BULLETIN OF THE ALLYN MUSEUM
THE ALLYN MUSEUM OF ENTOMOLOGY
Number 28 3 Sept. 1975
STUDIES ON NORTH AMERICAN PHILOTES
IV. Taxonomic and biological notes, and new subspecies
Department of Entomology, University of California, Davis, Calif. 95616
"Few butterflies are more completely 'one-plant insects' than members of the genus Philotes. Not only do
the larvae feed on the flower heads of Eriogonum, but the adults visit the flowers for nectar, so all of the food
requirements for both larvae and adults may be supplied by one species of plant" (Tilden, 1959: 261)
Philotes as applied in the broad sense is used here only for descriptive con-
venience to refer to battoides, rita, and enoptes on Eriogonum. The group formerly
known as the Nearctic "Philotes" has been shown (Shields, 1974b) in fact to be a
group of sibling species representing Philotes sonorensis (Felder & Felder, 1865),
Zizeeria speciosa (H. Edwards, 1876), Shijimiaeoides rita (Barnes & McDunnough,
1916), S. enoptes (Boisduval, 1852) and Caleta battoides (Behr, 1867). Since then,
however, a hybrid specimen of battoides glaucon X enoptes ancilla has come to my at-
tention, a male from Pattee Canyon, 3500', Missoula Co., Montana, VI-23-62 (J.
Scott). Face on, the left valve is like battoides and the right valve (though
somewhat shorter) is like enoptes! Also, two males from 8 mi. N. of Jarbidge,
Elko Co., Nevada, VI-28-67 (J. Lane) appear to have somewhat intermediate
genitalia between battoides glaucon and enoptes ancilla (i.e., both have bifurcate
valvae but not as deeply as is usual in battoides, and one has prominent teeth in a
row, as in enoptes, along the concave surface. Thus it would appear likely that these
three specimens are hybrids between b. glaucon and e. ancilla, two subspecies
that are nearly identical in adult faces and which utilize the same hosts,
Eriogonum umbellatum and E. heracleoides. Consequently, battoides should
belong in the genus Shijimiaeoides along with enoptes and rita, and should be
dropped from the genus Caleta where I had tentatively placed it. A close relation-
ship may still exist between Caleta and battoides, but it now appears doubtful
they should reside in the same genus.
P. sonorensis and Z. speciosa are readily distinguishable on adult facies
alone, but an occasional dwarf form of enoptes is mistaken for speciosa, and S.
rita, enoptes and battoides are often so similar in wing characters that only recourse
to their distinctive male (or female) genitalia will place them to proper species
(see Mattoni, 1957). In this study, it was first necessary to examine the male genitalia
of all named subspecies, to develop a recognition for the accompanying slight
facies differences and to associate each form with its proper Eriogonum host(s).
Similarly, genitalia of representatives of every new, unplaced population found
during the course of study had to be examined to avoid errors. Only after this
procedure was strictly adhered to for several years did I allow myself the luxury
of determining some populations on faces alone, having then a practiced eye for
subtle cues. Even at this stage, however, it was still necessary to examine genitalia
of exceptionally close members such as battoides glaucon-enoptes ancilla-e.
columbiae and b. comstocki-enoptes nr. enoptes (Tehachapi Mts.). Some 1250
male genitalia were determined and approximately 9,000 specimens were ex-
amined in all. Genitalia of representative specimens from many localities were
examined (exceptions are some enoptes mojave and rita spaldingi) by the procedure
outlined in Shields (1973).
The male genitalia of S. battoides and S. enoptes are adequately figured in
Langston (1963). (See also, for battoides, the figure 23 in Brown, 1970; pl. 17, figs.
3 & 4 of Barnes & McDunnough, 1917; and pl. V, fig. 3 of Williams, 1918. For enoptes,
see Williams, 1918, pl. 4, 5, fig. 6-10; and Barnes & McDunnough, 1917, pl. 17, figs.
1 & 2.) The male genitalia of rita are pictured in Barnes & McDunnough, 1917, pl.
17, figs. 5 & 7; Mattoni, 1965, figs. 2, 3, 4; and drawn in Tilden & Downey, 1955, pl.
11. Barnes & McDunnough, 1917, pl. 17, fig. 6, picture spaldingi's male genitalia
but at such an angle that the rudimentary cristae are not visible. Shijimiaeoides
r. spaldingi appears to be the link between the rita type of genitalia (with cristae
always present) and the enoptes type (its valve being close to enoptes ancilla and
e. dammersi-i.e., terminal end of valve with blunt teeth instead of spines, and
general shape of the valve as in enoptes). No enoptes subspecies possess cristae.
Clench's view (1967) that the pallescens-elvirae group and the rita-coloradensis
group may form distinct species is untenable in the light of recent discoveries
reported here. Mattoni (1954: 164) using genitalia differentiates female enoptes
from female battoides, and female enoptes from female rita (1965: 86).
The number of teeth on the terminal end (cucullus) of the valve in the rita-
enoptes group, for 59 populations and 17 subspecies sampled (N = 318), varied from
a low of 9 to a high of 35, the lowest being enoptes ancilla (averaging 11) and
the highest rita rita and rita coloradensis (averaging 26). The number of teeth
was diagnostic for each subspecies.
The terminology used in describing Philotes follows that of Nabokov (1944,
plate 5) and Mattoni (1954, 1965).
Some authors suggest combining the Nearctic species of Shijimiaeoides with
the Mediterranean-European genus Pseudophilotes (vicrama extends to Central
Himalaya and Tibet) by virtue of their similar faces. The two genera are no doubt
closely related, as the genitalia of vicrama is like enoptes with the "neck" twisted
on the valve (with bavius probably a vicrama derivative), and the baton genitalia
is quite similar to battoides except the bifurcation is upturned (with abencerragus
probably a baton derivative). This reverses my previous opinion (Shields, 1974b)
that was based on Hemming's drawings (Beuret's sketches are more instructive).
However, the length X width ratios of the aedeagus of the two genera differ con-
siderably (Pseudophilotes abencerragus 0.24 (Fig. 37) P. vicrama 0.68 (Fig. 34)
P. baton 0.50 (Fig. 36) P. bavius 0.25-0.33 (Fig. 35); Shijimiaeoides divina barine
1.73 (Fig. 40), S. enoptes 1.43) (Beuret, 1958). In this respect, Shijimiaeoides is
closer to Glaucopsyche (G. lygdamus 1.31, G. alexis cyllarus 1.77). Likewise, the
length X width ratios of the androconia differ in P. abencerragus (2.60) and P.
bavius (1.81-2.41), but are similar in P. baton (1.34-1.24), P. vicrama (1.19), S.
divina barine (1.16) and S. enoptes (1.10). The male androconial scales of S.
divina barine and S. enoptes enoptes are very similar in form, rib number, and knot
number but differ in size (barine = 32mm, enoptes = 19mm in width) ard are similar
to vicrama and baton but not to abencerragus and bavius (Beuret, 1958). In view of
these morphological differences in genitalia between Pseudophilotes and
Shijimiaeoides (and foodplant differences), I concur with Beuret in designating
them as distinct genera. On the basis of male genitalia, other genera that seem
closely related to Shijimiaeoides are Palaeophilotes, Praephilotes, Scolitantides
(orion), Sinia, Shijimia, Glaucopsyche (via atroguttata Obth. and coeligena Obth.),
Actizera (via panagaea H.-Sch.), Lycaena, Turanana (cytis), and Zizeeria (via
speciosa (Figs. 43-44). Brown (1971) places piasus in Glaucopsyche, a decision I
am in agreement with. (I think "Shijimiaeoides" lanty Obth. should belong in
Glaucopsyche because of the shape of the male valve.) Robinson (1971) reports
that the modal haploid chromosome number for Lycaenidae is n = 24, with
Glaucopsyche, 4 sp., n = 23; Lycaena, 7 sp., n = 24 (except heteronea and rubidus);
Pseudophilotes (baton, bavius, vicrama), n = 24; Scolitantides orion, n = 23;
Turanana panagaea, n = 24; and Zizeeria maha argia, n = 24.
Beuret (1958: 83) states that the uncus, tegumen, and lateral uncus hooks of
Sinia are near Shijimiaeoides (also the valve tip recalls enoptes, though much
reduced). However, it shows enough differences from Shijimiaeoides in length X
width ratios for the androconia and aedeagus to be properly kept as a separate
genus. In adult facies, Sinia clearly approaches Shijimiaeoides (see Beuret, 1958:
78; fig. 26-27). Sinia leechi Forst. (Fig. 36) is known from China (Setzschwan:
Kwanhsien) and is very local. Much confusion has centered about the names S.
leechi and Shijimia moorei Leech (Fig. 37). Beuret was unaware that all his dis-
cussion, figures, and statistics for Shijimia moorei in his 1958 paper referred instead
to Sinia leechi (=moorei auct., nec Leech), but after consulting Forster (1940),
he published a correction in 1959.
Jamides pura (Moore) has a very similar bifurcate valve to S. battoides (see
Eliot, 1973, fig. 95). Another candidate as a close battoides relative is the rare
Shijimia moorei. The male genitalia, as illustrated in Forster (1940: 878), has
strongly bifurcate valvae, with the rest of the genitalia somewhat similar also,
although apparently modified somewhat from the battoides' ground-plan. The
brief description in Seitz of the adult seems also to indicate that moorei belongs in
a separate genus from battoides. Shijimia moorei flies in central China (Kiukiang
and Chang-yang in June and July), southeast Chia (Fuchien: Kuatun), and
With Pseudophilotes on Thymus and Salvia, perhaps a nectar source became
a larval foodplant (from the unrelated Eriogonum). (This idea is attractive because
the Eriogonum serves as both larval foodplant and nectar source in Shijimiaeoides.)
There are several records of lycaenids imbibing nectar from Labiatae (Monarda,
Monardella) (Shields, 1972), although I have not seen any doing so on Salvia in
Larvae of Pseudophilotes vicrama cashmirensis (Moore) are found particularly
on the flowers of Thymus (Seitz, 1909:306). Colin Wyatt (in litt.) has found larvae of
Pseudophilotes bavius fatma (Oberthur) in the Middle Atlas of Morocco feeding
on the buds and flowers of a large Salvia species with hairy leaves spread out in
a rosette around the base of the plant; adults were "very local and only around the
foodplant." He reared a series of adults from the larvae.
Shijimiaeoides divina flies in Japan and Korea and feeds as larvae on
Sophora (Leguminosae) (Shields, 1974b). The species is likely Sophora flavescens
Ait., a perennial herb with many pale greenish-yellow flowers that blooms in
June and July (Ohwi, 1965). The other two Sophora species in Japan seem unlikely
hosts: S. franchetiana is very rare on Kyushu and China, and S. japonica is
a Chinese tree planted in Japan. S. flavescens is common on Honshu, Shikoku,
and Kyushu, and ranges to Korea, China, and Siberia (Ohwi, 1965). Today, five
species of Sophora inhabit the United States and Mexico, and 22 species are scatter-
ed through the warm parts of the world (Sargent, 1890: 59-60). Fossil Sophora date
back to at least Paleocene times in North America (Lamotte, 1952).
ERIOGONUM AND THE COLORADO PLATEAU
The plant genus Eriogonum (Polygonaceae) is a large and complex group
consisting of some 240 species. It occurs mainly in western North America from
northern Mexico to Alaska, with species on the islands in the Gulf of California,
in Baja California, and the islands off the coast of Baja and California to the
west (Reveal, 1969a,b). At least one species, E. tomentosum, occurs along the
eastern Coast Plain of the U.S., E. longifolium is found from Texas to Florida,
and E. allenii in Virginia and West Virgina. References to Eriogonum species in
South America are unfounded.
Most species occupy an arid or semiarid climate (Reveal, 1969b: 232).
"Eriogonum typically occurs in micro-environments where water is more abundant
than elsewhere over long periods of the season," such as road shoulders, washes,
and open exposed slopes (Reveal, 1968). Since polyploidy is a sign of ancestry,
Eriogonum probably originated early as Reveal notes (1969b) that "the vast
majority of the modern species are tetraploids." Eriogonum species lack an active
dispersal mechanism; the small seeds usually fall within a few meters of the
parent plants (Reveal, 1969b: 233).
The most primitive Eriogonum subgenus, Eucycla, has about 14 sections and
some 105 species (Reveal, pers. comm.). From a brief analysis of the first 10
sections (90 sp.), by far the greatest area of concentration is Utah with 21 species,
mostly endemics. (According to Stebbins & Major, 1965, p. 27, floras in arid regions
"are likely to be relatively stable, and most of the endemic species are ancient or at
least not recent.") Two-thirds of these 21 species plus four that also range into
western Colorado are located east of the major NE-SW Cordillera on or near the
Colorado Plateau in Utah, thus a likely place of origin. Going away from Utah,
the first 10 sections have species confined to other states but only 3-4 species each
(in Arizona, Colorado, Nevada, Baja California, Oregon, except that California
has ca. 8 species). The first 10 sections generally range over the western United
States west of the continental divide, including island forms, and Baja California,
Mexico. S. rita and S. enoptes subspecies are largely confined to Eucycla, as are
about half of the S. battoides subspecies.
