing apex acute, sharp, tornus rounded, outer margin slightly concave beneath apex; basic noctuiform pattern as in preceding group but even more extensively overrun by gold; reniform, apical patch, costal area, anterior end of PM line and basal area still exposed. Hind- wing white to fuscous; pattern obsolete. Thoracic vestiture of spatulate, den- tate scales, tufts normal, very well developed, tegular strongly erect.
Male genitalia. Tegumen moderately broad, neck short. Uncus long, cylin- drical, straight; setae short, numerous. Valve oblong, apex rounded, slightly upcurved; costa moderately expanded; distal sacculus division with a single, erect, basal, fingerlike lobe; coronal setae slender. Phallus vesica sometimes expanded sinistrad; cornuti very numerous, short, stout, spiculate basad, longer and spinelike distad, secondary group a patch of sharp spinelets. Anellus sclerites crescent-shaped. Juxta rectangular. VIII tergal sclerotization narrowly V-shaped, tips of arms expanded; sternal sclerotization U-shaped.
Female genitalia. Ovipositor lobe small, acuminate, pointed, convex and heavily sclerotized (no striae). Postapophysis and antapophysis subequal, both fairly stout. VIII pleural unsclerotized area V-shaped, no striae. Sterno-ostial concavity oval to flask-shaped. Lamella postvaginalis oval to ligulate, posterior margin eroded; 1. antevaginalis ligulate, bowed. Ductus bursae short, mostly membranous. Corpus bursae ellipsoid-oblong; lobe simple, ventral; sclerotiza- tion and ribbing strong.
Included species
The first seven species are difficult to separate on the basis of wing pattern. I suspect that there are actually fewer valid species.
68. territans Edwards, 1884. Papilio 4:45 (Basilodes) .
—arizona French, 1889. Canad. Ent. 21:161 (Plusia) . Synonymy of (?) Barnes and McDunnough, 1917. Check List Lepid. Boreal Amer., p. 71.
69 . anopis Dyar, 1918. U. S. Natl. Mus., Proc. 54:348-349 ( Chalco- pasta) .
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70. arianda Druce, 1889. Biol. Cent. Amer., Heter. 1:329, pi. 30, fig. 11 ( Plusia ) .
71. chalcophanis Dyar, 1918. U. S. Natl. Mus., Proc. 54:348 ( Chaleo - pasta ) .
72. pterochalcea Dyar, 1909. Ent. Soc. Wash., Proc. 11:24-25 ( Basi - lodes ) .
73. riandana Dyar, 1912. U. S. Natl. Mus., Proc. 42:69 {Chalcopasta) .
74. howardi Edwards, 1877. Pacific Coast Lepid., No. 25, p. 1 {Plusia). —ornata Ottolengui, 1898. Canad. Ent. 30: 106-107, pi. 5, fig. 1 ( Deva
?) . See remarks under genus for synonymy.
I5.fulgens Barnes and McDunnough, 1912. Contrib. Nat. Hist. Lepid. N. Amer. 1:25-26 {Chalcopasta) . Figured in an earlier number in the same volume of the same series, 1(4) :pl. 25, fig. 3. This species is incorrectly lab- elled as ellica by Draudt ( 19 19- 1939: pi. 44, row k).
76. sinuata Hampson, 1918. Nov. Zool. 25: 152-153 {Chalcopasta) .
77. restricta Hampson, 1918. Nov. Zool. 25: 152-153 {Chalcopasta) .
Species group XXV. C. chalcotoxa and allies Figure 29
Description
General. Size medium, expanse about 27-30mm. Frontal protuberance a circular ring inclosing a central, conical prominence. Foretibial “claw” shovel- shaped, arising from moderately produced tibial apex. Forewing apex acute, sharp, tornus rounded; pattern basically noctuiform: ground color irrorated, dark brown, extensive oblique inroad (not as extensive as in two preceding species groups) of brilliant metallic gold in median sector (also in marginal sector in acantha), otherwise as preceding species group. Thoracic vestiture as in preceding species group.
