EDITOR'S FILE COPY US. Department of Agriculture Forest Service United States Pacific Northwest Forest and Range Experiment Station Budworms General Technical Report PNW-100 Program February 1980 Nomenclature of

Nearctic Con ifer - Feedi ng- Choris tone ura (Lep ido pt e ra: Tort ric idae): HISTORICAL REVIEW AND PRESENT STATUS Jerry A. Powell

This file was created by scanning the printed publication. Mis-scans identified by the software have been corrected; however, some errors may remain. ABSTRACT ACKNOWLEDGMENTS

There have been 18 species-group I thank USDA Forest Service names proposed for Nearctic researchers V. M. Carolin Jr. , G. E. -feeding in the Daterman, R. E. Stevens, and T. R. . Of these, houstonana Torgersen; and W. J. A. Volney, (Grote) and its synonym, retana University of California, Berkeley, for (Walsingham) , apply to Cupress- review of a draft of the manuscript. aceae-feeding populations now assigned Although not all my nomenclatural to Obraztsov & Powell. The decisions are necessarily endorsed by remaining species are Pinaceae-feeders, any one reviewer, their comments and mostly members of two series: (1) the criticisms were helpful in clarifying Fumiferana complex, associated with the discussion and in developing the and (subfamily conclusions. Field travel during Abietoideae) and considered to consist 1966-70 and study at the British Museum of 5 allopatric species (with three in 1971 were in part funded by National synonyms) ; and (2) the Lamber- Science Foundation grants GB-4014 and tiana complex, feeding on pines GB-6813X. USDA Forest Service (subfamily Pinoideae) and consisting of personnel allowed me to use facilities two allopatric species, the western C. in the Modoc and Shasta National lambertiana (Busck) with three Forests in 1970-74 and provided field subspecies and the eastern C. pinus collections from the 1974 pilot study Freeman with two subspecies. A third of the effects of Dylox. group, the Carnana complex, is believed to be associated with Douglas- J. A. DeBenedictis and D. L. Wagner, (Pseudotsuga). Its populations occur University of California, Berkeley, in California, with two named races, assisted with field survey and sympatric with elements of both the laboratory rearing work during 1978-79, Abietoideae- and Pinoideae-feeding funded in part by the CANUSA Spruce complexes. One named entity, Budworms Program-Wes t (FS-PNW-G-5 9) . spaldingiana Obraztsov, in Utah, is of The maps were prepared by J. T. unknown affinities. Sorensen, University of California, Be r keley. Keywords: Nomenclature (), Chor istoneura. AUTHOR

Jerry A. Powell is Professor, Division of Entomology and Parasitology, College of Natural Resources, University of California, Be r k e ley. Possibly no species of Moreover, a revised checklist of has been more intensively studied North American Lepidoptera, which has during the past quarter, century than been several years in preparation by a the spruce budworm in , team of specialists, is nearly yet the nomenclature of the completed--the first updated list in 40 group has years (Hodges et al., in press). continued to be confusing. Although Acceptance of the nomenclature in this some problems are traceable to pre- list by the U.S. Department of mature or careless taxonomic work, the Agriculture and the Entomological principal source of turmoil lies in the Society of America (and therefore the complexity of variation among E.S.A. common names list--Sutherland et populations. An array of inter- al. 1978) is anticipated. This should re lat ionships exis t s : sympatric provide a considerably stronger base populations exhibit varying degrees of for stabilization of names of all North re productive i solation, and allopatric American Lepidoptera. I have con- populations show geographical tributed the text for the , variations in size and color of adults, aided by reviews and contributions from larvae, and pupae and in preferences severa 1 contempora ry spec iali s t s, for host plants. There are both including R. W. Brown, R. W. Hodges, polymorphic and continuously variable and W. E. Miller. Table I lists valid features, and characters conven- and synonymous names ,applied to tionally used by taxonomists in conife r -f eed ing Choristoneura and defining species in moths are not Cudonigera as they will be' treated in sufficiently differentiated in this Hodges' (in press) checklist. group to provide reliable morphological indicators of genetic and behavioral In general, the checklist is not compatibility among populations. envisioned as a vehicle for revising taxonomic work, but there will be many An increase in research on the nomenclatural changes. Names formerly Nearctic conifer-feeding Choristoneura applied in many taxa have become ob- as a result of the Canada/U.S. Spruce solete as a result of taxonomic study Budworms Program, launched in 1977, has of Nearctic or other faunas, parti- broadened interest in and the need €or cularly the Palearctic. Authors have better communication about these been expected to incorporate . De spite considerable fund ing nomenclatural changes where firsthand and intricate organizational planning , taxonomic expertise enables generic disagreement exists concerning the reassignments and synonymies. number, level of taxonomic distinct- Therefore , many new combinations and ness, and geographical distributions of new synonymies will be published in the species and races, as well as the names checklist without explanations. In applied to them. Thus, an updated most instances, explanations would be summary of nomenclature in this species of interest only to specialists, but in group and agreement among researchers a few cases the importance of the about correct application of names seem insects in agricultural entomology or e ssenti al. other applied fields is such that