The Colorado Plateau is a great, high-standing crustal block, uplifted 5000
feet since the Eocene, nearly square and 500 miles on a side, whose center is close
to the Four Corners area. By what process a block of the earth's crust the size of
the Colorado Plateau is uplifted a mile is quite uncertain. The crustal thickness of
the Plateau averages 45 km, greater than the average for the western United
States, so it has probably thickened, giving rise to the uplift (Gilluly, 1963: 156-157).
In eastern Utah, today a desert region, the Uinta Basin on the northern border of
the Plateau has Eocene overlying Paleocene sediments, and Paleocene sediments
along the Plateau's western border. However, the Plateau from the eastern and
southern borders of Utah to the edge of these Early Tertiary sediments consists
largely of thick Lower and Upper Triassic redbeds plus Upper Jurassic dune deposits
formed in a vast desert, with Upper Cretaceous plastic wedges in some areas,
overspread when the Plateau was near sea-level. Rocks of the stable crustal block
have been flexed or folded, but gently and on a grand scale, apparently due to the
Laramide Revolution, a time of mountain-building in the Rocky Mountain area
from Late Cretaceous to Oligocene times. The Colorado Plateau is a region of
plateaus, escarpments, and canyons laid out on a grand scale. The Colorado River
apparently did not cut through it until the Late Tertiary and has denuded the
Mesozoic-Tertiary sequence in the Grand Canyon. (Most of the above discussion
was extracted from King, 1959). Perhaps the subgenus Eucycla in eastern Utah
dates back to the Triassic-Jurassic desert.
Some authors (e.g., Hunt, 1946: 22-23; Gregory, 1938: 94; 1951: 94) reason that
Cenozoic formations have been largely stripped away from the interior of the Color-
ado Plateau by erosional processes. Hunt (1956: 71) estimates that about 80 per cent
of the Colorado Plateau has been eroded to formations below the middle of the Upper
Cretaceous, about 60 per cent is lower than the base of the Cretaceous, about 35
per cent is lower than the base of the Jurassic, and about 25 per cent is lower than the
base of the Triassic. The total Cenozoic degradation on the Plateau averages
about 10,000 feet (Hunt, 1956, p. 71), making the stripping of the Plateau lands a
major event in the geologic history of North America (Gregory, 1951: 94). "That
sediments and lavas of Tertiary age once covered all or part of this region [south-
eastern Utah] is suggested by their presence in adjoining regions--the Animas
and Mancos Valleys of Colorado, the Chuska Mountains of New Mexico and
Arizona, the Uinta Mountains and the High Plateaus of Utah" (Gregory, 1938:
63). Also, Cretaceous strata appear near the Henry and La Sal Mountains (Fisher,
Erdmann, & Reeside, 1960). Gregory (1938) notes that today most rains of the San
Juan country are short-lived, widely spaced, torrential downpours that run-off
from bare rock and are thus affected by evaporation (dry air, and spring and summer
high temperatures). The falling rain gathers at once into gullies and flows swiftly
to larger drainage channels. Frost is an active agent. Average annual precipitation
totals in the southeast Utah region are Hanksville (4200') = 5.41", Bluff (4200')
= 6.26", and Kayenta = 8.86". Precipitation falls in the winter and July-September.
Characteristic climatic features are the great range of diurnal temperature and
the prevalence of clear skies. There are years of heavy rain and years of drought. A
post-Eocene to Miocene (?) precanyon erosion cycle, as evidence by the "rock
stairway" topography from the Grand Canyon to the High Plateaus of south Utah,
is thought responsible for stripping away most of the Tertiary, much of the Mesozoic,
and some of the Paleozoic on the Colorado Plateau (Gregory, 1951: 83). Hunt
(1956: 71) states that the erosion rate of bedrock on the Colorado Plateau, as deter-
mined by sedimentation studies at Lake Mead from 1935-48, would take 21.5
million years to strip 10,000 feet of rock from the Plateau; degradation was dominant
during about half of the Cenozoic.
However, a case could also be made for Cenozoic strata never being present
over much of the Plateau, with Mesozoic strata eroded in situ during the Cenozoic.
The rate of chemical and mechanical weathering is extremely slow in the desert
because of the lack of moisture and takes place by sand-blasting (abrasion and
deflation), cloudbursts, frosts, and wide temperature fluctuations, so soils do not
form except for sand (Leet and Judson, 1965: 205-206). It thus appears incongruous
that two miles of sediments in 21.5 million years could be eroded away in a region
that in Mesozoic and Recent times was desert (and may also have been in the
Cenozoic). Axelrod (1972) notes that tropics and deserts are existing side by side
today in some regions and no doubt did so in the past. Why were the Paleocene-
Eocene strata that are exposed on the northern and northwestern edges of the
Colorado Plateau not eroded away as well? They are in closer proximity to the
moist Wasatch and Uinta Mountains where streams and rivers flow and where
rainfall is high. It would appear ironic that only Early Tertiary sediments close
to the mountains are preserved when erosion was stripping most of the Colorado
Plateau. Why are not the High Plateaus also denuded of their capping Tertiary
sediments, especially since they receive higher rainfall amounts? Perhaps the
Lower Jurassic and Lower Cretaceous sequences missing in eastern Utah were
periods of degradation as is the Cenozoic degradation perhaps occurring on a desert,
so that aggradation is not noted in these periods. I thus favor a view of desert
conditions persisting in southeastern Utah from the Mesozoic to the present.
Axelrod (1970) reasons on the basis of some desert floral links between North
and South America and from the fossil record that an archaic element was present
in American deserts prior to Tertiary and Late Cretaceous times, as tropical forests
and savannas had prevented any exchange between the Americas during the
Cenozoic. Raven (1963) notes two Chorizanthe species and one Oxytheca (both
close relatives of Eriogonum) with such a desert distribution today. Axelrod
(1970) argues for angiosperm links between dry areas of the Southwest U.S. and
Mediterranean being "the remnants of a dry flora that inhabited Gondwanaland
prior to its breakup by ocean-floor spreading near the close of medial Cretaceous
time" (p. 309).
The North Atlantic probably began to form in the Upper Jurassic; this age
is reported from the northwest coast of Africa and the coast of Spain (Ramsay,
1971, fig. 1). (For the fit of the continents around the Atlantic, see Bullard, Everett,
& Smith, 1965). Likewise, the North Pacific's oldest seafloor age appears to be
Upper Jurassic, at ca. 140 m.y. (Fischer, et al, 1970; Dietz & Holden, 1971). A major
expansion of the earth since Mesozoic times by seafloor spreading appears likely
(Meservey, 1969). For the reasoning behind the expanding-earth hypothesis, see,
e.g., Egyed (1957), Carey (1958) and Holmes (1965, Ch. 27). The presence of genera
related to Shijimiaeoides in eastern and western Asia suggests that this genus
complex had evolved prior to continental separation in the Upper Jurassic.
It is customary to view the Lepidoptera as quite young (Cenozoic), based on
their meager fossil record. However, Forbes (1932) cogently argues that the
Lepidoptera arose in the late Carboniferous or early Permian with the first flowers,
with butterflies (except Lycaenidae and Riodinidae) arising in the Jurassic with
Papilionidae, and Hesperiidae before that. Foodplants of some groups of primitive
Lycaenidae include lichens, algae, cycads, conifers, etc. and suggest to me that
their origin was early, in the late Paleozoic.
During the past two decades, much debate has hotly centered around the
worth of the subspecies concept. Even the validity of the species concept has been
called into contention by some of the more outspoken numerical taxonomists in
recent years. Perhaps the most sobering review of the subspecies "problem" is
that of Tilden (1961), who sagely remarks that "defenders of the subspecies are
more numerous among entomologists who work with the more plastic groups of
insects." I must say that from my training, personal experience, discussions with
fellow lepidopterists, and knowledge of the literature, I am strongly in favor of
naming subspecies although drawing the line short of extreme splitting such as
practiced by some European workers. I certainly do not hold with the negative
attitudes of certain numerical taxonomists toward species and subspecies con-
cepts, which if allowed to prevail unchecked would no doubt tear down the hierar-
chical superstructure of taxonomy. Numerical taxonomy can be used as an adjunct
to classical taxonomy (see, e.g., Dobzhansky, 1933; Lidicker, 1960), but hardly
as a replacement.
Downey (1956) was faced with much the same problem that one faces with
Philotes in dealing with the Plebejus icarioides complex, i.e., a highly variable
species to study and render more intelligible in evolutionary terms. His approach
was to analyze the adult characters and run population comparisons both
qualitatively and quantitatively, as he thought the variation too great for even
subspecific ranking. The result was the characterizing of populations by the sum-
mation of qualitative and quantitative differences. He admits that the statistical
differences in populations could not be used to approximate subspecific distinc-
tions. Perhaps had he tried correlating the adult variation with Lupinus foodplant
specificity, much of the complexities might have been resolved.
Bogert, et al (1943) discuss the different criteria for subspecies in various ver-
tebrates. In entomology the criteria for subspecies are usually not nearly as well
defined as for the vertebrates, with interpretations often varying depending on the
group (see Thorpe, 1930; Linsley, 1944; Remington, et al, 1951). For the purposes
of the present study, I have adopted Klots' (1936, p. 167) concept of "food-plant
subspecies." In Philotes, there appear to me to be numerous distinguishable sub-
species that are each specific to a particular species (or in some cases two or more
species) of Eriogonum, although there are rare exceptions. This rule of thumb
clears up almost all of the complex problems of variation in this group that the
taxonomist is otherwise faced with.
Problems in naming clinal variation are discussed by Mayr, Linsley, &
Usinger (1953), Hubbell (1954), Tilden (1961) and Cohn (1965: 9-10).
SIBLING SPECIES AND MIMICRY
Brown (1959, p. 80) defines sibling, or cryptic, species as reproductively isolated
populations that lack obvious distinguishing characters, and reviews the taxonomic
problems of closely related species. Sibling species (i.e, those that fit these criteria)
in Philotes are the Shijimiaeoides rita-S. enoptes-S. divina-S. battoides complex.
Brown further states that sibling species are like other species except they are
more difficult to segregate, especially when sympatric. In Shijimiaeoides, proper
identification can readily be made by examining the genitalia in cases where the
adult faces are nearly identical. Papers dealing with sibling species' examples
in Lepidoptera are Corbet (1943), Rawson & Ziegler (1950), Klots & Clench (1952),
Smith (1953, 1954), Remington (1953), dos Passos (1969), Carde, Shapiro, & Clench
(1970), Shapiro & Card6 (1970) and Heitzman & dos Passos (1974).
A recent review of the sibling species problem, with numerous examples, is
that of Mayr (1966: 33-58, 460-461). He states (p. 57) that recent work in genetics
has found much evidence for "a selective premium on the maintenance of the phe-
notype" in sibling species. This selective factor in Philotes presumably is a mimicry
condition, with the Plebejus acmon-lupini complex the model, and Shijimiaeoides
rita, S. enoptes and S. battoides the mimics. Here, S. rita does not greatly resemble
in faces its immediate ancestor, Z. speciosa, which uses other genera as well as
Eriogonum, but it does closely resemble Plebejus acmon-lupini (genitalically and
phylogenetically quite unrelated) and rita's derived species, S. enoptes and S.
battoides, both only on Eriogonum. Often two of the three species (battoides,
rita, enoptes) are sympatric and synchronic. S. rita spaldingi is distinct from all
the other Philotes and bears a strong resemblance to Plebejus melissa on the un-
derside (Langston, 1969), and the two are often sympatric though using different
foodplants. Striking resemblances also occur on the underside between S. battoides
intermedia and Plebejus lupini, and S. enoptes tildeni and Plebejus acmon,
which fly together on the same foodplant, respectively. Also, amazingly close in
undersurface faces are S. enoptes dammersi and S. rita rita (both on E. wrightii)
and S. battoides glaucon and S. enoptes ancilla (both on E. umbellatum), but in
both cases the species are allopatric. This close resemblance I feel is due to a common
origin, the former derived from the later in both cases. The selective effect of a
nearly identical environment may be responsible for some similarities, rather
than mimicry or common ancestry (see Hovanitz, 1941; Ali, 1969).