Male genitalia. Of two types: (1) Like preceding unit except for phallus vesica which is swollen distad; cornuti of short spines displaced dorsad {acan- tha), (2) Peculiar. Tegumen narrow, neck short. Uncus long, slender, cylin- drical, tapering and arched; setae sparse and short. Valve oblong, angular, wider distad; costa not expanded; distal sacculus division normal but without lobe as in preceding unit, instead produced distoventrad as a spurlike lobe (reduced on one valve in acema) ; corona absent. Phallus vesica tubular, coiled once; cornuti moderately long or long, secondary group absent. Anellus sclerites slender, crescent-shaped. Juxta elongate, rectangular. VIII tergal sclerotization nar- rowly V-shaped; sternal sclerotization circular, nearly solid in some (not acantha), only a subcentral, membranous area, a few small, lanceolate scales on posterior margin of latter (visible only in slide preparations).
Female genitalia. Generally the same as in preceding species group, but with two main peculiarities: (1) lobe of corpus bursae displaced cephalad so corner from which ductus seminalis leaves almost at middle {acantha), (2) corpus bursae fusiform {acema and chalcotoxa) .
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Included species
78. acantha Druce, 1889. Biol. Cent. Amer., Heter. 1:329; 3: pi. 30, fig. 13 ( Plusia ).
79. chalcotoxa Dyar, 1909. Ent. Soc. Wash., Proc. 11:25 (Basilodes) .
80. acema Druce, 1889. Biol. Cent. Amer., Heter. 1:330; 3 : pi . 30, fig. 14 (Plusia).
The last two species tend to intergrade and may actually constitute a single varied species.
Genus 8: BASILODES
1852, Basilodes Guenee. Hist. Nat. Ins. Spec. Gen. Lepid. Noc. II 6:358. Type species: B. pepita Guenee, 1852; Florida; monobasic.
1869, Deobriga Walker. Char. Undes. Lepid. Heter., p. 41. Type species: D. chrysopasa Walker, 1869; locality unknown; monobasic (= Basilodes pepita Guenee, 1852; Florida; synonymy of Hampson, 1910. Catalogue Lepid. Phalaenae Brit. Mus. 9:207).
1874, Stiria Grote. Buffalo Soc. Nat. Sci., B. 2:73. Type species: S. rugifrons Grote, 1874; Kansas, Colorado Territory; monobasic. Synonymy of Smith, 1882. Brooklyn Ent. Soc., B. 5:30, 35.
Remarks
As mentioned under the preceding genus, major phyletic line number 7 divides into two main sections on the basis of many features, principally the expression of either a basal or distal lobe on the distal sacculus division of the male valve. The dichotomy of brilliant, gold or dull-colored forewing is also important. The genus Chalcopasta represents the first section, the present genus Basilodes represents the second.
Basilodes antedates Stiria and is therefore the appropriate name for the genus. The tribe, however, retains the name Stiriini since according to the In- ternational Code of Zoological Nomenclature suprageneric names are based on priority and are not affected by synonymy.
One new species, B. inquinatus, is described herein (see appendix).
Description
General. A very compact, structurally uniform genus. Size medium to medium-large. Frontal protuberance varied, of two main types: (1) an oval to heart-shaped, shallow cup with a slightly convex ventral portion, (2) a heart-shaped ring inclosing a subcentral, truncate, low, conical prominence. Foretibial “claw” strong, setiform or shovel-shaped, arising from strongly pro- duced tibial apex. Forewing apex acute, sharp, tornus rounded or sharp, often with slight fringe tooth; outer margin crenulate, general outline often some- what sinuate and convex mesad; pattern primitively noctuiform, often with diffuse areas or general hue of brassy or yellow gold color, derived patterns varied but mostly of type similar to that in species group XXVII : ground color
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yellow, transverse lines, reniform and orbicular weak, a large hexagonal or square macula mesad in inner area, a small dot in reniform. Thoracic vestiture very rough, of spatulate scales; tufts normal and heavy, tegular tufts erect.
