BULLETIN OF THE ALLYN MUSEUM
THE ALLYN MUSEUM OF ENTOMOLOGY
Number 15 31 December 1973
STUDIES ON NORTH AMERICAN PHILOTES
II. The biology, distribution, and taxonomy of
Philotes sonorensis (F. & F.)
Department of Entomology, University of California, Davis, Calif. 95616
As I presently conceive it here, the Nearctic genus Philotes Scudder in-
cludes five basic species distinguishable on 3 genital valve features: sonorensis
(F. & F.), speciosa (H. Edw.), battoides (Behr), rita (B. & McD.), and enoptes
(Bdv.). Various authors (see, e.g., the "question" to Forbes, 1948) have noted the
difficulty in keeping these species all within the same genus. In a future paper I will
give in detail the reasons for making several generic reassignments within the
genus, based on comparative morphology, zoogeography, and foodplant relation-
ships. Mattoni (1954) proposed that sonorensis is not cogeneric with the rest
of the genus and tentatively assigned the Palearctic genus Turanana Bethune-
Baker to speciosa, battoides, [rita]and enoptes. Since sonorensis is the type species
of the genus Philotes (see Hemming, 1967), it would retain the genus name. Beuret
(1959) maintains that Turanana cytis Christ, approaches sonorensis in having well-
developed black spots on the upperside of the primaries and similar 3 genitalia
but considered these similarities not enough to combine them into a single genus.
In the event that T. cytis and P. sonorensis should become united into the same
genus upon further study, the older genus name is Philotes. T. cytis flies in Iran,
Turkestan, and Pamirs (Seitz, 1909: 305). It should be mentioned that the South
African Leptomyrina lara (L.) (see Clark & Dickson, 1957) somewhat resembles
sonorensis in undersurface color and pattern and feeds on Crassulaceae species
(e. g. Cotyledon); however, it differs from sonorensis in 8 genitalia, gross adult
morphology, and immature stages and thus is perhaps a case of convergent
In the type description of sonorensis (Felder & Felder, 1865), the type
locality is given as follows: "Habitat: Sonora. (Lorquin.) We agree equally that the
hitherto well-known Californian race is a full, isolated, stationary species"
(rough translation). Langston (1969) points out that no sonorensis are known from
the state of Sonora, Mexico, and refers to a paper by F. M. Brown which states
that the Felder type localities all come from within California. Grinnell (1904)
mentions that J. M. Lorquin took sonorensis in 1852 at San Diego, Calif, which
is hereby designated the type locality. I do not know where the type specimen
Several authors have commented on the relative lack of variation in
sonorensis throughout its range, (Martin, 1951; Langston, 1969). In series I have
examined, the extent of forewing orange spots varies considerably in 63, much
less so in 99. The orange spots on the upperside vary in intensity, sometimes
a red to very light orange or rarely pink. UFW row of black spots in 66 variable
in extent and size of spots, same on secondaries of UW. Also the amount of white
overscaling on the undersurface is variable. Six Baja California specimens
(8 from San Jacinto, La Guilla, Hamilton Ranch, San Telmo) appear to have a
slightly more whitish cast to the blue than the usually irridescent light blue.
Langston (1963) notes that central California specimens have a somewhat
duller shade of blue than southern California material. The long series of Los
Angeles Co. material viewed averages somewhat larger than central Calif.
populations. Also 99 from Los Angeles Co. are slightly more boldly marked on
the upperside, have a general darker aspect and frequently a wide dark band on
the underside, more weakly expressed in central California colonies. Mattoni
& Seiger (1963, fig. 1) figure the classes of variation of a sample from San Gabriel
Canyon. There was no variation in a short series sampled for valve teeth number
from Placerville (average = 5; 2 with 4,5 with 5, and 2 with 6 teeth) and Fish Canyon
at Azusa (average = 5; 2 with 4, 4 with 5, 2 with 6, and 1 with 7 teeth).
mean 41 male
S31.51 sonorensis from
I I desert area,
I \ San Diego Co.
Z 10. S.D.=1.71
E 9S. E.=
3 8 .267
ui 15 male coastal
O 7 sonorensis,
A 6- San Diego Co.
o 5 28.33
S4 Data and
0 S.D. =1.2 computations
Z S.E.i=.3 by F.T.Thorne
I I I i i i l l
27 28 29 30 31 32 33 34 35
SIZE MEASURED IN 1/64 INCH
Two variations of sonorensis have received names. Form comstocki
(Gunder, 1925) has the upperside spots in both sexes both above and beneath
fused into one large black spot on the primaries. Type locality is Duarte, San
Gabriel River, Los Angeles Co. Occasionally a specimen will look like this form on
the upperside but with undersurface spots normal (e.g. Tujunga Canyon, Los
Angeles Co., IV-13-60, 1&, leg. C. Goodpasture). Ova from a confined 9 comstocki
from Fish Canyon, Los Angeles Co., II-18-34, leg. C. Henne, produced one typical
6 sonorensis, emerged I-9-35 (Henne, in litt.). An "aberrant," sonoralba (Watson
& Comstock, 1920), has the two red spots on the a primary above and beneath a
cream-white (type locality = San Diego). Two specimens I have seen from Azusa
(III-12-32, III-27-49, leg. L. Martin) in the LACM collection have cream colored
spots on the upperside but with underside spots a definite yellowish-orange. In
a short series from Slide Lake, San Bernardino Co., III-26-60, leg. C. Goodpasture,
18 19 had cream spots on the upperside with light orange spots on the underside,
and la had the normal orange spots on both surfaces. Both comstocki and
sonoralba are figured in Comstock (1927).
