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Forest Service

U.S. Department of Agriculture Agriculture Handbook No. 514
NEARCTIC PINE TIP MOTHS

OF THE GENUS RHYACIONIA:

BIOSYSTEMATIC REVIEW

(Lepidoptera: Tortricidae, Olethreutinae)

by

Jerry A. Powell, Professor of Entomology

University of California, Berkeley, Calif. 94720

and

William E. Miller, Principal Insect Ecologist

North Central Forest Experiment Station, St. Paul, Minn. 55108

Agriculture Handbook No. 514

Forest Service

United States Department of Agriculture

Washington, D.C. April 1978
Powell, Jerry A., and William E. Miller.

1978. Nearctic pine tip moths of the genus Rhyacionia: biosystematic review (Lepidoptera: Tortricidae, Olethreutinae).

U.S. Dep. Agrie, Agrie. Handb. 514, 51 p., illus.

Knowledge of 13 previously known species of pine tip moths in

North America is reviewed, and 11 new species are described and

named.

KEYWORDS: biogeography, taxonomy, Pinus.

Library of Congress Catalog Number 77-600028

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402

Stock Number 001-000-03713-7

Class Number A 1.76:514
CONTENTS

Page

INTRODUCTION 1

BIOLOGY 2

Voltinism 2

Life Stages 2

Host Plant Associations 3

GEOGRAPHIC DISTRIBUTION 5

ALLOPATRIC SPECIES PAIRS 6

TAXONOMIC CHARACTERS AND

DIAGNOSIS 6

GENUS RHYACIONIA HÜBNER [1825] 7

SYSTEMATICS OF SPECIES 8

GROUP 1: BUOLIANA 8

Rhyacionia huoliana (Denis and Schiffermüller) . . 8

GROUP 2: RIGIDAN A, SUBTROPICA 9

Rhyacionia rigidana (Fernald) 9

Rhyacionia subtropica Miller 10

.GROUP 3: MULTILINEATA 11

Rh yacionia multilineata Powell, New Species. ... 11

GROUP 4: PASADENANA TO MARTINANA . 12

Rhyacionia pasadenana (Kearfott) 12

Rhyacioniazozana (Kearfott) 13

Rh yacionia neomexicana (Dyar) 14

Rhyacionia salmonicolor Powell, New Species ... 15

Rh yacionia monophylliana (Kearfott),

New Combination 15

Rhyacionia martinana Powell, New Species 16

Page

GROUP 5: ADANA, JENNINGSI 17

Rhyacionia adana Heinrich 17

Rhyacionia jenningsi PoweU, New Species 17

GROUP 6: BUSCKANA, BLANCHARDI 19

Rhyacionia husckana Heinrich 19

Rhyacionia hlanchardi Miller, New Species 19

GROUP 7: FUMOSANA 21

Rhyacioniafumosana Powell, New Species 21

GROUP 8: FRUSTRAN A TO AKTITA 22

Rhyacioniafrustrana (Comstock) 22

Rhyacionia bushnelli (Busck) 24

Rhyacionia sonia Miller 24

Rhyacionia aktita Miller, New Species 24

GROUP 9: SUBCERVINANA,

PALLIFASCIATA 26

Rhyacionia subcervinana (Walsingham) 26

Rhyacionia pallifasdata Powell, New Species. ... 28

GROUP 10: A UREOLA 29

Rhyacionia aureola Powell, New Species 29

GROUP 11: VERSICOLOR, FLAMMICOLOR. . . 30

Rhyacionia versicolor Powell, New Species 30

Rhyacionia flammicolor Powell, New Species.... 31

ACKNOWLEDGMENTS 31

LITERATURE CITED 32

MALE GENITAL CHARACTERS, Figures 3-40 34

FEMALE ABDOMINAL STRUCTURES,

Figures 41-61 43

ADULTS OF RHYACIONIA, Figures 62-85 49
INTRODUCTION

The genus Rhyacionia is widespread in the

Holarctic Region, ranging from Japan and Asia

to the Caribbean Antilles and Mexico. It attains greatest diversity in the mountainous

parts of the western and southwestern Nearctic. The genus is represented by about 35

described species worldwide; 24 are treated

here, including 11 previously undescribed.

Larvae of these moths inhabit growing tips of

pines and sometimes reach densities of economic concern in nurseries, in ornamental

plantings, and in natural and planted reforestation projects.

Except for several eastern species that are

multivoltine, most American Rhyacionia fly

only in early spring, so they have been infrequently sampled by general collectors. Most of

our knowledge about their biologies originates

from conspicuous larval damage in pure,

planted stands. Geographic distributions of

most species are poorly documented, being

dependent on sporadic collections. No one has

attempted to systematically survey these

moths in the Rocky Mountains, the Sierra

Madre Occidental of Mexico, and other ecologically diverse regions.

The taxonomy of North American Rhyacionia has rested largely on the treatment by

Heinrich (1923), which has grown increasingly

obsolete. Recent contributions by Miller (1961,

1967b) have updated knowledge in the frustrana complex, while studies by MacKay

(1959), Lindquist (1962), Miller and Wilson

(1964), Yates (1967), and Dickerson and Kearby (1972) have provided aids to identification

of larvae, pupae, and adults of selected species.

The purpose of the present work is to revise

comprehensively the classification of the

Nearctic Rhyacionia. We were prompted to

undertake the task by inquiries for identification and the resultant discovery of undescribed

forms.

Although we cooperated on various phases

of the work, responsibility for individual

species was divided: in general Miller handled

species of the eastern half of the continent and

Powell the western. Most of the specimens for

given species were studied by only one of us.

Photographs for all species were produced at

the North Central Forest Experiment Station

in St. Paul and drawings and maps at the University of California at Berkeley.

Material has been examined from the following collections; the abbreviations are used in

the text to indicate specimen repositories:

American Museum of Natural History, New

York,N.Y. (AMNH).

André Blanchard, Houston, Tex. (Blanchard).

British Museum (Natural History), London,

England (BMNH).

California Academy of Sciences, San Francisco, Calif. (CAS).

California State Department of Agriculture,

Sacramento, Calif. (CDA).

Canadian Forestry Service, Great Lakes

Forest Research Centre, Sault Ste. Marie,

Ontario (CFS-GL).

Canadian Forestry Service, Northern Forest

Research Centre, Edmonton, Alberta

(CFS-N).

Canadian Forestry Service, Pacific Forest

Research Centre, Vancouver, British Columbia (CFS-P).

Canadian National Collection, Ottawa, Ontario (CNC).

Charles P. Kimball, Barnstable, Mass. (Kimball).

Florida Division of Plant Industry, Gainesville, Fla. (FDPI).

Los Angeles County Museum of Natural

History, Los Angeles, Calif. (LACM).

Maine Forest Service, Augusta, Maine

(MFS).

Bryant Mather, Jackson, Miss. (Mather).

Michigan State University, East Lansing,

Mich. (MSU).

National Museum of Natural History, Washington, D.C. (NMNH).

Shirley Hills, Pensacola, Fla. (SH).
Gayle Strickland, Baton Rouge, La. (Strickland).

USDA Forest Service, North Central Forest

Experiment Station, St. Paul, Minn.

(USFS-NC).

USDA Forest Service, Pacific Southwest

Forest and Range Experiment Station,

Berkeley, Calif. (USFS-PS)

USDA Forest Service, Rocky Mountain

Forest and Range Experiment Station:

Albuquerque, New Mex. (USFS-RMA);

Bottineau, N. Dak. (USFS-RMB); Ft. Collins, Colo. (USFS-RMFC).

University of California, Berkeley, Calif.

(UCB).

University of Michigan, Ann Arbor, Mich.

(UM).

University of Wisconsin, Madison, Wis.

(UW).

Primary types are deposited in CAS on indefinite loan from UCB, in LACM, and in

NMNH in part as gifts from Blanchard, Kimball, and USFS-RMA. The letter n in text indicates number of specimens examined.

Copies of specimen label data and original

photographs are available at the California

Insect Survey, University of California, Berkeley, Calif., and at the North Central Forest Experiment Station, St. Paul, Minn.

BIOLOGY

The biologies of Rhyacionia species are

similar. Moths fly in spring, though some are

multivoltine. In species where host plants are

known, the larvae feed in developing pine

shoots during late spring and summer.

Voltinism

Most of the North American species appear

to be univoltine as seen in the following tabulation:

Status of knowledge No. ofspecies

Univoltine; life history investigated 6

Univoltine according to available

records 7

Multivoltine; life history investigated 4

Multivoltine according to available

records,at least in South 3

Available records too fragmentary to

permit conclusion 4

Adults 0Îpasadenana amd frustraría emerge as

early as late February or March in western and

southern coastal areas, and often in April or

May in other areas. Adults of several species,

including adana (Martin 1960), neomexicana,

jenningsi (personal communication with Daniel

T. Jennings), and monophylliana are active

while patches of snow are still on the ground.

On the other hand, versicolor, a montane

species, flies in late June and July, as does the

introduced buoliana. Members of the rigidana

and frustraría groups are bi- or multivoltine.

the number of generations varying with latitude. For example, frustraría annually develops three to six generations in the South and

only two in Pennsylvania and Ohio (Yates 1960,

Berisford 1974b).

A strong, obligate diapause has not been

clearly demonstrated for any species. In several, like pasadenana and frustrana, little if

any diapause seems to occur; pupae brought indoors in fall or winter complete development

and emerge.

Life Stages

Moths of most species are usually crepuscular or nocturnal, sometimes diurnal. This information is often lacking on the collection

record. Rhyacionia buoliana is the most extensively studied of the crepuscular species.

Observations by Pointing (1961) in Ontario

showed that adult emergence peaks at 6:00 to

7:00 a.m., with sporadic eclosión through the

day. Flight and mating occurs between 6:00

and 10:00 p.m., with greatest activity between

7:00 and 8:00 p.m. Most oviposition (numbers

of eggs per hour per individual and numbers of

individuals participating) takes place between

4:00 and 8:00 p.m., peaking at 6:00 to 8:00 p.m.

Based on numbers of males of frustrana and

rigidana attracted to caged, virgin females,

Berisford (1974b) found that female calling

periods are short and overlap only slightly. Ac-
tivity oífrustraría occurs mostly at 7:00 to 7:30

p.m., while that oirigidana was highest at 8:00

to 9:00 p.m. The major flight period oîneomexicana occurs at night, between 8:00 and 12:00

p.m. (Jennings 1975).

By contrast, spring generation hushnelli are

said to fly on warm days in May, although oviposition occurs at night (Swenk 1927). Males of

zozana, a species for which there are no light

attraction records, have been taken flying during afternoon, but Stevens (1966) states that

mating and presumably oviposition take place

at night. Adults of adana are mainly diurnal,

with copulation occurring between 1:00 and

8:00 p.m. when the air temperature is at or

above 16°C (Martin 1960).

Oviposition in all species recorded occurs irregularly, with eggs deposited singly or in

loosely organized groups of from 2 to 11. They

are placed on or between needles, on needle

sheaths, on bud scales, or on twigs near terminals. Development is related to temperature, usually lasting from 14 to 21 days but

from 8 to 14 days in huoliana (Pointing 1963),

and as short as 6 days in the summer brood of

hushnelli (Swenk 1927).

Newly hatched larvae of most species feed

first between needles, often entering a needle

and mining. Later instars feed inside needle

sheaths (suhcervinana) or buds and finally in

the growing shoots, completing growth by midsummer or fall. Rhyacionia huoliana, introduced from Europe, differs from native

American species in overwintering as a partly

grown larva in the buds. In spring, larvae of

huoliana resume feeding, moving to new terminals to complete development.

Pupation occurs in the larval gallery, on or in

the ground away from the tree bole, or in cocoons attached to the root collar just below soil

level. Most univoltine species spend the pupal

wintering period on the ground (zozana, neomexicana, adana, husckana), but pasadenana,

which lives in a maritime climate, pupates in

terminals. As noted, the introduced huoliana is

exceptional; the larva overwinters, and the

brief pupal stage takes place in the larval

gallery. By contrast, facultatively multivoltine

species usually spend the winter in terminals

(rigidana, suhtropica, frustrana), R. hushnelli

apparently uses both strategies; some or all

overwintering pupae occur in the ground, summer generation pupae in the larval galleries

(Swenk 1927).

Host Plant Associations

All Rhyacionia larvae for which data are

available feed on pine. Rearing records believed to represent native populations on

natural hosts in America north of Mexico are

summarized in table 1. Nomenclature of pines

throughout this work follows Critchfield and

Little (1966). Records involving introduced

trees, such as Pinus sylvestris, or unnatural

situations such as mixed native tree species in

nurseries, are excluded. There are no host data

for exclusively Mexican Rhyacionia species,

and the diverse pine flora of Mexico is omitted

from the table.

Table 1.—Known host plants for native North

American Rhyacionia, hased on rearing

records within native ranges of the pine

species

Pinus taxonomic group and

host species

Rhyacionia species occurring

on host

SUBGENUS STROBUS

Sect. Strobus

Sect. Parrya

Subsect. Cembroides

monophylla

SUBGENUS PINUS

Sect. Ternatae

Sect. Pinus

Subsect. Sylvestres

resinosa

tropicalis

Subsect. Australes

palustris

taeda

echinata

rigida

pungens

elliottii

caribaea

occidentalis

cubensis

Subsect. Ponderosae

ponderosa

monophylliana

jeffreyi

Subsect. Contortae

banksiana

contorta

virginiana

Subsect. Oocarpae

radiata

mûrieata

rigidana, adana, busckana,

subtropica

subtropica

rigidana, subtropica,

frustrana, aktita

rigidana, frustrana

rigidana, frustrana, aktita

frustrana^

subtropica, frustrana, aktita

sub trópica, frustrana

frustrana

frustrana

multilineata, zozana, neomexicana, jenningsi, busckana,

fumosana, bushnelli, subcervinana

zozana, subcervinana

adana, busckana, sonia

pasadenana\ zozana^

rigidana, frustrana

pasadenana

pasadenana^

Unconfirmed records.
Conclusions concerning host specificity must

be tentative because several species have been

reared at only one or a few localities. Records

from the better known species in the Eastern

United States and Canada suggest that a diversity of pines within the subgenus Pinus are

used by Rhyacionia whenever available. Similar host profiles are shown by the rigidana

group, adana, and the frustraría group in Subsections At^siraZes and Contortae. On the other

hand, the most widespread western pine of

Subsection Contortae y Pinus contorta, has

been definitely linked with only one Rhyacionia. Most western Rhyacionia are known

from only one or two closely related pines.

