VOLUME 57, NUMBER 2 153

FIGS. 1-4. Dichrorarnpha acuminatana. 1, Wings of male from Steuben, ME. 2, \\lings of male from Apetlon, Austria. 3, Genitalia of male from Steuben, ME. 4, Genitalia of male frum Apetlon, Austria. Further details are in the Specimens examined section of the text.

The species is widely distributed in western and genit. slide prep. WEM612012 (Fig. 4), foreWing central Europe (Razowski 1996). Two Palaearctic con­ length 5.5 mm. The four Palaearctic specimens are in geners, D. vancouverana McDunnough (=D. gue­ the U.S. National Museum of Natural History neeana Obraztsov) and D. petiverella (L.), were previ­ (USNM), Washington, D.C.; we thank J. W. Brown for ously reported in Maine (Roberts 1991), and loaning them. subsequent collecting there has revealed well estab­ lished populations of these species along the immedi­ LITERATURE CITED ate coastline wherever undisturbed stands of their na­ BENTINCK, G. A. & A. DIAKONOFF. 1968. De Nederlandse hladrollers (Tortricidae). Monogr. Nederlands. Entomo!' Ver. tive or naturalized foodplant Achillea millefolium L. No.3. 201 pp. (Asteraceae) occur. With captures of D. vancouverana KUZNETSOV, V 1. 1987. Family Tortricidae (tortricid ), pp. in the Pacific Northwest (Miller 1999), coastal distri­ 279-956. In Medvedev, G. S. (ed.), Keys to the of the European part of the USSR. U.S Dept. of Agr. & Nat!. Science bution patterns of the two hoi arctic congeners con­ Found. 991 pp. [translationl tinue to suggest they are immigrants, although the MILLER, W. E. 1999. A new synonymy in Dichrorampha that re­ possibility cannot be ruled out that they represent veals an overlooked immigrant record for North America (Tor­ trieidae). J. Lepid. Soc. 53:74-75. spotty relicts of circumpolar distributions. RAZOWSKI, J. 1996. Tortricidae, pp. 130-157. In Karsholt, O. & J. Specimens examined. 0, Steuben, ME (Fig. 1), Razowski (eds.), The of Europe: a distributional M. A. Roberts, 15/06/2001, genit. slide prep. checklist. Apollo Books, Stenstrup. 380 pp. ROBERTS, M. A. 1991. Two Pale arctic species of Dichrommpha dis­ MAR2027M (Fig. 3), forewing length 7.0 mm, in M. covered in Maine (Tortriddae). J. Lepid. Soc. 45:169- 171. A. Roberts collection, Steuben, ME; 0, Wangeroog, Ostfries. Inseln [ Germany], 07/09/1949, E. Jackh, MICHAEL A. ROBERTS, 367 Village Road, Steuben, genit. prep. on pin, forewing length 6.0 mm; 0, Kel­ Maine 04680, USA. Email: [email protected] AND heim, Obfrk. [ Germany], 03/08/1952, Jackh, genit. WILLIAM E. MILLER, Department of Entomology, Uni­ prep. on pin, foreWing length 6.0 mm; 0, Hannover, versity of Minnesota, St. Paul, Minnesota 55108, USA. Misb Moor [Germany], 29/05/1931, genit. slide prep. Email: [email protected] WEM 612011, forewing length 6.5 mm; 0, Apetlon, Received for publicatiun 8 April 2002; revised and accepted 11 No­ Burgenland [Austria] (Fig. 2), 11/09/1971, E. Jackh, vember 2002.

journal oftlte Lepidopterists' SOCiety 57(2). 2003, 153-156

HOST PLANT ASSOCIATIONS OF WESTERN SPECIES OF () WITH PARTICULAR REFERENCE TO THE APIACEOUS PLANTS Additional key words: host plants, Apiaceae, Asteraceae, polyphagy.

The Papaipema Smith (Noctuidae) is the seded only by the Holarctic genera Acronicta Ochs. (n largest noctuid genus endemic to North America has = 81 Nearctic species), Catocala Schrank (n = 110), long been a favorite among students of lepidopteran Lacinipolia McDunnough (n = 57), and Schinia Hub­ life histmy (e.g., Kwiat 1916, Hessel 1954). With 46 ner (n = 123 species in North America) (Hodges described species and at least 5 undescribed species of 1983). Papaipema currently includes 46 valid de­ which we are, Papaipema is the fifth most speciose scribed species, at least five undescribed species noctuid genus on this continent (Hodges 1983), super- (Quinter, in MS), and two valid subspecific entities, 154 J OURNAL OF THE LEPIDOPTERISTS' SOCIETY