2 CHRONOLOGICAL HISTORY OF THE SPECIES GROUP NAMES

published interpretation is war- fumiferana Clemens, 1865, Proc. ranted. Because the Canada/U.S. Spruce Entomol. Soc. Phila. 5:129 (Tortrix) Budworms Program is in its early phases, the spruce budworm complex is The earliest name applied to any such a case. North American conifer-feeding Choristoneura is fumiferana, described from "Virginia." A lectotype was Table 1--Checklist of valid and designated by Darlington (1947) , who synonymous names used for North indicted there are no data associated American conifer-feeding Choristoneura with the specimen, as is true of and Cudonigera (synonyms underlined)1/ Clemens' material generally (see Miller (1973) for a review of the history and Choristoneura Lederer, 1859 status of Clemens' types and associated f umiferana (Clemens, 1865) catalog numbers). Thus, the accuracy nigridia (Robinson, 1869) or any restriction of the type locality retiniana (Walsingham, 1879) cannot be determined. Treated in lindseyana Obraztsov, 1962 Tortrix or Harmologa, a New Zealand viridis Freeman, 1967 genus, in early literature, the species occidentalis Freeman, 1967 later was placed in (= Cacoecia) biennis Freeman, 1967 until Freeman (1947) assigned orae Freeman, 1967 fumiferana to the Palearctic genus carnana (Barnes & Busck, 1920) Choristoneura. c. californica Powell, 1964 lambertiana (Busck, 1915) 1. ponderosana Obraztsov, 1962 nigridia Robinson, 1869, Trans. Am. 1. subretiniana Obraztsov, 1962 Entomol. Soc. 2:268 (Tortrix) pinus Freeman, 1953 p. maritima Freeman, 1967 Described from "Ohio, Pa., and spaldingiana Obraztsov, 1962 Mass., " this is the only synonymous Cudonigera Obraztsov & Powell, 1977 name proposed for the widespread houstonana (Grote, 187 3) spruce budworm of eastern Nearctic re tana (Walsingham, 1879) boreal forests. Darlington (1947) mentioned a cotype at the Academy of 1/After Powell -in Hodges et al. Natural Sciences, Philadelphia; but (in press). apparently no formal designation of a lectotype and restriction of the type locality has been made. Representative specimens of type material of nigridia and fumiferana were examined by Fernald in the 1870's (Darlington 1947, p. 86), who concluded the two are synonyms (Fernald 1881, p. 63).

3 houstonana Grote, 1873, Bull. Buffalo retiniana Walsingham, 1879, illus. . Soc. Nat. Sci. :15 (Tortrix) Lepid. Het. Br. Mus. 4:12 ()

The next oldest name used for a The earliest name applied to any conif e r-f eeding former 1y western budworm is retiniana considered a member of Choristoneura is Walsingham, described from one specimen houstonana Grote, which was based on a collected at Mount Shasta, California, specimen from "Texas." The type in 1871. Failure to recognize its specimen is believed to be no longer status by most subsequent taxonomists e'xtant. A synonym, retana Walsingham, precluded use of this name in is available, but the species is literature on budworms. Generally, it distinctive, and Grote' s description was listed in the genus Archips by seems unequivocal so that neither earlier worker s, but Freeman (1958) suppression of it as a nomen dubium nor transferred retiniana to Choristoneura, designation of a neotype seems suggesting that it might be related to necessary. This species was included the pine-feeding species. Freeman in Choristoneura (Freeman 1958, Powell illustrated a specimen from Mono 1964); but recently Cudonigera County, California, as representing Obraztsov and Powell was proposed for ret iniana. houstonana, primarily on the basis of morphological features of the adult Obraztsov. (1962) recognized that which differ markedly from all other Freeman's asssumption was based on a Choristoneura (Powell and Obraztsov misidentification of moths now called 1977). Cudonigera houstonana feeds subretiniana Obraztsov, a pine-feeder. exclusively on Junipe rus (Cupressaceae) Obraztsov illustrated structural (Powell 1964, Heinrichs and Thompson features of the holotype of retiniana 1968) ; all other Nearctic conifer- and presented photographs of specimens feeding Choristoneura are restricted to from Oregon and New Mexico which he Pinaceae (Stehr 1967). supposed to be conspecific. Powell (1964) applied the concept of fumiferana Clemens broadly, to include retana Walsingham, 1879, illus. Lepid. all fir- and spruce-feeding Het. Br. Mus. 4:13 (Lozotaenia) populations, and recognized retiniana as a synonym because pale specimens of This name was ,recognized by Grote Abies-feeding populations in Modoc (1881, p. 9) as synonymous with County (about 150 km--90 miles--east of houstonana, a conclusion accepted by Mount Shasta) resemble Walsingham' s subsequent workers. The type locality, description and illustration of cited as "Texas," is Bosque County, retiniana. Texas, according to data on the lectotype (Powell and Obraztsov 1977).

