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Journal of The Lepidopterists' Society
Volume 53 1999 Number 1
Journal of the Lepidopterists' Society 53(1), 1999, 1-10
CONTRIBUTION TOWARDS THE STUDY OF THE PYRALINAE (PYRALIDAE): HISTORICAL REVIEW, MORPHOLOGY, AND NOMENCLATURE
M. Alma Solis
Systematic Entomology Laboratory, PS1, Agricultural Research Service, U.S. Department of Agriculture, c/o Nat. Mus. Nat. Hist., MRC 168, Washington, DC 20560-0168 USA
AND
Michael Shaffer
Entomology Department, The Natural History Museum, Cromwell Road, London SW7 5RD England
ABSTRACT. The monophyly of the Pyralinae and the two tribes, Pyralini and Endotrichini, is reviewed based on an analysis of previously used morphological characters of the adult and larva. Characters previously used to define these groups are plesiomorphic (i.e., they are not valid) or highly homoplastic (i.e., they are not reliable) to support the taxa as monophyletic, or both. We describe the male genitalia and present characters to support the monophyly of the Endotrichini, but the Pyralini is likely a paraphyletic taxon. Larval characters did not provide evidence to support or reject monophyly for either group. Rased on male genitalic morphology we reassign genera, and make additions or changes within these taxa in recently published checklists. In the Neotropical fauna: Perforadix Sein is transferred to the Pyraustinae and is a new synonym of Sufetula Walker; a lectotype is designated for Perforadix sacchari Sein; Micronix Amsel is transferred from the Pyralinae to the Cram-binae; and Micromastra Schaus and Taboga Dyar, revised status, remain in the Pyralinae. In addition, Sufetula pygmaea Hampson, presently in the Crambidae, is transferred to the Noctuidae: Nola pygmaea Hampson (Nolinae), new combination. In the Australian fauna Macna Walker is transferred from the Pyralinae to the Chrysauginae. A list of the subfamilies and tribes of the Pyralidae worldwide and of the species of the Pyralini of the Western Hemisphere are included.
Additional key words: Endotrichini, Pyralini, Neotropics, Australia, larval morphology.
Within the Pyraloidea, the Pyralinae are a large group of about 900 species that are more diverse in Africa and Asia than in the Western Hemisphere. This subfamily includes the worldwide stored-product pest species Pyralis farinalis Linnaeus, also known as the meal moth. A complete study to investigate the monophyly of the Pyralinae has never been conducted. However, Solis and Mitter (1992) proposed a character to define the Pyralinae and hypothesized it to be the sister group to the phycitine + epipaschiine clade (Table 1). In this paper we integrate previous findings in the Pyralinae and our observations to facilitate future studies on these moths.
Presently, there are two tribes in the Pyralinae, the Endotrichini and Pyralini (Table 1). The Endotrichini includes 7 genera, Endotricha Zeller being the largest genus with about 70 species. Based on our morphological and label data observations, the tribe is distributed
only in Asia and Africa. The Pyralini include 118 genera, with the vast majority of the species distributed in Africa and Asia, although some occur worldwide. The two tribes have been defined by two states of a hind-wing venational character (Endotrichini = Rs anastomosed with Sc+R^ Pyralini = Rs not anastomosed with Sc+R1) and they have been shifted between tribal and subfamilial rank based on the importance placed on this character by various authors (e.g., Ragonot 1891, Hampson 1896, Whalley 1961, Minet 1982). We dispute the validity of the use of the hindwing venational character at suprageneric levels. We also explore the literature and investigate the morphology of larval stage as an independent character set.
Recent publication of two checklists (Shaffer et al. 1996, Shaffer & Solis 1995) of the Pyralini and Endotrichini of Australia and the Neotropics, and the previous publication of the checklist of the Pyraloidea
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Table 1. Higher classification of the Pyralidae; current tribal names in use, although most tribes have not been shown to be monophyletic.