Male genitalia. Tegumen moderately broad, somewhat inflated ventrad in some, neck long and arched. Uncus moderately long, cylindrical, swollen mesad or not, straight or slightly curved; setae dense and moderately long. Saccus varied, usually rectangular, pointed or nippled cephalad. Valve oblong, apex rounded; costa barely expanded; basal sacculus division long, ending diffusely, lobe a low mound; distal division with a strong, curved, distal, spine- like lobe and weakly developed basal setose verrucae, rarely developed into a lobe; corona fully developed. Phallus vesica varied, of two types: (1) elongate, without secondary group of cornuti, (2) normally pear-shaped with secondary group of cornuti; primary group of cornuti varied in length and stoutness but always quite numerous. Anellus sclerites seculate. Juxta rectangular or kite- shaped. VIII tergal sclerotization circular, sternal sclerotization U-shaped or circular.
Female genitalia. Ovipositor lobe heavily sclerotized and rigid; deltoid to saggitate in outline. Postapophysis somewhat larger than antapophysis, both moderately strong. VIII pleuron with a V-shaped, median unsclerotized area, ventral margin strongly deflexed and arching; with opposite margin inclosing a depressed lamella postvaginalis and associated with it and 1. antevaginalis to form a special, complex sterno-ostial concavity. Lamella postvaginalis ligulate or crescent-shaped, eroded caudad; 1. antevaginalis ligulate, bowed. Corpus bursae elongate, slightly curved and swollen mesad; lobe simple, ventral; sclerotization and ribbing moderate to very strong.
Species group XXVI: B. chrysopis and allies Figure 30
Description
General. Size medium to medium-large, expanse about 30-40mm. Very robust facies. Frontal protuberance an oval to heart-shaped, shallow cup with slightly convex ventral portion. Foretibial “claw” moderately heavy, arising from strongly produced, straight tibial apex. Forewing apex acute, sharp, tornus rounded, outer margin crenulate, general outline somewhat sinuate; pattern noctuiform : ground color brassy or yellow-gold, brown median sector in most chrysopis, a small, eccentric, dark dot in reniform. Thoracic vestiture of spatulate, dentate scales; tufts normal, heavy.
Male genitalia. Tegumen moderately broad, inflated ventrad in inquinatus resulting in short vincular arms, neck long and arched. Uncus moderately long, cylindrical, swollen mesad, straight; setae dense, long, equal. Saccus rectangu- lar, pointed or nippled cephalad. Valve rhomboid-oblong, apex rounded; costa barely expanded; distal sacculus division with strong, curved, distal spinelike lobe, sometimes also a weak (large in pepita), irregular basal lobe. Phallus vesica elongate; cornuti short to long, longer distad, secondary group absent.
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Anellus sclerites seculate, approximate. Juxta rectangular. VIII tergal sclero- tization V-shaped, tips of arms expanded; sternal sclerotization U-shaped to circular.
Female genitalia. Ovipositor lobe elongate to broadly sagittate, heavily sclerotized (no striae). Postapophysis somewhat longer than antapophysis. VIII pleuron with V-shaped unsclerotized area. Sterno-ostial concavity round, width about equal the width of sclerotized, ventral half of neighboring pleuron. Lamella postvaginalis semilunar; 1. antevaginalis ligulate, bowed. Corpus bursae somewhat constricted at base of lobe; sclerotization and ribbing very strong.
Included species
81. chrysopis Grote, 1881. Papilio 1:154-155 (Basilodes) . This is an ex- tremely variable species, having many intergrading forms. I think the follow- ing is one of these forms and, therefore, synonymize it with the present species.
—catharops Dyar, 1911. Ent. Soc. Wash., Proc. 13:64 (Basilodes) . NEW SYNONYMY.
82. inquinatus Hogue, new species. (See appendix).
83. auratus Schaus, 1911. Ann. Mag. Nat. Hist., Ser. 8, 7:47 (Basilodes) .