4 lambertiana Busck, 1915, Proc. Entomol. pinus Freeman, 1953, Can. Entomol. Soc. Wash. 17:86 (Tortrix) 85 :122 (Choristoneura)

The earliest name proposed for any This name was provided by Freeman for pine-feeding population in Nearctic pine-feeding populations of eastern Choristoneura is lambertiana Busck, Canada, following extensive studies by described from Ashland, in southern several researchers on differences in Oregon (Jackson County). The locality biology, phenology, immature stages, was originally 'given as "Oakland," parasites, and isolating mechanisms Oregon, but the type series is from between this and the sympatric Ashland according to the labels and Choristoneura fumiferana. The type Hopkins' data. Some of the labels are locality is Beausejour, Manitoba. printed by machine, and some are handwritten; the latter were evidently misread by Busck. The species was spaldingiana Obraztsov, 1962, Am. Mus. assigned to Choristoneura by Freeman Novit. 2101:6 (Choristoneura) (1958), who listed the species also from California and New Mexico without This species was described from specific data. The concept of Provo, Utah, on the basis/of a single lambert iana was broadened by Obraztsov male specimen. (1962) to include populations in Idaho, Montana, Wyoming, and New Mexico. -sub retiniana Obraztsov, 1962, Am. Mus. Novit. 2101:9 (Choristoneura) carnana Barnes and Busck, 1920, Contr. Nat. Hist. Lepid. N.A. 4(3):214 Using specimens from "Monachee," ( Tortrix) Tulare County, California, 8,000 feet (2 450 m), Obraztsov described This distinctive, rust- and white- subretiniana. Presumably this is the colored moth was described on the basis site now called Monache Meadows, of specimens labeled "Camp Baldy, San situated near the headwaters of the Bernard ino Mountain s, Calif orn i a." The south fork of the Kern River, west of type locality presumably is at or near Olancha Pass in the southern Sierra . the place now known as Mount Baldy Post Nevada.. This race was treated as a Office (designated as Camp Baldy on subspecies of lambertiana by Powell roadmaps as recently as 1947), which is (1964) on the basis of phenotypic situated at about 1 300-m (4,250-foot) similarity of adults in populations of elevation near the Los Angeles- the central and northern Sierra Nevada San Bernardino County line in the Sari and the pine-feeding population of Gabriel Mountains. There have been no Warner Mountains, Modoc County, and verified records from the San because of the allopatric occurrence of Bernardino Mountains. The species was these populations in relation to recognized as a Choristoneura related lambertiana s. str. and californica to fumiferana by Freeman (1958), and Powell west of the Sierran crest. subsequently carnana was treated as a subspecies of lambertiana (Powell 1964).

5 ponderosana Obraztsov, 1962, Am. Mus. from a series of Choristoneura Novit. 2101:14 (Choristoneura) “fumiferana” (= retiniana) and these matched specimens reared from Abies in In broadening the concept of the Warner Mountains during 1950-57 lambertiana, Obraztsov recognized (Powell 1964, p. 176). similarity of moths in Colorado to those of the northern Rockies which he treated as lambertiana, and he californica Powell, 1964, Univ. Calif. designated a series reared from Publ. Entomol. 32:179 (Choristoneura) ponderosa pine at Sugar Loaf, Boulder County, as ponderosana. This name was applied to populations in the North Coast Range (type locality: Anderson Springs, Lake lindseyana Obraztsov, 1962, Am. Mus. County), southern Cascade Range, and Novit . 210 1: 16 (Choristoneura) west slope of the Sierra Nevada. Although none had been reared, larvae A collection of Choristoneura was of these populations were assumed to be made by A. W. Lindsey in the Davis pine-feeding , and californica was Creek area of the Warner Mountains, associated with lambertiana as a Modoc County, California, in 1922. subspecies on the basis of the Included were nearly 100 specimens allopatry with lambertiana in the representing each of two populations strict sense to the north, subretiniana now known to be fir-feeding and to the east, and carnana in southern pine-feeding species and referred to as California. retiniana and subretiniana, respectively. A portion of the. polymorphic Ab ies-feed Fng population -biennis Freeman, 1967, Can. Entomol. there consists of individuals with pale 99 :4 51 (Choristoneura) tan forewings without dark markings, and this form was selected by Obraztsov This species was distinguished to name. He supposed this to be a primarily on the basis of the larger subspecies of lambertiana, despite the average size and darker coloration of fact that specimens phenotypically adults and the fact that 2 years are similar to subretiniana had been required to complete the life cycle. collected by Lindsey in the Warner The type locality is 8 miles (13 km) Mountains, and others had been reared south of Cherryville (in the Monashee by USDA Forest Service researchers from Mountains near Vernon) in southern lodgepole pine in 1952-53 (Powell 1964, , where specimens were p. 183). In treating fumiferana reared from spruce (Picea sp.) and fir broadly, I relegatkd lindseyana to () . synonymy, recognizing the tan form as a polymorph ph’ase in samples reared from Abies (Powell 1964, p. 175).