A remarkable gynandromorph, left side 6, right side 9, was taken II-28-'00,
presumably at Pasadena, Calif. (Herr, 1900). I know of no other such examples.
Emmel, Trew, & Shields (1973) recently found considerable chromosome
variation in two central California sonorensis populations: N = 17 to 44, with
variation in number even within the same individual. We remark that this con-
dition has perhaps resulted from a stressful environment or polyploidy and hy-
bridization between host Dudleya species. San Gabriel Canyon is urgently in
need of sampling since sonorensis displays considerable variation there (e.g.
comstocki, sonoralba) and Dudleya lanceolata, D. cymosa, and D. densiflora all
grow there. Hovanitz (1967) notes that this colony has been drastically reduced
in the past few years by the ravages of man.
Central coastal California sonorensis records and a distribution map for
California appear in Langston (1965). Additional central coastal Calif. locals are
given in Langston (1963), and he figures the a genitalia. No distribution records
are listed here which are included in these two papers.
Philotes sonorensis records
BAJA CALIFORNIA: Agua del Refugio, IV-1-35 (Harbison, in Powell,
1958). Mesa W. of Cantillas Cyn., III-18-67, 1 a (J. Powell, CIS). Carrizo Canyon,
Valle de la Trinidad, III-15-36, 1 8 (C. F. Harbison, HR). Hamilton Ranch, IV-18-35,
1 a (LACM). La Grulla, IV-14-35, 1 $ (LACM). Llano de Hamalu, IV-15-35, 2 S
1 9 (LACM). Mesquite Spring, Valle de la Trinidad, III-16-36,1 a (C. F. Harbison, HR).
Punta Descanso, III-6-47, 1 6 (J. S. Garth, JG). Rio San Telmo, IV-14-35, 1 8
1 9 (LACM). Mouth of Rio Santo Tomas, 111-5-47, 1 S (J. S. Garth, JG). San Jacinto,
IV-19-35, 1 & (LACM). San Telmo, IV-15-35, 26 (LACM). Santo Tomas, 310 35'N.,
III-20-35, 1 6 (C. F. Harbison, HR). 7.6 mi. N. of Santo Tomas, in arroyo, IV-4-71,
7 6 (J. Donahue, D. Patterson, & H. G. Real, HR). 13.4 mi. S. San Vicente, IV-4-71,
1 S (H. G. Real, HR). In the Sierra Juarez (ca. 45 mi. SE of Tecate), IV-12 & 13-63,
2 8 2 9 (D. Dirks, OS). Sierra Juarez, 5500', 23 mi. S. of Tijuana-Mexicali Hwy.,
III-18-67, 1 6 (J. A. Powell, CIS). 35 mi. S. of Tijuana, III-9-46 (Provin, in Powell,
CALIFORNIA. Contra Costa Co.: Mt. Diablo (iA Williams & Grinnell,
1905). El Dorado Co.: Chili Bar, ca. 4 rd. mi. N. of Placerville, at crossing of S. Fk.
American River & Hwy. 193, 980', II-21-70, 16 a 3 9 (0. Shields, R. Wells, OS,
RW); II-22-70, 1 a 1 9 (0. Shields, OS); III-5-70, "12" (R. Wells, RW); III-16-70,
"26"; II-18-71, 2 1 9; III-15-71,15 ( 4 9 (all R. Wells, RW); II-17-73, absent (0. Shields).
Rock Creek Rd., S. Fk. American River, Placerville, 1200', II-26-68, 20 S 3 9 ;
II-29-68, 12 a 6 9; III-23-69, 2 a 4 9 (all D. L. Bauer, DB). Rock Creek, II-29-68, 2 a
1 9; 111-3-68, 1 a 2 9; III-6-70 (all P. Herlan, NSM). Rock Creek Road ca. 7 mi. NE
Placerville, 11-7-71, 16 a 1 9 (R. L. Langston, SB). Los Angeles Co.: Arroyo Seco,
V-20-07, 1 9 (V. L. Clemence, CAS). Arroyo Seco, above Pasadena, III-10-28,
3 a (J. S. Garth, JG). Arroyo Seco Canyon, San Gabriel Mts., III-22-32, 1 a; III-15-33,
6 6; IV-1-33, 2 a (all W. L. Swisher, WS); III-31-47, 1 a (D. L. Bauer, DB). Azusa,
IV-20-22, 1 a 1 9 (LACM); IV-21-22, 1 a (LACM); II-28-26, 3 3 1 9 (J. A. Comstock,
UCD, UI, LACM); III-3-27, 1 19 (Sternitsky, CAS); 1-29-28,1 6 19 (J. & C. Comstock,
AM); II-26-28, 1 9 (M. Doudoroff, CIS); II-16-29, 3 S 2 9 (CAS, LACM); VI-24-29,
2 6 (L. Martin, UW); II-16-30, 5 6 1 9 (C. M. Dammers, CAS, LACM); III-1-31,
1 S (LACM); III-14-31, 1 $ 3 9 (LACM); III-12-32, 13 a 3 9 (C. W. Anderson, L. M.
Martin, CIS, LACM); III-11-33,2 49 (CAS); II-24-35, 16 29 (J. A. Comstock, LACM);
III-10-35, 7 3 2 9 (CAS); III-29-36, 1 6 3 (CAS); III-20-37, 5 5 (MCZ); III-21-37, 2 6 1 9
(CAS); III-11-40, 1 a 1 9 (F. W. Furry, LACM); III-11-43, 1 S (CAS); III-7-45, 1 S (PO);
III-22-48, 1 4 9 (AM); III-6-49, 2 a (L. M. Martin, LACM); III-26-49,6 6 3 9 (LACM);
III-27-49, 1 S 3 9 (L. M. Martin, LACM); IV-5-49, 1 S 1 9 (AM); IV-11-49, 1 3 (C. A.