Pyralidae Latreille
Chrysauginae Lederer Galleriinae Zeller Galleriini Megarthridiini Tirathabini Cacotherapiini Pyralinae Latreille Pyralini Endotrichini Epipaschiinae Meyrick Phycitinae Zeller
Cryptoblabini
Phycitini
Cabniini
Anerastiini
Peoriini
of North America north of Mexico (Munroe 1983) have laid the groundwork for studies on the systemat-ics of the Pyraloidea. A large number of taxon transfers, and even misplaced taxa between superfamilies, have been documented in the recently published checklists. We herein explain how assignments in recent Australian (Shaffer et al. 1996) and Neotropical (Shaffer & Solis 1995) checklists were made based on our observations on male genitalic morphology and larval morphology. We also list corrections to the tribal and subfamilial headings of the recently published checklists, and list additions or changes made since their publication.
Materials and Methods
The collections at The Natural History Museum (BMNH), London, England; the National Museum of Natural History (USNM), Washington, D.C., USA; the Cornell University Collection (CU), Ithaca, New York, USA; and Zoologische Staatssammlung (ZSBS), Munich, Germany were studied to determine taxa not included in recently published checklists. Type specimens were examined and dissected when necessary. If the type specimen could not be located, the original descriptions and genitalic illustrations were used to place the species generically. Genitalia slides of non-type specimens were prepared, studied, and compared when type specimens were not available, or when type specimens were not in suitable condition for study.
Larvae from alcohol collections of the USNM and BMNH of Endotricha flammealis (Denis & Schiffer-muller), Pyralis farinalis Linnaeus, Aglossa caprealis Hiibner, and Herculia psammioxantha Dyar were examined with a stereomicroscope to verify the literature on larval morphology.
Adult and larval characters of the other subfamilies of the Pyralidae, Chrysauginae, Galleriinae, Phycitinae, Epipaschiinae, were used for out group comparison purposes based on a phylogenetic analysis by Solis and Mitter (1992) (Table 1).
Historical Review and Analysis of Previously Used Characters
Meyrick (1890) first brought the character of the veins 7 [=Rs] and 8 [ = Sc+Rj in the hindwing to attention, and since Ragonot (1891) the Pyralini and Endotrichini have been separated and defined primarily by two character states of the hindwing venation: in the Pyralini Rs and Sc+RL approach each other (Fig. 5), but do not anastomose; in the Endotrichini the two veins anastomose for at least part of their length (Fig. 6). We propose that this character is not reliable in the separation or definition of taxa at suprageneric levels in the Pyralinae and in the following historical review we use italics to draw attention to these two character states in descriptions.
Meyrick (1890) included Endotricha in the Pyralinae. He included the Pyralinae and Epipaschiinae in the Pyralididae and did not recognize them as separate subfamilies. Meyrick defined (and spelled) the Pyralididae as follows:
"Ocelli present, often concealed by scales. Tongue well-developed, or sometimes obsolete. Maxillary palpi well-developed, or rarely rudimentary. Fore wings with vein 1 usually shortly or obscurely furcate at base, sometimes simple, 4 and 5 closely approximated at base or often stalked, 7 and 8 out of 9. Hind wings without defined pecten of hairs on lower margin of cell, veins 4 and 5 closely approximated at base or from a point or stalked, 7 [=Rs] out of 6 near origin or rarely separate but closely approximated, free or sometimes anastomosing with 8 [=Sc+R ]" (Meyrick 1890:433) [italics ours].
He used head and wing venational characters occurring in other groups (i.e., plesiomorphic characters) to define the Pyralididae. The presence of the ocelli and maxilliary palpi are plesiomorphic characters, the proboscis is secondarily lost, the forewing venation is highly variable at lower taxonomic levels (and hence not used by most later workers), and the lack of a hindwing pecten is plesiomorphic.
Ragonot (1891) was the first to separate the Pyralinae and Endotrichinae based on the veins 7 [=Rs] and 8 [=Sc+Rj of the hindwing in a key: "Nervures 7 et 8 soudees aux inferieures, tres rarement separees" ["Veins 7 and 8 fused in the hindwings"] keyed to the Chrysauginae and Endotrichiinae and "Nervures 7 et 8 separees" ["Veins 7 and 8 separate"] keyed to the Pyra-lidinae (Ragonot 1891:446).
