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lanthus rigidus (Benth.) Jeps. (Orobanchaceae) (also a new food plant for the species) and Plant ago erect a during the springs of 1999 and 2001 (GFP pers. obs.). Recent molecular studies suggest these food plants (Cordijlanthus and Plantago) are more distantly related (separate families) than are Antirrhinum and Plantago (same family) (Olmstead et al. 2001). The food plant therefore may not be the most important character used to distinguish Euphydryas editha subspecies.
Alison Anderson, Greg Ballmer, Dave Hawks, Guy Bruyea, Chris Nagano, Brenda McMillan, and John Emmel have provided information on present and historical populations. Cecilia L. Pierce assisted the senior author on much of the fieldwork. We thank Fred Sproul for finding and providing information on the newly discovered San Vicente Reservoir population, Steve Meyers, Chet Mc-Gaugh, and Mike Wilcox for finding and providing information on a population north of Anza, and Sam Reed for providing information on a population southwest of Anza. We thank Andy Sanders for all of the plant identifications. We also thank Bob Luck for all of his help and support. This research was largely supported by a Fish and Game grant with the account title of CDFG FG7182ES LUCK 6/98.
Literature Cited
Garth, J. S. & J W Ttlden. 1986. California butterflies. University of California Press, Berkeley, California, 246 pp.
Journal of the Lepidopterists' Society 55(4), 2001.171-174
In 1840, J. A. Boisduval described a new species of Catocala Schrank (1802) as Catocala protonympha, as follows: "Species distinctissima antennis crassioribus dentatis; statura Callinympha; alae posticae fere ut apud jam dictam; anticae cinereo-fuscae fasciis vix an-gulosis" [A species very distinct, by thick dentate antennae; size of callinympha; hindwings about the same as with those I have already described; forewings ash grey-brown with bands slightly angled]. The type locality was listed as "P." [=Paris, France].
Guenee (1852:103) figured protonympha on his Plate 15, Fig. 2, comparing it to Catocala paranympha L., which is a synonym of the widespread Palearctic species Catocala fulminea (Scopoli 1763), and stated "Environs de Paris, en aout. Coll. Bdv. Un 6" [vicinity of Paris, in August. Collection of Boisduval. One male].
Mattoni, R., G. F. Pratt, T. H. Longcore, ]. F. Emmel & ). N. George. 1997. The endangered quino checkerspot butterfly Euphydryas editha quino (Lepidoptera: Nymphalidae). J. Res. Lepid. 34:99-118.
Olmstead, R. G., C. W Depamphilis, A. D. Wolfe, N. D. Young, W. J. Elisons & P. A. Reeves. 2001. Disintegration of the Scrophulariaceae. Amer. J. Rot. 88 (2):348-361.
Rogers, J. G. 1997. Endangered and threatened wildlife and plants: determination of endangered status for the Laguna Mountains skipper and quino checkerspot butterfly. Federal Register January 16, 1997 Vol. 62:2313-2322.
Singer, M.C. 1971. Evolution of food-plant preference in the butterfly Euphydryas editha. Evol. 25:383-389.
---------. 1972. Complex components of habitat suitability within a
butterfly colony. Science 176:75-79.
---------. 1982. Quantification of host preference by manipulation of
opposition behavior in the butterfly Euphydryas editha. Oeco-logia 52:224-229.
White, R. R. 1974. Food plant defoliation and larval starvation of Euphydryas editha. Oecologia 14:307-315.
Gordon F. Pratt, Department of Entomology, University of California, Riverside, California 92521, USA, Eric W. Hein, 1515 Cassidy Street, Oceanside California 92054, USA, and Douglas M. Krofta, 1361 Bluegrass Rd, Vista, California 92083, USA
Received for publication 8 January 2000; revised and accepted 22 October 2001.
Guenees original illustration of Catocala protonympha is reproduced here in Fig. lc. Guenee also remarked on the apparent extraordinary rarity of protonympha: "C'est jusqu'ici, une des plus grandes raretes. Pourtant, M. Begrand m'a affirme avoir vu, sur un mur, au bois de Boulogne, une certaine quantite de petites Catocala jaunes qui ne pouvaient appartenir qu'a cette espece, la paranympha, avec laquelle la confusion est impossible" [Up to now, it is one of the great rarities. However, Mr. Begrand is positive about having seen a certain number of small yellow Catocala on a wall in the Boulogne woods, they necessarily belonged to the species paranympha with which no confusion is possible].