Sekerman, CS); IV-13-49, 1 9 (AM); II-28-50, 2 9 (AM); III-18-51, 4 (L. Martin, UW);
III-20-53, 1 6; II-22-54, 2 6 1 9 ; II-24-54, 1 9; II-25-54, 3 9; III-1-54, 3 9 ; III-12-54,
1 1 9; III-7-55, 2 9 (all AM); III-22-58, 1 9 (L. M. Martin, LACM); III-12-59, 4 a
2 9 ; III-14-59, 1 6 5 9 (both K. Hughes, KH). Nr. Azusa, II-28-26, 1 6, (LACM);
III-24-40, 1 6; III-6-41, 1 a 2 9 (both C. I. Smith, CIS). Bell Canyon, III-30-57, 1 9
(J. Powell, CIS). Big Dalton Canyon, IV-17-21, 2 8 (Ingham, in Gunder, 1930).
Big Santa Anita, VI-19-07, 1 6 (CAS). Big Tujunga Canyon, III-7-48, 1 6 (R. L.
Langston, RL); II-12-61, 2 6 (J. Lane, C. Sekerman, JL, CS). Camp Baldy, IV-?-32,
2 6 (T. Craig, CAS, CIS). Camp Josepho, Rustic Canyon, IV-4-69, 1 S (G. A. Gorelick,
GG). Claremont, 1 6 1 9 (Peck, LACM). Cold Creek, 1200', Santa Monica Mts.,
II-23-61, 1 6 (C. Henne, CH). Dark Canyon, San Gabriel Mts., III-18-51, 1 9 (C.
L. Hogue, LACM). Eaton Canyon, San Gabriel Mts., V-4-47, 2 6 (R. L. Langston,
RL). Eaton's Canyon, Pasadena, VI-1-07, 1 S (V. L. Clemence, CAS). Elsmuir
Canyon, III-10-40, 1 6 (D. L. Bauer, DB). Fish Canyon, nr. Azusa, IV-20-22, 11 6
8 9 (LACM); III-7-24, 5 6 1 9 ; III-30-24, 1 6 (both J. A. Kusche, CAS, CIS); III-1-25,
2 8 2 9 (LACM); II-27-26, 40 6 24 9 (J. S. Garth, JG); III-23-27, 2 9 (E. Walter, EW);
IV-2-27, 25 & 7 9 (J. S. Garth, JG); III-7-31, 3 1 9 (CIS); III-19-31,1 9 (E. Walter, EW);
III-13-49, 4 9 (L. M. Martin, LACM). Fish & San Gabriel Canyons, III-
14-25, 50 8 10 9 ; III-12-27, 13 6 2 9 (both J. S. Garth, JG). Glendora,
II-16-30, 1 9 ; II-17-30, 2 9 ; II-23-30, 1 9 ; III-27-30, 2 II1-22-31, 3 6 (all AM);
III-8-31, 1 1 9 (YU, AM); IV-10-31, 1 6 2 9 (LACM); II-22-32, 1 6 (OS); II-22-33,
1 6 2 9 (OS). Glendora Mtn. Rd., off E. Fk. San Gabriel River, V-7-67, 2 3 2 9 (J. Lane,
C. A. Sekerman, JL, CS). Henniger Flats, San Gabriel Mts., III-5-60, 16 (C. L.
Hogue, LACM). Indian Springs trail from Big Tujunga Canyon, III-1-42, 23 (R. L.
Langston, RL). Inglewood, III-12-32, 13 19 (YU). Ladybug Canyon, Angeles Crest
Hwy., San Gabriel Mts. IV-24-65, 26 (J. Lane, C. A. Sekerman, JL, CS). La Tuna
Canyon, Verdugo Mts., III-17-42, 13; II-18-48, 16; III-5-48, 13 (all W. H. Evans,
AM); III-27-48, 16 (C. I. Smith, CIS); III-4-50, 16; III-17-51, 18; IV-9-51, 13, III-1-54,
1d; III-8-55, 13; III-25-55, 16; IV-4-55, 16; III-20-56, 16; III-24-57, 13 (all W. H.
Evans, AM); 1200', III-6-53, 19; 1360', IV-10-47, 19 (both W. H. Evans, AM). 1 mi.
E. of end of Paved Lat. Cn. Rd., 1350', main canyon, La Tuna Canyon, III-20-42,
19; III-26-42, 13 (both W. H. Evans, AM). Below spring, Wildwood Fire Rd., La
Tuna Canyon, III-6-42, 18; II-20-42, 16; III-3-42, 36; III-27-42, 23 (all W. H. Evans,
AM); III-11-59, 16; III-18-59, 13 (both C. Henne, CH). 1 mi. S. Little Rock Dam,
San Gabriel Mts., III-25-62, 19 (C. L. Hogue, LACM). 2 mi. S. of Little Rock Dam,
III-25-61, 19 (C. L. Hogue, LACM). Littlerock Res., III-20-65, 26 (J. Lane, C. A.
Sekerman, JL, CS). Los Angeles, "coll. Le Moult", 128 19 (AM); III-25-20, 18 (J.
A. Comstock, CAS). Malibu Road, Santa Monica, III-14-15, "44" (Haskins, in
Gunder, 1930). Millards Canyon, Pasadena, IV-8-07, 28 19 (V. L. Clemence, MCZ,
LACM, CAS); V-24-07, 36 (V. L. Clemence, CAS); III-22-08, 13 (LACM); III-14-09,
16 (F. Grinnell, Jr., CAS); II-27-18, 19 (J. A. Comstock, LACM); III-19-27, 13 (J.