Hampson (1896) included only the Phycitinae, Chrysauginae, Epipaschiinae, Endotrichiinae, and Pyralinae in his concept of the Pyralidae and used some of
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uncus
uncus arm
/.A^VUi
tegumen
gnathos
valva
vinculum
Fig. 1. Pyralis farinalis Linnaeus, type species; characteristic male genitalia of Pyralini. Fig. 2. Pyralis farinalis Linnaeus, aedeagus.
the same plesiomorphic characters as Meyrick (1890). But he grouped the Epipaschiinae, Endotrichinae and Pyralinae based on the following two characters:
"The three subfamilies of the Pyralidae, the Epipaschiinae, Endotrichinae, and Pyralinae, of which a classification is here attempted, all belong to the group of Pyralidae which have the median nervure of the hindwing non-pectinate on upperside, and vein 7 [ = Rs] of the forewing stalked with 8 [ = Sc+Rj" (Hampson 1896) [italics ours].
The lack of a hindwing pecten is plesiomorphic, and he used the same character (stalked veins 7 [=Rs] and 8 [=Sc+Rj) of Ragonot to define the Endotrichinae and Chrysauginae.
The relationship and definition of the endotrichines and pyralines was not addressed again until Whalley (1961), who did not provide characters to define the Pyralinae or the Pyralini. To define the Endotrichini he used the same plesiomorphic characters used by Hampson (1896), with the exception of the presence of the chaetosema, but this state is plesiomorphic as well. The Endotrichinae of Hampson (1896) was described as follows:
"Proboscis well developed; maxillary palpi present; build slender. Forewing with vein 7 stalked with 8, 9 (7 absent in Hendecasis).
Hindwing with median nervure non-pectinate; vein 7 [=Rs] anastomosing with 8" [italics ours].
The Endotrichini of Whalley (1961) was described as
follows:
"Proboscis well developed, maxillary palps present. Chaetosema present. Forewing with vein R5 stalked with R4 and R3. Hind wing with median vein non-pectinate. Vein Rs anastomosing with Sc+R" [italics ours].
Munroe and Shaffer (1980) revised three large genera in the Pyralini (Pyralinae). Their definition of the Pyralinae is basically a combination of Hampsons (1896) definition of the Endotrichinae and Pyralinae from a key with Whalley s (1961) rank of tribes. The Pyralinae of Hampson (1896) was described as follows:
"Proboscis usually well-developed; maxillary palpi present and usually filiform. Forewing with vein 7 stalked with 8,9. Hindwing with the median nervure non-pectinate; vein 8 [=Sc+R.}] free" [italics ours].
The Pyralinae of Munroe and Shaffer (1980) was described as follows:
"The three genera can now be defined as belonging to the Pyralinae from the following characters: chaetosema present; maxillary palpus present; proboscis well developed; fore wing with R5 stalked with R4 and R3; hind wing with Rs not anastomosed with Sc+R1 ( Pyralini), or
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uncus
uncus arm tegumen gnathos transtilla
valva saccular process
vinculum
Fig. 3. Endotricha flammealis (Denis & Schiffermiiller), type species characteristic male genitalia of Endotrichini. flammealis (Denis & Schiffermiiller), aedeagus.
FiG. 4. Endotricha
Rs anastomosed with Sc+R1 (Endotrichini); median vein non-pectinated" [italics ours].
Whalley (1961) recognized the problem with the definition of the hindwing character that separated the two groups: "In several cases they have been said to anastomose where, as close examination shows, they merely run very close together (e.g., Rostripalpus Hampson)." The lack of anastomosis of Rs and Sc+Rj varies in other groups within the Pyraloidea besides the Pyralini, and it has been documented as highly ho-moplasious at the generic level among the genera of the Pococera complex of the Epipaschiinae (Solis 1993) and at the species level (Shaffer & Solis 1994). Other groups where the majority of the taxa lack the anastomosis of Rs and Sc+RJ but where there are examples where the two veins barely anastomose have
been observed in representatives of the New World Cacotherapiini (Galleriinae) and some genera in the Crambinae (e.g., Pseudoschoenobius Fernald). Based on our observations of the distribution of this hind-wing character in other groups within the Pyraloidea, distribution of the hindwing character within the Pyral-inae, and lack of concurrence with the characters of the male genitalia, we propose that the hindwing venational character is not reliable in the separation or definition of taxa at suprageneric levels in the Pyralinae.