Subsequently, Catocala protonympha was listed with questionable provenience by Berce (1870:242): "est regardee comme douteuse par beaucoup de lepi-
PRECEDENCE OF CATOCALA LOUISEAE BAUER, 1965 AS A NOMEN PROTECTUM OVER CATOCALA PROTONYMPHA BOISDUVAL, 1840 (NOCTUIDAE)
ABSTRACT. The name Catocala protonympha Boisduval has heretofore been considered a junior synonym of the Palearctic species Catocala fulminea Scopoli. Examination of Bosiduvals type and relevant literature demonstrates that Catocala protonympha is in fact a disused senior synonym of the Nearctic species Catocala louiseae Bauer. Article 23.9.2 of the Code is invoked to give precedence to the established name Catocala louiseae, and maintain nomenclatural stability in the genus.
Additional key words: taxonomy, synonymy, types, Boisduval, Guenee, Staudinger, Culot.
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Journal of the Lepidopterists' Society
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Fig. 1. Catocala protonympha Boisduval 1840. a, male type in the Natural Histor)' Museum (London), approx. 3/4 size, b, specimen labels the type, c, painting of the type by Guenee (1852).
dopteristes" [is seen as doubtful by many lepidopterists], and the possibility of a hybrid or misattributed specimen was raised by Staudinger and Wocke (1871:138): "uno '": condita; an Eutychea al. posteri-oribus Hymenaeae adulterata? an sp. exotica?" [one male preserved; is it eutychea or otherwise from descendants of a cross with hymenaea?; or an exotic species?]. Despite the fact that Guenees illustration of protonympha does not resemble Catocala fulminea in either size or pattern, Staudinger and Rebel (1901:249) later placed protonympha as a dwarfed aberration of fulminea, believing then that Guenee s illustration was wanting: "t. 15, f. 2 [fig. non quadrat bene]; (sec. specim. typ. Stgr. ab. nana esse videtur)" [the figure does not agree well; according to the type specimen of Staudinger from which it seems to be dwarf]. Both Spuler (1908) and Hampson (1913) also treated protonympha as an aberration of fulminea, and in the Seitz (1913:319) volume on Palearctic noctuids, Warren treated protonympha as a form of fulminea. The Seitz figure of protonympha is nearly identical to the Seitz figure of fulminea (both appear on Plate 57 row D), the former differing from the latter only nominally in having the hindwing ground color lighter and the hindwing medial band somewhat reduced.
Culot (1913:201-203, Plate 77, Fig. 4) refigured Boisduval s type of protonympha, and correctly pointed out that the type did not at all resemble the Seitz figures of protonympha or fulminea. Although Culot clearly felt protonympha and fulminea were different, and that some sort of error might be involved, he left the taxonomic position of protonympha unchanged, as he was unable to resolve fully the puzzle that Boisduval s specimen presented:
"Quant a la soi-distant forme protonympha Bdv, elle parait etre fort peu connue des entomologistcs et ne correspond en aucune facon a l'exemplaire reproduit soue ce nom sur la Planche 57, ligne D, de l'ouvrage de Seitz. Pour ma part, je trouve la protonympha Bdv. si differente de fulminea qu'il
me parait temeraire de la rattacher specifiquement a cette derniere. Grace a la si large obligeance de mon ami M. Charles Oberthur, j'ai le plaisir de figurer ici le type protonympha de Boisduval. . . . En resume, je vois dans la protonympha Bdv. une espece tres particuliere, don't on ne connait que le seul exemplaire 6 figure ici et qui fut capture aux environs de Paris. C'est un cas evidemment fort rare en entomologie, mais a moins qu'il ne s'agisse d'une erreur don't j'ignorerais la source, je ne puis trouber aucune autre solution." [As for the so-called form protonympha Bdv, it seems to be mostly unknown to entomologists and in no way corresponds to the example presented under this name in Plate 57, line D of Seitz' work. As far as I am concerned, 1 consider protonympha Bdv. to be so different from fulminea that it seems farfetched to relate it specifically to that species. Thanks to the kindness of my friend Charles Oberthur, I have the pleasure to illustrate here the tspe of protonympha by Boisduval. ... In short, I see in protonympha Bdv. a very particular species of which only one male specimen, illustrated here, is known and that was captured around Paris. It is naturally a case seldom seen in entomology, but I cannot think of any other solution except for a mistake that I cannot explain].
