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Journal of The Lepidopterists' Society

Volume 44                                     1990                                     Number 3

Journal of the Lepidopterists' Society 44(3), 1990, 113-142

BODY SIZE AND DIET QUALITY IN THE GENUS CYDIA (TORTRICIDAE)

William E. Miller

Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108

ABSTRACT. I examine forewing length, a body-size index, relative to three quality classes of larval diet for more than 75 Cydia species. Quality of diet refers to protein concentration in the part of the food plant consumed. Mean crude protein percentages are near 25 in the high class, near 12 in the medium, and near 6 in the low. All data are from published sources. Forewings range in length from 4.0 to 10.5 mm among study taxa, and are longest in the high food-quality class, intermediate in the medium, and shortest in the low. The high food-quality class consists entirely of seed predators whose body sizes correlate positively with food-plant seed sizes. Medium and low food-quality classes consist mostly of nonseed feeders. Results imply that as Cydia colonize new food plants and plant parts of differing diet quality, body sizes evolve to those for which larvae can obtain sufficient nourishment. This interpretation withstands cladistic testing against an independent Cydia phylogeny.

Additional key words: Olethreutinae, evolution, cladistics, seed predation, food plants.

For organisms generally, body size is thought to be a quantitative adaptive trait (Bonner 1965, Calder 1984, Roff 1981). Much interest in body size derives from consequences of allometric relations among body components (Peters 1983). Body-size physiology and ecology are more tractable and better understood than body-size evolution, one facet of which is genesis of body-size diversity. Body size usually varies somewhat within species, but it does so around a genetically controlled norm. The norm represents an adaptive and fitness compromise in a given environment; analysis of norms and environments can yield evolutionary insights (Fisher 1930, Ridley 1983, Williams 1966).

Lepidopteran body-size diversity has just begun to be studied. Dietary factors are natural choices for independent variables: food quantity and quality are potent determinants. Among examined lepidopteran families, individual and population biomass correlate with density, size, and diet quality of food plants (Mattson 1977, 1980, Mattson & Scriber

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1987, Niemela et al. 1981). Within examined lepidopteran genera, subfamilies, and families, body size correlates also with breadth of diet (Niemela et al. 1981, Wasserman & Mitter 1978). Within lepidopteran families that mine leaves of Quercus (Fagaceae), body size correlates with leaf persistence, smaller-bodied species occurring on deciduous species (Opler 1978). If body size is viewed as an effect, then in evolutionary time it would seem to shift positively with diet breadth, food-plant size, and leaf persistence, and negatively with diet quality, although diet quality and plant size themselves correlate positively. Information is still too scant to yield well integrated generalizations about the genesis of lepidopteran body size diversity.

In this paper I examine body-size diversity relative to quality of larval diet in a large, worldwide sample of Cydia species (Tortricidae, Olethreutinae), defining food quality as protein concentration and testing analytical results cladistically.

For species of the subfamily Olethreutinae, a commonly available measure of body size is forewing length (L), values of which range from 4 to 20 mm (Miller 1987). Single genera cover large portions of this range. Cydia occupies more than half of it. Forewing length closely estimates dry body weight (W) of olethreutines by a power function summarized as W = 0.0085L3 (Miller 1977). This function reveals that small differences in forewing length denote larger differences in body mass. The low standard error of estimate of this function justifies using forewing length as an index of body size.

Cydia in the strict sense consists of ca. 250 species. Many species are important economically, a fact responsible for much information about the genus. The most famous species is the codling moth, C. pomonella (L.). The generic name is in flux; I follow Brown (1979) in using Cydia, but others argue for using Laspeyresia (Kuznetsov & Kerzhner 1984).

Materials and Methods

I devise three larval food-quality classes based on concentration of crude protein in the part of the food plant eaten, place each Cydia in an appropriate food-quality class, analyze inter- and intraclass differences in forewing length, and compare results with an existing phy-logeny.

I include only taxa conforming to the strict Cydia concepts of Ob-raztsov (1959) and Danilevsky and Kuznetsov (1968). Using broader generic concepts, some authors refer species to Cydia that are more strictly referable to Grapholita, Pammene, and other genera. Strict interpretation focuses and simplifies the study by limiting genetic heterogeneity to that of Cydia monophyly. In classification and nomenclature of Cydia subgenera, sections, species, and subspecies, I follow

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Danilevsky and Kuznetsov (1968) for taxa treated by them; for other taxa I apply their classification as far as I can do so confidently.