Minet (1982, 1985) was the first to maintain that the Pyralinae were paraphyletic because characters used by past workers were plesiomorphic. He stated: "Les Pyralinae semblent paraphyletiques par rapport a des taxa tels que les Endotrichinae, les Chrysauginae ou les Epipaschiinae (dont ils ne different que par un en-
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Fig. 5. Wing venation (idealized); arrow indicates lack of anastomosis. Fig. 6. Wing venation (idealized); arrow indicates anastomosis.
semble de caracteres plesiomorphes: palpes maxil-laires bien developpes, ailes anterieures sans ecailles herissees, etc.)," ["The Pyralinae appear to be para-phyletic in comparison with taxa such as the Endotrichinae, the Chrysauginae or the Epipaschiinae (in that they share a group of plesiomorphic characters: maxillary palpi well developed, forewings without raised scales, etc.)."] but he retained pyralines and en-dotrichines at the subfamily level. Whalley (1963), in his study of Endotricha, found that the retention of Ragonots concept of the Endotrichinae as a subfamily was not warranted and proposed the Endotrichinae as a tribe of the Pyralinae. He did not offer a reason or characters to support this conclusion. Solis and Mitter (1992) agreed with Minet that previous characters used to define the two taxa were plesiomorphic states, but they treated the endotrichines as a tribe within the Pyralinae according to Whalley (1961) because Minet (1985), in his study of the tympanal organs, presented no apomorphies for the Pyralinae, Pyralini, or En-dotrichini. Solis and Mitter (1992) proposed one character of the female genitalia as a synapomorphy for the Pyralinae, but proposed none for the Pyralini or En-dotrichini.
Results
Adult genitalic morphology. Previous authors (Whalley 1961, 1963, Munroe & Shaffer 1980) did not use genitalic characters to define the Pyralinae, Pyralini, or Endotrichini, although they used genitalic morphology at the species level for their studies. Solis and Mitter (1992) proposed a character of the female genitalia (corpus bursae barely extending cephalad beyond segment 7) to support the monophyly of the Pyralinae. This study, however, was based on a small sample size, a character that remains untested.
Pyralini (Figs. 1, 2, 7):
Description: Male: Uncus same width throughout or less narrow than the base, flat or spatulate, ventrally with spine clusters absent; uncus arms laterally not large and earlike; downcurved gnathos with arms strongly developed, with well-developed medial, narrow spike terminating in a small dorsally curved hook; tegumen strongly sclerotized; vinculum well developed; juxta simple, rarely heavily sclerotized, spiny catena (baso-medial portion of anellus) present or absent or laterally sclerotized, and heavily spined, sometimes anellus reflexed with heavy sclerotization; transtilla absent or, if present, membranous, rarely well developed and heavily sclerotized; valva variable in shape, same width to apex or more narrow distally, basal and costal process absent or present, if present well developed or not, without saccular process, ventral surface of valva bearing hairlike setae not arranged in radiating rows, costal setae absent; vesica of aedeagus with or without clusters of spinelike cornuti, vesica sometimes spined, reflexed with heavy sclerotization, or with broad bands of sclerotization its entire length. Female: Segment 8 and associated membranes either short, or long and extensible; apophyses anteriores and posteriores long, stout or slender; ostial end of ductus bursae membranous, with small, well-sclerotized to large, heavily sclerotized compact pouches present or absent; ductus bursae long and narrow with areas of minute spines immediately below antrum or other sclerotized areas; corpus bursae large, signum variable, absent, or if present from scobinate patches, usually within single large area, to long and spinelike.
Endotrichini (Figs. 3, 4, 8):
Diagnosis: Uncus broadest at apex; uncus arms laterally large, earlike; gnathos medially broad, spatulate, platelike.