I pointed out that Obraztsov’s lindseyana types had been extracted

6 occidentalis Freeman, 1967, Can. Freeman (1967, p. 453) listed Entomol. 99 :451 (Choristoneura) lindseyana among several forms thought to be pine-feeders. Considered to be the most widespread, dominant conifer-feeding Choristoneura in the western Nearctic, occidentalis maritima Freeman, 1967, Can. Entomol. was described from Klickitat County, 99:455 (Choristoneura) Washing ton, reared from Pseudotsuga. Populations believed to be conspecific This name was proposed as a were recorded from New Mexico through subspecies of , to the northern Rocky Mountain States to include populations from Massachusetts British Columbia and in northern south to Kentucky. The type locality California (Freeman 1967). is Blain (Perry County) ,in the Alleghe ny Mount a ins of south central Pennsylvania. Known hosts include orae Freeman, 1967, Can. Entomol. pines of the subsections Australes and 99:452 (Choristoneura) Contortae.

This name was applied to a population at Kitimat in north coastal British Columbia. Choristoneura orae consists of small individuals which are phenotypically similar to fumiferana. The type series was reared from Abies and Picea. Differences in polymorph phases (Stehr 1964) and characteristic physiological data (Harvey 1967) appear to be the primary distinguishing features of the population. viridis Freeman, 1967, Can. Entomol. 99:452 (Choristoneura)

On the basis of collections from Bidwell Creek, Modoc County, Calif or nia, Freeman renamed the fir-feeding population of the Warner Mountains viridis, disregarding the nomenclatural priority of lindseyana for that .population. Despite biological information documenting the status of lindseyana as the pale form of the fir-feeding Choristoneura and the occurrence of a sympatric- pine-feeding species (Powell 1964) ,

7 CURRENT STATUS OF NAMED FORMS

~~ ~~ There are two series of populations samples collected on Abies among North American Pinaceae-feeding concolor in the Warner Mountains, Choristoneura, one feeding primarily on California.l/ These included all spruces and firs of the genera Picea, ,larvae thought to differ from the Abies, amd Pseudotsuga (subfamily typical green budworm in preliminary Abietoideae; Ferre 1952), and the other sorting, as well as random subsamples almost exclusively on Pinus (subfamily of green budworm from each collection. Pinoideae) . For convenience, these are Of 5,368 moths reared, about 3,200 (60 referred to here as the Fumiferana percent) were C. retiniana and only complex and Lambertiana complex, about 5 (<1 percent) C. lambertiana respectively. In each there are subretiniana, which sometimes causes parallel patterns of geographical visible defoliation of lodgepole pine distribution, with resultant widespread at the same localities (Pierce and Hall sympatric pairs, one Abietoideae- 1974). Stehr (1967) , who attributes feeding and one Pinoideae-feeding, in most such records to dispersal of young eastern and western portions of the larvae in autumn or spring, states' that Nearctic. there are no known records of oviposition on the reciprocal host There do not seem to be documented leding to successful establishment of instances of sympatric species within larvae. Probably some published either the Fumiferana complex or records or data on labels of the Lambertiana complex. There is a third "wrong" host are the result of. group of populations, however, here misidentification of larvae or reared called the Carnana complex, for ,which adults. In general, the ecological insufficient data are available to subdivision appears to be remarkably precisely define relationships. distinct Formerly, these were thought to be members of the Lambertiana complex, In the Fumiferana and Lambertiana based- on allopatric distributions in complexe s , the pa rallel situ at ion relation to known pine-feeding exists wherein each is represented in populations (Powell 1964). Recent the eastern half of the continent by a evidence, however , indicates that the widespread species that is relatively Carnana complex consists of populations constant in biological and phenotypic feeding on Pseudotsuga, which are characteristics, but in the western parapatric (Mount Shasta) or sympatric portion is more complicated. (west slope of Sierra Nevada, San Entomologists agree that Choristoneura Gabriel Mountains) with Abies-feeding fumiferana is a monotypic form populations of the Fumiferana complex. occurring from the McKenzie River near

Instances of larvae feeding on other than the normal Abietoideae or Pinoideae host are occasionally lPowe11, J. A., and V. Donahue. observed but usually only in mixed 1974. Monitoring the effects of Dylox stands of conifer genera. For example, on microlepidopterous larvae. USDA in 1974 we reared a large number of For. Serv., Coop. Agreement Final Rep., from late 17 p.