84. pepita Guenee, 1852. Hist. Nat. Ins. Spec. Gen., Lepid. Noc. II 6:358; Noc.pl. 12, fig. 1 (Basilodes).
—chrysopasa Walker, 1869. Char. Undes. Lepid. Heter. pp. 41-42 (Deobriga) . Synonymy of Hampson, 1910. Catalogue Lepid. Phalaenae Brit. Mus. 9:207.
Species group XXVII: B. rugifrons and allies Figures 2-5
Description
General. Size medium to medium large, expanse about 28-40mm. Very robust. Frontal protuberance a heart-shaped ring inclosing a subcentral, trun- cate, conical prominence. Foretibial “claw” usually strong, shovel-shaped, arising from fairly strongly produced tibial apex. Forewing apex acute, sharp, tornus sharp with moderate fringe tooth, outer margin crenulate, general out- line undulate; pattern: usually, ground color yellow, a hexagonal or quadrate brown macula halfway along the inner margin, a brown area spreading in from outer margin, PM and AM lines weak, brown, reniform and orbicular obso- lescent, a small dot in former; this pattern modified strongly in a few: (1) all dark brown, markings barely showing, small silver discal spots ( dysnoa ), (2) wide suffused median sector, pattern generally obscure (tacky mora) . Thoracic vestiture of spatulate scales; tufts normal, heavy.
Male genitalia. Scarcely different from preceding species group. Uncus tending to be more curved; corona weaker. Phallus vesica of normal shape, not elongate; secondary group of cornuti present, a short row of spinelets. Juxta kite-shaped.
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Female genitalia. Ovipositor lobe deltoid to saggitate, heavily sclerotized, convex. Postapophysis somewhat longer than antapophysis, both moderately strong. VIII pleuron with a V-shaped unsclerotized area. Sterno-ostial con- cavity oval, width about twice the width of sclerotized ventral half of neigh- boring pleuron. Lamella postvaginalis crescent-shaped; 1. antevaginalis ligu- late. Corpus bursae normal; sclerotization and ribbing moderately strong.
Included species
85. dysnoa Dyar, 1912. U. S. Natl. Mus., Proc. 42:69 (Chalcopasta) .
—biforis Draudt, 1926, in Seitz. Macrolepid. World 7:308, pi. 44, row 1 (Chalcopasta) . Originally described as an aberration of dysnoa and considered as no more than a synonym by various authors but the synonymy needs re-examination.
86. consuelus Strecker, 1900. Lepid. Rhop. Heter., Suppl. 3, p. 34 ( Stiria ) .
The next six species are very similar and several may be synonymized upon future study.
87. rugifrons Grote, 1874. Buffalo Soc. Nat. Sci., B. 2:73 (Stiria).
88. ruficeps Draudt, 1927, in Seitz. Macrolepid. World 7:310, pi. 45, row b (Stiria).
89. colimae Draudt, 1927, in Seitz. Macrolepid. World 7:310, pi. 45, row b (Stiria). There is question as to the identity of this species. Apparently the type material was never labelled as such and its repository is unknown. Judging from the figure accompanying the original description, I guess that this species may be at most a variation or subspecies of the preceding species. However, I leave the problem to future studies.
90. intermixtus Dyar, 1918. U. S. Natl. Mus., Proc. 54:349 (Stiria).
91. ischune Dyar, 1912. U. S. Natl. Mus., Proc 42:70 (Stiria).
92. mouris Dyar, 1912. U. S. Natl. Mus., Proc. 42:70 (Stiria).
93. sisaya Dyar, 1912. U. S. Natl. Mus., Proc. 42:70 (Stiria) .
94. tachymora Dyar, 1914. U. S. Natl. Mus., Proc. 47:375 (Stiria).
95. dyari Hill, 1924. So. Calif. Acad. Sci., B. 23:183-184, pi. 3, fig. 3 (Stiria) .
96. sulphureus Neumoegan, 1882. Papilio 2:135 (Stiria).