S. Garth, JG); III-25-27 (Newcomb, in Gunder, 1930); III-16-40, 19 (J. S. Marsh,
RL); III-8-42, 16 (D. L. Bauer, DB). Mint Canyon, V-4-41, 13 (D. L. Bauer, DB);
III-15-20, 18 19 (J. A. Comstock, LACM). Summit of Mint Canyon, IV-15-28, 46
(Friday, in Gunder, 1930). Monrovia, II-24-24, 23 (E. Walter, EW). Mt. Lowe,
V-18-19, 13 (Malcolm, in Gunder, 1930). Mount Wilson Toll Road, III-16-41, 16
(R. L. Langston, RL). Pallett Crk., 4900', San Gabriel Mts., III-31-62, 16 (C. Henne,
CH). Pasadena, 26 (CAS); III-10-'98, 13 (LACM); III-28-01, 16 (Moeser, CAS);
"3/4", 15 (MCZ); "3/12", 19 (MCZ); "3-15", 19 (LACM); "2/25", 19 (MCZ); "2/28",
15 (MCZ). Rustic Canyon, Santa Monica Mts., II-25-47, 15 (W. A. McDonald,
LACM). San Gabriel, III-11-27, 15 (YU); III-20-27, 15 19 (YU); III-11-37, 15 (YU).
San Gabriel Canyon, 15 (T. Craig, CAS); III-4-?, 15 (W. Hovanitz, CAS); III-14-26,
45; III-18-26, 66 59 (both T. Craig, CAS); III-11-29, 18; III-19-29, 19 (both E. Walter,
EW); III-4-31, 568 109 (J. S. Garth, JG); IV-3-31, 45 19 (YU); III-19-32, 15 (PO);
III-4-35, 1( 19 (LACM); 11-9-36, 103 89 (LACM, OS); II-29-36, 38 39 (E. Y. Dawson,
YU); III-6-37, 28 19; II-13-38, 88 99 (both OS); IV-3-45, 45 49 (C. I. Smith, CIS); IV-
30-49, 25 19 (J. S. Garth, JG); III-24-56, 38 39 (UI); II-28-59, 25 (UI); III-19-60, 65
69 (J. Lane, C. A. Sekerman, JL, CS); IV-9-60, 25; 1-21-61, 65 (both J. Lane, C. A.
Sekerman, JL, CS); III-24-63, 25 (AM); II-22-64, 15 (AM); III-18-24, 19; III-27-64,
15 (both AM); V-14-65, 15 (R. E. Stanford, KH); III-22-68, 35 (C. C. Callaghan,
AM). Entrance to San Gabriel Canyon, II-12-68, 3 (LACM). Lower end San Gabriel
Canyon, 25 19 (J. Riddell, JJ). Mouth of San Gabriel Canyon, III-2-21, 495 219
(Malcolm, in Gunder, 1930); IV-15-33, "15" (W. L. Swisher, WS); II-28-34, 25 19;
III-4-34, 19; III-8-34, 23 19; III-21-35, 29; III-1-35, 15 59; III-14-35, 23; III-20-37, 19
(all C. Henne, CH); III-14-65, 46 (J. Lane, C. A. Sekerman, JL, CS). San Gabriel
Canyon, 2 mi. N. of Azusa, II-21-25, 245 29 (J. S. Garth, JG). San Gabriel Canyon,
near W. Covina, III-20-67, 145 69 (G. A. Gorelick, GG). San Gabriel Mts., III-22-08,
19 (V. L. Clemence, CAS); III-18-20, 85; IV-5-20, 15 (both J. A. Comstock, LACM);
IV-9-49, 15 (R. P. Allen, CAS). San Gabriel River, Azusa, III-26-49, 75 29; III-28-
49, 55 59 (both CAS). San Gabriel Wash, III-17-34,15 (LACM). Santa Anita Canyon,
III-14-31, 15 19 (CAS, CIS). Santa Monica Canyon, IV-2-17, "fairly common"
(M. E. Fountaine, in Fountaine, 1917). Shortcut Canyon, W. Fk. San Gabriel River,
V-17-42, 29 (R. L. Langston, RL). Summit Sierra Pelona Valley, IV-21-35, 78 19
(C. Henne, CH); IV-19-36, 1( (C. Henne, CH). Switzers, III-21-32, 18 (UCD).
Switzer's Camp, Aroyo Seco Canyon, San Gabriel Mts., IV-24-65, 105 (J. Lane,
C. A. Sekerman, JL, CS); IV-10-71, 13 (+ several others) (E. Walter, EW). Tie Canyon,
4700', III-17-50, 15 (W. H. Evans, AM). Vasquez Rocks, Mojave Desert, II-23-47,
55 (R. L. Langston, RL). Mariposa Co.: 4 mi. N. of Bear Valley, IV-15-61, 13 (P. A.