The original similarity to produce Miillerian mimicry arose by convergence,
then continued similarity is maintained by parallel evolution (Brower & Brower,
1972: 64), as perhaps has occurred in the Philotes mimic tending toward a Plebejus
model. Genera with somewhat similar faces to Shijimiaeoides but with wholly
different genitalia are Talicada nyseus Guer. (India, Ceylon, Burma), Chilades
laius Cr. (Indo-Oriental) and Luthrodes Druce (Indo-Australian). These genera
also perhaps served as models in the distant past for Philotes to mimic and have
since become extinct from the North American continent.
According to Janzen (1968), "A species of plant is an island in evolutionary
time to the insect species that feed on it." In Eriogonum with limited dispersal
ability and Philotes with the whole life cycle (reproduction, adult nectar source,
larval food requirements) of each subspecies usually confined to a particular
Eriogonum species, the direction of evolution in Philotes should parallel the evolu-
tionary development of the Eriogonum through time for those species it uses today
(i.e., after "jumps" have been recognized, from a perennial onto an annual most
frequently). "Allochronic isolation on unrelated plants with different fruiting
[or flowering] times" (Bush, 1969: 250) appears to have taken place from S. enoptes
dammersi on E. wrightii (fall) to enoptes mojave on E. pusillum (spring). In other
Lepidoptera that are not so host-specific, the evolutionary pattern would not be
nearly as discernable. For example, the moth Panaxia dominula L. uses many
different plant genera in 15 families (Cook, 1961). Downey (1962) discusses the
taxonomic usefulness of host-specific species.
Thorpe (1945) regards geographical, topographical, and ecological isolation
as three different scales of spatial isolation. He states, "Habitat change may lead
to topographical isolation just as topographical isolation may lead to habitat
change." As one example of how an initial separation or fission of a population
might take place, Thorpe gives a host-plant preference in which the progency
become selectively adapted and isolated on the new host-plant. Hovanitz has shown,
in a series of controversial experiments, that Pieris larvae reared on a variety of
crucifers preferred to oviposit on the particular larval host they were reared on
(see references in Shields, Emmel, & Breedlove, 1969). In Philotes, adaptation to
a new host would be manifest as foodplant "jumps" evolutionarily, while in most
cases a continuum of subspecies can be traced geographically which use phylo-
genetically related Eriogonum species, suggestive of an initial coevolutionary
development, with Philotes entering a new geographic, topographic, or habitat
niche as the Eriogonum did so. The resultant habitat isolation would then restrict
gene flow and a new subspecies would develop.
In no instance in my experience with Philotes were two or more species using
the same Eriogonum species at any given locality. Invariably where two species
were sympatric and synchronic, each occupied its own particular Eriogonum species
within the same habitat. However, in three instances, two Philotes occupied the
same Eriogonum species allopatrically. These were: S. enoptes ancilla (east) and
S. battoides glaucon (west) on E. umbellatum, E. wrightii with rita rita (east) and
enoptes dammersi (west), and enoptes smith (north) and battoides allyni (south)
on E. parvifolium. In the first case they come at least to within 30 air miles of
each other in extreme western Nevada, in the second case at least to within 5-10
miles in the vicinity of Cherry and Hillside, Yavapai Co., Arizona, and in the third
instance to within 50 miles.
This sort of host-use pattern sketched above strongly suggests that competitive
displacement or exclusion is taking place or has often already occurred. If otherwise,
at least some instances of sympatric-synchronic coexistence on the same Eriogonum
host by two or more species would be expected, as in fact exists between certain
Philotes populations and Apodemia mormo (for foodplants, see Opler & Powell,
1961), Plebejus acmon and P. lupini (for foodplants, see Goodpasture, 1974).
For a thorough review of the competitive displacement or exclusion concept, see
Hardin (1960), De Bach & Sundby (1963) and De Bach (1966). According to De
Bach (1966; 191), "Most cases of competitive displacement in nature have already
gone to completion."
According to Langston (1963), the appearance of Philotes adults is correlated
with the early full-bloom of the Eriogonum. However, it has been my experience
with S. rita pallescens at two Nevada localities that a good flush of fresh adults
occurs when plants of E. kearneyi are just coming into bloom, and most of the plants
are still in bud. Philotes adults seem most common when their Eriogonum
host is just coming into bloom to full bloom. By the time the Eriogonum plants mostly
reach the full bloom stage, many of the Philotes are worn. Adults are scarce or
lacking by the time the plants are in past bloom condition. Larvae feed on the flowers.
Pupae have been found in sand at the base of Eriogonum in one battoides, one
rita, and one enoptes colony. Each Philotes subspecies' life cycle from emergence
to pupation is synchronous with that Eriogonum's blooming period (Tilden, 1959:
261). When precipitation was recently rather great or abundant, it triggers the
Eriogonum to bloom and perhaps initiates the Philotes emergence as well (Reveal,
"The termination of diapause in response to environmental stimuli syn-
chronizes the appearance of reproductive forms at a favorable season" (Harvey,
1962). In Philotes, the synchrony is with the blooming time of the host Eriogonum,
different for each species. Stimuli producing the onset of diapause include photo-
period, temperature, and water, principally photoperiod (Harvey, 1962). Dia-
pause may be induced in any developmental stage, and all stages except the
pupa may be receptive to photoperiod (de Wilde, 1962). There is photoperiodism
in plants, and in one case with subterranean aphids living at 1'/ to 2 m in the
soil, photoperiod induction of sexual forms was from a "signalling" action
via the foodplant (De Wilde, 1962). A similar case has been reported for larvae
of the Cabbage-root fly, Hylemyia brassicae (Danilevskii, 1965: 80-81).
Perhaps a similar situation exists between the Eriogonum plant and Philotes
pupae beneath it in the soil. Cessation of diapause might be caused by hot tempera-
tures following a rainy period, or may also be triggered by some signalling action
of the plant such as chemical changes.
Philotes ovipositing females ranged from every hour between 8:30-3:30
(PST & MST, N = 36), with ca. 75% occurring in the morning. Females oviposited
mainly on unopened buds and inside flowers, frequently on the inside of the sepals,
after much abdominal probing, walking over the flowers, and turning around on
flowerheads. Nearly all day long, adults fly around, feed on, and alight to rest on
the Eriogonum flowers. They fly from plant to plant in a rapid, erratic manner
over the blooms. S. rita emmeli starts flying at 7:00-8:00 AM and seeks roosting
sites around 6:00 PM MST in southeastern Utah. Adults frequently roost on the
Eriogonum blooms for the night but also on other shrubs (Shields, 1973). Males and
occasionally females will come to moisture but do not seem to form mud-puddle
"clubs". Occasionally adults are to be found feeding on other flowers (including
other Eriogonum species, two yellow-flowered composite species, yarrow, aster,
a white-flowered crucifer, and Calyptridium sp.) especially when their host
Eriogonum is scarce that year or in past bloom, but this is rather exceptional. The
flower colors of the Eriogonum species used for foodplants (as well as for nectar
sources) include yellow (many)-cream-white (many)-pink-rose. Adults are often
common and could play a sizable role in Eriogonum pollination in some areas,
along with certain other butterflies, Hymenoptera, and some Diptera. Mating
apparently takes place on the Eriogonum plant; in copula pairs were noted at this
site at least from 9:00-4:30 (PST & MST), mostly from ca. 10:30-2:30 (N = 42)
(Shields & Emmel, 1973). Not uncommonly the female of an in copula pair is worn,
indicative perhaps of the occurrence of multiple-matings.
ABERRATIONS AND SEX RATIO
Various aberrants are met with not uncommonly in "Philotes" (est. less than
1 per cent). These include macules fused or absent, macules present on one side
and lacking on the other, wing-shape asymmetry, an occasional striking melanic
or "blond" phase, an extension of the aurora in one cell toward the wing base,
dwarfs and giants. Individual variation within a given "Philotes" population and
variation within the range of a given subspecies are often considerable, especially
on the undersurface (e.g.pallescens, glaucon, bernardino).
During this study one gynandromorph was discovered, an S. enoptes ancilla
from Sybille Canyon, 6500', Albany Co., Wyoming, 4 July 1966 (R. Hardesty).
The specimen is mostly a female with a strip of male blue in a narrow band on the
right hindwing upperside only, from the base to the margin, not present on the
underside. The wings otherwise are uniform brown except for the aurora.
A total of 51 males and 50 females of S. rita pallescens emerged over a month's
period under uncontrolled lab conditions (from 6:00-8:00 AM PST), from pupae col-
lected at Sand Mountain, Churchill Co., Nevada, by E. M. Perkins and myself in
early July, 1974. Similarly, 6 males and 7 females of S. battoides allyni emerged
from pupae (from ca. 10:00 AM-1:00 PM PST) collected at El Segundo, California,
by Perkins. These data help confirm field records of a probable 1:1 sex ratio in
TAXONOMY AND NEW SUBSPECIES
In this section, seven new subspecies are described. Two of these (battoides
allyni and b. comstocki) had been recognized by other workers (unpublished) as
distinct, while the remainder were either classed as known subspecies previously
or were discovered as new during the course of this survey. At least two other new
subspecies will be described in a future paper. The taxonomy of the parent species
battoides, rita, and enoptes is discussed, as well as spaldingi and mojave. Taxonomic
notes and biological and distributional data for the other known subspecies of
Philotes will appear in a future paper.
Shijimiaeoides battoides battoides (Behr), 1867
Behr. 1867. Proc. Calif. Acad. Nat. Sci. 1863-1867, 3: 282; no. 15.
Behr states, "This species was collected at an elevation of eleven thousand feet,
on the head-waters of the San Joaquin River, by the Geological Survey." At this
elevation in the Sierras, only S. battoides battoides and occasionally S. enoptes
enoptes fly, the former much more abundantly. The description in Latin refers
to quadrangular black spots on the underside, the characteristic square-spotting
of what today is referred to as S. battoides battoides. Further, McDunnough (1914)
notes that Behr very aptly compared it with the European battus (= Scolitantides
orion Pallas). Without more to go on, the type locality must lie between Yosemite
National Park and Kings Canyon Nat'l Park, Madera and Fresno Co., at 11,000',
where it flies abundantly at this elevation. Skinner (1911) presumed that the type
was destroyed in the 1906 San Francisco earthquake and fire. Williams (1918)
notes that a specimen in the Henry Edwards collection labelled "Lone Mountain,
Lake Tahoe" is marked "True to type" [of battoides] by Edwards, and agrees with
battoides figured by Barnes & McDunnough "except that it is not quite so strongly
marked below." Barnes & McDunnough (1916) falsely give the type locality as
"The headwaters of the Tuolumne River", and so corrected it in 1917. Langston
(1969) states the type locality to be "Mineral King, Tulare Co.,", surely another
error, as this lies two major river drainages to the south of the San Joaquin River.
Finally, Skinner (1911) erroneously gives "headwaters of San Joaquin Valley,
Shijimiaeoides battoides allyni, new subspecies
Male: Holotype forewing, 10 mm. UPPER SURFACE, Primaries: light blue
with slight violet tinge; marginal band wide, 1 mm, scalloped appearance (black
partway along outer veins); fringes white to dark; broad fuscous checkered at
Secondaries: marginal band variable, often broken into distinct interneural
spots; fringes white with little or no checkering at vein tips; aurora often prominently
light orange at CU1 and CU2 cells, faint in some; wing margin frequently concave
at CU1 and CU2.
UNDER SURFACE, Primaries: ground whitish cream; bold terminal line;
macules heavily marked and square; praeterminal mark at R4 faint or absent.
Secondaires: terminal line as primaries; aurora orange-brown, in continuous
band, usually 1 mm broad; macules bold with haloes; ground color cream, becoming
Female: Allotype forewing, 10 mm. UPPER SURFACE, Primaries: ground
dark brown; fringes as 3; discoidal macule faint or distinct.
Secondaries: aurora usually broad band from Mi to 2A, solid to terminal line;
concave wing margin at CU1 and CU2.
UNDER SURFACE, Primaries: as in 3. Aurora faint along semimacule
band or lacking.
Secondaries aB in 3.
HOLOTYPE3: CALIFORNIA: Los Angeles Co., El Segundo, VII-9-65
(R. E. Stanford).
ALLOTYPE 9: same.
PARATYPES: same, 56 69. Also, El Segundo, VII-30-72 (J. F. Emmel),
Disposition of type material: Holotype and allotype (AM), paratypes (KH,
Two males, Cedros Island, Mexico, III-18-39 (CIS) also fit this description
although with black markings slightly more pronounced; Rindge (1948) noted that
they were probably a new, undescribed subspecies.
DISTRIBUTION (viewed 86 gen. from 4 localities): CALIFORNIA: Los
Angeles Co.: El Segundo, 48, vii (CAS, PO, KH); El Segundo dunes, 106 19, vii,
ix (LACM, YU); El Segundo sand dunes, L. A. Airport, 206 209, viii (AM, LACM,
CIS); "N. Manhattan Beach", 2&, vii (LACM); Palos Verdes Estates, 19, ix (AM);
Redondo Beach, 16, vii (CIS).