Description: Male: Uncus broadest at apex, ventrally with spine clusters present or absent (uncus process of Whalley); uncus arms laterally large, earlike [socii of Whalley; socii, according to Klots (1956) are paired processes on either side of the base of the uncus; these structures are not socii, but the most lateral elements of the uncus arms of the Pyralidae (Solis & Mitter 1992)]; downcurved gnathos arms strongly or weakly developed, usually with a well-developed medial, broad, spatulate, and upturned central plate; weakly sclerotized tegumen; vinculum well developed; juxta simple, sometimes with spiny manica; transtilla present, usually heavily sclerotized; valva usually same width to apex, may bear basal process and saccular process; ventral surface bearing hairlike setae in rows pointing toward base of valva; prominent, reflexed, sometimes spear-shaped costal setae may be present arising from costa near apex; aedeagus with vesica bearing sticklike or clublike cornutus varying in shape and length. Female genitalia: Segment 8 and associated membranes long and extensible; apophyses anteriores and posteriores long and slender; ostial end of ductus bursae minutely spined, without pouches; antrum sclerotized; ductus bursae short, minutely spined, or very long and membranous; corpus bursae large with signum scobinate.
Although we can provide synapomorphies in states of the uncus and gnathos (see diagnosis above) for the Endotrichini in the male
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Fig. 7. Pyralis farinalis Linnaeus, type species; characteristic female genitalia of Pyralini. Fig. 8. Endotrichaflammealis (Denis & Schif-fermiiller), type species; characteristic female genitalia of Endotrichini.
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genitalia, we were unable to find synapomorphies for the Pyralini where the states are either shared with the Endotrichini or with related subfamilies. The genitalic characters of the Pyralinae are either variable at lower taxonomic levels or plesiomorphic, i.e., found in all related subfamilies. We provide a description of the genitalia for Endotrichini and Pyralini because a concept based on the morphology of the male genitalia was used to assign taxa in the Endotrichini and Pyralini in the Australian and Neotropical checklists.
Larval morphology. Comparison of the caterpillars of En-dot richa flammealis with Pyralis farinalis, Aglossa caprealis, and Herculia psammioxantha (with caterpillars of other subfamilies of the Pyralidae as outgroups) did not result in any apomorphic characters to support the monophyly of the Pyralinae, Pyralini, or Endotrichini. Historically, the Hasenfuss (1960) concept of the Pyralinae consisted of present-day galleriines, pyralines, and phycitines (he did not include chrysaugines or epipaschiines in his study); he considered Endotricha as a pyraline.
The larvae of Endotricha have a pinaculum ring on SD1 of A9, a synapomorphy for the Pyralidae (the plesiomorphic state, the absence of the pinaculum ring on SD1 of A9, occurs in the Cram-bidae). In sum, we found that E. flammealis larvae lack the unique characters assigned to other subfamilies and have the same plesiomorphic setal character states assigned to the larvae of the Pyralinae. The Epipaschiinae and Pyralinae both lack a pinaculum ring on any other segment other than A9 (in comparison to the presence of a pinaculum ring on T2 of the Phycitinae, T3 of the Chrysauginae, and Al of the Galleriinae; presence in each segment is the derived state, although the pinaculum has been secondarily lost in several genera and/or species of each subfamily). Based on work by Hasenfuss (1960) and Allyson (1977) the Epipaschiinae and Pyralinae are separated from each other by the distance between the ventral setae on A7 and A9. In the Epipaschiinae the two ventral setae are closer together on A7 than those on A9 and in the Pyralinae the two ventral setae on A7 and A9 are equidistant (the plesiomorphic condition shared by other subfamilies of the Pyralidae).
Taxonomic placement of genera. Recently, genera from southeast Asia and Australia previously placed in the Endotrichinae with anastomosed Rs and Sc+R, in the hindwing, but with genitalic characteristics of the Pyralini were transferred to the Pyralini in the Australian checklist (Shaffer et al. 1996) based on the genitalia morphology. Those genera transferred from the Endotrichinae to the Pyralinae were based on the genitalia morphology: Gauna Walker, Curena Walker, Arescoptera Walker, Scenedra Meyrick, Tanyethira Turner, Scenidiopsis Turner, Perisseretma Warren, and Ferula Mabille.