A perplexing evolutionary problem is

emerging from known host relations. Rather

than an even distribution oiRhyacionia species

across taxonomic groupings of the pines, nearly all records involve the subgenus Pinus. Only

Rhyacionia monophylliana definitely is associated with a member of the subgenus Strohus.

The whole Section Strobus, the white pines,

and Subsections Balfourianae and Sabinianae

are not known to harbor any Rhyacionia. In

sharp contrast, eight Rhyacionia species, 50

percent of those whose food plants are known,

are harbored by Pinus ponderosa. Thus, there

is an uneven distribution of moth species

among the conceivable ecological niches.

Berisford (1974a, b) showed isolating mechanisms and partial division of resource utilization through differences in pheromone

broadcast time and seasonal flight periods for

frustrana and rigidana living together. For

most Rhyacionia, however, too little is known

to explain how coexisting species accomplish

ecological or microgeographical subdivision of

the niche resource or whether observed isolating mechanisms are primary or secondary.

The concentration of eight species of Rhyacionia on ponderosa pine is particularly difficult to interpret. Except for busckana, each

has been reared only from members of the

Subsection Ponderosae. If any of the records

are incidental for these seven moths, the usual

hosts are unknown. Their known distributions

fall within that illustrated for Pinus ponderosa

by Critchfield and Little (1966). The host's

geographic range is partially subdivided; subcervinana is limited to Pacific coastal areas and

bushnelli, neomexicana, fumosana, and jenningsi to the eastern or Rocky Mountains (var.

scopulorum). The remaining two, zozana and

multilineatay are broadly sympatric in the west

with subcervinana, and in the eastern part of

the range with other members of the complex.

Moreover, within the eastern group, neomexicana and fumosana exhibit broad sympatry,

both narrowly overlapping the nearly allopatric bushnelli to the east. Data show local

sympatry among several pairs or triads:

zozana and multilineata in California;

bushnelli, neomexicana, and fumosana in

North Dakota; zozana and fumosana near Ft.

Collins, Colo.; multilineata, neomexicana, and

fumosana near Colorado Springs, Colo.; neomexicana, fumosana, a,nd jenningsi in Arizona.

At most of these places light attraction or

pheromone trap records indicate that two or

more species fly together on the same dates.

According to the equilibrium theory of

island biogeography (MacArthur and Wilson

1967), diversity increases for many organisms

with increasing geographic extent of habitat

(Sullivan and Shaffer 1975) or food-plant range

(Opler 1974, Strong 1974). A similar relation

applies to native North American Rhyacionia

and the genus Pinus when widespread subgenus Strobus pines are excluded. In the subgenus Pinus the number of Rhyacionia species

harbored (S) is positively correlated with area

of pine species distribution (A) (r^ = 0.74, P <

0.05, fig. 1). For all pine species on which one

or more Rhyacionia species were known to

feed, A was estimated as an index of km^ based

on range width x range length (right angle),

using maps of Critchfield and Little (1966). The

equation S=a+b A gives a closer fit than log S

and log A. The subgenus Pinus can be envisioned as an evolutionary stage for speciation

in Rhyacionia. Why the widespread species

Pinus strobus, albicaulis, and other members

of the subgenus Strobus as well as certain

members of the subgenus Pinus are not utilized remains unexplained.
NO. RHYACIONIA

SPECIES HARBORED (S)

8r

S =1.1 + 0.654A

r2 = 0.74

1 •

2 3 4 5 6

INDEX OF PINE AREA (km^/IO«) (A)

8

/igure 1. —Correlation between number of Rhyacionia species harbored per pine species (S) and index of the area of

geographic range (A) of individual pine species. Each point represents one North American subgenus Pinus species.

GEOGRAPHIC DISTRIBUTION

The genus Rhyacionia is widely distributed

in the northern hemisphere. Its range approximates the distribution of the genus Pinus

(Critchfield and Little 1966), although Rhyacionia have not been discovered in the southernmost extensions oiPinus in southeast Asia,

the Philippines, and Sumatra. There are 10

species assigned to Rhyacionia in the Palearctic iObraztsov 1964), a much larger area of

Pinus than that occupied by the 23 native

Nearctic species treated here. The greatest

concentration of Rhyacionia species occurs in

the Western United States.

Rhyacionia huoliana, introduced from the

Palearctic, is widely distributed in North

America (Map 1), and has become established

in South America, in Argentina, and in Uruguay (Obraztsov 1964). Most native American

Rhyacionia are distributed in either the

eastern or western half of the continent.

Although many species are widespread, only

busckana is known to occur transcontinentally.

Among native Nearctic species, 1 is widespread, 7 are restricted to the eastern half of

the continent, 12 are restricted to the western

half, and bushnelli occurs in the middle, sympatrically with members of the eastern and

western cohorts. Two species are described

from Mexico, and there are additional specimens insufficient for description that represent possibly two others, all from the Sierra

Madre Occidental in Sinaloa and Durango. No

endemic Rhyacionia have been described from

Eastern or Southern Mexico and Central

America, a region of diverse pine species. Two

Eastern North American species range into

the Greater Antilles, subtropica in Cuba and

frustrana on all the major islands, and into

southern Mexico and Central America.

The area of greatest concentration of species

5
is the southern Rocky Mountains; 10 species

are represented in Colorado, New Mexico, and

Arizona. Most of these are associated with

Pinus ponderosa, and at several places as

many as three have been taken. In addition,

versicolor, for which no biological data are

available, occurs sympatrically with members

of the ponderosa pine assemblage. Near Flagstaff, Ariz., and near Colorado Springs, Colo.,

they have been collected together. Four

species have been collected at Rock Creek near

Colorado Springs.

ALLOPATHIC SPECIES PAIRS

A parallel pattern of variation and distribution is shown by six pairs of Nearctic Rhyacionia species: rigidana-subtrópica (Map 2),

monophylliana-martinana (Map 6), adanajenningsi (Map 7), frustrana-bushnelli (Map

10), sonia-aktita (Map 11), and subcervinanapallifasciata (Map 12). Each pair is allopatric

with slight differentiation in size, color, minor

features of genitalia, and biology. Present

knowledge of Rhyacionia biology makes it arbitrary whether these are considered species,

subspecies, or unnamed components of polytypic species. We elected to treat each segregate as a species, partly because different

species of pines are used by the allopatric sibs

in each pair where such data are available.

Both members of some pairs are widespread

(rigidana-subtropica, frustrana-bushnelli,

sonia-aktita), while other pairs have one

widespread member or both members are

restricted in known range (monophyllianamartinana). Distribution patterns range from

widely disjunct (adana-jenningsi) to nearly

contiguous or parapatric (rigidana-sub trópica,

frustrana-bushnelli).

TAXONOMIC CHARACTERS AND DIAGNOSIS

Sixteen characters were used for determining species and defining species groups as seen

below. These may be grouped as follows: two

external features of both sexes, one external

feature of male, one sexual dimorphic, six male

genitalic (fig. 14), four female genitalic (figs.

59, 60) and two biological.

1. Labial palpus II segment length: 0.6 to

1.4x eye diameter.

2. Labial palpus III segment length: 0.2 to

0.4x II segment length.

3. Antennal pectination length (d): 0.3 to

2.Ox segment diameter.

4. Sexual dimorphism in size: d larger or 9

larger.

5. Pollex of valva (d): absent, rudimentary,

or present.

6. "Clasper'' (sacculus extension) of valva

(d): absent, rudimentary, or present.

7. Aedeagus form (d): short, thick (length 3

to 7x width), or elongate, slender (8 to 13x

width).

8. Terminal spur of aedeagus (d): present or

absent.

9. Subapical serration of aedeagus (d):

present or absent.

10. Number of cornuti (d): range 0 to 20+.

11. Sterigma form (9): multistate.

12. Accessory lobe below sterigma (9):

present or absent.

13. Sclerotized areas of ductus bursae (9):

present or absent.

14. Signa (9): absent, small, large.

15. Food plant: species of subgenus Strobus

or subgenus Pinus.

16. Overwintering stage: larva or pupa.

We found the genital characters, especially

valva and aedeagus in the male and sclerotization of sterigma and bursa copulatrix (including signa) in the female, to be the most reliable

for species definition. These features should be

employed in specimen determination. Valva

shape should be used cautiously as a diagnostic

character, as it is variable among individuals

depending partly on orientation in slide mounting. Thus we illustrate more than one valval

aspect for some species. The number, distribution, and relative size of the cornuti may prove

equally valuable, but we were unable to develop a technique for properly inflating the

vesica. Moreover, it is not known whether the
number varies within or between populations

or if cornuti are normally deciduous during the

life of the moth. We found cornuti occasionally

in corpus bursae, indicating transfer with spermatophores. Cornutus size may be correlated

with number; when 1 to 3 in number, they are

usually elongate and slender; when numerous,

they are short and stout.

Forewing color is often conveniently used in

making species identifications; it varies little

within species. However, pattern and color are

extremely similar among several species (multilineatay zozana, husckana, blanchardi,

fumosanüy adana, jenningsi). Absolute size may

also be useful in comparing species in some

cases, as we found narrow ranges within species, using forewing length, base to apex, as an

index. We have no evidence that size of specimens is affected by laboratory rearing (usually

from field-collected, full-grown larvae or pupae) and for this reason do not report whether

measured specimens were reared or field collected as adults.

We omit keys in the belief that the user can

identify a particular specimen more quickly

and more easily by comparing it with the wing

photos (figs. 62-85) and with the drawings of

male genitalia (figs. 3-40) and female genitalia

(figs. 41-61). To aid such comparison, illustrations of each kind for all species are grouped

together at the end of the text.

Where more than one species oi Rhyacionia

populate the same pine stand or tree, field

symptoms may not be distinctive enough to indicate the presence of several species. One frequent and well documented mixture is frustrana-rigidana (Miller and Wilson 1964, Baer

and Berisford 1975). Multispecies adult collections or rearings (present data) also include

aktita-b lanehardly aktita-frustrana, busckanasonia, bushnelli-fumosana, and jenningsineomexicana, which suggests Rhyacionia mixtures may exist on a larger scale than previously supposed.

Where mixtures are likely and population as

well as individual diagnosis is desired, enough

individuals must be diagnosed and findings interpreted statistically. Population composition

can be estimated from the size and composition

of a representative sample of individuals,

adults or immatures, using binomial confidence

intervals (Miller and Wilson 1964). Binomial

confidence interval tables are found in

Snedecor and Cochran (1967) and in many

other statistical references. Their use is illustrated here by two examples. If a sample of

10 individuals consists entirely of frustrana,

the population randomly represented by the

sample is between 69 and 100 percent frustrana (P<0.05). In a sample of 20 individuals, if

15 avefrustrana and 5 are rigidana, the population randomly represented is between 51 and

91 percent/r^5¿ra?2a and between 9 and 49 percent rigidana (P<0.05). Sample size is paramount; binomial confidence intervals show at a

glance the size of sample needed to give the

degree of precision desired for population

diagnosis.

GENUS RHYACIONIA HÜBNER [1825]

Type species: Phalaena Tortrix buoliana

Denis and Schiffermüller (1776) (selected by

Walsingham 1900:124).

Rhyacionia Hübner (1825:379); Obraztsov

(1964:6, synonymy).

Äeimia Guenee (1845:180).

Type species: Phalaena Tortrix buoliana

Denis and Schiffermüller 1776 (selected by

Fernald 1908:32).

A redescription of Rhyacionia and related

genera, together with thorough taxonomic

synonymy has been given by Obraztsov (1964).

Members of this genus differ from all other

Eucosmini by the following combination of

characters: forewing termen simple, not

notched, apex not falcate; veins Mz and M3

usually connate, sometimes separate but closely adjacent at base (rarely short stalked),

always nearer to one another at base than M3 is

to Cui; costal fold of male absent. In male

genitalia the socii are rudimentary or absent,

the cucuUus separated by a weak or moderate

constriction. The forewing features are shared

by members of Petrova Heinrich (1923) and

Barbara Heinrich (1923). These moths are

differentiated by their possession of well

developed, elongate socii and a deep constriction preceding the cucullus.

Biologically, Rhyacionia differ from all other

conifer-feeding Eucosmini including Petrova,

which feed beneath bark sometimes creating

pitch nodules or blisters, and Barbara, which
are cone moths. An exception occurs in pine

shoot borers of the sonomana-glorióla complex

of Eucosma. These are easily distinguished by

the well developed costal fold as well as other

adult and larval characters (Heinrich 1923,

MacKay 1959, Obraztsov 1964).

SYSTEMATICS OF SPECIES

GROUP 1: BUOLIANA

Female larger than male. Antennal pectination of d minute, < 0.1 segment diameter.

Labial palpus short, 11 = 0.9 eye diameter.

Male genitalia: sacculus without extensions or

poUex development; aedeagus elongate with

terminal spur, 3 to 7 elongate cornuti. Female

genitalia: sterigma simple; signa lacking.

Rhyacionia buoliana (Denis and

Schiffermüller)

(Figures 3, 4, 41, 62; Map 1)

Tortrix buoliana Denis and Schiffermüller

(1776:128).

Evetria buoliana; Bankes (1909:3).

Rhyacionia buoliana; Heinrich (1923:15);

Obraztsov (1964:141, synonymy).

The forewing is orange with silvery white

crossbands. The moth is medium sized: forewing length in males 8.0 to 9.5 mm (25 n);

females, 8.0 to 11.0 mm (25 n). Male genitalia

(figs. 3, 4): valva broad; aedeagus tapered distally, vesica with 3 to 7 elongate cornuti (25 n).

Female genitalia (fig. 41): sterigma funnel-like,

ductus bursae with a sclerotized spot, signa absent (21 n). Larvae and pupae have been differentiated from other Rhyacionia by MacKay

(1959), Lindquist (1962), and Dickerson and

Kearby (1972).

Type daia.—Vicinity of Vienna, Austria; no

syntypes are known (Obraztsov 1964).

Geographic distribution.—lnÍToá\xc,Qá in

North America (Map 1): Newfoundland south

to Delaware and west to Upper Michigan and

Missouri; also coastal British Columbia,

Washington, and Oregon (Pointing and Miller

1967, Carolin and Daterman 1974). It also occurs through much of Europe and in a small

area in South America (Obraztsov 1964, Miller

1967a).

LAMBERT AZIMUTHAL EQUAL-AREA PROJECTION

Map 1.—Geographic distribution of the European pine shoot moth, Rhyacionia buoliana, in North America (after Pointing

and Miller 1967, Carolin and Daterman 1974).

8
Flight period.—Msiinly June. The insect is

univoltine throughout its North American

range (Pointing and Miller 1967).

Hosts.—MaiTiy species of subgenus Pinus

(Miller 1967a).