HOST: Eriogonum parvifolium Sm. in Rees var. parvifoloium (Shields #56),
several adults reared from larvae collected on this plant, IX-8-69, El Segundo,
by J. F. Emmel (AM).
Subspecies allyni is on wing from early July to late September.
Occasionally a female within a battoides bernardino population appears
indistinguishable from b. allyni, as, e.g., La Mesa, San Diego Co., v (LACM),
and Cushenbury Springs., 5600', San Bernardino Mts., San Bernardino Co., iv
(YU). Also one female from Olancha, Inyo Co., x (CAS) appears to be b. allyni,
although E. parvifolium does not occur there.
The closest known relative of battoides allyni appears to be b. bernardino,
approaching allyni in size, coloration, overall appearance (though spots much
reduced by comparison), sympatry, and closely related foodplant. S. b. allyni seems
unrelated to S. battoides battoides, a subspecies also quite heavily marked on the
I take pleasure in naming this distinct subspecies for Arthur C. Allyn, whose
generous support has made this project possible.
Eriogonum parvifolium var. parvifolium occupies beaches and bluffs or
dunes along the coast, from Monterey Co. to San Diego Co., flowering from June
to October (Reveal, 1969a)..John Emmel and I checked this species on 18 Sept. 1968
at Solano Beach, San Diego Co. (J. F. Emmel #145) without success. Reveal (1969a
and in litt.) lists Eriogonum molle Greene, E. fasciculatum Benth. var.fasciculatum,
E. pondii Greene, E. wrightii Torr. ex Benth. in DC. var. taxifolium (Greene)
Parish, and E. intricatum Benth. for Cedros Island. Likely, E. molle serves as the
host there for battoides nr. allyni since it is in the same section of Eriogonum
(Fasciculata) as parvifolium, though E. pondii cannot be ruled out because it
resides in the adjacent section (Ectenomorpha). Both sections are in the subgenus
Eucycla. E. parvifolium is not known from the island. (It could be on E. fasciculatum
too, since battoides bernardino uses it).
Shijimiaeoides battoides comstocki, new subspecies
Male: Holotype forewing, 12 mm. UPPER SURFACE, Primaries: light sky
blue (about shade of bernardino); terminal band 1 mm wide, only slightly invading
veins; checkering broad, alternating with fuscous scales.
Secondaries: marginal fringe usually continuously white, some with mild
checkering at vein tips; 5 praeterminal marks distinct or partially fused to terminal
line; margin of Cul and Cu2 just slightly concave.
UNDER SURFACE, Primaries: ground with distinct yellowish-white cast
in fresh specimens; whole under surface has appearance of enoptes enoptes from
Sierras; macules distinct and rather small.
Secondaries: small first /2 scales of marginal fringe are white underside and
black upperside (except at vein ends on underside); aurora from Mi to Cu2
(sometimes tinge on Rs) and broken into distinct, small crescent-shaped spots;
aurora light orange-brown or yellow-orange.
Female: Allotype forewing, 111/ mm. UPPER SURFACE, Primaries:
aurora band sometimes prominent or trace, rarely absent; fringe checkering and
color as in 5; medium brown ground color; discoidal spot faint or somewhat distinct.
Secondaries: praeterminal macules distinct and form base of a continuous
auroral band from M1 to Cu2, 2 mm wide. Aurora to terminal band, or that space
filled with brown; sometimes blue scaling light basally on wing.
UNDER SURFACE, Primaries: as in a except auroral band often present.
Secondaries as in C.
HOLOTYPE 8: CALIFORNIA: Kern Co., Tehachapi, VII-22-18 (J. A.
ALLOTYPE 9: same.
PARATYPES: same, 183 279.
Disposition of type material: Holotype, allotype, and paratypes (LACM,
some paratypes at CAS).
Genitalia of three of the paratype males were determined. The upper and
underside and wing shape of both sexes superficially resemble enoptes nr.
enoptes from Blackburn Cyn., Tehachapi Mts., Kern Co., Calif., VII-10-
64 (N. La Due). Known only from the type locality and one 3 from Park City,
Summit Co., Utah, VII-2-'95 (CM); host Eriogonum unknown. Also at Tehachapi
in July flies battoides bernardino (15, CM).
The closest relative of b. comstocki is perhaps battoides intermedia. It resembles
comstocki on the underside but lacks the aurora on the undersurface of the primaries,
and male intermedia have a much broader black band on the upperside.
Shijimiaeoides battoides ellisii, new subspecies
Male: Holotype forewing, 12 mm. UPPER SURFACE, Primaries: irridescent
blue cyanic overlay; marginal band medium width, 3 mm wide; fringes white;
fuscous checkered at vein ends.
Secondaries: marginal band variable, often broken into distinct interneural
spots (= praeterminal marks showing through), sometimes solid band or nearly
so; fringes white with fuscous checkering; aurora faint or pronounced orange
spots in most, lacking in some.
UNDER SURFACE, Primaries: ground whitish gray; bold terminal line;
macules heavily marked; smoky suffusion toward lower half.
Secondaries: terminal line as primaries; aurora orange, variable from con-
tinuous band to nearly dissociated spots, usually broad, 1 mm wide.
Female: Allotype forewing, 12 mm. UPPER SURFACE, Primaries: ground
brown, usually dark or blackish; fringes as 6; discoidal macule generally definable
Secondaries: aurora usually broad and pronounced, from light to dark orange,
in extent from Cu2 to M2, solid appearance, to the praeterminal marks.
UNDER SURFACE, Primaries: as in 6; aurora occasionally present.
Secondaries: as in a.
HOLOTYPE 8: COLARADO: Mesa Co., W. Creek, Unaweep Canyon, 5400',
6 mi. E. of Gateway, VIII-25-67 (S. L. Ellis).
ALLOTYPE 9: same locality, IX-3-65 (S. L. Ellis).
Disposition of type material: (AM).
DISTRIBUTION (viewed 496 gen. from 25 localities): ARIZONA: Coconino
Co.: 7-8 rd. mi. ESE of jet. of Hwy. 389 & 89A, 5000', ESE Fredonia, "40", viii (AM),
Moenkopi formation (Lower Triassic); Hwy. 89A, 8 rd. mi. ESE Fredonia, 306
209, viii (KR). COLORADO: Mesa Co.: Unaweep Cyn., West Creek, 5400-6000',
"common", viii, ix (AM, SE, MF). UTAH: Grand Co.: Courthouse Wash, 4 rd.
mi. NW Moab, 16, viii (AM), Navajo sandstone (Jurassic?). Kane Co.: 5 rd. mi. W.
Paria River crossing U.S. 89, on U.S. 89, Fivemile Valley, 5000', 26, ix (AM).
San Juan Co.: 1 mi. E Indian Creek State Park, 16, viii (AM); 11 rd. mi. NNW
Mexican Hat, base of Cedar Mesa, along Hwy. 261, "19", ix (AM), Cedar Mesa
sandstone (Permian); 14 rd. mi. SW Mexican Hat, 5400', 26 59, ix (AM); Monument
Valley, 1&, viii (KT), Cutler formation (Permian); 8 rd. mi. W of jet. of State Hwys.
95 & 47, on road to Natural Bridges Nat'l Mon., 26, ix (AM), Morrison formation
(Upper Jurassic); Rainbow Bridge Canyon, Ariz.-Utah line, 16, ix (DB), Navajo
sandstone (Jurassic?). Uintah Co.: jet. of North Willow Creek turnoff & Ouray-
Rainbow Rd., ca. 30 mi. SE Ouray, 16, viii (AM); Willow Creek, 36 79, ix (AM).
Washington Co.: 1/2 mi. W West Entrance to Zion Nat'l Park, 16, ix (AM), Moenkopi
formation (Lower Triassic).
HOSTS AND DISTRIBUTION: Eriogonum corymbosum Benth. in DC var.
corymbosum. ARIZONA: Coconino Co.: 7 rd. mi. SE Fredonia, 5000', "73", viii
(AM), adult assoc. (Shield #49). UTAH: Duchesne Co.: 10 rd. mi. S of jet. Hwys.
53 & 216, along Hwy. 53, "45", viii (AM), adult assoc. (Shields #22). Kane Co.:
1 mi. E. Glendale on rd. to Fourmile Hollow, 6200', "38", viii, ix (AM), adult assoc.
(Shields #46) Upper Jurassic; 112 rd. mi. NNW Kanab, 5000', 26, viii (AM), adult
assoc. (Shields #48) Chinle formation (Upper Triassic). Uintah Co.: V2 mi. S Bonanza,
along Hwy. 45, 69, viii (AM), adult assoc. (Shields #17); 1 mi. S North Willow Creek
turnoff from Ouray-Rainbow Rd., ca. 31 mi. SE Ouray, "5", viii (AM), adult assoc.
(Shields #20); along Ouray-Rainbow Rd., 18 rd. mi. SW of jet. of 207 & 45, 19,
viii (AM), adult assoc. (Shields #19); along Utah State Hwy. 207, 10 rd. mi. SSE
Bonanaza, 26, viii (AM), adult assoc. (Shields #18). Eriogonum corymbosum Benth.
in DC. var. glutinosum (M. E. Jones) M. E. Jones. ARIZONA: Coconino Co.: 9.5
road mi. NE Winona, on Hwy. to Leupp, "37", ix (AM), oviposition (Shields #161);
3.7 rd. mi. S of turnoff to Wupatki Nat'l Mon., along Hwy. 89, "14", ix (AM),
adult assoc. (Shields #163). Navajo Co.: 2.5-3.0 rd. mi. SE Joseph City, along Hwy.
40, 13 19, viii (AM), adult assoc. (Shields #159). Eriogonum corymbosum Benth. in
DC. var. orbiculatum (S. Stokes) Reveal & Brotherson. UTAH: Grand Co.: 1.5
rd. mi. SE of jet. of Hwy. 128 & rd. to Castleton, along rd. to Castleton, "5", ix (AM),
adult assoc. (Shields #171); 12.5 rd. mi NE ofj ct. of Hwy. 128 & rd. to Castleton, along
Hwy. 128, "2", ix (AM), adult assoc. (Shields #172); 12 rd. mi. W and S of jet. of rd.
to Dead Horse Point State Park & U.S. Hwy. 160, 26, viii (AM), adult assoc.
(Shields #25); 14.8 rd. mi. NE of jet. 163 & 128, along Hwy. 128, 19, ix (AM), adult
assoc. (Shields #170). San Juan Co.: 10 rd. mi. S of Blanding, "4", viii (AM), adult
assoc. (Shields #32), Dakota (?) sandstone (Upper Cretaceous); 13 rd. mi. SW of
jet. of road to Canyonlands Nat'l Park (via Indian Creek State Park) & U.S. Hwy.
160, "4", viii (AM), adult assoc. (Shields #30); Grandview Point, Canyonlands
Nat'l Park, "3", viii (AM), adult assoc. (Shields #27); 12 rd. mi. N Mexican Hat,
along Hwy. 261 at base of canyon, N end Cedar Mesa, 5200', 23 29, ix (AM), oviposi-
tion (Shields #36; J. F. Emmel #139), Cedar Mesa sandstone (Permian); 10 rd. mi.
NW of jet. Hwy. 95 & Natural Bridges Nat'l Mon. turnoff, "4", viii (AM), adult
assoc. (Shields #37), Cedar Mesa sandstone; 19 rd. mi. NW of jct. of Hwy. 95 &
Natural Bridges Nat'l Mon. turnoff, 25, viii (AM), adult assoc. (Shields #38),
Cedar Mesa sandstone; 14 rd. mi. WNW of rd. to Needles Overlook & U.S. Hwy. 160,
"2", viii (AM), adult assoc. (Shields #29). Eriogonum corymbosum Benth. in DC.
var. velutinum Reveal. ARIZONA: Navajo Co.: 7.7 to 8.1 rd. mi. ENE of Kayenta
(from jct. Hwy. 464 & 164), just SE of Church Rock, "32", ix (AM), adult assoc.
(Shields #164), Chinle formation (Late Triassic). Eriogonum batemanii M. E.
Jones. UTAH: Uintah Co.: Ouray-Rainbow Rd., 22 rd. mi. SE Ouray, "12", viii
(AM), oviposition (Shields #21). See Maps 1 and 2.