According to the definition based on genitalic morphology given above, there are no known species of Endotrichini in the Western Hemisphere, but four genera, Perforadix Sein, Micronix Amsel, Micromastra Schaus, and Taboga Dyar have been historically placed within the Endotrichinae due to the anastomosing of Rs and Sc+R} in the hindwing. Perforadix, Micronix, and Micromastra were inadvertently excluded from the Neotropical Pyraloidea checklist (Shaffer & Solis 1995). Taboga was included in the Neotropical checklist, but needed to have its position in the Pyralinae verified. We found that Perforadix belongs in the Pyraustinae and Micronix belongs in the Crambinae, both hereby transferred, and, of the four, only Micro-
mastra and Taboga remain in the Pyralinae. Table 2 is a complete list of the Pyralinae (Pyralini) of the Western Hemisphere (Munroe 1983, Shaffer & Solis 1995).
Sein (1930) placed Perforadix sacchari Sein, commonly known as the Sugarcane root caterpillar, in the Endotrichinae. This species is a major pest of sugar cane in Puerto Rico and nearby islands. Sein (1930) illustrated the morphology of all life stages in great detail and described its biology and methods of control. The author failed to designate types or even list type specimens, but we found seven specimens each with a small label "P.R./Sein" and a red label "Cotype/Cornell U. No. 6087" at Cornell University. According to Sein (1930), W. T M. Forbes, who was at Cornell University at the time, identified the material and presumably he also labelled the material as cotypes. We designate one specimen (male) as the lectotype and the other 6 specimens as paralectotypes (material in poor condition, abdomens are missing), and they are labelled as such in the collection at Cornell University. We studied additional material collected by Sein, identified by H. G. Dyar, as stated by Sein (1930), and dissections by Carl Heinrich at the USNM and found that Perforadix is a synonym of Sufetula Walker, new synonymy, in the Pyraustinae. We discovered that based on the morphology of the tympanal organs (i.e., cram-bid "open" tympanal organs with a praecinctorium) it belongs in the Crambidae. Based on the external and genitalic morphology after comparison with other species in the genus, including the type species, it belongs in the genus Sufetela Walker. It is interesting to note that P. sacchari was originally identified for Sein by H. G. Dyar as Sufetula grumalis Schaus, a species presently placed in Sufetula (Munroe, 1995:76). We also examined another species, Sufetula pygmaea Hampson, and found it does not belong in the Pyraloidea, but in the Noctuidae (Nolinae): Nola pygmaea (Hampson), new combination.
Amsel (1956) described Micronix nivalis in the Endotrichinae. Nothing is known about the biology of this Venezuelan species. We were unable to locate the type, a male, but Amsel provided a photograph of the adult and poor line drawings of the male genitalia and wings. The hooded uncus and the costal process of the male genitalia characteristic of crambines are very evident in the illustration; therefore, we transfer this monotypic genus to the Crambinae. Although we cannot determine its placement within the Crambinae, we suggest that it belongs in the tribe Crambini.
Schaus (1940) placed Micromastra isoldalis in the Endotrichinae. Nothing is known about the biology of this Puerto Rican species. Dyar (1914) described Taboga inis in the Endotrichinae. The type series is from
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Table 2. Pyralini (Pyralidae: Pyralinae) of the Western
Aglossa Latreille, [1796]
Euclita Hiibner, [1825]
Agriope Ragonot, 1894 acallalis Dyar, 1908 baba Dyar, 1914 cacamica (Dyar, 1913) (Pyralis) caprealis (Hiibner, [\800-09])(Pyralis)
capreolatus Haworth, 1809
cuprealis Hiibner, [1825], missp.