Papaipema baptisiae baptisiae (Bird) and Papaipema notes of his son (archived at the American Museum of b. limata Bird (KL. Quinter, in Hodges 1983). Natural History), the late Junius Bird, the host plant Papaipema and its relatives form a putatively mono­ was described as a "large, Dill-like weed," suggesting an phyletic clade of endophagous plant borers in the apiaceous plant. The published association of P per­ (sensu Hodges 1983). Whereas most of the tincta with Lupinus polyphyllus Lindley (Fabaceae) species in this tribe are associated with monocotyledo­ (see Bird 1926) is curious because only two other Pa­ nous plants, species of Papaipema feed and specialize paipema species are associated with fabaceous plants: on members of between 22 and 25 plant families the western Papaipema angelica with Psoralea (Goldstein 1999). Though well studied, a number of macrostachya DC., and the eastern Papaipema bap­ questions remain eoncerning host plant associations in tisiae with Baptisia tinctoria (L.). this group, especially among the relatively few western Outside the complex, umbellifer­ species (Papaipema attains its highest regional diver­ feeding occurs in , a threatened sity in the eastern United States). In this paper, we species restricted to prairie wetlands where it speCial­ present life history data based on recent collecting and izes on , and in the P harrisi rearing efforts for species belonging to the Papaipema group, comprising P harrisi and P verona Smith). birdi (Dyar 1908) and Papaipema harrisi Grote Host records for P harrisi, whose distribution suggest species complexes, and summarize the known host as­ an association with Heracleum lanatum Michx. (Api­ sociations for the remaining western Papaipema aceae) along the Atlantic Coast and an association with species and those associated with Apiaceae regardless L. (Apiaceae) westward follow­ of geography. Our observations bear on the evolution ing the Great Lakes (Kwiat 1916, Hessel 1954, Jones of umbellifer-feeding in Lepidoptera, and Papaipema & Kimball 1943, Quinter unpublished data), Both of in particular and possibly the role of coumarin com­ these host species are apiaceous plants, In the North­ pounds in mediating the evolution of host association east, it is thought that P birdi and P harrisi segregate (e .g. , Berenbaum 1981, 1983). We also discuss collect­ themselves according to host plant, with P birdi con­ ing and rearing efforts on eastern umbel-feeding Pa­ fining itself to and P harrisi to Angel­ paipema species. All larvae encountered were reared ica atropurpurea (see Kwiat 1916, Hessel 1954), Pa­ on artificial diet, and adult specimens deposited at paipema verona, for which we do not report novel host AM"IH and FMNH. records, is a western species recorded primarily from Apiaceous host plant records for Papaipema species of the umbel genus Heracleum. species. A few species of Papaipema are known to Recent field collections. During 1995, we exam­ feed on apiaceous plants, and although at least three of ined several eastern USA sites for larvae of both P these (P birdi, P harrisii and P enjngii Bird) appear to birdi and P harrisi. Visits to wetlands in western Con­ be speCialists on Apiaceae, the others exhibit a broader necticut and Massachusetts with dense populations of range of apiaceous and non-apiaceous host use. The P Angelica atropurpurea yielded only two Papaipema bircli complex includes an eastern species (P birdi ) larvae (Papaipema harrisi has become deCidedly rare and two western species (P pertincta Dyar and P in­ in New England and is considered extirpated from sulidens [Bird] ), all of which are associated with the Massachusetts). However, visits to a calcareous sedge­ Apiaceae (= U mbelliferae). Papaipema birdi has been meadow complex in Otsego County, New York yielded considered oligophagous specialist on apiaceous more than two dozen Papaipema larvae from both An­ plants, its primary host being the water hemlock gelica atropurpurea and Cicuta maculata, All larvae Cicuta maculata L. Prior to the present study, other collected from C. maculata and A atropurpurea at the host records included Walt. (Apiaceae), upstate New York site proved to be P birdi. Although and "other umbellates" (Hessel 1954:60; treating P reports of "other umbellates" than Cicuta maculata birch as a synonym of P marginidens, of which there and Sium suave occur in the literature (e,g" Kwiat are no known host records), as well as several astera­ 1916), our collections appear to be the first documen­ ceous plants (Kwiat 1916). The two other species in tation of Angelica atropurpurea as a host for P birdi. th e P birdi complex, P pertincta and P insulidens, Although Kwiat (1916) reported non-apiaceous hosts each of which have been recorded from both apia­ for P birdi, it is conceivable that the taxonomic confu­ ceous and non-apiaceous plants, are western species sion that typically surrounds Papaipema has resulted in apparently separated by the Cascade Mountains, with erroneous reporting of hosts subsequent to that publi­ P pertincta to the west and P insulidens to the east. A cation, host of P insulidens was described by Bird (1921, Our findings in the northwestern United States ex­ 1931) as a species of Senecio (Asteraceae). In the field tended the known host ranges of P pertincta and P in- VOLUME 57, NUMBER 2 155

TABLE 1. Collecting information and host associations of western PapaipellUl species discovered during this study.