Discussion.—This is the well known European pine shoot moth, accidentally introduced

to North America in the early 1900's. Complete

taxonomic synonymy treating the Old World

literature is given by Obraztsov (1964).

The one-sentence original description is insufficient to distinguish between two similar

Rhyacionia occurring at the type locality. The

current identity of buoliana is due to Bankes

(1909) who called attention to genitalic differences and, in effect, assigned the names

buoliana and pinicolana. All subsequent

authors in Europe and North America have followed Bankes' designations. Rhyacionia pinicolana (Doubleday) does not occur in North

America.

Obraztsov (1964) and others have given

Schiffermüller as the sole author responsible

for the name buoliana probably because only

he is mentioned in the original work (copy examined at Niedersachsen State and University

Library, Göttingen, West Germany). According to Hagen (1862, 1863), both Denis and

Schiffermüller were responsible; other names

originating in the original work are given dual

authorship (as in Bradley and coworkers 1973).

The year of publication is taken as 1776 after

Dos Passos (1958).

This insect is the most studied member of

the genus. Miller (1967a) gives the most recent

comprehensive account of its biology and importance.

GROUP 2: RIGIDANA, SUBTROPICA

Female larger than male. Antennal pectination of d short, 0.4 segment diameter. Labial

palpus short, II = 0.6 to 0.8 eye diameter. Male

genitalia: sacculus extension rudimentary,

pollex not developed; aedeagus elongate, no

spur, cornuti absent. Female genitalia:

sterigma unmodified, ductus bursae with

sclerotized basal sleeve; signa tiny.

Rhyacionia rigidana (Fernald)

(Figures 5, 6, 7, 42, 63; Map 2)

Re finia rigidana Fernald, in Comstock

(1880:237).

Rhyacionia rigidana-, Heinrich (1923:19).

The moths have reddish-brown forewing

markings with a pale medial crossband as wide

or wider than the basal patch. The pattern is

similar to subtropica, but the pale areas are

darker in rigidana. Moths are moderate sized:

forewing length in males 5.5 to 8.0 mm (28 n);

females, 6.0 to 9.0 mm (33 n). Male genitalia

(figs. 5, 6, 7): valva broad, sacculus with a

distal triangular flange; emargination preceding cucuUus accentuated on the unflattened

valva (compare figs. 5, 6, 7); aedeagus straight

in dorsal-ventral aspect, sinuate and slightly

bent downward distally in lateral aspect (fig.

7a) (27 n). Female genitalia (fig. 42): sterigma

small, a nonsclerotized dorso-medial seam dividing lamella postvaginalis for 0.5 or more of

its length; ductus bursae sclerotized basally,

usually forming a partial ring, sometimes (8

percent of examples) a complete ring; two

small, unequal sized thornlike signa (24 n). Larvae are indistinguishable from sub trópica but

both were differentiated from frustrana by

Miller and Wilson (1964) and from neomexicana and buoliana by MacKay (1959). Pupae

were differentiated from those of three other

Rhyacionia by Yates (1967) and Dickerson and

Kearby (1972); Yates (1969) also differentiated

pupal sexes.

Type (Zaia.—Ithaca, N. Y.; lectotype female

70834 in NMNH designated by Miller (1970).

Geographic distribution. —{Maip 2) Maine

south to Florida and west to Missouri and

Texas (Miller and Neiswander 1959).

Flight peWocZ.—Mid-March to May and during July (Miller and Neiswander 1959, Berisford 1974a).

Hosts.—Vsivious species of subgenus Pinus

(Miller and Neiswander 1959) (table 1).

Discussion. —Moths of either sex of rigidana

are distinguishable from those of sub trópica by

the darker medial crossband and other pale

forewing areas; females by presence of the

dorso-medial seam of lamella postvaginalis,

which is absent in subtropica, and by thin

lateral margins of the sterigma, which are

thick in subtropica. No structural differences

between males are known.

Two small male examples taken in flight at

Pensacola, Fla. (11-3-62, SH), are tentatively

placed here. They are not distinguishable in
LAMBERT AZIMUTHAL EQUAL-AREA PROJECTION

Map 2.—Distribution records íovRhyacionia rigidana (triangles) andÄ. subtropica (circlesjT

genitalia, but possess longer antennal pectination and the forewing ground color is deep red

and hindwings are darker than typical specimens. The contrast in forewing hues is reminiscent OÎ subtropicüy but overall appearance is

more similar to rigidana. No matching females

are available.

Further study of male genitalia showed

polymorphism reported in the ventro-distal

area of the valva (Miller 1961) to be an artifact

of nonuniform microslide mounting.

Rhyacionia subtropica Miller

(Figures 8, 43, 64; Map 2)

Rhyacionia subtropica Miller (1961:231).

The moth has forewing markings similar to

rigidana but the contrast in hues is greater in

subtropica. The moths are moderate sized:

forewing length in males 5.0 to 8.0 mm (22 n);

females, 5.5 to 9.5 mm (31 n). Male genitalia

(fig. 8): similar to rigidana (10 n). Female

genitalia (fig. 43): sterigma small, lateral

margins forming a thick lip; ductus bursae

sclerotized basally, often forming a complete

ring, less often (33 percent of examples) a partial ring; two small, unequal sized thornlike

10

signa (21 n). Larvae are indistinguishable from

rigidana but both were differentiated from

frustrana by Miller and Wilson (1964). Pupae

were differentiated from two other Rhyacionia

by Yates (1967); Yates (1969) also differentiated pupal sexes.

Type daia.—Valparaiso, Fla., May (Gemmer); holotype female 65012 in NMNH.

Geographic distribution. —(Mdi^ 2) South

Carolina south to Florida and Cuba, west to

Mississippi and British Honduras (Miller 1961,

1965; Hochmut 1972).

Flight peWod.—December to March, May to

August, and September and October; in Florida the species is bivoltine with a partial third

emergence (Miller 1961, McGraw 1975).

jyo5¿5.—Several pines of subgenus Pinus,

but mainly P. elliottii var. elliottii and P. elliottii var. densa (Bethune 1963, Hochmut 1972,

McGraw 1975) (table 1).

Discussion.—^Q^diTdiiion of adults of subtropica and rigidana is discussed under

rigidana. Further study of male genitalia

showed polymorphism reported in the ventrodistal area of the valva by Miller (1961) to be

an artifact of nonuniform microslide mounting.
GROUP 3: MULTILINEATA

}^Q^-

Female slightly larger than male. Antennal whitish, legs sprinkled with dark gray extericiliation of d short, 0.3 segment diameter. orly. Forewing: Moderately broad, length 2.9

Labial palpus short, II = 0.9 eye diameter. to 3.0 times width; apex acute, termen strongMale genitalia: sacculus extension not defined, ly angled back, tornal angle distinct. Upperpollex weak; aedeagus short without arma- side dark gray to end of cell, orange-brown

ture; > 10 cornuti. Female genitalia: sterigma beyond, crossed by about 12 ill-defined transan open bowl, ductus bursae unmodified, signa verse lines, whitish on basal half of wing,

large. metallic bluish in distal part of gray, silvery

and shining orange in terminal area, the basal

half bluish to the unaided eye; the lines in midRhyacioniamultilineataPoweM^ew Species dorsal area enlarged, somewhat coalesced,

(Figures 9,10,44,65;Map 3) sometimes appearing as a pale blotch. Tornal

"ocellus" represented by a few to many black

scales roughly arranged along two transverse

rows. Fringe dark gray with a distinct black

V line at base. Underside dark gray mottled with

whitish in dorsal and terminal areas.

Hindwing: Slightly broader than forewing;

apex round, termen slightly convex, broadly

• curved to dorsum, tornal angle not discernible.

Ground color whitish basally, becoming dark

gray at distal margins, to dark gray brown.

Fringe pale gray. Underside whitish, the veins

dark gray. Abdomen: Dorsal scaling shining

dark gray, venter pale, genital scaling whitish.

Genitalia (figs. 9, 10, drawn from paratypes

JAP preps. 3452, Colorado, and 3462, California; 7 n): valva broad without marked constric-

^ tion before cucullus, pollex short, aedeagus

abruptly attenuate distally; cornuti 11 to 12.

Female.—Length of forewing 6.4 to 7.1 mm

(9 n). Essentially as described for the male, but

darker and more vividly colored. Forewing

^^ ^^ .1 slightly broader than in male, length 2.8 to 2.9

^ times width; terminal area banded pale and

Map3.--r)istributionrecordsforÄÄ2/adomamu/h7meaia. dark orange between the transverse shining

lines. Hindwing darker than in male. Genitalia

(fig. 44, drawn from paratype, Blodgett Forest,

A small moth with forewing pattern JAP prep. 3472; 4 n): ductus bursae with a

zozana-like, bluish with reddish termen, large- sclerotized band at junction of ductus semly obscured by many whitish transverse lines. inalis; signa broad, curving.

MaZe.—Length of forewing 6.5 to 7.0 mm (7 Holotype male and allotype female: Californ). Head: Labial palpus moderately short, II nia, Buck Creek Ranger Station, 1,660 m elevasegment length 0.9 eye diameter. III 0.4 as tion, Modoc County, June 5 to 7, 1970, at light

long as II; scaling broadly spreading. III seg- (P. A. Opler and J. Powell) (CAS). Fourteen

ment nearly obscured, pale gray tinged darker paratypes, as follows. California: same data as

exteriorly toward base. Scaling of front and holotype. Id; Blodgett Forest, 21 km E.

crown whitish gray, slightly to extensively Georgetown, El Dorado Co., 19 VI-24-67, at

tinged with dark gray, the scales white-tipped. light (J. Powell); Middletown, Lake Co., 19

Thorax: Dorsal scaling darker than head, the III-23-64, emerged from Pinus ponderosa log

scales dark gray, white-tipped. Venter shining (D. Kinn). Wyoming: 10 km NW Newcastle,

11
Weston Co., 4d, 59 VI-23-25-65 (R. W.

Hodges). Colorado: Rock Creek Canyon, Colorado Springs, El Paso Co., Id V-15-59, 19

VI-3-59 (M. May) (Kimball, NMNH, and UCB).

Additional specimens subsequently examined: Texas, Guadeloupe Mts., Smith Canyon,

Culberson Co., 1,753 m, 2cJ, 39 V-22-73 (R. W.

Hodges); Blodgett Forest, Calif., 6c?, 19 VI-4,

5-75, at light (P. A. Rude and J. Powell).

The Wyoming, Colorado, and Texas specimens differ from California examples in having

the hindwing slightly darker (a distinction

more evident in the male) and in the more

uniformly colored forewing, the basal half less

bluish and terminal area less reddish.

The single rearing record from Pinus ponderosa probably represents pupation in bark;

the moth emerged from logs caged for bark

beetle studies. The March emergence date is

abnormal, presumably the result of laboratory

conditions.

GROUP 4: PASADENANA TO MARTINANA

Male larger than female. Antennal ciliation

of d short to moderate, 0.3 to 0.7 segment

diameter. Labial palpus normal, H = 0.9 to 1.0

eye diemeter. Male genitalia: sacculus and

pollex weakly to well developed, aedeagus

short to moderately elongate with terminal

spur; 1 to < 20 cornuti. Female genitalia:

sterigma a broad ring, ductus bursae unmodified; signa tiny to large.

Rhyacioniapasadenana (Kearfott)

(Figures 13, 45, 66; Map 4)

Evetriapasadenana Kearfott (1907:3).

Rhyacionia pasadenana; Heinrich (1923:16).

A pale moth with extensive reddish orange

over the distal half of the forewing, gradually

blending with the grayish basal area. Forewing length of males 7.0 to 9.1 mm (15 n);

females 6.5 to 8.0 mm (10 n). Male genitalia

(fig. 13): pollex and distal projection of sacculus weakly developed; aedeagus short,

blunt, vesica with about six slender cornuti (3

n). Female genitalia (fig. 45): sterigma a simple, broad ring, ductus bursae sclerotized

basally; signa tiny (3 n).

Type cZaia.—Three specimens, Pasadena,

Calif., W. G. Dietz; Alameda Co., Calif., March;

Carmel-by-the-Sea, Monterey Co., Calif., A. H.

Vachell. Lectotype male. Alameda County, in

AMNH designated by Klots (1942) based on

the selection of Heinrich (1923). Type locality

restricted here to the city of Alameda for

reasons discussed below.

Geographic distribution. —{Map 4) Immediate coastal areas of California. The native

range is unknown, but it may have been limMap 4. —Distribution records for Rhyacionia pasadenana

(triangles) andR. zozana (circles).

ited to the Monterey peninsula, the endemic

distribution of Pinus radiata. Now pasadenana

occurs from Humboldt County (Koehler and

Tauber 1964) south to Los Angeles County on

ornamental and feral pines.

Flight period.—End of March to beginning

of July according to combined year collections

at light in the east San Francisco Bay area (25

n), and as late as August at Eureka, the northernmost locality. These records represent a

single generation per year (Koehler and

Tauber 1964).

12
Hosts.—Pinus radiata in naturalized stands

and P. contorta and P. muricata (Lange 1937),

the last two possibly only in ornamental situations. A specimen labelled "reared from cypress twigs," (UCB) probably is mislabelled or

involves a misidentified host.

Discussion. —This species closely follows the

"fog belt" distribution pattern, which limits

many insects to areas near the coast in California (Keifer 1935). R. pasadenana was not taken

during a 5-year light trap survey at Walnut

Creek in an area with many Pinus radiata of all

ages. This station is only 17 airline km east of

the Berkeley Hills, which harbor populations of

the moth yet serve as a barrier to inland influence of the fog belt. Thus it is assumed that

the species was originally limited to a coastal

pine, either Pinus radiata or P. muricata.

There are no records from native stands, but

adults were collected before 1907 at Carmel,

within the natural range of these pines as well

as at Alameda and Pasadena where no native

pine occurs.

The selection of "Alameda County" as the

type locality is inadequate because it raises the

possibility of Pinus sabiniana serving as the

native host at higher, inland locations. The

present coastal distribution of the moth,

together with negative evidence from sampling P. sabiniana fauna of inland Alameda

County (Burdick and Powell 1960), suggests

that ornamental pine on the San Francisco Bay

coast was the source of the type specimen.

Moreover, the label on the type is from the

Koebele collection, and Koebele made his headquarters at Alameda during 1885 to 1888

(Essig 1931:674). He must have reared a pinefeeding olethreutid there because the tachinid

Urophyllopsis retiniae (Coquillett 1897) was

described from "Alameda, Calif., bred April

17, 1888, from chrysalids oiRetinia sp. by Mr.