—demaculatus Strand, 1916. Arch. Naturges. 81: 165 (Stiria). De- scribed originally as an aberration of sulphur ea and considered as such by various authors.
GENERIC STATUS UNDETERMINED
97 . prepontendyta Dyar, 1914. U. S. Natl. Mus., Proc. 47:375 (Anta- plaga).
98. primulina Druce, 1889. Biol. Cent. Amer., Heter. 1:303; 3:pl. 28, fig. 10 (Metoponia) . Hampson has given this species autonomous generic
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Definition and Classification of Stiriini
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standing ( Xanthiria Hampson, 1910. Catalogue Lepid. Phalaenae Brit. Mus. 9:244-245). It would seem best to retain this combination until the species is better known and further studied.
99. iticys Dyar, 1914. U. S. Natl. Mus., Proc. 47:375-376 ( Stiria ).
100. arivaca Barnes, 1907. Canad. Ent. 39:66-67 ( Lythrodes ). The streaked forewing pattern is similar to certain members of the genus Cirrhop- hanus but a relationship is yet to be determined on a structural basis.
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TABLE I
SUMMARY OF CLASSIFICATION AND COMPOSITION OF PHYLETIC UNITS
GENERA
1 . Plagiomimicus Grote, 1873
MAJOR
PHYLETIC
LINES
1
PRIMARY
PHYLETIC
UNITS—
SPECIES
GROUPS
L
II.
III.
IV.
V.
VI.
VII.
VIII.
2 IX.
X.
SPECIES
1. resolutus Dyar, 1909
2. ochoa (Barnes, 1904)
3. laodamia (Druce, 1890)
4 . jalada (Schaus, 1898)
5. aureolus (Edwards, 1882)
6. bajae Hogue, new species
7. olvello (Barnes, 1907)
8. raglena (Dyar, 1912)
9. unicus (B. & Benj., 1926)
10. chalcospilans (Dyar, 1918)
11. laverna (Druce, 1898)
12. manti (Barnes, 1904)
13. dimidiatus (Grote, 1877)
14. corazonci (Schaus, 1898)
1 5. astigmatosus (Dyar, 1921)
16. curiosus (Neum., 1883)
17. spumosus (Grote, 1874)
18. mavina (B. & McD., 1910)
19. hilli (B. & Benj., 1923)
20. expallidus Grote, 1882
21. triplagiatus Smith, 1890
22. pityochromus Grote, 1873
23. concinnus (Dyar, 1909)
24. tepperi (Morrison, 1875)
25. hutsoni (Smith, 1907)
26. olivalis (B. & McD., 1916)
27. hachita (Barnes, 1904)
28. sexseriatus (Grote, 1881)
29. biundulalis (Zeller, 1872)
30. alesaea (Dyar, 1918)
31 . psamathochromus (Dyar, 1909)
32. argyropolius (Dyar, 1914)
33. phalaenoides (Dyar, 1918)
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Definition and Classification of Stiriini
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GENERA
2. Chrysoecia Hampson, 1910
3. Gorgora Dyar, 1914
4. Chichimeca new genus
5. Cirrhophanus Grote, 1872
TABLE I (continued)
MAJOR
PHYLETIC
LINES
3
PRIMARY
PHYLETIC
UNITS—
SPECIES
GROUPS
XI.