Wing measure of Cydia species is usually published as a range. I therefore use midrange, a statistic approximating the mean. Wingspan (S), the maximum distance between tips of spread wings, more often appears in the literature than forewing length (L). I convert span to length by the proportionality constant L = 0.458S (Miller 1977). All midranges are from published sources, mostly detailed faunal works (Appendix). Where forewing-length sample sizes are small, and a best source unavailable, I combine data from several sources. When mid-range could not be based on at least two individuals, I excluded the taxon. Using standard approximate methods, I estimate standard deviation (SD) of the midrange for all taxa where explicit sample sizes are nine or more individuals. I work with midranges expressed to three decimal places, but record them here to only one place (Appendix and elsewhere).

Food-plant parts eaten by Cydia larvae differ in food quality. Protein is a major component of food quality (Mattson 1980, Mattson & Scriber 1987, Scriber & Slansky 1981). "Crude protein", a standard food-science term, refers to the mathematical product of Kjeldahl nitrogen percentage and a multiplier, usually 6.25 (Crisan & Sands 1978, Williams 1984, and others). Crude protein percentage of food-plant parts used by Cydia forms a continuum. Below, I divide this continuum for study purposes into three food-quality classes: high, medium, and low. The letter n (or N) denotes number of analyses in this section of the text, number of observations in later sections.

High (mean near 25%)—seeds of Leguminosae; Pinus, Picea, Abies (Pinaceae); Malus and Pyrus (Rosaceae). Nutritional superiority of seeds for seed eaters over other plant parts is evident in the survey by Mattson (1980). It may be further documented with Leguminosae, one of the two plant families most used by Cydia: in forage species (6n), mean percentage crude protein is 24 in seeds, 16 in foliage, and 7 in pod husks (Skerman 1977). In other surveys, percentage crude protein in seeds ranges from 4 to 62 in Leguminosae (>1000n), 6 to 38 in Pinus and Picea (2In), and 18 to 49 in Malus and Pyrus (6n) (Barclay & Earle 1974, Dickmann & Kozlowski 1969, Earle & Jones 1962, Haut 1938, Jones & Earle 1966, Katsuta & Satoo 1964, McCarthy & Matthews 1984, NAS 1979, Pulliainen & Lajunen 1984, Rader-Roitzsch 1957, Short & Epps 1976, Skerman 1977, Winton & Winton 1932, 1935, Yoon et al. 1983).

Medium (mean near 12%)—inner bark, fungus-infected woody parts, photosynthesizing bark, foliage, flowers; seeds of Acer (Aceraceae), Cor-ylus (Betulaceae), Cryptomeria (Taxodiaceae), and Quercus. Fungi are

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equally as proteinaceous as the seeds comprising the high food-quality class (Crisan & Sands 1978, Mattson 1980), but fungi only supplement Cydia underbark diets. Photosynthesizing bark occurs in the food-plant genus Populus (Salicaceae) (Shepard 1975). Percentage crude protein in seeds is 7 to 32 for Acer (1 In), 4 to 32 for Quercus (2In), and 11 to 26 for Corylus (5n) and Cryptomeria (In) (Anderson & Kulp 1921, Barclay & Earle 1974, Earle & Jones 1962, Jones & Earle 1966, McCarthy & Meredith 1988, Schmidt-Hebbel & Pennacciotti M. 1979, Short & Epps 1976, Wainio 1941, Winton & Winton 1932).

Low (mean near 6%)—habitations of other insects, fruits or other seed-containing parts, sapwood, outer bark; seeds of Araucaria (Arau-cariaceae), Castanea, Fagus (Fagaceae), and Palmae. Wood and seed-containing parts are among the plant parts lowest in protein (Mattson 1980, Skerman 1977). Percentage crude protein in seeds is 4 to 16 in Araucaria (2n), Castanea (8n), Fagus (3n), and Palmae (20n) (Barclay & Earle 1974, Cardemil & Reinero 1982, Earle & Jones 1962, Jones & Earle 1966, McCarthy & Meredith 1988, Schmidt-Hebbel & Pennacciotti M. 1979, Wainio 1941, Winton & Winton 1932).