Albert Koebele." This fly has been reared from

R pasadenana in San Francisco (JAP lot 61B2,

det. C. W. Sabrosky). Thus the type locality

probably is the city of Alameda with Pinus

radiata the probable host.

Rhyacionia zozana (Kearfott)

(Figures 14,15, 46, 67; Map 4)

Evetria zozana Kearfott (1907:2).

Petrovazozana; Heinrich (1923:26).

Rhyacionia zozana; Lange (1937:25, 31).

Evetria matutina Meyrick (1912:35) (unnecessary substitute name).

Evetria montana Busck (1914:147). New synonymy.

Rhyacionia montana; Heinrich (1923:18).

The moths have a dark bluish-gray thorax

and basal two-thirds of the forewing, marked

with whitish transverse striae. The head,

palpi, thorax anteriorly, and distal one-third of

the forewing are reddish orange, the forewing

becoming dark red at the margin. The species

thus resembles jenningsi, described further

on. However, in that species the adults are

smaller, with gray palpi and thorax, and the

orange of the forewing is less extensive. R.

zozana also is superficially similar to certain

Petrova, especially P. edemoidana (Dyar), and

was thought to be a Petrova until the male was

discovered and the biology studied (Lange

1937).

Forewing length in males 8.7 to 11.0 mm

(usually 10 to 11 mm) (27 n); females, 7.8 to 9.0

mm (14 n). Male genitalia (figs. 14, 15): poUex

moderately elongate, less defined when valva

flattened (compare figs. 14 and 15); vesica with

a dense bunch of 12 to 15 cornuti (13 n). Female

genitalia (fig. 46): sterigma a heavy ring with

posterior protruding enlargement, base of ductus bursae with a broad sclerotized sleeve; corpus bursae with a sclerotized band on proximal

half; signa small with broadly sclerotized bases

(8n).

Type daia.-Placer Co., Calif., April 20 (A.

H. Vachell); female type in AMNH (Klots 1942)

(zozana), Elliston, Mont., on Pinus contorta,

issued December 10, 1913 (J. Brunner); male

type in NMNH (montana).

Geographic distribution. —(Map 4) Widespread in the Western United States, from

Tuolumne County, Calif., northward on the

west slope of the Sierra Nevada to Oregon,

southern British Columbia, western Montana,

and northern Colorado.

The species was also recorded in eastern

Washington by Stevens (1966), but Stevens

(personal communication) states that this was

based on a record from Grant County sent him

from NMNH. We examined the NMNH specimen from Grant County, and it proved to be R.

busckana Heinrich.

Flight penod.—Mid-March to Mid-May in

California and Oregon, based on field collected

adults (10 n) and Stevens (1966).

13
Hosts.—Pinus ponderosa and P. jeffreyi in

California (Lange 1937, Stevens 1966) and P.

contorta in Montana (Busck 1914). Pines of

both subgenera serve as hosts under conditions of mixed planting (Stevens 1966).

Discussion.—H\ie synonymy of montana

Busck under zozana was suggested by Lange

(1937) but not followed by subsequent workers. We did not locate further material from

Montana or adjacent States, but comparison of

the type male with specimens from northern

California and Oregon reveals no appreciable

differences. The montana type is slightly

darker than typical zozana, duller golden reddish in the terminal area, while some specimens from northern California and Oregon are

intermediate. The male genitalia of the montana type show no differences when compared

to El Dorado County, Calif., material.

Further information is needed to clarify the

status of the following populations from Colorado, Utah, and New Mexico, which are tentatively referred to zozana.

Two samples from near Ft. Collins, Colo.

(USFS-RMFC), differ from typical zozana and

from one another: Id, 19,16 km NW of Ft. Collins, collected at light in May 1973, have the

whitish striae of the forewing obscured by

dark ground color and the terminal orange

duller than typical. The female appears to lack

the sclerotization of the corpus bursae (although the preparation is heavily red-stained,

which may obscure a light sclerotization).

Ninedd, Ft. Collins, taken at pheromone traps

between May 27 and June 20, 1975, have an excessive number of cornuti (20 to 24) and a

slightly narrower pollex (4 n). The moths are

larger than 66 from California (forewing

length 11.1 to 12.1 mm) and have a more restricted, less defined, and paler apical orange

blotch on the forewing. This results in a larger

dark portion, which has distinctive, whitish

cross striae.

Four öd reared at Albuquerque, N. Mex.,

from ornamental ponderosa pine in 1974

(USFS-RMA) are indistinguishable from the

typical California phenotype, unlike the darker

Colorado examples discussed above. Whether

they represent an introduction from nearby or

distant populations is unknown. If native New

Mexico components of the species exist in this

phenotype, a polytopic interpretation of the

variation will be necessary. One 9 reared from

Pinus mugo (a Palearctic species) at Bountiful,

Davis Co., Utah, in 1971 (NMNH) is atypical.

The reddish terminal of the forewing is reduced in extent and brightness, with corresponding expansion of the dark ground and

pale transverse striae, so the forewing somewhat resembles multilineata. The sterigma

form is less defined, especially the posterior

median protrusion.

Rhyacionia neomexicana (Dyar)

(Figures 11,12, 47, 68; Map 5)

Evetria neomexicana Dyar (1903:286).

Rhyacionia neomexicana; Heinrich (1923:16).

The pinkish buff covering the distal area of

the forewing, above the tornus and beyond the

cell, distinguishes adults ol neomexicana. This

area has no transverse striae and is marked only by two thin black lines from the end of the

cell through the terminal area. There is

marked sexual dimorphism, the females

smaller and brighter colored, with more

distinct markings. Forewing length in males

12.5 to 13.5 mm (10 n); females, 8.7 to 10.0 mm

(10 n). The males are the largest moths in

Rhyacionia. Male genitalia (figs. 11, 12): pollex

and distal flange of sacculus moderately well

defined; cornuti, three slender spines (3 n).

Female genitalia (fig. 47): sterigma a broad

ring with subtending sclerotized enlargement,

ductus bursae unmodified; signa large, moderately broad cones with broadly sclerotized

bases (3 n). Larvae were distinguished from

other Rhyacionia by MacKay (1959).

Type data.—Las Vegas, San Miguel Co.,

N.M., "bred from pine" (T. D. A. Cockerell);

holotype male in NMNH.

Geographic distribution. —{Map 5) North

central Arizona, New Mexico, Colorado,

Nebraska, southern Montana, and southwestern North Dakota.

Flight period. —From late March to late May

in the south (Jennings 1975) to June in the

north (Keen 1952). There are specimen records

(NMNH) for mid-March (1 n) and late June

(6 n) in northern Arizona and July in Colorado

(18n).

Hosts.—Pinus ponderosa (Arizona,

southwestern Colorado, and Nebraska)

(NMNH, Jennings 1975) and P. ponderosa var.

scopulorum (Colorado Springs) (NMNH).

14
••

• •

c^r:

Map 5. —Distribution records for Rhyacionia neomexicana

I circles) and i?. salmonicolor (triangle).

Discussion.—The original specimens were

reared from ''imported pine trees" in a plantation, but Cockerell (1901) stated that the same

caterpillars were found in Pinus ponderosa

var. scopulorum at nearby Las Vegas Hot

Springs, N.M.

The species was also reared at Halsey,

Nebr., from 1928 to 1933 from ponderosa pine

plantings, in a manmade forest planted near

the turn of the century (Pool 1953). This colony

may have resulted from short-distance

spreading—less than 100 km—rather than as

an introduction from a distant population

because ponderosa pine was present in the

western half of Nebraska before 1900.

Rhyacionia salmonicolor Powell, New Species

(Figures 16, 69; Map 5)

A small moth with salmon-colored forewings

with a thin longitudinal dark line through the

cell.

Ma¿e.—Length of forewing 7.2 to 7.7 mm (4

n). Head: Labial palpus short, II segment 0.8

eye diameter. III less than 0.3 length of II.

Scaling of palpus grayish, becoming reddish

distally. Scaling of front and crown pale

orange, becoming whitish posteriorly. Thorax:

Dorsal scaling orange brown anteriorly, becoming whitish gray posteriorly. Underside

white, legs dark gray, metathoracic leg paler.

Forewing: Length 3.4 times width, costa nearly straight, termen strongly angled back. Upperside pale salmon colored; base blackish,

bordered at immediate base and outwardly

with whitish (bluish to unaided eye); a thin,

dark reddish brown line in cell from just

beyond basal patch to end of cell; termen narrowly dark red, bordered outwardly at base of

fringe with whitish; fringe distally bright rose

pink. Underside dark gray, dorsal area

whitish, fringe as above. Hindwing: Pale gray,

fringe whitish. Underside white. Abdomen:

Shining sordid whitish. Genitalia (fig. 16,

drawn from JAP prep. 3587; 2 n) similar to

neomexicana except distal projection of sacculus less prominent; poUex narrower and

slightly curved; apical spur of aedeagus relatively smaller; cornuti three long spines.

Female, —Unknown.

Holotype male.—Texas, Mt. Locke, 2,161 m,

Davis Mtns., May 19, 1971 (A. & M. E. Blanchard) (NMNH). Three paratype males, same

data (Blanchard and UCB).

This moth strikingly resembles the widespread mistletoe feeding pyralid, Dasypyga

alternos quammella Ragonot. Rhyacionia

salmonicolor seems closely related to neomexicana, judging from wing color and morphology. Thus the female may be expected to be

smaller with less extensive salmon color on the

forewing. The host plant of salmonicolor is

unknown, but a likely suspect is the isolated

segregate of ponderosa pine in the Davis

Mountains.

Rhyacionia monophylliana (Kearfott), New

Combination

(Figures 17, 70; Map 6)

Evetria monophylliana Kearfott (1907:1).

Petrovamonophylliana; Heinrich (1923: 26).

The moths are paler than any other Rhyacionia, having bluish gray basal, reddish terminal areas, on the forewing. The whole basal

one-fourth and the basal half of costal areas are

pale blue gray with whitish transverse striae.

The remainder of the wing is pale ochreous except that the terminal margin and fringe are

dark reddish. Forewing length in males 7.9 to

15
8.4 mm (5 n). Male genitalia (fig. 17): pollex

moderately elongate, slender, sacculus with a

weak distal ridge; vesica with two or three

slender spines (2 n).

Type data.-Coso Valley, Calif. , May 1891

(Koebele); lectotype male in AMNH, designated by Klots (1942), who followed Kearfott

in stating that this locality is in Kern County.

Recent maps do not show a Coso Valley. Presumably the name refers to the area west of

the Coso Range near Coso Junction, or to the

east between the Coso Range and Argus Mountains. Part of the type series came from the

Argus Mountains. All of these places are in

Inyo County.

Geographic distribution. —{M8ip 6) Known

only from the margins of the Mojave Desert in

California, in southern Inyo and northwestern

Ventura counties.

Flight period.—May; a series from the west

base of Frazier Mountain at 1,670 m elevation

was collected May 7 to 9, a few days after the

last snowfall in the area.

Hosts.—Pinus monophylla (Kearfott 1907).

Discussion.—The species was originally described from four specimens, but by 1923 all

four had lost their abdomens. The only later

collection we have seen is a series of males

taken in 1959 (UCB); we have not been able to

study female genital characters.

Rhyacionia martinana Powell, New Species

(Figures 48, 71; Map 6)

The forewing of this small moth has the

basal area and costal half to the end of the cell

bluish gray, sharply contrasted with the

salmon-ochreous dorsal-terminal area.

MaZ^.—Length of forewing 7.2 to 8.8 mm (14

n). Head: Labial palpus short, II segment

length 0.7 to 0.8 eye diameter. III 0.3 to 0.4 as

long as II; scaling spreading, nearly obscuring

III segment, brick red exteriorly, white interiorly, III blackish. Scaling of crown pale rust

brown. Thorax: Dorsal scaling ochreous and

dark rust red on collar, becoming pale ochreous to whitish posteriorly; venter shining

whitish, legs banded with dark brown, metathoracic leg paler. Forewing: Narrow, length

3.6 to 3.8 times width; termen steeply angled

back, evenly curved to dorsal margin. Color

pattern sharply divided, basal one-fourth and

costal half to end of cell white with scattered

16

Map 6. —Distribution records for Rhyacionia monophylliana (triangles) andR. martinana (circle).

gray scales, bluish to the unaided eye; basal

area with indistinct darker, transverse strigulae; dorsal half beyond one-fourth immaculate

ochreous salmon becoming salmon pink in terminal area; termen narrowly deep rust-red and

a tinge of rust red in costal area beyond cell;

outer costa and base of fringe narrowly white,

fringe distally dark rust red. Underside dark

gray in costal half through cell, blending to

whitish distally; whitish and red spotting of

costa and terminal fringe repeated. Hindwing:

About 1.1 wider than forewing; costa moderately concave distally, apex narrowly acute.

White, lightly tinged with brownish gray on

veins distally and at margins. Fringe white.

Underside white, lightly tinged with brownish

along veins on costal half. Abdomen: Dorsal

scaling pale brownish gray, venter whitish.

Genitalia similar to monophylliana (fig. 17),

pollex slightly broader, not recurved; aedeagus with one to two cornuti (3 n).

Female.—hength of forewing 6.3 to 8.3 mm

(7 n). Essentially as described for male except

eye slightly smaller, labial palpus II segment

0.8 to 0.9 eye diameter. Forewing broader,

length 3.3 to 3.6 times width. Genitalia (fig. 48,

drawn from paratype, JAP prep. 3944; 2 n):

sterigma a slightly raised ring, produced posteriorly; ductus bursae broad, unsclerotized;

signalarge.
Holotype male and allotype female: Arizona,

8 km north of Prescott, elev. 1,660 m, Yavapai

Co., May 7, 1973, and May 1, 1974 (L. M. Martin) (LACM). Paratypes (19): Id, same data except V-7-73; 12ci, 69, same data except IV-28

to V-11-74 (LACM, NMNH, and UCB).

This species is closely related to R. monophylliana. It differs from monophylliana by

having the bluish-gray area of the forewing

sharply delimited and restricted to the basal

area and costal half of the cell. The salmonochreous area is darker and is extended over

the dorsal half more broadly and distinctly

than in monophylliana. In male genitalia the

pollex of monophylliana is more slender and

slightly recurved, and there are two to three

cornuti compared w^ith one to two in martinana. In female genitalia the new species is similar to zozana, with the sterigma less produced,

the signa larger, and ductus bursae without a

sclerotized band. Females of monophylliana

and salmonicolor, when found, probably will be

similar to martinana.

The close relation to monophylliana and to

distributions of pinyon pines (Critchfield and

Little 1966) suggest that Pinus edulis is the

host plant oiR. martinana.

This moth is named for Lloyd M. Martin,

dean of Arizona Lepidoptera collectors.