aenalis (Costa, 1836)(Pyralis)
domalis Guenee, 1854
incultella (Walker, [1866])(Acrobasis)
enthealis (Hulst, 1886)(Tetralopha)
cuprialis Heinrich, 1931, missp. costiferalis (Walker, 1886) (Pyralis)
costigeralis (Walker, [1865] (Pyralis), preocc. (Walker, 1862) cuprina (Zeller, 1872) (Pyralis) disciferalis (Dyar, 1908) (Pyralis) electalis Hulst, 1866 furva Heinrich, 1931 gigantalis Barnes & Benjamin, 1925 oculalis Hampson, 1906 pinguinalis (Linnaeus, 1758)(Pyralis)
marmorella (Geoffroy, 1785)(Tinea)
marmoratella (Villers, 1789)(Tinea)
pinguiculatus (Haworth, 1809)(Crambus)
guicciardii Constantinio, 1922 Arispe Ragonot, 1891
Uscodys Dyar, 1909 cestalis (Hulst, 1886)(Anerastia)
atalis (Dyar, 1908) (Uscodys) concretalis Ragonot, 1891 ovalis Ragonot, 1891 Catocrocis Ragonot, 1891
Catacrocis Ragonot, 1892, missp. lithosialis Ragonot, 1891 Dolichomia Ragonot, 1891
amoenalis (Moschler, 1882) (Asopia)
isidora (Meyrick, 1936)(Pyralis) binodulalis (Zeller, 1872)(Asopia) craspedalis (Hampson, 1906) (Tegulifera) datames (Druce, 1900)(Pyralis) decetialis (Druce, 1900)(Pyralis) graafialis (Snellen, 1875)(Asopia) impurpuratalis (Dognin, 1910)(Pyralis) nigrapuncta (Kaye, 1901)(Pyralis) olinalis (Guenee, 1854)(Pyralis)
trentonalis (Lederer, 1863)(Asopia)
himonialis (Zeller, 1872)(Asopia)
infimbrialis (Dyar, 1908)(Hercidia) phanerostola (Hampson, 1917)(Paractenia) planalis (Grote, 1880)(Asopia)
enniculalis (Hulst, 1886)(Asopia)
occidentalis (Hulst, 1886)(Asopia) plumbeoprunalis (Hampson, 1917')(Herculia) resectalis (Lederer, 1863)(Asopia) thymetusalis (Walker, 1859)(Botys) New combination
devialis (Grote, 1875)(Asopia) vernaculalis (Berg, 1874) (Asopia) Herculia Walker, 1859
Buzala Walker, 1863
Cisse Walker, 1863
Bejuda Walker, [1866]
Bleone Ragonot, 1890
Herculea Amsel, 1956 [index], missp. tabidalis (Warren, 1891)(Pyralis)
Hypsopygia Hiibner, [1825]
costalis (Fabricius, 1775)(Phalaena)
fimbrialis ([Denis & Schiffermuller], 1775)(Pyralis)
purpurana (Thunberg, 1784)(Tortrix)
hyllalis (Walker, 1859)(Pyralis) Mapeta Walker, 1863
Homalochroa Lederer, 1863 cynosura Druce, 1895 omphephora Dyar, 1914 schausi Druce, 1895 xanthomelas Walker, 1863
aestivalis (Lederer, \863)(Homalochroa)
divisa (Boisduval, 1870)(Pyralopsis) Micromastra Schaus, 1940
isoldalis Schaus, 1940 Neodavisia Barnes & McDunnough, 1914
Davisia Barnes & McDunnough, 1913, preocc (Del
guercio, 1909 [Hemiptera]) melusina Ferguson, Blanchard, & Knudson, 1984 singularis (Barnes & McDunnough, 1913)(Davisia) Ocrasa Walker, [1866]
Parasopia Moschler, 1890 nostralis (Guenee, 1854)(Pyralis)
helenensis (Wollaston, 1879)(Pyralis)
tenuis (Butler, 1880)(Pyralis)
dissimilalis (Moschler, 1890)(Parasopia)
sordidalis (Barnes & McDunnough, 1913)(Herculia)
psammioxantha (Dyar, 1917)(Herculia)
venezuelensis (Amsel, 1956)(Herculia) tripartitalis (Herri ch-Schaffer, 187l)(Asopia) Pseudasopia Grote, 1873
cohortalis (Grote, 1878)(Asopia)
florencealis (Blackmore, 1920)(Herculia) intermedialis (Walker, 1862)(Pyralis)
sodalis (Walker, 1869)(Pyralis)
squamealis Grote, 1873 phoezalis (Dyar, 1908)(Herculia) Pyralis Linnaeus, 1758
Aletes Rafinesque, 1815, nom. nud.
Ceropsina Rafineques, 1815, nom. nud.