Subspecies ellisii flies from mid August to early September (Fig. 29)
S. battoides etlisi is a distinctive subspecies, although populations in the
vicinity of Flagstaff and Joseph City, Arizona, approach b. centralis from northern
New Mexico on the undersurface somewhat. The width of the aurora is highly
variable in ellisii, sometimes as narrow as in battoides intermedia, sometimes as
broad as in battoides martini (the latter appear very much like martini though are
larger). Specimens of a rare fall-flying, small battoides asp. from the Clark and
Providence Mts. of eastern California appear somewhat similar to ellisii (see Emmel
& Emmel, 1973, pl. 8, fig. 7), and occur on varieties of E. heermannii, while another
population in central Nevada on a different variety of E. heermannii in June (to
be discussed in a later paper) appear closer to bernardino than to the Mojave fall
form. Near Church Rock, Navajo Co., Arizona, b. ellisii and rita emmeli occur
together. Here, the adults are associated with their proper Eriogonum species
even when roosting; i.e., in pure stands of each Eriogonum, only the proper Philotes
sp. flew, and where there was a mixed stand, both flew over their respective
This subspecies is named for my good friend Scott L. Ellis, who initially
Reveal (1967) gives extensive locality records for Eriogonum corymbosum
varieties corymbosum, glutinosum, orbiculatum, and velutinum. According to Reveal
at present, E. corymbosum contains six varieties, distributed from southwestern
Wyoming southward through eastern and southern Utah and adjacent western
Colorado into northern Arizona and northwestern and central New Mexico,
flowering from July to October (Reveal, 1969a). John Emmel and I found no
S. battoides ellisii in late August 1969 on E. corymbosum var. erectum Reveal &
Brotherson, E. c. var. davidsei Reveal, E. duchesense Reveal (Shields #15), E.
lancifolium Reveal & Brotherson, and E. saurinum Reveal, all in theE. corymbosum
complex, in eastern Utah, and none on E. c. var. glutinosum near Cedar City and
Shivwits, Utah, although this was used in northern Arizona. Also, Ellis, Toliver,
and I found none at the type locality of E. c. var. velutinum near San Antonio,
New Mexico, on 21 August 1970.
Eriogonum batemanii occurs in northeastern Utah and adjacent northwestern
Colorado, flowering from June to September (Reveal, 1969a).
Shijimiaeoides battoides, new unnamed subspecies
On July 29, 1969, John Emmel and I found 8 mature larvae of Philotes feed-
ing on the flowers of Eriogonum shockleyi S. Wats. var. shockleyi (J. F. Emmel
#199) on sand dunes 1 air mile south of Blind Spring, 5800', southeast end of Baking
Powder Flat, west of Snake Range, T11N, R67E, White Pine Co., Nevada. All
the larvae were unusually small when they pupated. All the pupae subsequently
died, but one that had matured inside the pupa was dissected out and genitalically
determined as battoides. I feel that collecting in the last two weeks of June at this
locality should disclose the presence of a new subspecies.
Eriogonum shockleyi var. shockleyi occurs from eastern California to western
Utah and northward to southeastern Idaho, flowering from May to July (Reveal,
Shijimiaeoides battoides baueri, new subspecies
Male: Holotype forewing, 11 mm. UPPER SURFACE, Primaries: light sky
blue cyanic overlay; marginal band medium width, /2 mm wide; fringes white;
fuscous checkering at vein ends at M3, Cul, Cu2, and 2A only (sometimes others);
black scales from marginal band diffusing out into blue ground slightly, forming
patchy spots between veins (= praeterminal mark on upper side).
Secondaries: marginal band as in upper surface; small black dots at praeter-
minal marks distinctly detached from marginal band and somewhat diffuse;
fringes white with fuscous checkering; no aurora showing through.
UNDER SURFACE, Primaries: ground pale whitish (snowy white), bold
terminal line, 1/2 mm wide; smoky suffusion from Cu2 to 2A; macules moderately
Secondaries: terminal line as primaries; aurora light orange, band thin
(lacking any orange spot in cell 2A), aurora 2/3 mm wide.
Female: Allotype forewing. 11 mm. UPPER SURFACE, Primaries: ground
color light sky blue with terminal band very broad, sometimes extending nearly
to discoidal macule; discoidal macule distinct to absent; black suffusion over blue
heavily along veins, lighter between veins.
Secondaries: aurora in medium width band to small spots or absent; prae-
terminal marks prominent; aurora not extending to terminal line; blue ground color
extensive or blackened in toward Sc and Rs; distinct whitish blue narrow band
between praeterminal marks and terminal line.
UNDER SURFACE, Primaries.as in 3.
Secondaries as in 3.
HOLOTYPE 6: CALIFORNIA: Inyo Co., W side Gilbert Pass, 6200', V-23-67
(D. L. Bauer).
ALLOTYPE 9: same.
PARATYPES: same, 66 99.
Disposition of type material: Holotype and allotype (AM), paratypes (DB).
DISTRIBUTION (viewed 126 gen. from 8 localities): CALIFORNIA: Inyo
Co.: pass 4-5 mi. ENE Deep Springs, 18, v (DB); Waucoba Cyn., 6000', 16 mi.
SE Saline Valley Jet., 18, v (GG); Westgard Pass, 29, v (UCD). NEVADA: Churchill
Co.: Buffalo Canyon, Desatoya Mts., 56 29, v (NSM); Buffalo Cyn., 5500', 12 mi.
S. of Eastgate on road to lone, W. of O'Donnell Summit, 246, v (CAS). Clark Co.:
Arrow Canyon Range, 16 19, v (NSM). Douglas Co.: Pine Nut Mts., E. side, 36
19, v (AM). Esmeralda Co.: 5 mi. S Tonopah, 5500', 16, v (DB). Humboldt Co.:
Winnemucca, 56 39, v (CAS, CIS); 10 mi. N. Winnemucca, 2(, v (CIS). Lander Co.:
4.0 road mi. E Carroll Summit, 6700', 16, vi (AM). Lincoln Co.: Oak Springs, Delamar
Range, 46 39, v (NSM). Ormsby Co.: Clear Creek Cyn., 5 mi. SW Carson City,
16, v (PO).
HOSTS AND DISTRIBUTION: Eriogonum ovalifolium Nutt. var. ovalifolium.
CALIFORNIA: Inyo Co.: ca. 2 rd. mi. NE Westgard Pass, 7000', 26, v (AM),
adult assoc. (Shields #76), growing mixed in with the abundant E. caespitosum
Nutt. (Shields #77) that was not used. Eriogonum ovalifolium Nutt. var. multiscapum
Gand. NEVADA: Humboldt Co.: 10 rd. mi. N Winnemucca, on Hwy. 95, sand dunes
area between hwy. & Little Humboldt River, 4350-4450', "few", v (AM), adult assoc.
(J. F. Emmel #286). Subspecies baueri is in flight in May (Fig. 30).
At Westgard Pass near the type locality of b. baueri, a distinctive subspecies
of battoides (near glaucon, to be described later, on E. umbellatum) flies a month
later and appears quite different. However, at the Buffalo Canyon and vicinity of
Winnemucca localities, b. baueri closely approaches the subspecies glaucon, and
at Steamboat Hot Spgs., 5 rd. mi. N jet. of Hwy. 3B & 395, on Hwy. 395 SE of Reno,
Washoe Co., Nev., VI-19-71, 18 battoides glaucon was on Eriogonum ovalifolium
Nutt. var. nivale (Canby) M. E. Jones (Shields #196) at an alkali spring.
This subspecies is named for David L. Bauer, its original discoverer.
Eriogonum ovalifolium var. ovalifolium occupies dry slopes and flats,
mostly 5000-7000', east slope Sierra Nevada of California, n. and e. to Alberta,
Rocky Mts.; blooms from May to July. Richard Funk and Kilian Roever took a
short series of S. battoides ssp. (gen. det.) on E. ovalifolium Nutt. var. ovalifolium
(det. by W. B. McDougall and Reveal, Funk #160, deposited at the Museum of North-
ern Arizona, Flagstaff), on V-18-68, Hwy. 89A, 8 road mi. ESE Fredonia, Coconino
Co., Arizona. The phenotype is not very similar to b. baueri and probably deserves
a name. This most intriguing population appears to be not closely related at all
to b. ellisii and E. corymbosum var. corymbosum that flies at the same locality
some three months later.
Shijimiaeoides rita rita (Barnes & McDunnough), 1916
Barnes & McDunnough. 1916. Canad. Ent., 48: 223-224.
From their type description and the paratype illustration in Barnes & Mc-
Dunnough, 1916, pl. XI, figs. 3 & 6, there is no question that this is what is currently
called rita rita from southern Arizona. S. enoptes dammersi, with which it might
easily have been confused, is not known to enter southern Arizona, and their il-
lustration (1917, pl. 17, figs. 5 & 7) of rita's genitalia clearly establishes it as such
(see also Mattoni, 1965: 86). The type locality in the original description reads,
"Types-Three 3's, S. Arizona (Poling); one S, Santa Rita Mts., Ariz.; one S,
Rio Verdi Mts., Ariz.; three 9's, S. Arizona (Poling) in Coll. Barnes." Thus the de-
termination of an exact type locality becomes problematical, unless it is assumed
that the first mention ("S. Arizona (Poling)") was their intention. If so, Comstock
(1953) sheds light on a precise location: "Mr. [Lloyd] Martin has concluded, from
a study of various records and dates on material collected by Poling, that the 'S.
Arizona' series were taken in the Huachuca Mountains near Ramsay [sic]
Canyon." Comstock, Martin, Ford, and Thorne subsequently found rita rita at
the foot of Ramsay [sic] Canyon in abundance (700 taken!) on IX-1-53, acting on
this hunch. I have determined the genitalia of 1763 from this series as rita rita.
Unless evidence to the contrary eventually emerges, the type locality is hereby
designated as near Ramsey Canyon, Huachuca Mts., Cochise Co., Arizona. As
noted by Clench (1967), the Rio Verde Mts. are located northeast of Phoenix,
based on two Barnes' specimens, and are not synonymous with the Huachuca
Mts. as Mattoni (1965; 86) surmised.
Shijimiaeoides rita emmeli, new subspecies
Male: Holotype forewing, 11 mm. UPPER SURFACE, Primaries: light blue-
lavender (i.e. light purplish blue cyanic overlay); terminal fuscous band V/2 mm wide;
fringes broad, white, with row of black scales at base, black checkering at Cu2
and 2A vein ends (not pronounced); some black scales along extremities of veins
and scattered in cells lightly.
Secondaries: terminal band broad in outer V/ of Sc, outer 1/3 (or less) of Rs,
V2 mm wide in Mi and M2, becoming distinct spots (praeterminal marks) in M3,
Cul, Cu2, with light band between these spots and terminal band (sometimes
faint or absent); pink aurora usually prominent, from M3 to Cu2 or only Cul
and Cu2; outer quarter of veins darkly scaled; wing margin of Cu2 slightly concave.
UNDER SURFACE, Primaries: ground dull pearly white to gray-white;
thin terminal line; semimacules and praeterminal marks somewhat diffuse,
other macules distinct; macules with haloes.
Secondaries: aurora light gold-orange or yellow-orange, prominent band 1 mm
wide; extending from M1 to 1A, occasionally auroral spot in 2A; macules not quite
as bold as in coloradensis; prominent haloes.
(Male differs from coloradensis in smaller size, lighter blue color, less width
of terminal band, less pronounced underside macules, and lacking slight melanic
suffusion of underside primaries.)
Female: Allotype forewing, 101/2 mm. UPPER SURFACE, Primaries: ground
color medium brown (usually lighter than coloradensis), slightly lighter in basal
1/3 with faint blue-gray scaling; fringe as in 6; brown color is yellowish (= "orange-
green") at ca. 450 to light (same in coloradensis); discoidal macule distinct.
Secondaries: pronounced orange aurora from M2 to 1A (sometimes all or
part of MI); orange occasionally to terminal line but more often stopping at
praeterminal marks (filling between); aurora 1 1/3 mm wide.
UNDER SURFACE, Primaries same as in 6, though ground color
usually more gray-white.
Secondaries same as in 6, though ground color usually more gray-
HOLOTYPE 6: UTAH: Emery Co., summit of road to 11/ mi. E., of Little
Flat Top, 5500', T26S, R13E, ca. 10-11 mi. SE of Utah Hwy. 24, VIII-26-69 (J. F.
Emmel & O. Shields).
ALLOTYPE 9: same.
PARATYPES: Same, 44 adults.
Disposition of type material: holotype, allotype, and paratypes (AM).
Number of teeth on terminal end (cucullus) of valve = 20 (6), 21 (6), 22 (3),
23 (2), 26 (1); N = 18, average = 20-22, 4 localities = Tuba City, Calf Creek, Dead
Horse, Klondyke Bluffs.