In intraclass analyses, I examine Cydia forewing length relative to food-plant seed size measured as weight of one seed. Mean seed weights for a given plant species do not vary much (Harper et al. 1970). I take most seed weights from published sources (Appendix, Table 2), using sample means and midranges in that order of preference. Where reported weights are few, I combine data from several sources if available. For seeds also consumed by man, I use seed weights of wild-type food plants if known. For other domesticated seeds, I use the smallest values reported. Because such plants have long been cultivated or bred for large seeds (Smartt 1980), the lowest weights are probably nearer to wild-type values. For those few species for which only seed dimensions are available, I estimate weights from one or more relatives of known seed weight and similar seed dimensions. Where one main food-plant species is not apparent, I use the mean of available seed weights for the appropriate number of food-plant species. In botanical nomenclature, I give only original authors (Appendix), following Schopmeyer (1974) and Krussmann (1978) in that order for food plants treated by them, and following source authors for others. I work with seed weights expressed to four significant figures, but report them here to three figures (Appendix, Table 2).

I use both nonparametric and parametric statistics because sample sizes are sometimes unequal, and distributions are sometimes more important than parametric values. Nonparametric methods treat data by ranks rather than by values. Hence differences between means may actually refer to differences between underlying rank distributions.

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Because forewing-length equations involve dimensions, I use nonlinear power functions for them (Peters 1983) solved by ordinary least squares.

For one-third of Cydia taxa, the part of the food plant eaten is known, thus enabling estimation of the quality of larval diet. This fraction of the genus comprises the study sample. There are 82 observations in all, one each for 78 species, 2 other than nominotypical subspecies, and 2 additional populations that behave like separate taxa in using food plants different from those of allopatric sister populations. Observations are grouped into high, medium, and low food-quality classes within which they appear alphabetically by Cydia species-level taxon in the Appendix. The 34 taxa and populations in the high food-quality class, all of which are seed predators, are from six continents; the 34 in the medium class feed mostly on nonseed parts, and are from three continents; and the 14 in the low class feed mostly on nonseed parts, and are from five continents. Corresponding numbers of food-plant families are 3, 12, and 9, respectively. Food-plant families and percentages of Cydia study species using them are, for the high class: Pinaceae—47%, Legumi-nosae—47%, Rosaceae—6%; for the medium class: Pinaceae—35%, Fa-gaceae—16%, Aceraceae—10%, Leguminosae—9%, Salicaceae—9%, others—21%; for the low class: Pinaceae—21%, Salicaceae—21%, Fa-gaceae—14%, others—44%.

I provisionally include five Cydia species whose strict generic affinities are unconfirmed: araucariae (Pastrana), palmetum (Heinrich), staphiditis (Meyrick), stirpicola (Meyrick), and tonosticha (Meyrick). Some species whose strict generic affinities require exclusion are Ful-crifera torostoma (Clarke), F. tricentra (Meyrick), Grapholita deshaisi-ana (Lucas), Leguminivora glycinivorella (Matsumura), L. ptychora (Meyrick), Matsumuraeses critica (Meyrick), and M. fabivora (Meyrick).

Results

Forewing lengths range from 4.0 to 10.5 mm (Appendix, Table 1). The main assumption in this study is that body-size diversity really exists in this sample range. If such diversity is present, variance (SD2) in forewing length among taxa should exceed that within taxa; if it is absent, the two variances should not differ. Comparison shows that variance among taxa (1.611, 82n) exceeds weighted average variance within taxa (0.156, 38n) (1.611/0.156 = 10.3, P < 0.001, variance ratio test), thus confirming the presence of diversity.

Body-size and Food Quality

Forewings are longest in the high food-quality class (6.8 mm), intermediate in the medium class (6.4 mm), and shortest in the low class

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Table 1. Body size of Cydia taxa and populations relative to quality of larval diet.