8 66 N, southeastward in a broad arc faunas which were isolated during through southern Canada and adjacent. glacial advances. parts of the United States, to the Maritime Provinces and Newfoundland. Choristoneura pinus is sympatric along Western Components of the the southern part of this range in the Fu m if erana Com p lex region, extending farther south than does fumiferana, in the Present taxonomic concepts consider northern Plains States, and in the this array to comprise four mainly Allegheny-Appalachian region, where allopatric species, the names of which populations are referred to as the have been applied through historical subspecies -C. pinus maritima. chance. The oldest name was given to the only representative available a West of the continental divide, century ago (Walsingham 1879) ; more populations of the Fumiferana and recent names were proposed for samples Lambertiana complexes are also of sporadically collected adults sympatric in many areas, but there is (Obraztsov 1962) or were applied to the considerably greater varability. Among populations which were best known western components of both the through laboratory studies a decade ago Abietoideae- and Pinoideae-feeding (Freeman 1967). groups, a great deal of polytypic (interpopulational) variation exists, Choristoneura retiniana (Wlsm.).-- not only in adult and larval I Clarification of the status of morphologies, but in degree of genetic polymorphism, physiological traits, and retiniana is pivotal to nomenclatural preferences for host plants. Part of assignments. Surveys were conducted in the complexity can be attributed to a northern California and southern Oregon relatively fragmentary status of in 1966-70, toward this goal.2/ Only sampling compared with eastern areas. sporadic rearing records were-obtained, but sufficient population samples of In general, however, the situation moths taken at lights were accumulated reflects real biological phenomena: to assess variation. I compared the intermingling of forest tree representative specimens with the species, small-scale patterning of pure holotype of retiniana at the British stands, and geographical complexity of Museum in 1971 and established its western forest regions (Eyre 1954, p. identify with samples from McBride 9; Stehr- 1967). The isolation or Springs on Mount Shasta and Donner semi-isolation .of populations by the Summit (Placer County), California islandlike distribution of mountain (reared from ). ranges and host tree types presumably Obraztsov's (1962) description of the has led to divergence and changes in mechanisms that isolate species. Probably the biological instability 2Powell, J. A. 1970. Taxonomic and cor responds generally to the more biological investigations on California recent, postglacial colonizing of these microlepidoptera. Univ. Calif. Agric. areas and the consequent reestablish- Exp. Stn. proj. 2063. Prog. Rep. 1970, ment of contact between floras and 3 P.

9 retiniana type specimen as "badly Gabriel Mountains and at lower denuded and damaged'' is in error. The elevations (below 1 700.m-- specimen is in good condition. 5,500 feet) in the Sierra Nevada, these retiniana-like Choristoneura are T. D. Eichlin of the California State sympatric with the monomorphic, Department of Food and Agriculture and Pseudosuqa-feeding C. carnana, and at I carried out further surveys. in the higher elevations, near the Sierran Modoc-Siskiyou area in 1974, partly in crest (e.g ., Lake Tahoe) , with Pinus- collaboration with the USDA Forest feeding C. lambert iana subre t iniana. Service, which provided samples of larvae from nine sites in Modoc County Northward, C. retiniana is replaced from a pilot control project of Dylox by occidentalis, either abruptly, in on the Modoc budworm (see footnote 1). northern Oregon as indicated by Freeman Larval collections from Abies on Mount (1967) and Stehr (1967) , or gradually Shasta produced moths confirming in a series of blend zone populations. similarities in phenotype, polymorphic V. M. Carolin (personal communication) variation, and the weak sclerotization states that a Choristoneura with green of larval and pupal integument (source larvae identical to retiniana of Modoc of the common name "green budworm") County, California, occurs in the King among populations there and in Modoc Mountain area near Burns, in east- County. Therefore, I believe the name central Oregon, whereas a fir-feeding ret iniana applies to Ab ies-f eeding form in south-central Oregon west of populations in the southern Cascade Lakeview is heterogeneous, showing a Range, at Mount Shasta, and in the gradation between retiniana and Warner Mountains. If, in the future, occidentalis morphs in both larvae and the Warner Mountains population is adults. Currently under investigation deemed sufficiently distinguishable to by W. J. A. Volney and W. E. Waters, war rant nomenclatural recognition, the University of California, Berkeley, name lindseyana Obraztsov, 1962, has populations in this region are critical priority over viridis Freeman, 1967. to a more thorough understahding of the systematic and biological relationships Southward, in the Sierra Nevada, of C. retiniana, C. carnana Tehachapi Range, and mountains of californica, and -C. occidentalis. southern California, there are populations of Choristoneura that Choristoneura occidentalis exhibit similar genetic polymorphism, (Freeman) .--As presently conceived , having a morph with pale tan, unmarked this name applied to all Abietoideae- forewings. Though the adult phenotype feeding populations from northern is much less variable than that of Oregon, Washing ton, southern British retiniana in Modoc County, these Columbia, and the Rocky Mountain States populations appear to be a counter- south to New Mexico and eastern Arizona part of retiniana. Their distribu- (fig. 1). Freeman (1967) also listed tion is entirely in association with California in the range of Abies, and larvae reared in 1979 from occidenta1is; and Stehr (1967) , in his -.Abies in El Dorado County were of the figure 1, showed a symbol for green, unmarked form. In the San occidentalis in the vicinity of Lake Tahoe, both without indication as to