Despite Culot's misgivings, in later systematic works the name protonympha has also been listed as a synonym of fulminea (e.g., Lhomme 1923-1935, Poole 1989, Hacker 1990), or has been omitted from the synonymy of fulminea, even in works covering France or nearby areas (e.g., Bergmann 1954, Forster & Wolfliart 1971, Leraut 1980). The most recent use of the name protonympha as a valid species name appears to be by Staudinger and Wocke (1871).
In 1965, Bauer described Catocala louiseae as a new Nearctic species from Florida. This local and generally uncommon Catocala occurs primarily in the southeastern United States, from North Carolina to Florida and
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westward along the Gulf Coast to Texas. The name louiseae has been used exclusively for this species, having appeared in catalogues treating Nearctic (Hodges 1983) and Holarctic Noctuidae (Poole 1989), a book devoted to Catocala (Sargent 1976), a moth field guide (Covell 1984), the experimental zoological literature (Gall 1991), regional surveys and compilations (Cro-martie & Schweitzer 1997), and in numerous shorter reports on Catocala distributions and life histories in the News of the Lepidopterists' Society and the Southern Lepidopterists' News (e.g., Baggett 1994, Neal 1999).
We recently located Boisduval's type specimen of Catocala protonympha at the Natural History Museum, London in one of the "miscellaneous" type drawers. The male type (Fig. la) bears the following labels (Fig. lb): "Catocala/protonympha/Paris, type"; "Vu par/Staudinger/Catalogue 1900"; "Fig. par J. CU-LOT/Noct et Geom d'Europe/Pl. 77 Fig. 4"; "EX MUSAEO/BOISDUVAL"; "Ex Oberthur Coll/Brit. Mus. 1927-3." The type is in fact a specimen in good condition of Catocala louiseae, and not an aberrant dwarfed specimen of Catocala fulminea.
Because the name protonympha actually refers to a local and generally uncommon Nearctic Catocala species, this explains prior lepidopterists' difficulties in placing protonympha as a Palearctic taxon, and the absence of any Palearctic specimens other than Boisduval's type. The type locality of Paris, France for protonympha is undoubtedly erroneous, and likely the result of a labeling error or other misattribution. Boisduval's type could even be one of John Abbots 18th or early 19th century Lepidoptera specimens from Georgia, as Catocala louiseae inhabits the counties in Georgia where Abbot worked, and Abbot did depict louiseae in one of his unpublished watercolors in the Oemler compilation at the Houghton Library at Harvard University (the Francillon compilation of Abbot s unpublished watercolors at the Natural History Museum, London does not contain a painting of louiseae). Note that even though Guenee's illustration of protonympha is stylized, it matches the type well, and is nevertheless recognizable as louiseae. Staudinger and Rebel's (1901) mischaracterization of Guenee's illustration seems all the more peculiar in light of the data label that indicates Staudinger examined Boisduval's specimen, and especially since Staudinger and Wocke (1871) initially felt that protonympha might not be a Palearctic species.
Thus, the name protonympha Boisduval (1840) has been mistakenly tabulated throughout the 20th century in the Palearctic Catocala literature as a junior subjective synonym of fulminea Scopoli (1763). The name protonympha has never appeared in the Nearctic Catocala literature, and reintroducing the name
protonympha for louiseae Bauer (1965) would upset established nomenclatural usage. Since the provisions of both Articles 23.9.1.1 and 23.9.1.2 of the Code appear to be met, the name Catocala louiseae Bauer (1965) is hereby given precedence per Article 23.9.2 as a nomen protectum over its disused senior subjective synonym Catocala protonympha Boisduval (1840), which becomes a nomen oblitum.