Species Locality Life stage Host plant Dates P. pertincta Oregon: Tillamook Co.: Rt. 101, 4 larvae Heracleum maximum 8 July 1995 1 mi S. of Wheeler Oregon: Clatsop Co.: Rt. 101, 9 larvae, 1 pupa Cicuta douglasii 8 July 1995 8 mi. S. of Astoria (at jet. Rts. 101 & 30) Oregon Tillamook Co.: Rt. 101, 8 larvae Senecio vulgariS 9-10 July 1995 1-2 mi. N. of Manzanita 8 larvae Heracleum maximum 4 larvae Ligusticum apifolium 1 larva Daueus sp. 8 larvae Cirsium sp. 3 larvae ErechWes minima Oregon: Lincoln Co: , E. Devil's 1 larva Cicuta douglasii 11 July 1995 Lake Rd. , 0.7 mi E. of Jet. Rt. 101 Oregon: Tillamook Co.: Rt. 101, 6 larvae, 1 pupa Heracleum maximum 11 July 1995 12 S. of Jet. Rt. 22 Oregon: Lincoln Co: Three Rocks 1 larva Heracleum llUlximum 1,3 July 1995 Rd. , l..'5miW Rt.10l l' sauzalitae Oregon: Lincoln Co: Three Hocks Ilmva Cirsium sp. 13 July 1995 Rd. , 1.5 mi W. Rt. 101 P. insnlidens Washington: Whitman Co: Steptoe 13 larvae Heracleum maximum 15 July 1995 Butte, el. 2500'-3000' sulidens. These represent the least well-known Pa­ nius Bird indicating large apiaceous host plants in paipema species for which published host records ex­ Whitman County, Washington. Near localities visited ist; their dose resemblance to P birdi as well as the in­ by Junius Bird, several stands of Conium maculaturn formal description by Junius Bird of a "large, dill-like" L. (Apiaceae), which matched his description and host for P insulidens suggested that apiaceous plants sketch, were checked without success. Thirteen larvae might fall within the host spectra of one or both of of P insulidens were discovered and reared from Her­ these two western species in the birdi complex. An ad­ acleum maximum Bartr. (Apiaceae) at Steptoe Butte at ditional southwestern species, Papaipema angelica an elevation of 2500-3000'. Although we could not ver­ Smith, 1899 has remained uncollected in recent ifY many published host associations of the two western decades despite our knowledge of its life history and members of the Papaipema birdi complex, we did take host affiliation (Bird 1931). Although several dozen them on other hosts, apiaceous and otherwise. One specimens of P pertincta exist in the Oregon State possibility is that the dill-like host plant of P insulidens University collection (which now includes the referred to hy Junius Bird was not apiaceous at all , but private collection of the late Elmer Griepentrog), we the introduced tansy ragwort Senecio vulgaris, one of have been unable to verify the association of P pert­ the host plants from which we reared P pertincta. incta with any species of Lupinus. We thoroughly ex­ The reported host associations of the western P amined the botanical holdings at OSU to identify sites sauzalitae (Grote) are atypically diverse for Pa­ likely to support strong populations of various western paiperna , and include memhers of the asteraceous lupines, but we found no Papaipema at any of these, plant genera Arctium, Cirsium, and Cynara as well as and apparently no western collectors have observed or Castilleja (Scrophulariaceae), and Rumex (Polygo­ reared P pertincta from Lupinus since Bird's (1926) naceae) (Crumb 1956). Peter McEvoy (pers. com. ) of second-hand account of the association. However, sev­ Oregon State University reports an association of eral dozen P pertincta were reared from a variety of P sauzalitae with Senecio (Asteraceae) as well. Our plants, mostly apiaceous, at six sites in Tillamook, Clat­ collecting efforts yielded but a Single specimen, from sop, and Lincoln counties (Table 1). Like its eastern the exotic Cirsium vulgariS. However, California ma­ relative P birdi, P pertincta appears to feed primarily terial at the Essig Museum includes specimens from In­ in apiaceous plants; but unlike its eastern associate, P verness Park (Marin Co.) where larvae were observed in pertincta also feeds in non-apiaceous plants. Heracleurn maximum, Artemisia douglasiana Bess. in In three weeks of field work in eastern Washington Hook. (As teraceae), and Ribes sp. (Grossulariaceae), and Idaho, we failed to collect P insl1lidens from its suggesting that P sauzalitae may be polyphagous (J. recorded host, Senecio hydrophilus Nutt. We exam­ Powell pers. com.). If this is the case, the member ined sites suggested by the literature, museum labels, species of each of the primary umbellifer-feeding Pa­ and the hand-written field notes and sketches of Ju- paipema species groups (the han'isi-verona-sauzalitae 156 JOURNA L OF THE LEPIDOPTERISTS' SOCIETY complex and the birdi-pertincta-insulidens complex), ---. 