Forewing-length midrange (F) (mm)

--------------------------------------------------------------------------- Estimated mean

Food-quality class                     N                          Mean ± SDa                             Range                 dry wt. (W) (mg)b

High                        34                   6.8 ± 1.4                   4.0-10.5                   2.6

Medium                  34                   6.4 ± 0.9                   4.8-8.2                     2.2

Low                         14                   5.8 ± 1.3                   4.4-8.5                     1.7

a Means differ statistically (F-test, P < 0.05); their rank distributions differ in all possible comparisons (Kruskal-Wallis and Mann-Whitney tests, P < 0.001). b W = 0.0085 F3.

(5.8 mm) (Table 1). Estimates of body weight (Table 1) differ more markedly among the classes because forewing length is being multiplied by a power function. On average, members of the high class are 1.16 times heavier than those of the medium class, and 1.57 times heavier than those of the low; members of the medium class are 1.35 times heavier than those of the low.

Members of the high food-quality class have the greatest range and variability in forewing length (Table 1). Further scrutiny and analysis of this group reveals a major source of intraclass body-size variation. In this class, forewing length correlates with food-plant seed size; the larger the seeds, the longer the fore wings in both the pinaceous and leguminous subsets (Fig. 1). In a control analysis of nonseed-feeders from the medium and low food-quality classes, no such correlation appears between forewing length and food-plant seed size (Fig. 2: line A).

Size is only part of the seed factor accounting for body-size variation in the high class. Food-plant seed weight in this class traverses three orders of magnitude; the smaller the seed, the greater the number required to nourish a larva, the number increasing from 2 to more than 30 in taxa for which such data are available (Fig. 3, Table 2). Thus the product of seed size and number required underlies the correlation of forewing length and seed size shown in Fig. 1.

In the medium and low food-quality classes, range and variability of forewing length are narrower than in the high class (Table 1), and sources of intraclass body-size variation are less evident. In pooled medium and low classes, mean forewing length of seed predators (7.2 mm) is greater than that of nonseed feeders (5.9 mm) (P < 0.001, Student t- and Mann-Whitney tests). Forewing length of seed predators in these classes also correlates with seed size (Fig. 2: line B). Secondary plant chemistry could be a factor in intraclass variation in the medium and low food-quality classes because a greater diversity of food plants is involved: 12 and 9 food-plant families, respectively, compared with 3 in the high class.

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Forewing length (mm) (F)

12-

10

6H

.001

F=10.1±0.08S°-11±0-020 r=0.71 (P<0.01

.01

Seed wt. (g) (S)

Fig. 1. Cydia body size as related to food-plant seed size in the high food-quality class. Each point represents a Cydia taxon or population enumerated in the Appendix. Solid triangles depict the pinaceous subset; open squares, the leguminous subset; open circles, others. The symbol ± denotes standard error.

Table 2. Number of seeds eaten or destroyed per Cydia larva in the high food-quality class. (Ditto is abbreviated do.)

a Brackets denote weights absent or different in the Appendix. Bracketed weights for Pinus from Krugman & Jenkinson

(1974).

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Foravving length (mm) (F)

.0001               .001                 .01                   .1                     1                     10                  100

Seed wt. (g) (S)

Fig. 2. Cydia body size as related to food-plant seed size in pooled medium and low food-quality classes. Each point represents a Cydia taxon or population enumerated in the Appendix. Line A (closed circles) depicts the nonseed-feeding subset; line B (open circles), the seed-predator subset. The symbol ± denotes standard error.

Cydia taxa with one recorded food plant and those with more than one are analogous to the specialist and generalist diet-breadth classes of Niemela et al. (1981) and Wasserman and Mitter (1978). For "specialist" Cydia taxa, mean forewing length is 6.4 mm (37n), and for "generalist" taxa, 6.6 mm (42n). The difference, 0.2 mm, is statistically significant nonparametrically (P < 0.001, Mann-Whitney test), but not parametrically (Student t-test, P > 0.40). The first result matches findings of the above authors. When the high food-quality class is examined separately, however, results are reversed: mean forewing lengths are 6.9 mm (17n) for specialists and 6.6 mm (16n) for generalists. This reversal suggests that forewing length correlates more strongly with seed size than with diet breadth.