The behavioral adaptation of rita emmeli to a sandstorm is described in
DISTRIBUTION (viewed 386 gen. from 18 localities): UTAH: Emery Co.:
along rd. 1/ mi. S. of Goblin Valley turnoff from highway 24, 1 mi. SE along this
rd. from hiway, 5100', T25S, R12E, 39, viii (AM), Upper Jurassic; 10 rd. mi. SE of
Hwy. 24, Little Flat Top, Green River Desert, "12", ix (AM), Upper Jurassic; 15.3
rd. mi. SE of Hwy. 24, on road to Orange Cliffs via Little Flat Top, 19, ix (AM),
Upper Jurassic; San Rafael Desert, 66 59, viii (AM). San Juan Co.: 20 road mi. S
Blanding, 16, viii (AM). Wayne Co.: 4.0 road mi. S Hanksville, on road to Henry Mts.,
19, ix (AM).
HOSTS AND DISTRIBUTION: Eriogonum leptocladon Torr. & Gray var.
leptocladon. UTAH: Emery Co.: 10 road mi. NE of Goblin Valley turnoff from Hwy.
24, on Hwy. 24, Green River Desert, "19", ix (AM), adult assoc. (Shields #169),
Upper Jurassic. Garfield Co.: Poison Spring Cyn., 1 mi. S Wayne Co. line, along
Hwy. 95, 18 road mi. SE Hanksville, 38 19, viii (AM), adult assoc. (Shields #39).
Grand Co.: 1'/ mi. N of Courthouse Wash, along U.S. Hwy. 160, ca. 17 road mi.
NW Moab, 18 69, viii (AM), oviposition (Shields #23) Morrison formation (Upper
Jurassic); sand hillocks 5 rd. mi. W of jct. of rd. to Dead Horse Point State Park
& U.S. Hwy. 160, NW of Moab, "53", viii (AM), oviposition (Shields #24); 3 rd. mi.
S & W on road to Klondyke Bluffs from jet. of this rd. with rd. to Devils Garden,
Salt Valley, "7", viii (AM), adult assoc. (Shields #28). San Juan Co.: 8 rd. mi. S
of juct. of rd. to Dead Horse Point State Park & Canyonlands Nat'l Park, 18,
viii (AM), adult assoc. (Shields #26); 8 rd. mi. NW past turnoff ot Dugout Ranch,
along rd. to Canyonlands Nat'l Park, via Indian Creek State Park, "13", viii
(AM), adult assoc. (Shields #31). Wayne Co.: 6 rd. mi. SE Hanksville, "9", viii
(AM), adult assoc. (Shields #40); 20.7 rd. mi. SE of Hwy. 24, on road to Orange Cliffs
via Little Flat Top, "6", ix (AM), adult assoc. (Shields #167) Upper Jurassic.
30.6 road mi. SE of Hwy. 24, on rd. to Orange Cliffs via Little Flat Top,
"19", ix (AM), oviposition (Shields #168) Upper Jurassic. Eriogonum leptocladon
Torr. & Gray var. ramosissimum (Eastw.) Reveal. ARIZONA: Coconino
Co.: 5.0 rd. mi. NE Sunrise Trading Post, N of Leupp, 19, ix (AM),
adult assoc. (Shields #162); 14 rd. mi. NE Tuba City, "15", viii (AM),
adult assoc. (Shields #50); 9 rd. mi. SW of jet. with Hwy. 89, & rd. thru Wupatki
Nat'l Mon., 16 19, viii (AM), adult assoc. (Shields #51).Navajo Co.: 21/2 rd, mi, N
Joseph City, 5200', "11", viii (AM), adult assoc. (Shields #160); 7.7 to 8.1 rd. mi.
ENE Kayenta (from jet. Hwy. 464 & 164), just SE Church Rock, "14", ix (AM),
adult assoc. (Shields #165), Chinle formation (Late Triassic). UTAH: Kane Co.:
8 rd. mi. SSE Mt. Carmel Junction, 5800', along Hwy. 89, 26, viii (AM), adult assoc.
(Shields #47). San Juan Co.: at SW base of a bluff, 3 rd. mi. N & 1 rd. mi. E of Bluff,
5000', T40S, R22E, "17", viii (AM), adult assoc. (Shields #33), Morrison formation
(Upper Jurassic); 5 rd. mi. WSW Bluff, "20", viii (AM), adult asssoc. (Shields #34),
San Rafael group (Upper Jurassic); 6.8 rd. mi. SW Mexican Hat, along Hwy. 163,
"29", ix (AM), adult assoc. (Shields #166); 2 rd. mi. NW of jet. Hwy. 261 & 47,
along Hwy. 261 (5 mi. N. Mexican Hat), "5", viii (AM), adult assoc. (Shields #35),
Figures 1-16: new Shijimiaeoides battoides subspecies. 1-4: S. b. allyni, new
subspecies; Holotype 5 upper (1) and under (2) surfaces; Allotype 9 upper (3) and
under (4) surfaces; CALIF.: Los Angeles Co.: El Segundo, vii.9.1965 (R. E. Stanford).
5-8: S. b. comstocki, new subspecies; Holotype a upper (5) and under (6) surfaces;
Allotype 9 upper (7) and under (8) surfaces; CALIF.: Tehachapi, vii. 22.1918 (J. A.
Comstock). 9-12: S. b. ellisii, new subspecies; Holotype S upper (9) and under (10)
surfaces; Allotype 9 upper (11) and under (12) surfaces; COLORADO: Mesa Co.:
Unaweep Canyon, viii.16.1969 (S. L. Ellis). 13-16, S. b. baueri, new subspecies;
Holotype a upper (13) and under (14) surfaces; Allotype 9 upper (15) and under (16)
surfaces; CALIF.: Inyo Co.: W side Gilbert Pass, 6200', v.23.1967 (D. L. Bauer).
Pennsylvanian. Wayne Co.: 5 rd. mi. NE Hanksville, 4600', "4", viii (AM), adult
assoc. (Shields #44), Upper Jurassic. Eriogonum leptocladon Torr. & Gray var.
papiliunculi Reveal. UTAH: Garfield Co.: 5 rd. mi. SSW Calf Creek Recreation
Area, 12 rd. mi. E Escalante on Hwy. 54, 5800', T35S, R4E, 6c 59, viii (AM), ovi-
position and 19 reared from larva (Shields #45). See Map 3.
Subspecies emmeli is on wing in late August and early September.
In the vicinity of the type locality on VIII-26 & 27-69, at 1/2 mi. E. Little Flat Top,
Upper Jurassic, 24 adults were onE. leptopcladon var. ramosissimum (Shields #41).
Some adults were on E. leptopcladon var. leptocladon (Shields #42) and E. smithii
Reveal (oviposition, Shields #43) at the summit (Fig. 31-32). About 1 miles east is
a mixed stand where all three Eriogonum's occur together.
S. rita emmeli in parts of northern Arizona occasionally bears a strong re-
semblance to rita rita, and in northwestern New Mexico populations of rita that
are clinal between rita rita and rita coloradensis, on E. effusum, also approach
emmeli. Reveal (1971: 2) states that recently in Kane Co., Utah, intergrades were
discovered between E. leptocladon var. ramosissimum and E. kearneyi, the later
being the principle foodplant of rita pallescens (Shields, unpubl.), while on the
eastern flank var. ramosissimum is approached by the New Mexican phase of
E. effusum var. effusum. Thus, rita emmeli appears to be related to the rita rita-r.
coloradensis series on the one hand, and (perhaps rita pallescens on the other.
In valve teeth number it lies intermediate between the two (averages, rita rita
Figures 17-28: new Shijimiaeoides. 17-20: S. rita emmeli, new subspecies;
Holotype & upper (17) and under (18) surfaces; Allotype 9 upper (19) and under (20)
surfaces; UTAH: Emery Co.: 1.5 mi. E Little Flat Top, ca. 5500', viii.26.1969 (J. F.
Emmel & 0. Shields). 21-24: S. rita mattonii, new subspecies; Holotype a upper
(21) and under (22) surfaces; Allotype 9 upper (23) and under (24) surfaces; NEVADA:
Elko Co.: 4 mi. W Charleston Reservoir, 6600', vii.24.1969 (J. F. Emmel & O. Shields).
25-28: S. enoptes langstoni, new subspecies; Holotype a upper (25) and under
(26) surfaces; Allotype 9 upper (27) and under (28) surfaces; CALIF.: Mono Co.:
1.6 mi. N Inyo-Mono Co. line on Hwy. 395, 5700', v.18.1970 (0. & R. Shields).
and r. coloradensis = 24-26, rita emmeli = 20-22, rita pallescens = 19-21).
This subspecies is named for Dr. John F. Emmel, my companion on many
summer trips, in appreciation for his efforts with this subspecies.
For Eriogonum leptocladon var. leptocladon and var. ramosissimum,
specific localities are listed in Reveal (1965). They range from SE Utah, NE Arizona,
extreme SW Colorado, and adjacent New Mexico, flowering from June to October.
White-flowered var. ramosissimum is confined mostly to white "blowsand",
while yellow-flowered var. leptocladon is confined to red "blowsand" (Reveal,
1969a). The var. papiliunculi, with its white flowers and glabrous stems, ranges
from Garfield Co., Utah, southward to northern Coconino and Apache Cos.,
Arizona (Reveal, 1974). E. smithii is known only from the vicinity of Little Flat
Top, Emery Co., Utah, flowering from late July to September (Reveal, 1967).
John Emmel and I separately have checked a closely related species from Imperial
Co., Calif., E. deserticola S. Wats., that blooms from November to February,
Shijimiaeoides rita mattonii, new subspecies
Male: Holotype forewing, 10 mm. UPPER SURFACE, Primaries: lavender
blue cyanic overlay; marginal line 1/2 mm wide, black scales extending ca. outer
1/3 of veins and scattered black scales over rest of veins (veins almost outlined
thinly in black in several), blue scales lightly dusted with black scales over ground;
fringe white with black checkering at vein tips (but usually lacking for MI and M2);
small 1/3 length fringe scales black, uneven.
Secondaries: fringe white except Cul, Cu2, small row black and uneven;
terminal band partially or nearly filling cells Sc and Rs; praeterminal macules
fused on lower 1/2 to terminal line, giving scalloped appearance to band from Mi
to Cu2, or separate; some black scales dusting over blue ground overlay; aurora
faintly present in Cul, Cu2 of one, absent in the others; black scaling on outer 1/3
of veins M2 to Cu2, boxing in the praeterminal macules, veins M1 and Rs completely
covered with black scales.
UNDER SURFACE, Primaries: ground is snowy white with slight grayish
cast in several; terminal line thin; macules heavily marked (except for faint
praeterminal marks), slight or extensive smoky suffusion in cell Cu2.
Secondaries: aurora golden orange or yellowish orange, 1/3 mm wide, con-
tinuous band from Mi to Cu2 (except one discontinuous), scalloped appearance;
praeterminal marks separate from aurora; macules smaller than those of forewing.
Female: Allotype forewing, 11 mm. UPPER SURFACE, Primaries: ground
dark brown, solid; discoidal spot distinct.
Secondaries: aurora 3% mm wide, from M, to Cu2; fringe white without checkering;
solid dark brown ground.
UNDER SURFACE, Primaries as in S except smoky suffusion below Cu2.
Secondaries: as in S except aurora 1 mm wide.
HOLOTYPE 5: NEVADA: Elko Co., ridge 4 air mi. W. of Charleston Reservoir,
6600', SW corner of T43N, R57E, SW of Jarbidge Mts., VII-24-69 (J. F. Emmel &
ALLOTYPE 9: same.
Disposition of type material: (AM).
Number of teeth on terminal end (cucullus) of valve = 15, 18, 19, N = 3,
from Charleston Reservoir and Beaver Creek.
DISTRIBUTION: NEVADA: Elko Co.: West Fk., Beaver Creek, 23, vii
HOST: Eriogonum microthecum Nutt. var. laxiflorum Hook. At the type
locality, oviposition (Shields #10).
Subspecies mattonii flies in July.
Named in honor of Dr. R. H. T. Mattoni's major contributions to this complex
S. rita mattonii appears to have no known close relatives, though perhaps it
is anciently related to rita coloradensis on E. effusum by virtue of microthecum
and effusum both residing in the E. microthecum complex. Certainly the two rita
subspecies, though, are today quite disparate. Certain rita forms, as yet unplaced,
from central Nevada and eastern California, also use E. microthecum varieties
but bear no obvious resemblance to rita mattonii.
Eriogonum microthecum var. laxiflorum is widespread and common in
western U.S., from 5,000-10,500', flowering from June to October (Reveal, 1971;
see fig. 6). Hence, rita mattonii may prove to be much more widespread than is
Shijimiaeoides rita spaldingi (Barnes & McDunnough), 1917
Barnes & McDunnough, 1917. Contrib. Nat. Hist. Lepid. N. Amer., 3: 215-216;
262, 264; pl. 16, figs. 9-11; pl. 17, fig. 6.