GROUP 5: ADANA, JENNINGSI

Male larger than female. Antennal ciliation

of d short. Labial palpus normal, II = 1.0 eye

diameter. Male genitalia: sacculus extension

not developed, pollex moderately broad;

aedeagus elongate, slender; two to four cornuti, elongate. Female genitalia: sterigma a

broad ring, ductus bursae unmodified; signa

small.

Rhyacionia adana Heinrich

(Figures 18, 72; Map 7)

Rhyacionia adana Heinrich (1923:18).

Like several other Rhyacionia^ the basal

three-fourths of the forewing has alternating

light and dark gray crossbands. The adult cannot be reliably separated from husckana, blanchardi, and jenningsi except by genitalic

characters and the long antennal pectination of

males of the first two. The moths are small to

medium sized: forewing length in males 6.5 to

9.0 mm (47 n); females, 5.5 to 8.5 mm (42 n).

Male genitalia (fig. 18): valva moderately constricted preceding cucullus; aedeagus gradually attentuated, vesica with two to four spines

(10 n). Female genitalia (6 n) similar to jenningsi (fig. 49), distinguished under that

species. The larvae have been differentiated

from three other Rhyacionia by Lindquist

(1962).

Type data.-Fovest Hills, Mass., IV-5-1910

(Raff); holotype male 24786 in NMNH.

Geographic distribution. —(M3,p 7) Massachusetts south to Virginia and west to Ontario

and Mississippi.

Flight period.—MaiTch through May in the

North where the insect is known to be univoltine (Martin 1960). A Mississippi specimen was

captured flying in January.

Hosts.—Pinus resinosa^ P. banksiana, and

P. sylvestris in the North (Martin 1960).

Rhyacionia jenningsi Powell, New Species

(Figures 19, 20, 49, 73; Map 7)

A western species resembling adana and

zozana, with the base of the forewing bluish,

distal one-third orange and reddish.

MaZe.—Length of forewing 7.5 to 8.5 mm (18

n). Head: Labial palpus moderately long, II

segment length 1.0 eye diameter; III segment

0.4 as long as II; scaling broadly spreading,

hoary gray, the scales dark, tipped with white.

Antennal ciliation slightly more than 0.5 segment width. Scaling of front and crown hoary

gray, slightly to broadly tinged with reddish

posteriorly. Thorax: Dorsal scaling entirely

hoary gray to slightly tinged with reddish

orange anteriorly. Venter and legs paler gray,

the scales white-tipped. Forewing: Length 3.2

to 3.3 times width; costa evenly curved,

termen strongly angled back. Ground color

dark gray on basal half, in cell and beyond in

costal area, crossed by whitish transverse

striae; apex, terminal area beyond cell and tornal area from well before end of cell pale

orange, becoming dark red at margins. Fringe

dark gray, sometimes with basal scale row

dark reddish. Underside dark gray, some

17
?^^ií$---

.^^

Map 7. —Distribution records ior Rhyacionia adana {circles) andR. jenningsi (triangle).

whitish marks along costa. Hindwing: Slightly

broader than forewing; apex bluntly rounded,

termen slightly concave, tornal angle distinct.

Ground color dark gray, slightly darker

toward margins. Fringe slightly paler. Underside pale gray, the veins darker. Abdomen:

Scaling dark gray, genital scaling paler.

Genitalia (figs. 19, 20, drawn from paratypes,

JAP prep. 2671, 3640; 4 n) similar to adana (fig.

18), the pollex usually slightly broader,

aedeagus stouter, abruptly narrowed at the

slender, curved tip in jenningsi; cornuti three,

elongate.

FemaZe.—Length of forewing 6.4 to 7.8 mm

(20 n). Essentially as described for male, but

eye slightly smaller, labial palpus length 1.0 to

1.2 eye diameter. Genitalia (fig. 49, drawn from

paratype, JAP prep. 3745; 4 n) similar to

adana, the sterigma plate, especially bordering

the ostium, distinctly defined, shape variable

but relatively shorter (length 0.45 to 0.55

width in jenningsi, 0.55 to 0.65 in adana); with

anterior margin excavated laterally, less triangular than in adana; base of ductus bursae

more heavily, broadly sclerotized in jenningsi;

signa variable, one small, one large, slender.

18

Holotype male and allotype female: Arizona,

Chevelon Ranger District, Sitgreaves National

Forest, Coconino Co., reared from Pinus ponderosa, April 2 and 8,1969, Hopkins U.S. 37276

(D. T. Jennings) (NMNH). Paratypes (36), all

with the same data except as follows: 8d, 39,

Hopk. U.S. 37257, 37274, III-28-31-69; 2d,

149, Hopk. U.S. 37276, ni-31 to IV-17-69; 6d,

29, Hopk. U.S. 37322, ni-26 to IV-7-70; Id

Hopk. U.S. 37323 IV-22-70 (NMNH, UCB,

USFS-NC, USFS-PS, and USFS-RMA). The

emergence dates may not reflect the true

flight period.

This species is closely related to adana. In

addition to distinguishing features in the genitalia, the new species has longer ciliation on

the antennae in the male (setae length slightly

more than 0.5 the segment diameter, 0.3 in

adana). The above diagnosis of females is

based on specimens of adana from Ontario and

Michigan. It should be reassessed when specimens become available from areas near the

type locality of adana (Massachusetts). Pinus

ponderosa is the only known host iorjenningsi,

while adana has been recorded on pines of

other subsections, P. resinosa and P. banksiana.
We saw worn specimens from one locality in

the geographical gap between adana and jenningsiy the man made forest near Halsey,

Nebr. (Pool 1953). These were collected in 1927

and 1928, and some were reared and bear two

host labels each, P. ponderosa and P. banksiana. The condition of these specimens

prevents definite identification, but they have

jeimingsi antennal and aedeagus characteristics. Whether a long-distance introduction

occurred is unknown. It may have been a temporary establishment because we have no evidence of collection records there in the subsequent half century.

The Arizona specimens originated from an

area that burned in 1956 west of Chevelon

Canyon, 72 km SW of Winslow. In places that

did not regenerate naturally, plantings of

ponderosa pine were colonized by Rhyacionia

(Jennings 1975).

The species is named for Daniel T. Jennings

who conducted biological studies on Rhyacionia in the Southwest and supplied material

for our study.

GROUP 6: BUSCKANA, BLANCHARDI

Male larger than female. Antennal ciliation

of d long, two times segment diameter. Labial

palpus normal, II = 1.0 eye diameter. Male genitalia: sacculus extension and pollex well

developed, aedeagus stout with terminal spur

and apical dentation, two to eight cornuti,

short. Female genitalia: sterigma a convex

ring with convoluted folds, ductus bursae with

basal sleeve; signa tiny.

Rhyacionia busckana Heinrich

(Figures 21, 22, 23, 50, 74; Map 8)

Rhyacionia busckana Heinrich (1923:17).

Like several other Rhyacionia^ the basal

three-fourths of the forewing has light and

dark alternating grayish crossbands. One conspicuous distinguishing feature is the elongate

antennal ciliation of the male. Individual setae

are about twice segment width, proportionately much greater than in all other Nearctic

Rhyacionia except blanchardi. Otherwise the

moth is similar in appearance to adana, jenningsiy and zozana, though there are marked

genitalic differences. Adults of busckana are

medium sized: forewing length in males 5.5 to

9.5 mm (25 n); females, 4.5 to 9.0 mm (28 n).

Male genitalia (figs. 21, 22, 23): valva shape

variable depending on orientation, pollex

broad; vesica with two to eight cornuti (13 n).

Female genitalia (fig. 50): sterigma ringlike

with posterior convolutions, ductus bursae

enlarged distally (26 n). Larvae have been

differentiated from three other Rhyacionia

(Lindquist 1962).

Type daia.—Bellmore, Long Island, N. Y.,

IV. 1913. Holotype male 24785 in NMNH.

Geographic distribution—(Map 8) The most

widely distributed North American Rhyacionia. It occurs from Maine south to Florida

and west to British Columbia, Oregon, Colorado, and Texas. Records west of a line between Manitoba and Texas are based on seven

specimens. R. busckana ranges to 55° N. lat. at

Walker Lake, Manitoba (not mapped).

Flight period.—MsiTch to June; also taken in

flight in November in Levy Co., Fla. (Heppner

1975).

Hosts.—Pinus banksiana in Ontario (Lindquist 1961). Specimens from Ontario were

reared from P. resinosa and P. sylvestris

(CFS-GL). Specimens from British Columbia

were reared from P. ponderosa (CFS-P).

Discussion.—The seven western specimens

mentioned above are larger than eastern ones.

With more specimens and biological information additional taxa might have to be recognized. One of the western specimens is the

basis of an early erroneous record of the European Rhyacionia (Retinia) duplana (Hbn.) in

North America (Walsingham 1879:77). The

specimen is Walsingham 91964 (NMNH) collected in Grant Co., Oreg., during the 1871 to

1872 expedition (Essig 1941) and labeled by

Walsingham as a voucher for the published

record.

Rhyacionia blanchardi Miller, New Species

(Figures 24, 51, 75; Map 8)

A small moth with forewing pattern similar

to busckana and several others.

ikfaZe.—Length of forewing 5.5 mm (1 n).

19
Head: Antennal setae twice as long as the

width of antennal segments. Labial palpus

short, length of II segment 0.9 eye diameter,

III 0.4 as long as II and nearly obscured by

scaling of II; white interiorly, pale grayish

brown exteriorly. Scaling of front and crown

pale grayish brown, the scales white-tipped.

Thorax: Dorsal scaling similar to head. Venter

shining whitish, metathoracic legs whitish,

pro- and mesothoracic legs pale grayish brown

exteriorly, tarsi white banded. Forewing: Narrow, length 2.8 times width; apex acute,

termen strongly angled back. Ground color of

basal third grayish brown crossed by irregular

whitish striae; apical two-thirds reddish

orange crossed by several irregular, pale shining blotchlike fasciae. Middle third between

cell and costa dark bluish. Fringe gray with

basal scale row dark red. Underside dark gray,

several white marks along costa. Hindwing:

Slightly wider than forewing; apex round,

termen slightly convex, broadly curved to dorsum, tornal angle not discernible. Ground color

brownish gray. Fringe pale gray. Underside

brownish gray. Abdomen: Dorsal scaling shining dark gray, venter pale, genital scaling

whitish. Genitalia (fig. 24, drawn from JAP

prep. 3627; 2 n): Pollex curved interiorly, neck

of valva deeply excavated interiorly, aedeagus

with about 12 short or rudimentary spurs; vesica with 2 to 3 cornuti; posterior margin of

uncus oval in outline.

Female.—hengiYi of forewing 5.5 to 6.5 mm

(7 n). Essentially as described for male but

more vividly colored. Forewing slightly broader, length 2.9 to 3.0 times width. Genitalia (fig.

51, drawn from JAP prep. 3628; 5 n): sterigma

a broad plate encircling ostium and with a central posterior lobe, tiny dual signa.

Holotype male and allotype female: Texas,

Conroe, Montgomery Co., March 10, 1968,

genitalia slide WEM 1219733; and March 9,

1971, genitalia slide WEM 1219732, respectively (A. and M. E. Blanchard) (NMNH). Eight

paratypes as follows: Id, 19 same data as holotype, genitalia slides JAP 3627, 3628; 19, same

data as holotype except March 13, 1969, genitalia slide WEM 125743; 499, same data as

allotype, genitalia slide WEM 125744; 19,

Town Bluff (Dam B), Jasper Co., Tex., March

11, 1966, genitalia slide JAP 3631 (Blanchard,

NMNH, and UCB).

This species most resembles husckana in

wing shape, color pattern, and genitalia of both

Ç p^

y ' 1

\

>„

...

Map 8.—Distribution records for Rhyacionia husckana (circles) and R. blanchardi (triangles). The northernmost record for

husckana, at Walker Lake, 55° N., is near the geographic center of Manitoba, beyond the limits of this map.

20
sexes. In genitalia males differ by their oval

posterior margin of uncus; females differ by

their broader sterigma.

This species is named in honor of the lepidopterist André Blanchard who collected the

types as well as many other Rhyacionia.

GROUP 7: FUMOSANA

Male larger than female. Antennal ciliation

of d short. Labial palpus normal, 11=1.1 eye diameter. Male genitalia: sacculus extension

weakly developed, poUex distinct; aedeagus

moderately elongate with terminal spur and

extensive dorsal dentation; six to eight cornuti, short. Female genitalia: sterigma complex with posterior lobe, ductus bursae weakly

sclerotized at base; signa tiny to a trace.

Rhyacionia fumosana Powell, New Species

(Figures 25, 26, 27, 52, 76; Map 9)

A large, narrow-winged species like neomexicana, the forewing pattern obscured by smoky

infuscation, usually with dull orange in the terminal area.

MaZe.—Length of forewing 9.4 to 11.0 mm

(12 n). Head: Labial palpus II segment length

1.1 eye diameter. III 0.3 as long as II; scaling

moderately spreading, hoary gray, the scales

white-tipped. Scaling of head and crown dark

to light hoary gray. Thorax: Dorsal scaling

hoary gray, the scales more broadly white distally on tegulae and on scutellar area. Underside grayish white; legs dark gray exteriorly.

Forewing: Moderately narrow, length 3.3 to

3.5 times width; costa nearly straight, termen

strongly angled back, tornal angle not distinct.

Ground color dark gray from base to beyond

cell, becoming dull orange brown in terminal

area, the pattern completely obscured by

smoky infuscation to moderately distinct with

numerous transverse whitish striae in middle

third of wing, giving a bluish-gray appearance

to basal two-thirds. Fringe gray with a whitish

basal scale row. Underside gray, paler distally.

Hindwing: 1.2 wider than forewing; apex

broadly rounded; termen slightly concave.

Ground color smoky gray, margins and fringe

darker. Underside pale gray, the veins slightly

darker. Abdomen: Scaling entirely gray. Genitalia (figs. 25, 26, 27, drawn from paratypes,

JAP prep. 3564 Arizona, 3621 Colorado and

WEM prep. IV-21-71 North Dakota; 16 n) simi-

.r

Map 9.—Distribution records for Rhyacioniafumosana.

lar to neomexicana, the poUex outwardly

recurved distally, inner margin of sacculus

with 3 or 4 flaplike projections forming an irregular ridge; dorsal surface of aedeagus particularly at sides with many fine spurlike

teeth; vesica with six to eight cornuti.