10 Figure 1.--Distribution of Western United States members of the Abietoideae- feeding Fumiferana and Carnana complexes: Choristoneura occidentalis Freeman (closed circles), C. retiniana (Wlsm.) (open circles), C. carnana carnana (B. & Bsk.) (open triangles), and C. carnana californica Powell (closed trian- gles). Half-closed circles in southern Oregon and northern California indicate localities with possible hybrid populations. (Data from Stehr (1967) and specimens examined.) 11 the basis of the record. If Carnana Complex occidentalis and, retiniana were sympatric in northern California, it Choristoneura carnana (Barnes and would not be possible to Busck).--The typical form of this morphologically differentiate them species is known in the literature on except by degree of integumental from the type series and from one sclerotization in the larva and . specimen collected at Mount Lowe in t Probably the record for occidentalis in San Gabriel Mountains (Powell 1964, p California refers to a population 184). Recently it has been collected discussed here under C. carnana in the Mount Baldy area in associatio Califor nica. with Pseudotsuga macrocarpa. TO the north, along the west slope of the 'Over the broad range of occidentalis Sierra Nevada, there are populations there is considerable interpopulational exhibiting a similar , monomorphic variation in ecological and geographi- phenotype, but individuals have more cal features (such as host preferences extensive , darker rust-orange forewing and phenology), as well as in phenotype markings and darker hindwings. Sever the adults, particularly in of collections of the latter have been existence and frequencies of poly- made since my summary of the morphic phases. Thus, many populations distribution (Powell 1964: map 9) , an could be described in which the adults its occurrence appears to be restrict, differ from typical occidentalis of to elevations below 1700 m (5,500 eastern Washing ton. feet), co'rresponding to the range of Pseudotsuga menziesii. Larvae were Cho r i stoneura bienni s Freeman -- . collected and reared from that host i The 2-year life cycle form replaces C. El Dorado County in 1979. occidentalis in interior British Columbia and northward. Freeman (1967) Cho ristoneura carnana Californ ica and Stehr (1967) listed two Picea Powell. --The Coast Range populations species and Abies lasiocarpa as host northern California we re conside red plants, not specifying which are used along with carnana to be probable in various parts of the range-- pine-feeders- because of their allo- reported to include mountains of patric displacement of C. lambertiana western Alberta and southwestern Yukon and subretiniana (Powell 1964). Territory. Subsequently, however, C. carnana ' californica has been reared from Choristoneura Freeman.-- Apparently, there have been no further data on the geographical range of this species or its relationship with occidentalis to the south and biennis in the interior of British Columbia at the same latitude.

12 Pseudotsuga menziesii in Napa and Lake Western Components of the Counties, near the type locality. Larvae of these and the El Dorado Lambertiana Complex County populations are heavily marked, brick red, similar to certain forms Choristoneura lambertiana lambertiana described for C. occidentalis (Harvey (Busck) .--Other than the original and Stehr 1967) , contrasting markedly series, reared from sugar pine in the with the green Abies-feeding larvae of vicinity of Ashland, Oregon, the retiniana. The Choristoneura recorded typical race of this species is known as californica from lower elevations only from one collection, from Pinus around Mount Shasta (Powell 1964, p. lambertiana in adjacent Siskiyou 181) has been reared from Pseudotsuga County, California. Freeman (1958, in the Salmon River area of the p. 36) listed lambertiana from Siskiyou Mountains by Carolin, Volney, California and New Mexico and cited and Waters. Juniperus (Cupressaceae) as a food plant, without specific data--the last These records, together with the probably based on misidentified Pseudotsuga-associated distribution of specimens of Cudonigera houstonana. the Sierran west slope populations and Stehr (1967, p. 461) mentioned a form carnana carnana, indicate this complex with pale, washed out forewings within is primarily associated with populations of this species (e.g., Abietoideae, not Pinoideae as was Obraztsov 1962: figs. 7 and 8) but previously supposed. This is the first equated this pale form with known instance of two sympatric species ponderosana, populations of which using Abietoideae. C. carnana (in the consist entirely of a more extreme broad sense) occurs within a few washed out phenotype (Obraztsov 1962: kilometers of Abies-feeding retiniana figs. 30-31) . at Mount Shasta City (about 600 m-- 2,000 feet--lower in elevation) , and Northern Rocky Mountain moths are the two species together at the somewhat intermediate, exhibiting a same sites in mixed Pseudotsuga-Abies mixture of color pattern features of forests on the west slope of the Sierra typical lambertiana, subretiniana of Nevada (e.g., El Dorado County at eastern Oregon, and ponderosana in 1 300 m--4,250 feet) and in the San Colorado. The concept of lambertiana Gabriel Mountains (fig.1). was expanded to include these populations in Idaho, Montana, and Wyoming by Obraztsov (1962) and Stehr (1967), and some Montana and Wyoming populations were included in C. 1. ponderosana by Stevens et al. (1977). There appears to be clinal change in phenotype in a broad arc through the northern Rocky Mountain States (fig. 2).