Opler, RL). Briceburg, 111-7-70, 65; II-14-71, 35 29; III-5-71, 65 19 III-20-71, 215
59 (all J. Mori, JM); II-18-73,29 (J. Brock, JB). Briceburg, to ca. 1 mi. E. of, along RR
bed, III-11-72, "15" (J. Brock, J. Mori, O. Shields, & W. Steele). Jerseydale, along
Skelton Creek (canyon to Devil's Gulch), 19E04.1 & 19E04.2, 8.5 air mi. NE of
Mariposa, III-29-70, "common" (J. F. Emmel & O. Shields, OS); IV-12-70, 35 19
(J. Mori, O. Shields); III-27-72, "6" (M. O'Brien, O. Shields, OS). Monterey Co.:
Big Sur, V-4-52, 2 9 (T. W. Davies, RL); II-25-54, 5 8 (AM); IV-?-55, 1 (M. Doudoroff,
MCZ). 10 mi. S. Carmel, IV-14-63, 1 5 (D. W. Conrad, CIS). Partington Canyon,
IV-15-52, 5 5; IV-29-52, 1 C (both AM); V-4-52, 1 5 (T. W. Davies, YU); IV-21-56,
5 3 (P. A. Opler, PO). Partington Canyon, 9 mi. S. of Big Sur, IV-9-68, 13 6 5 9
(R. L. Langston, RL, CAS, LACM); V-10-68, 1 3 1 9 (R. L. Langston, LACM);
V-11-64, 2 C (R. M. Brown, RL). Orange Co.: Aliso Canyon, Laguna Beach, II-16-35,
1 5 (LACM); III-6-35, 6 5 (D. Meadows, RL); III-10-35, 3 6; III-16-35, 1 (; III-1-36,
1 5 (all LACM). Black Star Canyon, III-8-64, 2 5; 111-19-64, 2 5 (both E. Walter,
EW). Irvine Park, III-7-34, 4 9; III-11-45, 1 5 1 9 (both E. Walter, EW). On the hills
ca. 1 mi. N. of Irvine County Park, in Santiago Canyon, Santa Ana Mts., III-17-32,
2 5; III-18-33, 1 5 1 9; III-3-45, 1 5 (all J. W. Johnson, JJ). Ladd Canyon, Santa Ana
Mts., II-16-53, "numerous" (J. Mori, JM); III-21-53, 1 5 (Cooper, UW); IV-18-53,
1 5 (Cooper, UW). In hills to N. of Olive, III-2-34, 1 5 1 9 (E. Walter, EW). San Juan
Canyon, 700', Cleveland Nat. Forest, V-5-55, 1 9 (C. Henne, CH). Lower Santa Ana
Canyon, IV-1-29, 1 9 (LACM). Santiago Canyon, III-15-24, 3 (; III-15-25, 2 5
III-10-31, 2 5, III-19-33, 2 3; III-15-64, 2 & (all E. Walter, EW). Silverado Canyon,
III-31-28, 2 8 (T. Craig, CAS, CIS); II-16-65, 1 6; III-29-65, 1 5 (both E. Walter,
EW). Silverado Canyon rd. from Irvine Park to Silverado Canyon, at fork to
Modjeska Canyon (R. Wells). Trubuco River, IV-8-60, 1 5 (J. F. Emmel, OS).
Placer Co.: near Auburn, III-2-65 (N. La Due, RL); III-25-67, 2 5 (C. Goodpasture,
OS). Auburn, Jet. North & Middle Forks of American River, 600', II-29-68, 7 5
3 9 (D. L. Bauer, DB). American River E. of Auburn, III-15-69, absent (0. Shields);
1-27-70, absent (R. Wells); 11-4-70, absent (R. Wells). 3 mi. NE of Auburn on Hwy.
49, Middle Fk. American River, 580', l/2 mi. upstream from bridge, II-21-70, 4 &
3 9 (0. Shields, R. Wells, OS, RW); II-23-70, 2 9 (R. Wells, RW); 1-30-71, absent (R.
Wells); 11-2-71, absent (R. Wells); II-10-71, 2 6 2 9 (R. Wells, RW); III-8-70, 1 a
(+ many eggs) (R. Wells, RW). Rocks above & W. of Murderer's Bar, Middle FK.
American River, %-/-1. mi. E. of juct. of Middle & North Fk. of American River,
II-6-71, 2 (6; II-13-71, 58 6 12 9 (both O. Shields, R. Wells, OS, RW); II-14-71, 17 8
89 (R. Wells, RW); II-20-71, 14( 19; II-23-71, 63; III-16-71, 46 19 (all R. Wells, RW);
II-12-72, absent (0. Shields, R. Wells); II-19-72, "41" (J. Mori, R. Wells,
JM, RW); II-20-72, 203 49 (0. Shields, R. Wells, OS, RW); II-17-73; 2 eggs,
no adults (S. Mattoon, O. Shields, S. Sims); II-23-73, 73 (0. Shields, S.
Sims, OS). North Fk. American River, 3 mi. NE of Auburn, III-
23-65, 13 29 (R. L. Langston, CIS); III-23-68, 13 (R. L. Langston,
RL); II-22-70, 11 a 1 9 (R. L. Langston, RL, NSM). Riverside Co.: L. S. C., Arlington,
II-22-42, 1 3 (D. L. Bauer, DB). The Gavilan, IV-2-30, 2 9 (C. M. Dammers, CAS);
III-9-41, 7 6 (C. Dammers, LACM). Gavilan Hills, 2200', III-14-59, 1 19 (C. Henne,
CH). La Sierra College, III-12-39, 1 9; III-19-39, 1 9 (both D. L. Bauer, DB). Mocking-
bird Canyon, III-20-45, 1 9; III-23-46, 1 9 (both W. H. Evans, AM). Perris, III-19-
53, 1 8 (LACM). Pinon Flat, Santa Rosa Mts., IV-23-51, 2 6 (C. D. MacNeill, RL).
Railroad Canyon, 4 mi. E. of Elsinore, IV-17-65, 1 9 (J. Powell, CIS). Sage, III-
20-51, 6 3 2 9 (R. H. Reid, LACM). Snow Creek, III-12-33, 1 6 (CAS). Upper San
Andreas Canyon, III-22-31, 3 3 1 9 (J. S. Garth, JG). Snow Creek Canyon, III-30-
29, 2 8 (J. S. Garth, JG). Whitewater, III-25-30, 1 9 (C. M. Dammers, CAS). San
Benito Co.: Chalones Crk. trail to the Balconies, Pinnacles Natl. Mon., III-18-72,
6 ( (R. Langston, P. Sisk, RL, SB). Fremont Peak (in Tilden & Mansfield, 1941).