Cladistic Test of Results

Statistical analyses like the foregoing must be interpreted cautiously. Related taxa may simply inherit a given trait from a common ancestor rather than evolve it independently. If this happens, the assumption of statistical independence of observations is violated (Felsenstein 1985, Ridley 1983). It therefore seems necessary to test the assumption of independence, which is equivalent to the assumption that Cydia body-

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No. seeds/larva (E)

30 H

20 H

101

E=1.4±0.32S-°-43±008 r=-0.86(P<0.01)

.001

■ '■ I

.01

.1

1

Seed wt. (g) (S)

Fig. 3. Number of seeds eaten or destroyed per larva as related to food-plant seed size in the high food-quality class. Each point represents a Cydia taxon or population enumerated in Table 2 and the Appendix. The symbol ± denotes standard error.

size diversity represents many rather than a few evolutionary events. Below, I follow to the extent possible the test methodology advocated by Coddington (1988). I map food-quality classes indicated by the analysis on the Cydia phylogeny of Danilevsky and Kuznetsov (1968), then check for congruence. Congruence implies inheritance; lack of congruence, independence.

Danilevsky and Kuznetsov recognize three Cydia subgenera: Cydia, Kenneliola, and Endopisa. Their phylogeny to subgenera has the first two as sister taxa, with Endopisa in an outgroup position. Within each subgenus, Danilevsky and Kuznetsov further define a number of sections or species-groups, each of which represents a likely monophyletic lineage. The phylogeny to species is not resolved, but in three sections, four pairs of sister species are evident among study taxa.

Assignments to subgenus are made for 66 species in the study, 50 by Danilevsky and Kuznetsov, 16 by me. Food-quality class of each of these mapped on the phylogeny to subgenera produces the distribution in Table 3. All food-quality classes and associated body sizes occur among all subgenera, although most Endopisa species (12/15 = 0.80) are in the high class, and most Kenneliola species (13/14 = 0.93) are not. Cydia outgroup genera such as Grapholita also contain species

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Table 3. Distribution of 66 Cydia species by food-quality class and subgenus.

assignable to high, medium, and low food-quality classes (Danilevsky & Kuznetsov 1968). Therefore, within Cydia outgroups as well as subgenera there is evidence that food-quality classes and associated body sizes evolved independently of subgeneric lineage.

Within subgenera, at the section or species-group level, the positions of 50 study species are available, almost entirely assigned by Danilevsky and Kuznetsov (1968). Two or more food-quality classes and associated body sizes occur in 7 of 15 sections overall, and in all 5 sections containing three or more species (Table 4). The detailed distribution of food-quality classes among sections shows 9 within-section shifts in food-quality class and associated body size out of 18 possible ones (Table 4). Thus evolutionary shifts in body size are likely to have occurred within sections.

Table 4. Distribution of 50 Cydia species by food-quality class and section, and inferred numbers of evolutionary shifts in food-quality class within sections.

No. species in class                                       Minimal no. shifts

Section                                 High                Medium                 Low                   Possible              Evident

Subgenus Cydia pactolanae strobilellae pomonellae illutanae servillanae duplicanae cosmophoranae

Subgenus Kenneliola splendanae maackianae trasias exquisitanae

Subgenus Endopisa suceedanae nigricanae adenocarpae semicinctanae

Total          18                 9

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Table 5. Food-quality classes and associated body sizes of four Cydia sister-species pairs. Data are from the Appendix. (Ditto is abbreviated do.)

The four pairs of sister species mentioned earlier represent two sections (strobilellae and pomonellae) each containing only two species, and one section (cosmophoranae) containing four species that can be resolved into one Palearctic sister pair and one Nearctic sister pair. The four pairs, their food-quality classes, and forewing lengths are shown in Table 5. The members of the first pair belong to different food-quality classes, and differ in size by 25% ([6.5 — 4.9J/6.5 = 0.25). The remaining pairs all belong to the high class, but members of the third differ in size similarly to or more than members of the first, as expected from their differing food-plant seed sizes ([7.6 — 5.7J/7.6 = 0.25; [7.6 — 4.0J/7.6 = 0.47). Members of the second and fourth pairs differ least, from 10 to 13%; inherited similarity in their body sizes cannot be ruled out.