SPECIES
34. scira (Druce, 1889)
35 . gladiola (Barnes, 1907)
36. requies (Dyar, 1909)
XII. 37. atrolinea (B. & McD., 1912)
38. dela (Druce, 1894)
39. stigmatosa (Dyar, 1912)
40. hemicrocea (Dyar, 1912)
XIII. 41. morga Dyar, 1914
4 XIV. 42. thoracica (Edwards, 1884)
43. spreta (Draudt, 1927)
44. subfumosa (Dyar, 1909)
45. anaemica (Draudt, 1927)
4 6. salacon (Druce, 1895)
47. simplicia (Dyar, 1926)
48. dulcita (Schaus, 1898)
49. pseudovarra (Dyar, 1926)
50. varrara (Dyar, 1918)
51. muricolor (Dyar, 1926)
5 XV. 52. chryseochilus (Dyar, 1909)
XVI. 53. dyari Cockerell, 1899
54. dubifer Dyar, 1907
55. nigrifer Dyar, 1907
56. magnifer Dyar, 1907
57. triangulifer G rote, 1872
XVII. 58. plesioglaucus (Dyar, 1912)
59. comstocki (Hill, 1924)
XVIII. 60. papago Barnes, 1907 61. miaiphona Dyar, 1912
XIX. 62. hoffmanni Hogue, new species
XX. 63. discistrigus (Smith, 1903)
XXI. 64. compositus (Edwards, 1884) 65. hyper ion (Dyar, 1918)
XXII. 66. chalcocraspedon (Dyar, 1913)
6. Cuahtemoca new genus
6
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TABLE I (continued)
|
PRIMARY |
|||
|
MAJOR |
PHYLETIC |
||
|
GENERA |
PHYLETIC |
UNITS— |
SPECIES |
|
LINES |
SPECIES GROUPS |
||
|
7. Chalcopasta |
7 |
XXIII. |
67. ellica Dyar, 1915 |
|
Hampson, 1910 |
XXIV. |
68. territans (Edwards, 1884) |
69. anopis Dyar, 1918
70. arianda (Druce, 1889)
71. chalcophanis Dyar, 1918
72. pterochalcea (Dyar, 1909)
73. riandana Dyar, 1912
74. howardi (Edwards, 1877)
75. fulgens B. & McD., 1912
76. sinuata Hampson, 1918
77. res trie ta Hampson, 1918
XXV. 78. acantha (Druce, 1889)
79. chalcotoxa (Dyar, 1909)
80. acema (Druce, 1889)
8. Basilodes Guenee, 1852
XXVI.
XXVII.
Generic Status Undetermined XXVIII.
( Xanthiria XXIX.
Hampson, 1910)
XXX.
8 1 . chrysopis Grote, 1881
82. inquinatus Hogue, new species
83. auratus Schaus, 1911
84. pepita Guenee, 1852
85. dysnoa (Dyar, 1912)
86. consuelus (Strecker, 1900)
87. rugifrons (Grote, 1874)
88. ruficeps (Draudt, 1927)
89. colimae (Draudt, 1927)
90. intermixtus (Dyar, 1918)
91. ischune (Dyar, 1912)
92. mouris (Dyar, 1912)
93. sisaya (Dyar, 1912)
94. tacky mora (Dyar, 1914)
95. dyari (Hill, 1924)
96. sulphureus (Neumoegan, 1882)
97. prepontendyta Dyar, 1914
98. primulina Druce, 1889
99. iticys Dyar, 1914
XXXI. 1 00. arivaca Barnes, 1 907
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Definition and Classification of Stiriini
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APPENDIX: DESCRIPTIONS OF NEW SPECIES 6: Plagiomimicus bajae Hogue, new species
Figures 31b-c, 32a-e, k
Description
Adult male. Size medium-small, forewing length 13.5-14 mm. Upper sur- face of forewing with mixed medium grey-brown, brown and white scales; scales predominantly brown in center of wing forming a large, oblique, pear- shaped macula (apex directed toward apex of wing) ; macula bounded immedi- ately distad by a thin, white PM line bounded cephalad (along costal area) and basad by light scales; marginal sector of mixed light and dark scales giving a finely irrorated, grey-brown shade generally blending with fringe; orbicular and reniform and other markings entirely absent. Upper surface of hindwing predominantly white, grading to grey-brown distad and blending with fringe; exterior line faint. Lower surface of forewing dirty, grey brown, slightly lighter toward distal and costal margins. Lower surface of hindwing evenly, dirty white. Vestiture of head (including palps) and thorax of spatulate scales, light basad, becoming dark distad with white tips; a few white scales on antenna, es- pecially at base. Abdomen evenly, dirty light-brown. Frontal protuberance an oval ring