Lime Kiln Canyon, 5 mi. W. Paicines, III-24-66, 1 8 3 9 (R. L. Langston, J. Powell,
CIS); III-2-68, 1 3 (R. L. Langston, LACM). San Bernardino Co.: 5 air mi. SW of
Lucerne Valley, mts. S. of Rabbit Dry Lake, IV-14-64, 1 3 (R. L. Langston, CIS).
Along bank of Mojave River, 6 mi. SE of Hesperia, IV-15-62, 1 6 (R. M. Brown, RL).
San Bernardino, 1 9 (Wright, CIS). Slide Lake, III-26-60, 2 6 1 9 (C. Goodpasture,
OS). Upland, IV-18-58, 2 6 (T. M. Blackman, LACM). San Diego Co.: Bennington
Cemetary, Point Loma, II-16-30, 27 6 5 9; II-22-32, absent; II-28-32, absent; II-
12-33, 3 2 9; II-19-33, 10 4 9; III-4-33, 4 7 9; II-12-34, 4 1 9; II-26-34, absent (after
several heavy rains); II-12-35, 1 6 1 9; III-1-36, 20 3 4 9; II-22-37, 12 6 2 9 (all F. T.
Thorne, FT). Boiling Springs, Laguna Mts., IV-29-34, "1" (F. T. Thorne, FT).
S. slope of Dictionary Hill, in large canyon, IV-5-62, 1 9 (F. T. Thorne, FT). Division
Street, about 5600 to 5900 blocks, National City, II-3-48, 1 6 (R. W. Breedlove,
SD); II-21-50, 46' 19; III-7-50, 16 89; III-12-50, 216 59; II-8-51, too early; II-19-51,
13 (all F. T. Thorne, FT); II-24-51, 18 19 (R L. Langston, RL); II-25-51,
126 39, (F. T. Thorne, FT); III-4-51,16 (R. L. Langston, RL); III-14-51, "Santa Ana con-
dition, adults common"; III-3-52, 26 19; III-6-52, 26; II-6-53, 46 19; II-7-53,
7 3 19; II-15-53, 2 6 3 9; IV-9-54, absent (all F. T. Thorne, FT); III-14-55, 2 3 (R. W.
Breedlove, SD); III-3-56, 4 6 2 9; III-7-57, absent; II-21-58, absent (all F. T. Thorne,
FT); III-21-59, 1 9; 11-26-61, 2 6 1 9 (both R. W. Breedlove, SD); III-4-62, 1 6 (F. T.
Thorne, FT); IV-1-62, 1 (R. W. Breedlove, SD); II-5-64, absent; II-7-68, absent (both
F. T. Thorne). Encanto, III-11-50, 5 6 1 9; II-24-51, 2 6 1 9 (both J. Powell, CIS);
III-28-52, 1 6 1 9 (M. J. McKenney, CIS). Summit of Garnet Peak, 6000', Laguna
Mts., V-17-69 (F. T. Thorne, RL); V-8-70, 1 3 (R. Priestaf, RL). 3 mi. W. of Jacumba,
3200', III-11-60, 1 9 (0. E. Sette, RL). Julian, IV-13-'99, 1 3 (LACM). Laguna Mts.,
"6/3/51", 1 9 (M. J. McKenney, CIS); VI-1-56, 2 3 (R. E. Stanford, KH). Hills E.
of La Jolla, III-2-40, 7 ( 1 9; III-3-40, 4 6 2 9; III-5-40, 2 6 (all O. E. Sette, RL). Back
of La Jolla country club, II-23-41, "2" (F. T. Thorne, FT). La Pressa Canyon, SE
of Dictionary Hill, III-5-55, 1 6; III-12-55, 1 6; III-15-55, 2 3; III-19-55, 1 6; III-1-
58, 1 3 (all R. W. Breedlove, SD). Lemon Grove (in Wright, 1908). Mine Canyon,
Pinon Mts., Anza Borrego State Park, II-14-71, 7 6 4 9 (S. K. Dvorak, SD). N. side
of Mission Valley, in canyon about opposite Co. Hospital, III-1-42, "8" (F. T. Thorne,
Fowler, FT). Monument Peak, Laguna Mts., IV-16-33, 1 6 (F. T. Thorne, FT).
Hills 1 mi. S. of Otay, III-17-51, 4 6 (R. L. Langston, RL, CAS, CIS). "Paradise
Hills", 6 mi. SE of San Diego, III-3-56, 5 a 1 9 (0. E. Sette, RL). S St. and Otay Rd.,
III-3-56, 2 6 (F. T. Thorne, FT). San Diego, II-27-14, 1 6 (G. H. Field, YU); "2
July 1917", 1 9 (YU); II-15-47, 1 9 (J. Powell, CIS); II-24-51, 3 6; III-11-51, 3 6 (both
M. J. McKenney, CIS); II-13-53, 3 6 (J. Powell, CIS). San Felipe Creek, Anza
Desert, III-5-60, 2 S (R. W. Breedlove, SD). Canyon off Warner leg of Scissors
Crossing, III-15-49, 3 ( 1 9 (F. T. Thorne, FT). Near Scissors Crossing at base of
Grapevine Mts., III-25-51, 1 6 + 10 eggs; III-20-52, "1" + eggs & 1 larva (both F. T.