There is no doubt from the original description that spaldingi refers to a form
similar to enoptes in male genitalia, with the primaries of both sexes possessing
a broad submarginal orange band on the underside, and with the female upperside
having the subterminal orange band of the secondaries continued on the primaries
for half the length. The type series is mixed: "our types are 26, 19 from Provo,
Utah (Aug. 1-7), 1 from Silver Lake, Utah (July 24-30) and 16 (very large) from
Stockton, Utah, the latter in the collection of Mr. Spalding." Plate 16, fig. 9 & 11,
are of the a and 9 "Type" from Provo, Utah, hereby designated the type locality.
In the past, much confusion has centered about the names glaucon and spaldingi,
apparently stemming from the fact that the genitalia of glaucon was unknown
(see McDunnough, 1914; Williams, 1918). In the original description of glaucon,
the male being pruinose blue (like Everes comyntas), underside gray-brown,
etc. would also fit spaldingi. Barnes & McDunnough (1918: 79), however, note that
the type description of glaucon does not apply to spaldingi in certain particulars.
Both Barnes & McDunnough (1918: 77-79) and Brown (1970: 406-409) come
to the same conclusion, apparently independently, that Henry Edwards' glaucon
type was number "251" and has a battoides male genitalia. They note that this
number referred to Storey Co., Nevada, from "mountain pastures". Barnes & Mc-
Dunnough further restrict this locality to "the hills around Virginia City",
though on what basis is unclear. Brown (1970: 407), also found an entry in Hy.
Edwards' catalogue mentioning Virginia City. Likewise, Williams (1918: 101)
says the type specimens probably came from near Virginia City. Unless evidence
to the contrary is brought to light, near Virginia City, Storey Co., Nevada, is hereby
designated the type locality, for glaucon. S. b. glaucon has been taken at 7 mi.
Canyon, Virginia City, V-18-69, leg. D. L. Bauer (gen. det.). I feel that part of the
confusion surrounding glaucon is due to the type description being a composite
of specimens of what today are known as glaucon, enoptes ancilla, and spaldingi;
a similar opinion was expressed by Barnes & McDunnough (1916: 117; 1917: 215;
1918: 79). Williams (1918: 101) indeed thought that glaucon and spaldingi were
synonymous. A male spaldingi specimen I have seen in the MCZ Collection, bears
the labels "Colorado. glaucon Edw. Col. Gift of H. C. Fall." I have proposed (1973:
2) that spaldingi be considered a subspecies of rita. This decision is based on the
presence of cristae on the male genitalia of spaldingi, a trait universally present
in rita and lacking in enoptes forms.
The number of teeth on terminal end (cucullus) of valve = 11 (3), 12 (3), 13 (8),
14 (10), 15 (8), 16 (1), 17 (1), 18 (1), N = 35, average = 13-15, localities = North Rim,
Indian Hollow, Robbers Roost, Bee Spring, Schilling's Spring (most), Alamosa
Reservoir (many), Swain Creek.
DISTRIBUTION (viewed 376 gen. from 17 localities): ARIZONA: Coconino
Co.,: Bee Springs, Kaibab Plateau, 56, vi, vii (DB, LACM); Bee Spring, 7800',
Kaibab Plateau, 36, vii (KR) Kaibab formation (Upper Permian); Indian Hollow,
6800', Kaibab Plateau, 38, vii (KR) Kaibab formation; North Rim, Grand Canyon,
28, vii (LACM); Robbers Roost Spr., 8200', N. Rim, Grand Canyon Nat'l Park,
26, vii (KR) Kaibab formation. Navajo Co.: Gomez Crk., Upper Log Rd., 6 mi. E.
Hwy. 73, Fort Apache Indian Res., 16, viii (AM). COLORADO: Archuleta Co.:
2 mi. S Pagosa Springs, vi (RM); Trujillo, vii (NL). Conejos Co.: 1/ mi. N of E end
Alamosa Reservoir, 36, vii (JS); W end Alamosa Res., large series, vii (JS); Aspen
Glade Recreation Area, Rio Grande Nat. For., "1", vii (CC); Schilling's Spring,
large series, vii (JS); 1/ mi. SE Schilling's Spring, large series, vii (JS). Dolores Co.:
3 mi. E Dove Creek (MF). Gunnison Co.: Almont, 8000', 26, vii (SE); 1.5 mi. N Almont,
Hwy. 135, 1$, vii (RL); Jack's Cabin, 36, vii, viii (CIS, YU). La Plata Co.: Rockwood,
7360', vii (in Brown, 1955). Montezuma Co.: Mesa Verde Park, 19, viii (LACM);
Morefield Canyon, Mesa Verde Nat'l. Park, 56, vii (CIS, AM) Mesa verde sandstone
(Upper Cretacecus); Prater Canyon, Mesa Verde Nat'l Park, 46 39, vii (RL)
Mesa verde sandstone (Upper Cretaceous). Montrose Co.: Black Canyon, 16,
vii (RL); Cimarron, 7000', 26, vii (SE). San Miguel Co.: Wilson Draw, 8000', 6 mi.
SE Egnar, San Juan Nat'l For., 126, vii (SE). NEW MEXICO: Catron Co.: nr.
Mangas, Apache Nat. For., "1", vii (CC). McKinley Co.: 4 mi. S Fort Wingate, 16,
vii (CAS). Rio Arriba Co.: 4 mi. S Tierra Amarilla (RL). San Juan Co.: Shiprock,
16 59, viii (LACM). Torrance Co.: Capillo Peak Summit, 9200', Manzano Mts.,
19, vii (CM); 1 mi. S. Capillo Peak, 9000', Manzano Mts., 16, vii (RH). UTAH:
Garfield Co.: Bryce Nat'l Park, 18, vii (CAS); near Panguitch, vii (KR). Juab Co.:
Eureka, 7500', 26, (CM). Kane Co.: nr. Strawberry Creek, 18, vii (KT); nr. Swain
Creek, Hwy. 14 mile 36, 16 19, vii (KH, JL). Tooele Co.: S. Willow Cr., Stansbury
Mts., 13, vii (SE). Utah Co.: Springville, left fork Hobble Creek, 16, vii(DB). Wasatch
Co.: Provo Canyon, 1S, vii (LACM). Washington Co.: Pine Valley C. G., Pine Valley
Mts., 16, vii (KR). See Map 4.
HOST: Eriogonum racemosum Nutt. COLORADO: Conejos Co.: W end
Alamosa Res., NW of Capulin, viii (no adults; JS took large series here) (Shields
#140). Montezuma Co.: hillside ca. 1 mi. W Park Point Fire Lookout (with water
tank at summit), Mesa Verde Nat'l Park, "8", viii (AM), adult assoc. (Shields
#136). Also sympatric here was Plebejus melissa, 9 adults, flying around legumes,
feeding on flowers other than racemosum, or roosting on sagebrush. Though
sympatric, spaldingi and melissa had different flight patterns and behaviors.
Scott, Ellis, & Eff (1968) also record spaldingi's foodplant as racemosum. Subspecies
spaldingi is in flight from latest June to mid August (Map 4).
Brown (1955) gives a good phenotypic description of spaldingi. Garth
(1950: 36) lists four North Rim, Grand Canyon records.
The relationship of spaldingi to the other rita subspecies remains unclear.
However, the general heavily sclerotized valves, their overall form, and teeth
number on terminal end of valve of spaldingi closely approach enoptes ancilla
and e. dammersi. S. e. dammersi feeds on Eriogonum elongatum and E. wrightii,
both related to E. racemosum. S. rita rita also feeds on E. wrightii but appears to
intergrade with rita coloradensis on E. effusum, of the Sect. Corymbosa, a taxon
quite different from Racemosa, and both rita rita and r. coloradensis average
26-28 valve teeth, exactly double that for spaldingi. The subspecies with teeth
number nearest spaldingi found so far is the Mojave Desert r. elvirae on E. pluma-
tella, but elvirae facies do not resemble those of spaldingi in the slightest. Perhaps
a connecting link between spaldingi and the known rita subspecies will eventually
turn up on the species most closely related to E. racemosum: E. panamintense,
E. rupinum and E. zionis. Another possibility is that rita rita-r. coloradensis were
the ancestral subspecies to spaldingi (r. rita has reduced cristae, as does spaldingi;
both are comparable to spaldingi in size; rarely there is a trace of an aurora on the
underside primaries of r. emmeli, rita rita and r. coloradensis; males are similar
on the upperside of rita rita, r. coloradensis and r. spaldingi), in which an aberra-
tion in genitalia isolated spaldingi (i.e., heavy sclerotized parts, fusion of teeth
to form '/2 the original number). It would have been maintained because of its
selective advantage in mimicing Plebejus melissa. S. e. dammersi and S. rita rita,
except for genitalic differences, appear nearly identical in fades, and both use
E. wrightii. Although E. wrightii and E. effusum (the latter being the host of rita
coloradensis) are widely separated by Reveal, these two Eriogonums have a
generally similar appearance and E. effusum is largely replaced by E. wrightii
in New Mexico, but occupy almost an identical ecological niche. S. r. spaldingi
does overlap the range of enoptes ancilla and approaches the range limits of
enoptes dammersi south of the Grand Canyon, but is temporally isolated from both.
In Utah, spaldingi and ancilla are sympatric in the Stansbury Mts. and the
vicinity of Eureka. In the Wasatch Range, ancilla comes down as far as Alta
(8600'), Salt Lake Co., and spaldingi comes up to the vicinity of Provo and Spring-
ville, Utah Co. There are no ancilla records for Arizona and New Mexico, where
spaldingi flies. In Colorado, the two are known to be sympatric in the vicinity
of Almont, Gunnison Co., and at Mesa Verde National Park. One female spaldingi
from nr. Swain Creek, Kane Co., Utah, has a strong tendency toward ancilla
on both dorsal and ventral surfaces.
Eriogonum racemosum ranges (5000-10,000') from central and eastern
Nevada to most of Utah, northern Arizona, northern New Mexico, and south-
western Colorado, flowering from June to September. In view of the distribution
of E. panamintense and E. rupinum, perhaps two old California spaldingi speci-
mens I have seen in the MCZ Coll. are authentic, viz: 18, "Ariz. or Cal./L. battoides
Behr/C. J. Paine Collection," and 16, "Cal. '89, Ariz_ [struck out in pencil]
battoides/C. J. Paine Collection."
Shijimiaeoides enoptes enoptes (Boisduval), 1852
Boisduval, 1852. Ann. Soc. Ent. France, (2) 10: 298-299, no. 36. (From article:
Lfpidopteres de la Californie.)
According to Williams (1918), Oberthir figured the types (Etud. Lep. Comp.
IX. P1. 237, fig. 19486, 19499) in 1913 and sent a photograph to Williams of the male
genitalia of the type in his collection (Plate IV of Williams' article). It possesses
ca. 17 valve teeth and is certainly an enoptes-type of genitalia. The original
description (as translated by Skinner, 1911) compares favorably with what today
is known as enoptes enoptes; "it is found in May in the dry [or barren] places"
(California). Perhaps an exact type locality may never be known. Barnes & Mc-
Dunnough (1917: 215) say their specimens from Truckee, Calif., exactly match
Oberthir's figures of the types, and that a series from Mineral King, Calif., is
nearly identical. The Oberthiir figures to me appear fairly typical of Sierran
Shijimiaeoides enoptes langstoni, new subspecies
Male: Holotype forewing, 11 mm. UPPER SURFACE, Primaries: ground
uniform light bright blue cyanic overlay; terminal band narrow, 2/3 mm wide;
fringe white with pronounced black checkering at vein tips and row of small
dark scales of uneven length.
Secondaries: praeterminal macules small but distinct, from M1 to 2A; cell
Sc nearly black with light blue scales scattered basally, cell Rs partially so.
UNDER SURFACE, Primaries: ground whitish gray; terminal line very
thin; checkering at vein tips, rest of fringe white; macules distinct, semimacules
well-developed; smoky suffusion in cells 1A and 2A (partial or complete).
Secondaries: aurora golden orange, from M1 to Cu2, 1/ mm wide, crescents
fused or separate; praeterminal marks distinct and separate from aurora; terminal
line thin; ground uniform over wing.
Female: Allotype forewing, 12 mm. UPPER SURFACE, Primaries: ground
black with sky blue cyanic overlay, varying from none (1) to partial to nearly
complete except outer quarter, blue diffuse toward outer perimeter (one = diffuse
throughout); fringe as in a; whitish blue band just inside terminal line (in 4 of 6);
discoidal macule rather distinct to absent.
Secondaries: blue overlay variable (lacking in one), mostly black in Sc and Rs;
thin whitish blue band just inside terminal line (in 4 of 6); fringe white with black
scales for basal 1/ at vein tips; aurora golden orange and continuous with praeter-
minal macules (separate in one).
UNDER SURFACE, Primaries: trace of aurora in lower '/A or absent; macules
distinct and prominent; fringe as in 6.