13 Figure 2.--Distribution of western Nearctic members of the Pinoideae-feeding Lambertiana complex: Choristoneura lambertiana lambertiana (Bsk.) (closed circles), C. lambertiana subretiniana Obr. (open circles), C. lambertiana ponderosana Obr. (triangles). Half-closed circles indicate populations re- ferred to lambertiana s.str. by Obraztsov (1962) or Stehr (1967) and/or repre- sented by more recent material of uncertain relationships. (Additional data are from Stevens et al. (1977) and specimens examined.) Shaded areas on figures 1 and 2 depict major topographic features; areas above 5,000 feet (1 530 m) are shaded for all but the central coastal area of Cali- fornia (2,000 ft--600 m) and Great Basin region (7,OOO ft--2 150 m), matching fairly well the distribution of coniferous woodland.

14 Choristoneura lambert iana Uncertain Status subretiniana Obraztsov.--This race occurs in eastern Sierra Nevada Choristoneura spaldingiana lodgepole pine forests from Tulare Obraztsov.--This form is too poorly County north to Plumas County, in the known to associate with any of the Warner Mountains, and, in a slightly above defined complexes. Described different phenotype, in Harney County from Provo, Utah, at the western edge of eastern Oregon (fig. 2). It has of the Wasatch Range, data are too been reared from lodgepole pine in sketchy to indicate a probable host. A Nevada County (Stark and Borden 1965) typical phenotype of C. occidentalis and Warner Mountains (Powell 1964, p. occurs at higher elevations in the 183; Pierce and Hall 1974). Northward Wasatch Range, but no population of the from the California border , the Lambertiana complex is recorded there. phenotype loses the gray color, Adults which are phenotypically similar especially on the forewing interstices to spaldingiana have been collected in between the .rust-red banding, and the Schell Creek Range of eastern begins to resemble the form prevalent Nevada. The moths were taken in a pure in Idaho and Montana. Harvey and Stehr stand of Abies concolor, which suggests (1967), who followed Obraztsov in the possiblity that the name treating subretiniana as a species, spaldingiana represents a pallid form cited six larval collections, including of the Fumiferana complex in the three from pine (Pinus contorta, p. foothills or dry ranges of the Great jeffreyi, and p. washoensis) and one Basin. from Abies concolor, apparently all from the area of Lake Tahoe.

Chor istoneur a lambert i ana ponder osana Obraztsov.--This distinctive race, in which the forewings have no rust or reddish markings, has been recorded at numerous sites along the eastern flank of the in Colorado, in extreme southern Colorado near Durango, as well as in Montana, South Dakota, and Wyoming (Stevens et al. 1977). Pinus ponderosa is the host in each recorded instance except in Montana, where Pinus flexilis is used in the same area from which Stehr (1967: fig.1) assigned populations to , lambertiana s.str. (fig. 2).

15 CONCLUSIONS

It is important to understand that Entomologists should realize that names applied to both Abietoideae- and there is every reason to expect that a Pinoideae-feeding Choristoneura, given population of either the particularly in western North America, Fumiferana complex or Lambertiana are little more than signs posted at complex will differ in some respects irregular intervals on mosaics of from typical populations representing differing populations. Any attempt to available species names. For example, define species as reproductively many populations assigned to C. isolated entities of comparable rank or occidentalis, especially those of the genetic integrity on the basis of far north, Great Basin ranges, and present knowledge will be artificial. southern Rocky Mountains, vary Adults vary phenotypically between phenotypically to a degree comparable populations in average size, color, with that to which named "species" extent and kind of wing markings; differ from typical occ identalis and larvae and pupae vary in intensity of from one another. Thus, nomenclatural integumental color; populations differ recognition could have been given to in whether or not genetic polymorphism many additional populations had there is expressed and, if so, in the been physiological and genetic studies proportions of the polymorph phases. or were there a need for communication Physiological and behavioral traits about them. Taxonomists should agree also vary in response to ecological and that application of any further names geographical factors, so that host to this complex at our present state of preferences, seasonal timing of stages, knowledge would be absurd. cold-hardiness, and other features differ from place to place independently of the phenotype.