Thorne, FT). Scissors Crossing, II-16-54, 7 6; II-20-54, 10 a 19; II-21-54, 14 6;
II-22-54, 14 6 2 9; II-19-55, 1 (; II-21-55, 2 9; III-12-55, 4 6 3 9; II-19-56, absent; III-
4-56, 6 8 3 9; II-26-57, 7 a 1 9; 1-11-58, 7 S (all F. T. Thorne, FT); 1-28-58, 16 6 4 9
(R. Albright, F. T. Thorne, FT); III-29-58, 1 6; II-19-61, 1 9 (both O. Shields, LACM);
III-11-51, 1 6 (R. L. Langston, CIS); III-12-55, 1 & (F. T. Thorne, OS); II-17-60,
"3"; III-11-61, 4 eggs, no adults; II-18-62, 1 3; II-16-63, 1 6 2 9; II-18-63, 3 6; II-22-
64, 2 8 (all F. T. Thorne, FT)..6 mi. E. of Scissors Crossing, III-12-60, 1 (0. Shields,
LACM). 3 mi. E. of Scissors Crossing, III-11-51, 43 1 9 (R. L. Langston, RL, CIS).
1 mi. W. of Scissors Crossing, II-13-61, 1 9 (0. Shields, LACM). "Two Mile Hill",
2500', W. of Scissors Crossing, III-1-64, 19 (0. Shields, LACM). Hills NE of Scissors
Crossing, 7 mi. E. of Banner, III-4-56, 15 6 3 9 (0. E. Sette, RL). 1 mi. NE of Scissors
Crossing, San Felipe Valley, III-19-66, 2 6 1 9 (S. K. Dvorak, SD). 2 mi. N. of Scissors
Crossing, II-1-58, 3 3 3 9; III-6-60, 3 6 (both D. Dirks, O. Shields, LACM). 3 mi. N.
of Scissors Crossing, II-1-58, 26 1 9 (D. Dirks, O. Shields, LACM). Tributary canyon
in middle of Sentenac Canyon, II-8-46, 2 3 1 9; II-24-46, 3 3 1 9 (both F. T. Thorne,
FT). Sentenac Canyon, III-14-53, 9 6 3 9 (J. Powell, CIS); III-22-53, 1 9 (R. A. Mackie,
UI); III-22-53, 12 3 2 9 (J. Powell, CIS); III-24-53, 1 8 (M. J. McKenney, CIS);
II-26-57, 7 6 2 9; III-2-57, "1" (both F. T. Thorne, FT); 1-11-58, "2"; 11-8-58, "2"
(both F. T. Thorne, FT); II-9-58, 2 6; 11-6-59, 1 6 (both K. Hughes, KH); II-27-59,
2 3 1 9; III-8-59, 2 6 (both F. T. Thorne, FT); III-12-60, 16 (S. K. Dvorak, SD); III-
30-60, absent (F. T. Thorne); II-19-61, 3 9 (0. Shields, LACM); III-11-61, 1 6; III-17-
62, 2 6; 1-9-63, 1 9 (all S. K. Dvorak, SD); II-16-63, 17 3 1 9 + "18" (R. W. Breedlove,
S. K. Dvorak, F. T. Thorne, FT); II-18-63, 4 6; II-22-64, 2 3 (both F. T. Thorne, FT);
2000', III-1-64, 1 3 (0. Shields, AM). Sentenac Canyon & Two Mile Hill, II-14-65,
16 6 2 9 (R. W. Breedlove, FT). Sweetwater Lake, III-19-60, 1 9 (D. Dirks, O. Shields,
LACM). Sweetwater Valley near Higgins Ranch, Bonita, III-3-40, 3 6; III-17-40,
1 6; II-4-41, 2 6; III-6-41, 3 6; II-15-42, absent; III-8-42, 4 ( 4 9; II-12-44, absent;
IV-2-44, 1 3; III-10-49, absent; 1-29-53, absent; II-25-53, absent (all F. T. Thorne,
FT). Telegraph Canyon, SE of Chula Vista, 1-1-18, 3 6 1 9; III-8-18, 1 9; III-9-18,
1 6; III-16-18, 3 3; IV-30-18, 1 9; III-3-56, 2 3 (all O. E. Sette, RL). N. entrance to
Torrey Pines State Park, 50', S. of Del Mar, II-21-64, 1 6 (0. Shields, LACM).
"Two Mile Hill", 2500', 4 mi. E. of Banner, III-4-56, 1 9 (0. E. Sette, RL); II-26-57,
"2"; II-9-58, 1 9; III-11-61, absent (all F. T. Thorne, FT); II-18-68, 10 3 19 (R. W. Breed-
love, FT). Vallecito Mts., III-6-49, 1 ( (D. L. Bauer, DB). San Luis Obispo Co.:
Atascadero, III-6-32, 5 8; III-7-32, 1 8; III-8-32, 1 3; III-13-32, 3 6; III-16-32, 2 6;
III-18-32, 4 8; IV-4-32, 1 3; III-14-35, 1 $; III-26-35, 1 6; IV-5-35, 1 6 1 9 (all V. L.
Clemence, LACM). Cerro Alto Canyon, IV-16-61, 1 8 (V. Chang, OS). Vicinity of
Montana De Oro State Park, S. of Morro Bay, ca. II-25 or 26-72, "about a dozen"
(T. Tolman, WS). Oso Flaco sand dunes, 1 mi. SE of Oso Flaco Lake, III-1-72,
16; III-9-72, 1 3 (both W. L. Swisher, WS). Paso Robles, III-10-'94,1 6 (W. G. Wright,
CIS). Santa Barbara Co.: Nr. Davy Brown Campground, San Rafael Mts., III-24-
73 (R. C. Priestaf, R.P). Mission Canyon, Santa Ynez. Mts., III-?-62, 1 6
(J. W. Tilden, AM); II-23-71 (R. C. Priestaf, RP). Oso Creek, San Rafael Mts.,
early III-69 (R. Priestaf, R.P.). Refugio Pass, III-11-61, 5 6 (J. S. Garth, JG).