Female.—Length of forewing 7.9 to 9.8 mm

(9 n). Essentially as described for male but

labial palpus more elongate, II segment length

1.4 eye diameter; and forewing broader, length

3.1 to 3.2 times width. Genitalia (fig. 52, drawn

from paratype, JAP prep. 3599; 9 n): sterigma

a broad convex plate, produced posteriorly into an irregular or notched lobe; one signum

very small, the other absent or a speck.

Holotype male and allotype female, Colorado, Rock Creek Canyon [Colorado Springs],

El Paso Co., May 2, 1959, and April 20, 1960

(Margot May) (NMNH). Paratypes (19) as

follows. South Dakota: Hardy W. C. (T3N, RIE

S30) [29 airline km SW Lead, Lawrence

21
Co.], 3d VI-18-65 (R. W. Hodges). Colorado:

same data as holotype, 19 V-20/VI-7-57; Id,

29 V-13/15-59; 3d, 49 IV-19/26-60; 19

V-12-62. Arizona: Apache Co., Greer, Id

VI-11-36 (G. H. and J. L. Sperry). Coconino

Co., Chevelon Ranger Distr., Sitgreaves Nati.

Forest, 2d IV-12-71 (netted at night) (D. T.

Jennings); Fort Valley, elev. 2,240 m, 12 km

NW Flagstaff, Id VI-19-61 (R. W. Hodges)

(AMNH, Kimball, NMNH, USFS-RMA, and

UCB).

Additional specimens subsequently examined: North Dakota, Sec. 11, T. 136 R. 102,

Slope Co., Id, 29 Vn-9-70 "collected from

Pinus ponderosa'' (Hopk. U.S. 54060-G, with

pupal shells) (A. D. Tagestad); same locality,

Id Vn-20-72 (Hopk. U.S. 56507-L) (M. E.

McKnight); same locality, 18d, 79 V-15-73

"collected from Pinus ponderosa'' (McKnight

and Tagestad); same data. Id VI-7-73 (Hopk.

U.S. 54176). Colorado, Blue Mtn. Cyn., Pike

Nati. For., Park Co., 2,438 m, 2d VI-9-73 light

trap (R. E. Stanford); Rist Cyn., Bellvue,

Larimer Co., Id, IV-21-71 "ponderosa pine

stand" (J. Bodenham); 16 km NW Ft. Collins,

Larimer Co., 7d V-1973 "light" (J. Bodenham).

New Mexico, Fenton Lake, Jemez Mts., Sandoval Co., 19 V-2-74 at U.V. light (M. E.

Toliver) (NMNH and USFS-RMFC).

The variation is in part geographical, there

being a decrease in melanistic coloration from

north to southwest. About half the individuals

from North Dakota and those from South Dakota have smoky blackish forewings, while the

rest of the North Dakota series, most of the

Colorado specimens, and eastern Arizona examples are intermediate, with a dull orangebrown termen and variable development of the

whitish transverse striae. The moths from

Coconino County, Ariz., exhibit the most

distinct color pattern, with little smoky infuscation.

Rhyacionia fumosana resembles neomexicana in size, wing shape, and sexual dimorphism in size, though the last is more pronounced in neomexicana. The male genitalia

are closest to neomexicana, differing by the

recurved pollex, ridged sacculus, and dentate

dorsal surface of the aedeagus. The female

genitalia of fumosana are markedly different

from neomexicana, with a larger, more convex,

posteriorly produced sterigma, less sclerotization in the ductus and a small single signum.

The signa in neomexicana are double and

large.

The two species have been collected sympatrically in Slope County, N. Dak., at

Chevelon Ranger District, Ariz., and at Rock

Creek Canyon, Colo., and adults of the two

were taken on the same nights in April and

May.

GROUP 8: FRUSTRANA TO AKTITA

Female larger than male. Antennal ciliation

of d minute to short. Labial palpus normal,

n = 0.8 to 1.0 eye diameter. Male genitalia: sacculus projection well developed, pollex elongate; aedeagus stout with terminal spur; cornuti absent. Female genitalia: sterigma a

simple ring, sometimes broadened laterally;

ductus bursae without sclerotized structures;

signa tiny.

Rhyacionia frustraría (Comstock)

(Figures 28, 29, 53, 77; Map 10)

Retiniafrustrana Comstock (1880:236).

Rhyacionia frustrana; Heinrich (1923:20);

Miller (1965:58; 1967b:591)

Rhyacioniapseudostrobana Amsel (1962:396)

Rhyacionia schwerdtfegeri Miller (1967b:594),

nomennudum.

Like several other Rhyacionia, the moth has

mottled rusty red forewing markings and a

dark basal patch bordered by a lighter crossband that is narrower than the basal patch.

Adults can be reliably distinguished from sonia

and aktita by genital characters and from

bushnelli only by nonstructural means as discussed below. The moth is the smallest North

American member of the genus: forewing

length in males 4.0 to 7.0 mm (103 n); females,

4.0 to 7.5 mm (108 n). Male genitalia (figs. 28,

29, drawn from CDW prep. 410, 411; 17 n):

valva with a moderately pronounced emargination preceding cucullus, deaccentuating the

sacculus projection; pollex slender, variable in

22
200 400 600 800 1000 1200 14-00 KILOMETERS

LAMBERT AZIMUTHAL EQUAL-AREA PROJECTION

Map 10. —Distribution records iov Rhyacioniafrustraría (circles) andÄ. bushnelli (triangles).

shape; aedeagus stout with a rectangulate,

preapical thickened area. Female genitalia (fig.

53, drawn from JAP prep. 3716; 25 n): sterigma ringlike, poorly defined posteriorly; signa

tiny or reduced to a speck. Larvae have been

differentiated from other Rhyacionia by

MacKay (1959) and Miller and Wilson (1964)

and pupae by Yates (1967) and Dickerson and

Kearby(1972).

Type daia.—Virginia (near Washington,

D.C.). Lectotype female in NMNH designated

and illustrated by Miller (1967b).

Geographic distribution. —(Map 10) The

second most widely distributed native North

American member of the genus, R. frustrana

occurs from Massachusetts south to Florida,

Dominican Republic (Etheridge 1971), Cuba

(Hochmut 1972) and Jamaica, west to Missouri,

Oklahoma, Texas, Oaxaca State, Mexico,

Guatemala (Schwerdtfeger 1962), Honduras,

and Nicaragua (Becker 1973). The species was

discovered in San Diego and Kern Counties,

Calif., where it appears to be recently introduced from Eastern U.S. (Brown and Eads

1975).

Flight period.—From two to six generations

per year are known in the U.S. (Yates 1960,

Berisford 1974b). Voltinism is probably greater in parts of the Caribbean area; Hochmut

(1972) found larvae and pupae every month of

the year in Cuba. Schwerdtfeger (1962) postulated four generations per year in Guatemala

at an elevation of 1,500 m.

Hosts.—Many species of subgenus Pinus

(Yates and Beal 1971, and others) (table 1).

Discussion.—This is the well known Nantucket pine tip moth. Daviault's (1932) reference to R. frustrana in Quebec was in error

and actually pertained to Eucosma glorióla

Heinrich (Daviault 1942).

Differentiation of frustrana from bushnelli

rests on size (fig. 2), geographic distribution,

and place of hibernation. Overwintering/rí¿5-

trana have been found on the host tree rather

23
than on the ground (Miller 1967b, Brown and

Eads 1975). Further discussion of differentiation appears under bushnelli.

^

R. frustrana

^^^^^^^^^^^^^^^

R. bushnelli

$

4 5 6 7

Forewing length (mm)

8

Z)¿5Cí¿55Í07i.—Separation of bushnelli and

frustrana rests on size (fig. 2), geographic distribution, and place of overwintering (Miller

1967b). R. bushnelli occurs in a region where

the large-budded Pinus ponderosa var. scopulorum predominates. Heinrich (1923) believed

the larger size of bushnelli to be host induced.

Size integrity of the two forms remains to be

examined in a manner that rules out host as an

accountable variable. Information that bushnelli overwinters on the ground is due to

Swenk (1927) whose studies should be repeated elsewhere to clarify whether place of

overwintering is fixed or variable relative to

latitude and other factors.

Figure 2.—Forewing lengths of male and female Rhyacionia frustrana and R. bushnelli. Short vertical lines indicate means, horizontal bars one standard deviation, and

horizontal lines total range. Based on 102 to 118 measurements per sex per species, made to nearest 0.5 mm.

Rhyacionia bushnelli (Busck)

(Figure 78, Map 10)

Evetria bushnelli Busck (1914:144).

Rhyacionia frustrana bushnelli; Heinrich

(1923:20).

Rhyacionia bushnelli) Miller (1967b:593).

Like several other Rhyacionia, this moth has

mottled rusty red forewings and a dark basal

patch bordered by a lighter crossband that is

narrower than the basal patch. Adults can be

reliably distinguished from sonia and aktita by

genital characters but irova frustrana only by

nonstructural means. The moths are moderate

sized: forewing length in males 5.0 to 7.5 mm

(118 n); females, 5.0 to 8.0 mm (102 n). Male

and female genitalia as in frustrana (40 n and

41 n, respectively).

Type data.—Fort Bayard, N. M. Holotype

female 18439 in NMNH.

Geographic distribution. —(Msip 10) Montana and North Dakota south to New Mexico,

Kansas, and Missouri (Miller 1967b, McKnight

1973).

Flight period.—One or more generations

develop per year depending on latitude

(McKnight 1973). Where two annual generations occur, moths fly during May and from

June to late July (Swenk 1927).

Hosts.—Pinus ponderosa var. scopulorum

and other members of the subgenus Pinus

(Graham and Baumhofer 1930) (table 1).

Rhyacionia sonia Miller

(Figures 30, 31, 54, 79; Map 11)

Rhyacionia sonia Miller (1967b: 592).

The moth is similar in forewing color and

pattern to aktita, frustrana, and bushnelli. It

can be separated reliably only on genital characters. The moth is small: forewing length in

males 4.5 to 6.5 mm (17 n); females, 4.5 to 6.0

mm (20 n). Male genitalia (figs. 30, 31, drawn

from prep. CDW 271 and WEM 307732; 11 n):

valva with a deep emargination preceding cucullus, sacculus with a broad flange and small

terminal extension, pollex irregular in shape;

aedeagus stout with a long apical spur. Female

genitalia (fig. 54, drawn from prep. CDW 273;

15 n): sterigma a broad ring, widened laterally;

signa as in aktita (fig. 55). The larvae have

been differentiated from three other Rhyacionia (Lindquist 1962).

Type data.—BuTwash, Ontario, Canada,

reared from Pinus banksiana, 1959; holotype

female in CNC.

Geographic distribution. —(Msip 11) Ontario,

Manitoba (Miller 1967b), and Upper Michigan.

Flight period. —April and May.

Hosts.—Pinus banksiana (Sippell and Lindquist 1961, McDowell and Wong 1962).

Discussion. —Re-examination of specimens

identified as sonia from Maine (Miller 1967b)

and South Carolina (Fox and Harris 1969)

shows they are aktita.

Rhyacionia aktita Miller, New Species

(Figures 32, 33, 55, 80; Map 11)

Rhyacionia sonia Miller (1967b:592) (part); Fox

and Harris (1969).

24
Map 11. —Distribution records for Rhyacionia sonia (triangles) andi?. aktita (circles).

A small moth with sonia-\ike forewing pattern, color, and genitalia.

MaZ^.—Length of forewing 4.0 to 7.0 mm (18

n). Head: Labial palpus short, II segment

length 0.9 eye diameter. III 0.3 as long as II;

scaling spreading, partly obscuring III segment; scaling of palpus, front and crown gray

with white tips. Thorax: Dorsal scaling darker

than head, white tipped. Venter shining white;

legs gray with white bands exteriorly sometimes sprinkled with rust; metathoracic legs

much paler. Forewing: Narrow, length 3.0 to

3.3 times width; apex acute, termen strongly

angled back, tornal angle distinct. Basal

quarter gray, forming a faint basal patch. Four

or five blotchlike irregular orange to rust

bands from a few to many scales wide partly or

entirely crossing wing. Basal patch and crossband just beyond middle sometimes appearing

bluish to the unaided eye. Costa with 8 to 10

white marks, usually arranged in pairs. Apex,

including fringe, heavily dusted with brick-red

scales. Fringe, except near apex, dark gray

with a black line extending around termen.

Underside gray except for a sprinkling of

brick-red scales near costa. Hindwing: Slightly

25
broader than forewing; apex round, termen

slightly concave, tornal angle discernible.

Ground color pale grayish brown. Fringe paler,

nearly white. Underside light gray. Abdomen:

Dorsal scaling shining light gray, venter paler.

Genitalia (figs. 32, 33, drawn from paratypes

New Jersey, WEM prep. 118731, 287733; 11 n):

valva constricted before cucullus; caudal margin of cucullus sculptured interiorly to form a

broadly pointed process; pollex longer than cucullus, spatulate; aedeagus terminating in a

fingerlike spur.

Female.—Length of forewing 5.0 to 6.0 mm

(17 n). Essentially as described for male. Genitalia (fig. 55, drawn from paratypes North

Carolina, WEM prep. 29673c, d; 9 n): ostium at

center of sterigma, sterigma depressed caudally around ostium, flattened laterally; small

dual thornlike signa.

Holotype male: New Jersey, Lakehurst, 5

May 1962 (R. W. Hodges) genitalia slide WEM

277734; allotype female: North Carolina, Currituck Co., Knotts Is., July 8, 1962 (D. R.

Davis) genitalia slide WEM 118732 (NMNH).

Thirteen paratypes as follows: same data as

holotype, 2d, slides WEM 118731 and 287733

(UCB, NMNH); 4d same data except May 4

to 6, 1962, slides WEM 88732, 88733, 72742,

and 72743; North Carolina, Id and 29, same

data as allotype, slides WEM 29673a, c, and

125742 (UCB, NMNH); Maine, Id, 29, Pine

Point, March 1963 (C. A. Millet) genitalia

slides JTE 49, 50, and WEM 28743; Texas, Id,

Montgomery Co., Conroe, 9.ni.71 (A. and M.

E. Blanchard) slide JAP 3630 (Blanchard).

This species is similar to sonia from which it

differs in genital characters such as narrower

valva with less extensive basal opening, interior process on cucullus, more curved apical

spur on aedeagus, and small details of sterigma. It overwinters in shoots (Fox and Harris

1969) whereas sonia overwinters on the ground

(Sippell and Lindquist 1961).

Previously published records for sonia in

Maine (Miller 1967b) and South Carolina (Fox

and Harris 1969) apply to aktita. The confusion

of the two species arose from their close similarity coupled with previously limited localities

and numbers oí aktita specimens.