Spyrella Rafinesque, 1815, repl. name
Asopia Treitschke, 1828
Sacatia Walker, 1863
Eutrichodes Warren, 1891 farinalis Linnaeus, 1758
domesticalis (Zeller, 1847)(Asopia)
fraterna Butler, 1879
manihotalis.- Matsumura, 1900 (not Guenee, 1854)
meridionalis Schmidt, 1934
orientalis Amsel, 1961 manihotalis Guenee, 1854
vetusalis Walker, [1859]
gerontesalis Walker, [1859]
laudatella (Walker, 1863)(Sacatia)
despectalis Walker, [1866]
miseralis Walker, [1866]
achatina Butler, 1877
haematinalis (Saalmiiller, \880)(Asopia)
gerontialis (Meyrick, 1888)(Asopia), emend.
centripunctalis (Gaede, 1916)(Endotricha)
pupalis Strand, 1919
compsobathra Meyrick, 1932 Taboga Dyar, 1914 inis Dyar, 1914
Volume 53, Number 1
9
Panama. Study of the genitalia of the type series at the USNM of both of these species confirm their placement within the Pyralini, and not in the Endotrichini.
In the Australian checklist (Shaffer et al. 1996:173) the headings of the Endotrichini and Pyralini were mislabeled and difficult to change at proof stage. The Endotrichinae should have been titled the Endotrichini and placed under the heading of the Pyralinae. In addition, the genus Macna Walker was inadvertently included in the Pyralinae (Pyralini), but it should have been placed in the Chrysauginae. In the Neotropical checklist (Shaffer & Solis 1995:80) the Pyralini should have been included as a subheading under Pyralinae to indicate the tribal placement of the genera found in the Western Hemisphere.
Discussion
The Pyraloidea, one of the larger superfamilies of the Lepidoptera, has over 15,000 described species, yet much remains to be done in taxonomy, and, more so, with the phylogenetic relationships. A taxonomic study usually begins with a checklist or a catalogue of described species as an inventory to document those that have already been described. A checklist may refine the placement of taxa and can clearly mark taxon transfers, as well as provide other information, such as misplaced taxa. By definition, a checklist or catalog does not adequately state or discuss the taxonomic problems solved or those that remain to be solved.
We have described the morphological reasons for the placement of taxa in the Pyralini or Endotrichini in two checklists (Shaffer et al. 1996, Shaffer & Solis 1995). We have also summarized the taxonomic and phylogenetic status of the two tribes included in the Pyralinae. The genitalia of the Endotrichini are clearly different from those of the Pyralini, but authors have dealt only with a few genera in both taxa and, as we have shown, have used the same plesiomorphic or ho-moplasious characters since Meyrick (1890) to define higher level taxa. We retain the two tribes in the classification system for the sake of stability and retention of character information, but acknowledge that the Pyralini is likely a paraphyletic group. Moreover, a preliminary study of an independent character set, the larval stage, provides no obvious synapomorphies for the Pyralinae or the Pyralini.
Our observations of the genitalia and larvae of the Pyralini and Endotrichini are made with the expectation that they may prove useful in a future phylogenetic study of the Pyralini genera that includes an entire suite of adult and immature characters. Any future study should also include pupal and perhaps behav-
ioral characters, although caution is suggested concerning the latter due to the convergent nature of behavioral characters. A phylogenetic analysis of the genera of the Pyralini would be the first attempt to test the paraphyly of the Pyralini with respect to the Endotrichini, with the possibility that results may also invalidate the traditional tribal concept. Such a study may also provide characters to support the monophyly of the Pyralinae.
Acknowledgments
We thank for their hospitality: E. R. Hoebeke, Cornell University, Ithaca, New York, USA, for help while studying the Sein material and A. Hausmann, Zoologische Staatssammlung, Munich, Germany, while studying the Amsel material. We also thank Linda Lawrence, scientific illustrator, Systematic Entomology Laboratory, U.S. Department of Agriculture, for the figures. The following provided reviews or helpful comments: Vitor O. Becker, Marianne Horak, Douglas Ferguson, James Pakaluk, and David Smith.
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Received for publication 15 February 1998; revised and accepted 29 September 1998.
MANUSCRIPT REVIEWERS, 1998
The merit of a scientific journal depends on the quality of its reviewers as well as its authors, but the former are usually unknown to the readers of the published articles. The Journal relied on the expertise of 60 reviewers last year to provide 66 evaluations of manuscripts. It is with much gratitude that the Journal acknowledges the services of the people listed below, from whom manuscript reviews were received in 1998. Those who reviewed two or more manuscripts are denoted by asterisks.
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