1983. Coumarins and caterpillars: a case for coevolution. have broadened their host usage to include both com­ Evolution 37:163-179. BIRD, H. 1921. New species and life histories in Papaipema Sm. posites and umbels on the west coast. (Lepidoptera). No. 20 Can. Entomol. 53:79-81. The association of P pertincta with apiaceous and - --. 1926. New life histories and notes in Papaipema (Lepi­ non-apiaceous plants is noteworthy for two reasons. doptera). No. 24. Can. Entomo!' 58:249-284. ---. 1931. The Papaipema species of the Pacific Coast (Lepi­ First, this species parallels P insulidens in having a doptera). Can. Entomol. 63:183- 187. wider range of non-apiaceous recorded hosts than ex­ CRUMB, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. pected, given the apparently tighter associations of Agric. Tech. Bull., no. 1135. U.S. Government Printing Office, Washington, D.C. 356 pp. their eastern relatives (P birdi and P sauzalitae, re­ GOLDSTEIN , P. Z. 1999. Molecular systematiCS and the macroevolu­ spectively) with umbels. Second, based on available tion of host plant use in the endophagous genus Papaipema Smith 1899. Ph.D. Dissertation, University of Connecticut. DNA sequence data, P pertincta is nearly indistin­ H ESSEL, S. A. 1954. A guide to collecting the plant-bOring larvae of guishable from P birdi (Goldstein 1999). We can not, the genus Papaipema (Noctuidae). Lepid. News 8:57-63. therefore, rule out the possibility of P pertincta's feed­ HODGES, R. W. et al. (eds.). 1983. Check list of the Lepidoptera of ing facultatively on Lupinus, though we were unable America north of Mexico. London, E. W. Classey Limited and The Wedge Entomological Research Foundation. 284 pp. to recover larvae from any fabaceous plants and it is JO NES, F. M. & C. P. KIMBALL. 1943. The Lepidoptera of Nan­ clear that the species is not thus restricted. tucket and Martha's Vineyard Islands, Massachusetts. Publica­ Although both facultative and obligate association tions of the Nantucket Maria Mitchell Association IV. Nan­ tu cket. 217 pp. with asteraceous plants is common among Papaipema KWIAT, A. [=A. K. WYATT). 1916. Collecting PapaiperrUle (Lep.) species, umbel-feeding is less common. The parallel Entomol. News 27:228-234. variation in host breadth among umbel-feeding Pa­ QUI NTER, E. L. 1983. Papaipema , pp. 138-139 (in part). In Hodges, R. W. et aI. (eds.), Check li st of the Lepidoptera of paipema species is thus noteworthy, and suggests a America north of Mexico. London, E. W. Classey Limited and profitable line of inquiry for further work on host use The Wedge Entomological Research Foundation. specialization in this group. PAUL Z. GOLDSTEIN, Division of Insects, Field Mu­ We thank P. McEvoy, E. Griepentrog, and J. Lattin for access to seum of Natural History, 1400 S. Lake Shore Drive, specimen material and assistance in locating collecting sites. J. Pow­ ell loaned material from the Essig Moseum and shared his observa­ Chicago, Illinois 60605, USA and Committee on Evo­ tions based on collecting and rearing efforts. We thank Kenneth and lutionary Biology, University of Chicago, Culver 401 , Elaine Bernard for lodging and permission to collect in their farm in 1025 East 57th Street, Chicago, Illinois 60637, USA Otsego Co., NY. This work was supported by an NSF doctoral dis­ sertation improvement grant, an EPA graduate fellowship, and an AND ERIC L. QUINTER, Division of Invertebrate Zool­ AMNH Theodore Roosevelt Memorial Fund travel grant to PZG. ogy, American Museum of Natural History, Central Park West @ 79th, New York , New York 10024, USA LITERATURE CITED BERENBAUM, M. 1981. Patterns of furanocoumalin distribution an d Received f or publication 25 August 2002; revised and accepted 20 i:lsect herbi VOry in the Umbelliferae: plant chemistry and com­ December 2002. munity structure. Ecology 62:1254-1266.