Rhyacionia aktita is coastal in distribution;

it is allopatric to sonia which occurs inland in

the North. Label data suggest two generations

of aktita per year, at least in the South. Adults

have been reared from pupae overwintering in

terminals of Pinus rigida (Miller 1967b), P.

taeda and P. elliottii var. elliottii (Fox and

Harris 1969). Adults were found in the same

capture series with frustrana and hlanchardi.

GROUP 9: SUBCERVINANA, PALLIFASCIATA

Female larger than male. Antennal ciliation

of d moderately long. Labial palpus normal,

11 = 0.8 to 1.0 eye diameter. Male genitalia:

without sacculus projections, pollex variable;

aedeagus moderately elongate with terminal

spur; one to two cornuti, elongate. Female

genitalia: sterigma funnel-like with enclosed

narrow fold; ductus bursae with small sclerotized spot or unmodified; signa small.

Rhyacionia subcervinana (Walsingham)

(Figures 34, 56, 81; Map 12)

Retinia subcervinana Walsingham (1879:25).

Evetria subcervinana; Fernald (1903:449).

Rhyacionia subcervinana; Heinrich (1923:21).

This is a small Pacific Coast species with

pale forewings and dark hindwings. The forewing has transverse bands of lead-colored

scales that are lost on some specimens, leaving

whitish bands alternating with pale rust-tan

ones. As the species has been known only from

the original description based on worn females,

it is redescribed as follows:

MaZe.—Length of forewing 5.4 to 7.0 mm (9

n). Head: Labial palpus short, II segment

length 0.8 to 0.9 eye diameter. III 0.4 as long as

II; scaling moderately spreading, pale ochreous to whitish with a few brown scales on

outer side. Scaling of front pale ochreous to

whitish, crown posteriorly darker, dark ochreous to brown. Thorax: Dorsal scaling concolorous with posterior of crown. Venter shining

whitish, legs brownish exteriorly. Forewing:

Moderately narrow, length 3.5 to 3.6 times

width; costa nearly straight, apex acute, termen strongly angled back, tornal angle not

distinguished. Ground color pale rust brownish, or ochreous tan, crossed by about five

evenly spaced broad, transverse bands of

26
''^ " ^ <i „

Type data.—Oregon, ''Rouge River, June

1872." The lectotype female (BMNH) by present designation bears the data "Rogue R.,

Josephine Co., Oreg., 7.V.1872 Wlsm. 91965," a

label probably written after 1900 by Durrant,

based on the Carrier diary of Walsingham's expedition (Essig 1941) (BM slide 11787).

Geographic distribution. —{Msip 12) Southern British Columbia to southern Sierra

Nevada in California. In addition to the type,

the following specimens were examined:

Canada, British Columbia, Vernon, 6d, 19

VI-5-12-61 r.f. P. ponderosa (61-5101-01 FIS);

Lytton, 19 VI-6-72 "P. ponderosa' (72-289-07

FIS). Oregon, Rogue River Nati. For., 3 km. N

Bull Gap, Jackson Co., 19 VII-3-70 (R. E. Dietz

and P. A. Rude). California, Ash Cr. Rgr. Sta.,

15 km. E. McCloud, Siskiyou Co., Id VI-11-74

^ at light (Chemsak and Powell); Grant Grove,

Sequoia Nati. Park, 4d, 19 IV-28-59 (pupae)

r.i.P.jeffreyi (Hopk. U.S. 37575 A) (R. C. Hall)

(CNC,UCB,USFS-PS).

Map 12.-Distribution records for Rhyacionia subcer- ^¿-^^^ period.-E^vly June to early July

vinana iirieingles) andR.pallifasciata {circles). . -, c- ^ i n i i i i^ /r» \

based on iield-coUected adults (2 n).

Hosts.—Pinus jefferyi in California, P.

metallic lead-colored scales, darker, brownish ponderosa northward. Specimens reared from

at costa; distal two bands irregular, less de- jeffreyi were recorded as having been colfined and broken. Fringe shining gray. Under- lected as pupae in "needle sheath mines," and

side uniform brownish gray, base of fringe British Columbia specimens are pinned with

with a pale line. Hindwing: Narrow, apex bases of needle bundles where larvae pupated,

acute, termen strongly angled back and slight- Discussion.—The type specimen is worn,

ly convex. Ground color dark brownish gray; Our identification with recent material is

fringe paler gray. Underside as on forewing. based on examination of the type by Miller, a

Abdomen: Dorsal scaling brownish gray, ven- Kodochrome photo in the Obraztsov file at

tral and genital scaling paler. Genitalia (fig. 34, AMNH, and the type genitalia slide (BMNH

drawn from plesiotype. Sequoia Nati. For., 11787). The slide preparation is distorted and

Calif., JAP prep. 3612; 3 n): valva broad with overly cleared, but diagnostic features appear

moderately deep emargination preceding cu- to match recent slides from British Columbia,

cullus, pollex moderately elongate with a Oregon, and California. The type lacks all

spurlike expansion on inner side; aedeagus metallic scaling on the forewing (as does our

with fine dentation near apex; vesica with one single Oregon example) and has more rust-tan

elongate cornutus. ground color bands than our specimens, which

Female.—Length of forewing 5.7 to 6.4 mm are ochreous tan in Oregon, brownish tan in

(3 n). Essentially as described for male, but California. The reduction of metallic scaling

with metallic scaling of transverse bands on may be due to age.

forewing less developed, reduced to shining The precise type locality and date of coUecwhitish bands, with or without scattered lead- tion are unknown. The only known account of

colored scales, brownish at costa. Genitalia the Oregon part of the expedition (Essig 1941)

(fig. 56, drawn from plesiotype, Rogue River is a copy at the British Museum made by DurNatl. For., Oreg., JAP prep. 3604; 4 n): ster- rant in 1902, and this corroborates the May 7,

igma with enclosed sclerotized fold, subtended 1872, label data on the type but the "Rouge"

by slight enlargement and sclerotization of spelling is repeated. Presuming "Rouge" to be

ductusbursaebasally; signa tiny. a misspelling of Rogue by Carrier, Walsing27
ham, and Durrant (there is no Rouge River in

southern Oregon or Northern California), it

may be assumed that the type locality was

near Grants Pass, Josephine County, according to the itinerary. If the original June date is

correct, then the locality may have been further south, in the area of the present Rogue

River National Forest in Jackson County

where the Walsingham party spent the first 2

weeks of June 1872.

Rhyacionia pallifasciata Powell, New Species

(Figures 35, 57, 82; Map 12)

Small moths with narrow forewings

transversely banded with pale rust tan and

whitish.

Male.—hength of forewing 5.9 to 8.3 mm (24

n). Head: Labial palpus short, II segment 0.8 to

1.0 eye diameter. III 0.3 as long as II; scaling

loosely spreading, whitish, tinged with pale

brownish exteriorly. Scaling of upper front and

crown spreading, roughened, pale brownish

tipped with white. Thorax: Dorsal scaling as on

head, ventral shining whitish, legs brownish.

Forewing: Narrow, length 4.0 to 4.4 times

width; costa nearly straight, termen strongly

angled back, tornal angle not evident. Ground

color shining pale whitish with about six pale

rust-tan transverse bands, more or less evenly

spaced, the two nearest base narrower, the

two beyond middle somewhat broken and irregular; each brown at costa, giving a more

distinctly banded appearance to costal area;

the pale intervening bands of ground color

overlaid with brownish, which is mostly lost in

older specimens. Fringe whitish, the scales

tipped with brownish. Underside pale brownish, darker on veins; fringe as above. Hindwing: Narrow, only slightly broader than

forewing; apex acute, almost lanceolate, termen strongly angled back, slightly concave.

Dorsal scaling pale brown, darker at distal

margin. Fringe whitish. Abdomen: Dorsal scaling pale brown, ventral whitish. Genitalia (fig

35, drawn from paratype. Rustler Park, JAP

prep. 3571; 4 n): valva with slight emargination before cucullus, pollex broad, short with a

secondary extension on inner side; aedeagus

with fine dentation near apex, vesica with two

elongate cornuti.

Female.—Lenth of forewing 5.7 to 7.4 mm (7

n). Essentially as described for male except

labial palpus more variable in size, II segment

length 0.7 to 1.1 eye diameter; forewing

broader, length 3.7 to 4.0 times width. Genitalia (fig. 57, drawn from paratype, Fort

Valley, JAP prep. 3570; 2 n): sterigma with

subtending sclerotized folds, ductus bursae

sclerotized at base with weak enlargement;

signa small, spinelike.

Holotype male and allotype female, Arizona,

Fort Valley, 2,240 m elev., 12 km northwest of

Flagstaff, Coconino Co., July 10 and 20, 1961

(R. W. Hodges) (NMNH). Paratypes (29) all

Arizona, as follows: same data as holotype, 20d

VII-10-19-61, 29 VII-23-61; Hart Prairie,

elev. 2,590 m, 16 km NNW of Flagstaff, Coconino Co., 19 Vn-12-61 (R. W. Hodges); West

Fork, elev. 1,980 m, 26 km SW of Flagstaff,

Coconino Co., Id, 19 VII-15-61 (R. W.

Hodges); Madera Canyon, Santa Rita Mts.,

Santa Cruz Co., Id VII-27/30-47 (L. M. Martin); Rustler Park, elev. 2,590 m, Chiricahua

Mts., Cochise Co., Id, 29 VII-12/14-72, at

light (J. Powell) (LACM, NMNH, and UCB).

This species appears closely related to subcervinana, from which it differs by having narrower wings, paler colors (especially the hindwing), and by the more distinctly banded

forewing pattern. In subcervinana the pattern

consists of narrower, more irregular and less

defined lead colored and tan to orange bands.

There are differences in the sterigma as well.

Labial palpus size in pallifasciata appears to increase allometrically with increase in eye diameter and forewing length; this may be more

exaggerated in females, but the small sample

size limits conclusions.

In addition we examined two female specimens from Mexico representing a population

that is closely similar to pallifasciata and subcervinana. The specimens were collected 16

km west of El Salto, Durango, elevation 2,680

m, July 18, 22, 1964 (J. A. Chemsak and J.

Powell). They resemble pallifasciata in size,

wing shape, and color pattern but are much

darker, with dark gray hindwings as in subcervinana and bronze-brown forewings crossed by

rust-colored bands. The genitalia (1 n) are nearly indistinguishable irom pallifasciata.

28
GROUP 10: AUREOLA

Male larger than female. Antennal ciliation

of d short. Labial palpus elongate, II = 1.2 to

1.3 eye diameter. Male genitalia: sacculus

without projections, pollex broad, stout;

aedeagus elongate with apical spur and distal

dentation; 8 to 10 short cornuti. Female genitalia: sterigma funnel-like, with enclosed narrow fold, ductus bursae with slight accessory

lobe: signa small, one tiny , one larger.

Rhyacionia aureola Powell, New Species

(Figures 36, 37, 59, 83)

A Mexican species with rust-orange forewings crossed by many fine silvery striae, burnished golden to the unaided eye.

MaZe.—Length of forewing 7.0 to 8.4 mm (10

n). Head: Labial palpus moderately elongate,

II segment 1.2 to 1.3 as long as eye diameter,

III 0.3 to 0.4 as long as II; scaling broadly

spreading, whitish tan blotched with orange

brown exteriorly; scaling of upper front and

crown spreading into upraised tufts, whitish

tan tinged with brown. Thorax: Dorsal scaling

bright rust orange anteriorly becoming whitish posteriorly. Underside shining white, anterior and middle legs rust orange exteriorly.

Forewing: Moderately narrow; length 3.5 to

3.8 times width; costa gently curved, termen

strongly angled back, tornal angle scarcely

discernbile. Ground color rust orange with

about 15 evenly spaced, somewhat broken and

irregular, transverse striae of shining metallic

silver, reflecting rosaceous, whitish at costa,

crossing full width of wing. Fringe whitish, the

scales tipped with tan. Underside pale brownish gray, becoming whitish distally. Hindwing:

Slightly broader than forewing; apex acute,

termen slightly concave. Ground color pale

gray, slightly darker toward margins. Fringe

whitish. Underside white; fringe scales pale

ochreous at their base. Abdomen: Scaling entirely shining whitish. Genitalia (figs. 36, 37,

drawn from paratypes, JAP prep. 3568, 3636;

3 n): valva broad with a deep emargination

preceding cucullus not apparent on the unflattened valva (figs. 36, 37); cornuti a cluster

of 8 to 10 stout spines.

FemaZe.—Length of forewing 6.6 to 8.2 mm

(6 n). Essentially as described for male but

labial palpus slightly larger, II segment length

1.3 to 1.5 eye diameter; and forewing slightly

broader, length 3.3 to 3.6 times width. Genitalia (fig. 59, drawn from paratype, JAP prep.

3567; 3 n): ductus bursae sclerotized and enlarged basally; signa small, one twice the size

of the other, proportionately broader.

Holotype male and allotype female, Mexico,

13 road km west of El Palmito, 1,950 m elevation, Sinaloa, August 8 through 12, 1972 (J.

Powell, D. Veirs, and C. D. MacNeill) (CAS).

Paratypes (14) same locality: Id VII-29-64 (J.

A. Chemsak and J. Powell), 9d, 59 same data

as holotype (NMNH and UCB). All specimens

were taken at lights.

The host is unknown; the type series was

taken in a mixed madrone-pine woods. Three

species of pines were present in the immediate

vicinity, Pinus lumholtzii, P. montezumae,

and P. oocarpa, the last most abundant.

Four additional specimens (UCB) were examined but not designated as paratypes:

Five road km east of Revolcaderos, Durango. Id 19 VIII-11-72 (J. Powell). These specimens are in good condition and are indistinguishable on external features from typical

aureola. However, there are appreciable

differences in both male and female genitalia

(figs. 38, 58) especially the long, slender

aedeagus, which lacks the terminal spur, bearing instead a row of small teeth on its venter.

The female has a more elongate sclerotized

area at the base of the ductus. Both individuals

are slightly larger than the largest aureola

of the corresponding sex (forewing length in

each 8.7 mm). Based on knowledge of other

Rhyacionia these would seem to be speciesdistinguishing structural differences, but it

seems best to defer nomenclatural designation

until more information is available. This locality is only 30 road km (probably half that distance airline) from the type locality oí aureola,

and probably less than 300 m higher in elevation. There are nonetheless appreciable differences in the habitat and moth fauna sampled

at lights, so that a different pine host may be

involved.

Forty-eight km west of Durango, Dgo., 29

VII-31-64 (J. A. Chemsak and J. Powell).

These females are very worn, but the genitalia

appear to be essentially the same as the

29
Revolcaderos female (fig. 58). This locality is

125 km east of the Revolcaderos site, and

although higher (2,560 m) is well into the dry.

east or rainshadow side of the Sierra Madre

Occidental. The habitat is considerably different, an oak-juniper-scrub pine association.