16 LITERATURE CITED

Barnes, W., and A. Busck. Grote, A. R. 1920. Notes and new species. Contri- 1881. "North American Tortricidae" , butions Nat. Hist. Lepid. N.A. by Lord Walsingham, London, 1879. 4 (3) :209 -278. Papilio l(1) :8-9. Darlington, E. P. Harvey, G. T. 1947. Notes on certain types of 1967. On coniferophagous species of Lepidoptera described by Bracken- Choristoneura (Lepidoptera: Tortri - ridge Clemens. Trans. Am. Entomol. cidae) in North America. V. Second Soc. 73 (2) :85-104. as a species character. Eyre, F. H. (Chairman) , W. A. Dayton, Can. Entomol. 99 (5) :486 -503. D. DenUyl, and others. Harvey, G. T., and G. W. Stehr. 1967. Forest cover types of North 1967. On coniferophagous species of America (exclusive of Mexico). Re- Choristoneura (Lepidoptera: Tortr i- port of the Committee on Forest cidae) in North America. III Types. 67 p. Soc. Am. For., Some characters of immature forms Washing ton, D. C. helpful in the identification of Fernald, C. H. species. Can. Entomol. 99 (5) : 1881. Oviposition in the Tortri- 4 64-4 81. cidae. Am. Nat. 15(1):63-66. Heinrichs, E. A., and H. E. Thompson. Ferre, Y. de. 1968. The biology of Choristoneura 1952. Les formes de jeunesse des- houstonana (Lepidoptera: Tortri Ab i e tacee s, Ontogenie- Phy logeni e. cidae) , a pest of Juniperus -In Etudes Dendrologiques. 1. Les species. Can. Entomol. 100 (7) : Gymno spe rme s-ge'ne'rali te's , 75 0 -76 3. vol. 3(1). Traveaux Lab. For. , Hodges, R. W. , D. R. Davis, W. D. Toulouse. 284 p. Duckwo rth , and other s. Freeman, T. N. [In press. ] Checklist of Lepidoptera 1947. A new generic assignment for of America north of Mexico (Micro- Archips fumiferana (Clemens) , the lepidoptera Families). E. W. Classey spruce budworm. Can. Entomol. Ltd. and Wedge Entomol. Found., 79 (1): 21. London. Freeman, T. N. Miller, W. E. 1958. The Archipinae of North America 1973. Clemens types of Olethreutinae (Lepidoptera: Tortricidae) . Can. (Lepidoptera, Tortricidae) . Trans. Entomol. 90. (suppl. 7) , 87 p. Am. Entomol. Soc. 99 (3) :205 -234. Freeman, T. N. Obraztsov, N. S. 1967. On coniferophagous species of 1962. New species and subspecies of Choristoneura (Lepidoptera: Tortri- North American , with cidae) in North America. I. Some notes on other species (Lepidop- new forms of Choristoneura allied tera, Tortricidae). Am. Mus. to C. fumiferana. Can. Entomol. Novit. 2101:1-26. 99 (5) :449-455.

17 Pierce, J. R., and R. C. Hall. Stehr, G. W. 1974. A biological evaluation of 1967. On coniferophagous species of sugar pine tortrix on the Modoc Choristoneura (Lepidoptera : Tortri- National Forest in 1974. 10 p. cidae) in North America. 11. Geo- USDA For. Serv., San Francisco. graphic distribution in accordance Powell, J. A. with forest regions. Can. Entomol. 1964. Biological and taxonomic 99 (5) :456-463. studies on tortricine moths, with Stevens, R. E., T. K. Borg, and T. 0. reference to the species in Cali- Thatcher. fornia. Univ. Calif. Publ. 1977. Notes on a pine-feeding Entomol. 32, 318 p. budworm, Choristoneura lambertiana Powell, J. A., and N. S. Obraztsov. ponderosana (Lepidoptera: Tortri- 1977. Cudonigera: A new genus for cidae), in the Colorado Rockies. moths formerly assigned to Chorist- Can. Entomol. 109(9): 1269-1274. oneura houstonana (Tortricidae) . Sutherland, W. S., W. .F. Barr, C. A. J. Lepid. Soc. 31(2) :119-123. Barlow, and others. Stark, R. W., and J. H. Borden. 1978. Common names of insects and re- 1965. Life-history of Choristoneura lated organisms. Entomol. Soc. lambert iana subretiniana Obraztsov Am. 132 p. rev. Spec. Publ. 78-1. ( Lepidopt era : To r t r icida e) College Park, Md. attacking lodgepole pine. Can. Walsingham,. Lord Thos. de Grey. Entomol. 97 (7) :684-690. 1879. North American Tortricidae. Stehr, G. W. Illus. Lepid. Heterocera Br. Mus. 1964. The determination of sex and 4, 84 p. Br. Mus., London. polymorphism in microevolution. Can. Entomol. 96 (1-2) :418-428.

GPO 990-268

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