In sum, independence and number of degrees of freedom are no doubt overestimated, but only mildly. Cydia body-size evolution seems sufficiently independent of subgeneric, section, and species phylogeny to uphold rather than refute results of the statistical analysis.

Discussion

Even though I consider only one independent variable, it relates well to body size (Table 1, Fig. 1). The correlation of body size and food-plant seed size among seed predators of all food-quality classes ultimately results from finite size and finite numbers of seeds. Seeds occur within seed-bearing parts of lesser food quality (Mattson 1980, Skerman 1977). The smaller the seed, the more feeding disruptions larvae experience as they finish one seed and seek another or eat seed-bearing tissue or both; the more feeding disruptions, the smaller the body size.

Potential number of seeds per fruit is reduced by frequent failure of some seeds to develop (Stephenson 1981, Tripp & Hedlin 1956) and

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by other seed predators. Intraspecific predation occurs in the high class (Bovey 1966, Coyne 1968, Hedlin 1967, Tripp & Hedlin 1956, Kraft 1968, Putman 1963) as well as in the medium and low classes (Bovey 1966). Intraspecific predation promotes net survival when the seed supply in one fruit or the nourishment in one large seed is not sufficient for all inhabitants.

I assume that food-plant seed size precedes Cydia body size, not the reverse. Evidence for this assumption is scant, except that many factors besides seed predation determine seed size (Harper et al. 1970).

In the medium and low food-quality classes, larvae probably eat more grams of food to support a given body size than in the high class. Smaller body size may lessen food needs of a larva. At least two members of the medium class, C. zebeana and C. milleniana, have evolved lengthened life cycles enabling their larvae to feed a second season (Kuznetsov 1987, Postner 1978). Forewing lengths of these species (7.0-7.3 mm) are greater than the class mean (6.4 mm) (Appendix, Table 1).

Among seed predators in the medium and low classes, food-plant seeds are mostly large and borne singly. Correlation between food-plant seed size and forewing length in seed predators of these classes (Fig. 2: line B) may exist because most such larvae use but one seed (Bovey 1966) through a range of seed sizes.

In conclusion, I interpret the correlation between Cydia body size and diet quality to reflect mechanisms whereby (a) larvae evolve body sizes for which they are able to obtain sufficient nourishment, and (b) body-size diversity arises as lineages colonize new larval food plants and plant parts of differing diet quality.

Acknowledgments

I thank D. Prokrym, G. Terlemezian, D. A. Andow, M. Elling, and S. Lazzari for translating help, and D. L. Wagner, D. G. McCullough, W. J. Mattson, V. I. Kuznetsov, and two anonymous referees for reviewing the manuscript. Discussions I had with D. L. Wagner and A. H. Porter greatly aided the study.

Appendix. Enumeration of Cydia study taxa and populations. Asterisks denote foo probable food plants; braces ({ }), seed weights estimated from relatives of similar s Kuznetsov (1968) is abbreviated D & K (1968); ditto, do.

Species Subgenus Section

Sample area

Food plant(s)

Main part eaten (sourc

High food-quality class (diet ca. 25% crude S Europe                       Adenocarpus spp., Seed (D & K 196

Cytisus scopar-ius (L.)*

adenocarpi (Rago-not) Endopisa adenocarpi anaranjada (Miller) Cydia

blackmoreana

(Walsingham) Endopisa nigricanae bracteatana (Fer-nald) Cydia

color ana (Kearfott)

conicolana (Hey-laerts) Cydia illutanae

SE North America Pinus elliottii En- do. (Hedlin et al. gelm.*                            1981)

N Africa

Retama monosper- do. (D & K 1968) ma (L.)

W North America Abies bracteata D. do. (Hedlin et al.

Don, A. concolor          1981)

(Gord. &

Gland.),* A.

magnifica A.

Murr.* SW North America Pinus edulis En-           do. (do.)

gelm.*

Britain

P. sylvestris L.,* P. nigra Arn.*

do. (Bradley et al 1979)

Appendix. Continued.

Endl.!

Appendix. Continued.

Appendix. Continued.

Appendix. Continued.

Species Subgenus Section

Sample area

Food plant(s)

Main part eaten (source)

Appendix. Continued.

Appendix. Continued.

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Received for publication 10 February 1990; revised and accepted 11 July 1990.