We thank Martin Honey for his hospitality at the Natural History Museum (London), and for providing access to specimen material there. Julie Harvey, Michael Pogue and John Rawlins chased down several literature leads. Louis Hanfield assisted with the French translations, and Stephanie Spaulding, Leo Hickey and Victor Bers with the Latin. Don Lafontaine offered helpful comments on the manuscript.
Literature Cited
Baggett, H. D. 1994. Current zone reports. Zone IV Florida.
News So. Lepid. Soc. 16:14. Bauer, J 1965. A new Catocala from Florida (Lepidoptera: Noctuidae). Entomol. News 76:197-198. Berce, J. E. 1870. Faune Entomologique Francaise Lepidopteres.
Quatrieme Volume: Heteroceres, Noctuae. Chez Deyrolle Fils,
Paris. 256 pp. Bergmann, A. 1954. Die Grosschmetterlinge Mitteldeutschlands.
Band 4/2. Eulen. Urania-Verlag, |ena. 479 pp. Boisduval, J. B. A. D. de. 1840. Genera et Index Methodicus Eu-
ropaeorum Lepidopterorum. Roret, Paris. 238 pp. Covell, C. V. 1984. A field guide to the moths of eastern North
America. Houghton Mifflin, Boston. 496 pp. Cromartie, J & D. F Schweitzer. 1997. Catocala louiseae, C.
grisatra and C.jair (Lepidoptera: Noctuidae) in North Carolina. Entomol. News 108:389-390. Culot, J. 1913-1917. Noctuelles et Geometres d'Europe. Premiere Partie. Noctuelles. Vol. 2. Oberthur, Rennes. 243 pp. Forster, W. & T. A. Wohlfart. 1971. Die Schmetterlinge Mit-
teleuropas. Eulen (Noctuidae). W. Keller, Stuttgart. 329 pp. Gall, L. F. 1991. Evolutionary ecology of sympatric Catocala
moths (Lepidoptera: Noctuidae). III. Experiments on female
opposition preference. J. Res. Lepid. 29:217-233. Guenee, A. 1852. Histoire Naturelle des Insectes. Species General
des Lepidopteres. Tome Septieme, Noctuelites, Tome 3. Roret,
Paris. 441 pp. Hacker, H. 1990. Systematische und synonymische Liste der
Nocktuiden Deutschlands und der angrenzenden Gebiete. Es-
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the British Museum. Volume 12. Taylor & Francis, London.
626 pp. Hodges, R. W. 1983. Check list of the Lepidoptera of America
north of Mexico. Univ. Press, Cambridge. 284 pp. Leraut, P. 1980. Liste systematique et synonymique des lepidopteres de France, Belgique et Corse. Alexanor 13 (supple-
ment):l-334. Lhomme, L. 1923-1935. Catalogue des Lepidopteres de France et
de Belgique. Volume I. L. Lhomme, Paris. 800 pp. Neal, T. 1999. State Coordinator reports. Florida. News So. Lepid.
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(N. S.),Fasc. 118:1015-1314. Sargent, T. D. 1976. Legion of night: the underwing moths. Univ.
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Varietates Methodo Linnaeana. Joannis Thomae Trattner,
Wien. 418 pp. Seitz, A. 1909-1914. Die Gross-Schmetterlinge der Erde. Die
Gross-Schmetterlinge des Palaearktischen Faunengebietes.
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gemeiner Teil. Spezieller Teil. E. Schweizerbarche, Stuttgart.
385 pp. Staudinger, O. & H. Rebel. 1901. Catalog der Lepidopteren des
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Staudinger, O. & M. Wocke. 1871. Catalog der Lepidopteren des Europaeischen Faunengebiets. [No publisher given], Dresden. 426 pp.
Lawrence F. Gall, Entomology Division, Peabody Museum of Natural History, Yale University, New Haven, Connecticut 06520, USA, and David C. Hawks, Department of Entomology, University of California Riverside, Riverside, California 92521, USA
Received for publication 7 November 2001; revised and accepted 21 November 2001.