GROUP 11: VERSICOLOR, FLAMMICOLOR

Female larger than male. Antennal ciliation

of ó moderately long. Labial palpus elongate,

II = 1.3 to 1.7 eye diameter. Male genitalia:

sacculus projection weakly developed, pollex

narrow and truncate or broad; aedeagus elongate without armature, cornuti none to eight.

Female genitalia: sterigma a broad ring with

transverse fold, ductus bursae with sclerotized

accessory lobe at base; signa tiny.

Rhyacionia versicolor Powell, New Species

(Figures 39, 60, 84; Map 13)

A large moth with shining pale forewings irregularly blotched with rust orange and tinged

with irridescent rose purplish in the distal

third.

MaZe.—Length of forewing 8.4 to 10.0 mm

(13 n). Head: Labial palpus elongate, II segment 1.3 eye diameter. III 0.4 as long as II;

scaling of palpus, upper front and crown

roughened, spreading, white tinged with gray

exteriorly. Thorax: Dorsal scaling shining

white; ventral whitish, prothoracic legs tinged

with gray. Forewing: Length 3.3 times width;

costa nearly straight, termen strongly angled

back, gradually curved to dorsum. Ground color whitish, irregularly overlaid with golden in

transverse striae and rust orange in broad

blotches, concentrated near base and beyond

middle, leaving an ill-defined, broad transverse band of parallel, whitish striae before

middle; an irridescent blotch of rose purplish

or pink on outer area of cell and surrounding

area below costa (paler and less discernible on

older specimens), blended with orange in its

basal half; costa white with brownish at origin

of the golden striae in outer half. Fringe and

underside whitish, lightly tinged with brownish. Hindwing: Pale gray, veins slightly

darker, fringe whitish. Abdomen: Scaling

probably shining whitish (greasy). Genitalia

(fig. 39, drawn from paratypes, Arizona,

NMNH slide 183, JAP prep. 3600; 3 n): valva

with broad, shallow emargination preceding

cucullus, sacculus with elongate, protruding

flange, pollex broad, round; aedeagus elongate, slender, tapered to a point on dorsal

edge; cornuti, eight stout spines.

Female.—Length of forewing 10.2 to 11.6

mm (5 n). Essentially as described for male except larger and forewing slightly broader,

length 3.0 to 3.1 times width. Some patches of

salmon-colored scales along Cu fold and beyond

in fresh examples; the golden banding generally broader and more contrasting than in male.

Genitalia (fig. 60, drawn from paratype,

Arizona, JAP prep. 3561; 2 n): sterigma a

broad rim around ostium, with a transverse

flap and subtending pouchlike base of ductus

heavily sclerotized; signa small spurs with

broad sclerotized bases.

Holotype male, Arizona, Fort Valley, elev.

2,240 m, 12 km NW Flagstaff, Coconino Co.,

June 29, 1961 (R. W. Hodges) and allotype

female, Arizona, Vail Lake Rd., 1,980 m, 15 km

SE Flagstaff, VIMl-61 (R. W. Hodges)

(NMNH). Sixteen paratypes: Arizona, same

data as holotype, 3d VII-10/12-61; Id West

..(>

Map 13.—Distribution records ior Rhyacionia versicolor.

30
Fork, elev. 1,980 m, 26 km SW Flagstaff, Coconino Co., VII-19-61 (R. W. Hodges); 2d Alpine

Divide Camp, 6 km N Alpine, elev. 2,590 m.

Apache Co., VII-15-65 (F., P. & M. Rindge);

49 Rustler Park, elev. 2,590 m, Chiricahua

Mts., Cochise Co., VII-3/20-72 at light (J.

Powell). Colorado, 6d Rock Creek Cyn., Colorado Springs, El Paso County, VI-23-59, VI27-60 (M. May) (AMNH, Kimball, NMNH, and

UCB).

This species resemblesflammicolor in external features (size, form of labial palpus, antennal pectination, wing shape) and in female

genitalia, but the male genitalia in versicolor is

distinct from all known Rhyacionia.

Rhyacionia flammicolor Powell, New Species

(Figures 40, 61, 85)

A large Mexican moth with shining dark red

forewings.

FemaZe.—Length of forewing 8.2 to 9.6 mm

(7 n). Head: Labial palpus elongate, II segment

length 1.5 to 1.7 eye diameter, III 0.3 as long as

11; scaling of palpus, front and crown broadly

spreading, grayish, the scales pale basally,

darker apically. Thorax: Dorsal scaling gray,

tinged with red brown anteriorly. Underside

shining whitish, anterior and middle legs gray

brown exteriorly. Forewing: Broad, length 3.1

to 3.2 times width; apex acute, termen strongly angled back, tornal angle evident. Ground

color shining metallic red, basal half with illdefined transverse grayish striae; discal area

beyond middle to beyond cell with a rosepurplish shade, terminal area deep red, becoming ochreous orange below costa and above tornus. Fringe gray. Underside gray, a pale line

around terminal margin. Hindwing: Apex

acute, termen strongly angled back, slightly

concave. Ground color dark gray; fringe pale

gray. Underside whitish gray, the veins

darker. Abdomen: Dorsal scaling dark gray,

ventral whitish gray. Genitalia (fig. 61, drawn

from paratype, JAP prep. 3644; 2 n): sterigma

a complex broad ring with transverse fold

anteriorly.

MaZe.—Length of forewing 9.2 mm (1 n). A

single, worn specimen essentially as described

for female except labial palpus relatively

shorter (eye larger), II segment length about

1.4 eye diameter. Genitalia (fig. 40, drawn from

allotype, JAP prep. 3432; 1 n): valva weakly

emarginate preceding cucullus, pollex well

defined, short, subtending an emargination in

the distal edge of the cucullus; aedeagus elongate, slender, strongly curved, cornuti absent.

Holotype female and allotype male, Mexico,

16 km west of El Salto, 2,680 m elevation,

Durango, August 18, 1964 (J. A. Chemsak and

J. Powell) (CAS). Six paratype females, same

data VII-18-30-64 (NMNH and UCB).

The host is unknown. The locality is a mixed

conifer and oak forest with 12 pine species, of

which Pinus durangensis and P. cooperi are

the most abundant (Thomas 1966).

The external features, particularly the elongate labial palpus, broad wings, purplish blotch

in the discal area of the forewing, and female

genitalia suggest this species is close to versicolor, but the male genitalia characters

differ. These two species show phenetic similarity in 12 characters (86 percent), dissimilarity in 2 (with no comparison in the 2

biological features) among the 16 characters

used in this study.

ACKNOWLEDGMENTS

We thank G. E. Daterman, D. T. Jennings,

M. E. McKnight, R. E. Stevens, and A. D.

Tagestad of the Forest Service U. S. Department of Agriculture, and W. H. Kearby of the

University of Missouri, who provided specimens and data from field studies. Cooperation

by André Blanchard, Houston, Tex.; Bryant

Mather, Jackson, Miss.; Gayle Strickland,

Baton Rouge, La.; John Heppner, Gainesville,

Fla.; and Charles Kimball, Barnstable, Mass.;

as well as authorities of institutional collections listed in the Introduction, enabled use of

specimens in their care. W. B. Critchfield,

Forest Service, U.S. Department of Agriculture, provided identifications of Mexican

pines.

The Forest Service provided facilities for

field work by California Insect Survey personnel at several ranger stations between 1959

and 1974, resulting in significant Rhyacionia

collections. Travel in Mexico during 1964, when

J. A. Chemsak and Powell collected Rhyacionia

31
as a byproduct to other field work, was funded

by the Naitonal Science Foundation grant,

"Carpenter Bees of the World" (P. D. Hurd,

Jr., principal investigator).

The maps were prepared by R. E. Dietz and

the drawings by Celeste Green, University of

California, Berkeley; their help is gratefully

acknowledged.

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33
MALE GENITAL CHARACTERS, FIGURES 3-40

Figures 3 through 7.—Male genital characters oíRhyacionicu (3) buoliancL, ventral aspect, valvae spread and (3a) aedeagus,

lateral; (4) buoliana, valva inner aspect, flattened; (5-7) rigidana, valvae inner aspect, flattened in (7), and (7a) aedeagus

venterolateral.

34
^rZf'íÍTnf '^;.~;^'^' ^"''^'"^ characters of Rhya^ionia: (8) ...6¿ropica, valva inner aspect and (8a) aedeaei

lateral; (9. 10) multilmeata, valvae inner aspect and Í9a) íipH^^anc lof^.oi ,„;.u .:*....• ....^T^^ ,,.1 ..... ^^^^^ ^^P®^^ ^'^^ ^^^^ aedeagus lateral with cornuti dehisced- (11 12) valvae inner aspect and (12a) aedeagus lateral. uemscea, ui.i^;

us venteroneomextcana,

35
Figures 13 through 16.—Male genital characters ofRhyacionia: (13) pcLsadenancL, valva inner aspect and (13a) aedeagus ventral; (14) zozanoj ventral aspect, valvae spread and (14a) aedeagus, lateral; (15) zozana, valva inner aspect, flattened;

(16) salmonicolor, valva inner aspect and (16a) aedeagus, lateral, ae = aedeagus, cZ = "clasper," en = cornuti, p = pollex,

V = valva.

36
Figures 17 through 20.-Male genital characters in Rhyacionia: (17) monophylliana, valva inner aspect and (17a) aedeagus,

dorsolateral; (18) adana, valva inner aspect and (18a) aedeagus lateral; (19, 20} jenningsi valvae inner aspect and (19a)

aedeagus lateral.

37
Figures 21 through 24.-Male genital characters in Rhyacionia: (21) busckana, ventral aspect, valva spread and (21a)

aedeagus lateral; (22,23) busckanoy valvae inner aspect; (24) blanchardi ventral aspect, valva spread and (24a) aedeagus,

lateroventral.

38
Figures 25 through 29.—Male genital characters in Rhyacionia: (25) fumosana, valva inner aspect and (25a) aedeagus,

lateroventral; (26,27)/iimosawa, valva inner aspect and (26a) aedeagus, dorsolateral; (28,29)/rusirana, valva inner aspect

and (29a) aedeagus lateral.

39
Figures 30 through 34.-Male genital characters in Rhyacionia: (30,31) sonia, valvae inner aspect and (30a) aedeagus lateral;

(32,33) aktita, valvae inner aspect and (32a) aedeagus lateral; (34) subcervinana, valva inner aspect and (34a) aedeagus

lateroventral.

40
Figures35 through 37-M I

41
vv^\^\^\vv \'

^'ÏZT ^^ ?r"f/ir^^'* f""^' characters in Rhya^ionia: (38) Revolcaderos "aureola." valva inner aspect and (38a)

42
FEMALE ABDOMINAL STRUCTURES, FIGURES 41-61

Figures 41 through 43.-Female" abdominal structures in Rhyacionia, VII sternite. sterigma and ductus bursae, ventral

aspect: (41) buoliana; {42) rigidana. with signa below; (43) subtrópico.

43
Figures 44 through 47.-Female abdominal structures in Rhyacionia, VII sternite, sterigma, ductus bursae. ventral aspect,

and signa shown separately 2x enlarged: (44) multilineata; (45) pasadenana, sterigma and ductus bursae only; (46)

zozana, including corpus bursae; (47) neomexicana.

44
51

Figures 48 through 51.—Female abdominal structures in Rhyacionia, VII sternite, sterigma, ductus bursae, ventral aspect,

and signa shown in corpus bursae or separately enlarged 2x: (48) martinana; U9) jenningsi; (50) busckana; (51)

blanchardi.

45
Figures 52 through 55.—Female abdominal structures in Rhyacionia, VII sternite, sterigma and ductus bursae, ventral

aspect: {52)fumosana; {5S)frustrana; (54) sonia; (55) aktita, with signa shown separately 2 x enlarged.

46
57

56

Figures 56 through 58.—Female abdominal structures in Rhyacionia: (56) subcervinana, VII to IX sternites and associated

genital structures with signa shown separately 3 x enlarged; (57)pallifasciata, VII sternite, sterigma, ductus bursae and

signa shown separately enlarged 2 x ; and (58) Revolcaderos ''aureola,'' VII sternite, sterigma, ductus bursae and signa

shown separately enlarged 2 x .

47
staJ.~i~^j--Â

59

Figures 59 through 61.-Female abdominal structures in Rhyacionia: (59) aureola, VII to IX sternites and associated genital

structures with signa shown separately 2 x enlarged; (60) versicolor, VII sternite, sterigma and ductus bursae- (61 )/Zammicolor, VII sternite. sterigma and bursa copulatrix. aJ. = accessory lobe, c.6.= corpus bursae, d6.= ductus bursae

sg. = signa, st. = sterigma.

48
ADULTS OF RHYACIONIA, FIGURES 62-85

Figures 62 through 69.—Adults of Rhyacionia: (62) buoliana d, Livingston Co., Michigan; (63) rigidana <3, Spring, Harris

Co., Texas; (64) subtropica 9, Everglades National Park, Fla.; (¡65) multilineata <3,10 km NW Newcastle, Wyo.; (66)poiodenaTui Ö, Lobos Cr.,San Francisco, Calif.; (67) zozana d, 5 km NE Tuolumne City, Calif.; (68) neomexicana 9, Sitgreaves

National Forest, Ariz.; (69) salmonicolor d, Davis Mts., Tex.

49
Figures 70 through 77.—Adults oíRhyacionia: (70) monophylliana <S, Chuchupate Ranger Station, Ventura Co., Calif.; (71)

martinana cJ, 8 km NW Prescott, Yavapai Co., Ariz.; (72) adana <S, Higgins Lk., Mich.; (73) jenningsi 9, Sitgreaves

National Forest, Ariz.; (74) busckana 9, Bluemont Jet., Va.; (75) blanchardi 9, Conroe, Montgomery Co., Tex.; (76)

fumosana 9, Slope Co., N. Dak.; (11)frustraría, Marysville, Pa.

50
Figures 78 through 85.—Adults oíRhyacionia: (78) bushnelli d. Fort Bayard, N. Mex.; (79) sonia 9, holotype; (80) aktita d,

Conroe, Montgomery Co., Tex.; (81) subcervinana 9, Vernon, British Columbia; {82}pallifasciata d, Fort Valley, Coconino

Co., Ariz.; (83) aureola d, 13 road km W El Palmito, Sin., Mexico; (84) versicolor d, Fort Valley, Coconino Co., Ariz.; (85)

flammicolor 9,16 km W El Salto, Dgo., Mexico.

51

irU.S.GOVERNMENT PRINTING OFFICE; 1978 0-244-306