Secondaries: macules distinct to reduced; aurora in continuous band or
separate spots, 1 mm wide; fringe mostly white with basal 1/2 all white or black-white,
some slight checkering at vein tips.
HOLOTYPE 8: CALIFORNIA: Mono Co., 1.6 rd. mi. N of Mono-Inyo Co.
line, Hwy. 395, ca. 6 rd. mi. S of Sherwin Summit, ca. 5700', V-18-70 (0. & R. Shields).
ALLOTYPE 9: same.
PARATYPES: same, 23 69.
Disposition of type material: (AM).
DISTRIBUTION (viewed 45 gen. from 2 localities): CALIFORNIA: Inyo
Co.: Carroll Crk., 9 mi. SW Lone Pine, 36 19, v (CIS).
HOST: Eriogonum kennedyi Porter ex Wats. var. purpusii (Brandg.) Reveal
in Munz. At type locality, oviposition (Shields #81). Subspecies langstoni is on wing
in May (Fig. 33).
Number of teeth on terminal end (cucullus) of valve = 14, 15, 16, N = 3,
from Carroll Creek. Named for Robert L. Langston, a keen student of the genus.
S. enoptes langstoni's closest relative is doubtless e. mojave, from which it
differs in its larger size, different foodplant, and females with prominent aurora
and extensive blue on the upperside.
Eriogonum kennedyi var. purpusii occupies dry granitic flats and slopes,
5000-8000', east slope of Sierra Nevada from Mono Co. s. to Argus and Coso Mts.,
Inyo Co., flowering May-June.
Shijimiaeoides enoptes mojave (Watson & W. P. Comstock), 1920
Watson & W. P. Comstock, 1920. Bull. Amer. Mus. Nat. Hist.; 42: 455-456; no
This was originally described as a subspecies of enoptes but seems to have
been referred to as a separate species by various workers (W.J. Holland, Mattoni,
dos Passos, Langston, J. A. Comstock) ever since for no apparent reason. After
viewing many specimens of enoptes mojave from different localities, examining
the male genitalia of 23 specimens, and comparing the adults with the other
subspecies of enoptes, I am convinced it belongs as a subspecies of enoptes and
not as a separate species. The original authors note that the male genitalia of
e. mojave is definitely related to enoptes and not rita (see also Mattoni, 1954: 157),
while both sexes bear a strong resemblance in facies to enoptes enoptes but are
only three-fourths the size. The subspecies is adequately pictured in Emmel &
Emmel (1973: pl. 8, figs. 17-19). The type specimen was collected in the "Mojave
Desert, California, April 18, 1913." "These three specimens were obtained through
a dealer and no further information than that given above was available." Thus
a more precise type locality does not appear forthcoming.
Number of teeth on terminal end (cucullus) of valve = 11 (1), 12 (4), 13 (3),
14 (3), 15 (7), 16 (2), 17 (1), 19 (1), 20 (1), N = 23, average = 3-15, localities =
Jawbone Canyon, Pinyon Crest, Rock Corral, and most from Juniper Hills,
Bob's Gap, and Littlerock. A detailed life history description for this subspecies
appears in Comstock (1966). He records the larval foodplant as Eriogonum pusillum
Torr. & Gray.
DISTRIBUTION (viewed 236 gen. from 8 localities): CALIFORNIA: Inyo
Co.: Argus Mts., 36, iv, v (LACM, CAS); Argus Mts., 6000' [N. of Argus Peak],
13, v (LACM). Kern Co.: Dove Well, 18, v (CIS); Last Chance Canyon, El Paso
Mts., 26, iii (AM); E. branch Last Chance Canyon, E. off Hwy. 6, 4 mi. N Ricardo,
18, iv (CIS); Mojave, 16, iv (LACM); nr. Randsburg, Mojave Desert, 28, iv(LACM);
16 mi. S Weldon, 16, iv (CIS). Los Angeles Co.: Bob's Gap, 166 29, iii, iv (PO,
KH, JL); Bob's Gap (Holcomb Ridge), nr. Llano, 4000', 106 29, iii-v (AM); Juniper
Hills, 3600-3700', Mojave Desert, nr. Pearblossom, 183, iii-v (LACM, KH, AM);
Littlerock, 86, iii, iv (PO, JL); 1 mi. W Little Rock, 16, iv (CIS); Littlerock Dam 86
39, iii, iv (KH, GG). Riverside Co.: 6 mi. S Cottonwood Spr., Joshua Tree Nat'l
Mon., 29, iv (CIS); Pinyon Crest, 12 rd. mi. SW Palm Desert, 156 69, iv (CIS, LACM);
Pinyon Crest Jet. on Hwy. 74, 13 mi. SW Indio, 3400', "24", iv (LACM); E. entrance
nr. 29 Palms, Joshua Tree Nat'l Mon., 19 iii (GG). San Bernardino Co.: Apple Valley,
16, v (CIS) W. of Barstow, 19, iv (LACM); Bighorn Canyon, 4000', 43 mi. ESE
Victorville, 16, iv (RL); Bonanza King, Providence Mts., 16, iv (LACM); Bonanza
King Cyn., 5000', Providence Mts., 19, iv (PO); sect. 1, extreme NE corner of T2S,
R15E, N end Coxcomb Mts., 2000', 16, iii (AM); Kramer Hills, 16, iv (LACM);
5 mi. S Lucerne Valley, 4500', 23 mi. ESE Victorville, 19, iv (RL); Monarch Flat,
4200', 29 mi. ESE Victorville, 19, iv (RL); 4 mi. SSW Rabbit Dry Lake, 4300',
16 mi. SE Victorville, 56 39, iv (RL); mts. S Rabbit Dry Lake, 5 air mi. SW Lucerne
Valley, 56 29, iv (CIS); Rattlesnake Canyon [NW of Yucca Valley], 36 19, iv (LACM);
Rock Corral, ca. 20 mi. NW Yucca Valley, "abundant", iv (CS, KH); 1 mi. SE
Sheephole Summit, Sheephole Mts., 29, iv (CIS); Sunflower Wash, 3200', Old
Woman Mts., 56 39, iv (CIS); 21 mi. ESE Victorville, 4100', 16, iv (RL); 22 mi. SE
Victorville, 4700', iv (NL).
HOSTS: Eriogonum pusillum Torr. & Gray. CALIFORNIA: Los Angeles
Co.: Bob's Gap, 3800', reared 23 39 from larvae coll. v (TE). Riverside Co.: Pinyon
Crest Jet., "series", iv (FT), adult assoc. Eriogonum nudum Dougl. ex Benth. var.
pubiflorum Benth. in DC. CALIFORNIA: Kern Co.: canyon 2.2 rd. mi. SW Jawbone
Canyon turnoff on St. Hwy. 14, 2600', 26, iii (AM), adult assoc. (J. F. Emmel #51);
Last Chance Canyon, El Paso Mts., 36, iv (AM), adult assoc. (J. F. Emmel, "A").
See Map 5.
Subspecies mojave flies from mid March to mid May.
S. enoptes mojave probably is a close relative of S. enoptes dammersi, as
it is sympatric (but allochronic) with it in part of its range, both possess similar
male valve teeth number counts (mojave av. = 13-15, dammersi av. = 14-15), and
both have rather similar facies. Its nearest relative, however, is probably enoptes
Eriogonum pusillum ranges from southern California northward to SE
Oregon and SW Idaho, and across southern Nevada to SW Utah and western
Arizona, flowering from March to July (Reveal, 1969a). It is noteworthy that so
far enoptes mojave is only known from southern California (Map 5). One of Z.
speciosa's larval foodplants, E..reniforme, is closely related to pusillum. I suspect
that e. mojave has secondarily come on to E. pusillum and certainly is quite un-
related to speciosa.
I thank the following individuals for making available loans and/or records
(* = in litt.): (AM) Allyn Museum of Entomology, Lee D. Miller; (DB) David L.
Bauer; (CAS) Calif. Acad. Sci., Paul Arnaud & Thomas W. Davies; (CIS) Calif.
Insect Survey, Robert L. Langston & Jerry A. Powell; (CM) Carnegie Museum,
Harry K. Clench; (CC) *C. R. Cushing; (SE) Scott Ellis; (TE) *Thomas C. Emmel;
(MF) Michael Fisher; (GG) Glenn A. Gorelick; (RH) *Richard Holland; (KH)
Keith Hughes; (NL) Noel La Due; (JL) John Lane; (RL) *Robert L. Langston records;
(LACM) Los Angeles County Museum of Nat. Hist., Julian P. Donahue; (RM)
*R. H. T. Mattoni; (MCZ) Museum of Comparative Zoology, John Burns; (NSM)
Nevada State Museum, Peter Herlan; (PO) Paul A. Opler; (KR) Kilian Roever;
(JS) James A. Scott; (FT) Fred T. Thorne; (KT) Kenneth B. Tidwell; (YU) Yale
Univ. Coll., Douglas C. Ferguson & Charles L. Remington. Special thanks are
due Dr. James L. Reveal for identifying the Eriogonum species, and for placing
voucher specimens on file at the United States National Arboretum (NA) in Wash-
ington, D.C. John F. Emmel, Stanley K. Dvorak, David P. Levin, Scott L. Ellis,
and Michael Toliver assisted me in the field work. Scott Ellis and Scott L. Ellis
kindly provided some pertinent geology literature. This study was supported by
grants from the Allyn Museum of Entomology (Arthur C. Allyn and Lee D. Miller),
the Los Angeles County Museum (Julian P. Donahue, Charles Hogue, and Lloyd
M. Martin), John M. Burs, Theodore J. Cohn, and Charles L. Remington, and an
N.S.F. graduate traineeship. This aid enabled me to cover ca. 40,000 miles in four
summers, in the states of Calif., Oreg., Nev., Ariz., Utah, Colo., and New Mexico,
in quest of Philotes and Eriogonum. All my personal Philotes material will be
deposited with the Allyn Museum of Entomology. Paddy McHenry made available
some needed type descriptions. J. F. Emmel and my brother Dick took the photo-
graphs. Drs. J. F. Emmel, W. H. Lange, J. L. Reveal, and R. W. Thorp reviewed the
paper and offered helpful suggestions. This series of papers is submitted in partial
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Figures 29-33: habitat and Eriogonum photographs. 29: Habitat of S. battoides
ellisii, new subspecies, with its foodplant Eriogonum corymbosum var. corymbosum
in foreground; ARIZ.: Coconino Co.: 7 mi. SE Fredonia, 5000'. 30: Habitat of S.
battoides baueri, new subspecies, where its foodplant E. ovalifolium var. ovalifolium
grows; CALIF.: Inyo Co.: 2 mi. NE Westgard Pass, 7000'. 31: Habitat of S. rita
emmeli, new subspecies, 8 mi. NW past turnoff to Dugout Ranch along road to
Canyonlands Natl. Prk, San Juan Co., UTAH; E. leptocladon var. leptocladon
growing on red Entrada blowsand. 32: Another foodplant of S. rita emmeli, new
subspecies, E. smithii, at type locality for insect. 33: foodplant of S. enoptes lang-
stoni, new subspecies, E. kennedyi var. purpusii, at type locality of insect.
Figures 4-7: a genitalia of Pseudophilotes (all after Beuret, 1958). 34: P. vicrama
Moore. 35: P. bavius Eversmann. 36: P. baton Bergstrisser. 37: P. abencerragus
Figures 38-40: a genitalia of Philotes relatives. 38: Sinia leechi Forster
(after Beuret, 1958). 39: Shijimia moorei Leech (after Forster, 1940). 40: Shi-
jimiaeoides divina barine Leech (after Beuret, 1958).
Figures 41-42: 6 genitalia
Moore. 42: Z. knysa Trimen.
of Zizeeria (after Beuret, 1955). 41: Z. karsandra
Figures 43-44: 8 genitalia of Zizeeria speciosa Hy. Edwards; NEVADA:
Churchill Co.: S end Stillwater Range, 28-29 road mi. SE Fallon. 43: entire view of
genital capsule, approx 100x; cristae not visible. 44: same specimen showing detail
of valve teeth, 30 in number, approx 280x. SEM photographs courtesy R. O.
Map 1. Distribution in Utah, Colorado, Arizona, and New Mexico of Eriogonum
corymbosum, var. corymbosum (C) with short dash lines showing the form of the
variety represented by the type, the long dash lines showing the desert form, and
the dotted line which represents the low mountain and southern form; var.
erectum (E); var. davidsei (D); var. albogilvum (A); var. orbiculatum (0); var.
velutinum (V); and var. glutinosum (G) (from Reveal, 1967, Map 3).
Map 2. Distribution of Shijimiaeoides battoides ellisii (three Arizona localities
Map 3. Distribution of Shijimiaeoides rita emmeli.
Map 4. Distribution of Shijimiaeoides rita spaldingi.
Map 5. Distribution of Shijimiaeoides enoptes mojave.