Old San Marcos Pass Rd., Santa Ynez Mts., II-21-71 (R. C. Priestaf. RP). Santa
Barbara, IV-14-33, 1 6 (UI); II-8-34, 1 9 (LACM). Santa Ynez Mts., "early III-70"
(R. Priestaf, RL), "reappeared early Mar. after disastrous 1964 fire." Santa Clara
Co.: Alma, II-?-?, "2" (Grundel, in Coolidge, 1908). Alum Rock, II-23-47, 3 6
(YU). Alum Rock Canyon, III-16-40, 1 6 (CAS). Alum Rock Park, III-1-42, 1 3
1-30-48, 2 a (both AM); II-19-50, 3 6 (T. W. Davies, YU); II-21-54, 1 6 (P. A. Opler,
RL); III-7-54, 1 6 (LACM); III-11-56, 2 & 2 9 (P. A. Opler, PO); III-21-57, 3 9 (0. E.
Sette, RL); II-21-59, 1 S (P. A. Opler, PO); II-27-59, 1 6 (R. E. Stanford, KH); III-
8-59, 1 9 (P. A. Opler, PO); III-14-60, 1 6 (R. E. Stanford, KH); IV-2-60, 1 6 (0. E.
Sette, RL); II-12-64, 12 6 5 9 (R. L. Langston, RL, CIS, CAS, LACM, AM); II-21-64,
1 8; II-22-64, 7 a 1 9; III-7-64, 1 6; II-11-65, 1 6; II-13-65, 1 a (all P. A. Opler, PO);
II-14-65, 2 S (A. J. Slater, CIS); II-20-65, 4 8 1 9 (W. J. Steele, RL); III-1-65, 5 6
3 9 (P. A. Opler, PO); 11-21-67, 13 a 8 9 (R. L. Langston, RL, CAS, LACM, AM);
II-26-67, 7 1 9; II-28-67, 7 6 9 (both G. A. Gorelick, GG); III-1-67, 3 (C. D. MacNeill,
RL); 1-30-71, 5 6 1 9 (R. L. Langston, RL, SB); II-14-73, 31 S 9 9; II-23-73, 7 a (both J.
Brock, JB). Coyote Crk. nr. China Hole, Henry W. Coe State Park, III-20-71,
1 5 (G. Frisch, RL); III-21-71, 1 5 (P. Sisk, RL). Gilroy (in Coolidge, 1908). Soda
Springs Creek, Henry W. Coe State Park, III-20-71, 4 5 (R. L. Langston, RL, SB).
Santa Cruz Co.: Gavilan Mt., 2400', Santa Cruz Mts., IV-18-18, 1 9 (W. E. Groth,
CAS). Stanislaus Co.: At mouth of Adobe Cyn., on Del Puerto Cyn. road, 1320',
ca. 21/2 rd. mi. W. of Frank Raines Park, S. 15, T.6S., R.5E., II-23-70, 3 (J. F. Emmel,
OS). 18 road mi. W. of Patterson, ca. 1700', Del Puerto Canyon, III-7-70, 1 8
1 9 (0. Shields, OS). Del Puerto Canyon, 21 mi. W. of Patterson, III-15-69, 1 5
1 9 (R. L. Langston, RL); II-27-72, 2 5 2 9 (R. L. Langston, C. Henne, RL, SB).
Del Puerto Canyon, 22 mi. W. of Patterson, III-5-63, 1 9; IV-30-63, 1 5; II-22-64,
4 3 4 ; III-12-66, 1 9 (all R. L. Langston, RL, CIS, CAS); III-30-68, 1 (G. A. Gorelick,
GG); III-15-69, 2 8 (R. L. Langston, LACM); 111-29-69, 7 3; IV-10-69, 2 6 (both G. A.
Gorelick, GG); 1200', II-20-70, 2 8 2 9; II-25-70, 7 3; II-7-71, absent (all J. Mori, JM);
III-11-73, 1 3 1 9 (0. Shields, OS). Del Puerto Canyon, 23 mi. W. of Patterson, II-
19-67, 1 5; III-25-69, 5 3 2 9 (both J. Powell, CIS); 1500', II-27-72, 2 3 2 9 + 7 eggs
(C. Henne, R. L. Langston, RL). 0.3 road mi. SSE of mouth of Hideout Cyn., 1550-
1600', Del Puerto Canyon road, S. 16, T.6S, R.5E., II-23-70, 2 3 (J. F. Emmel,
OS). Tuolumne Co.: N. Fk. Tuolumne River, 3 mi. NE of Tuolumne, IV-2-61,
1 6; II-20-65, 1 3; IV-25-65, 1 5 (all M. R. & S. H. Lundgren, CIS, CAS); III-23-66,
4 6 (R. L. Langston, CIS). Ventura Co.: Derrydale Canyon, 3600', Los Padres Nat.
For., IV-19-58, 1 5 (C. Henne, CH). 1 mi. S. of Lake Sherwood, Santa Monica
Mts., III-17-67, 2 a (P. Opler, J. Powell, CIS). 2 mi. E. of Lake Sherwood, III-16-
67, 1 6 (J. Powell, CIS). Port Hueneme, 10 mi. E. of, II-26-'94, 2 6 (W. G. Wright,
CIS, CAS). County undetermined: Agusta, III-16-46, 1 5 (L. M. Martin, UW).
Fork Canyon, III-11-24, 1 a (CAS). Mouth of San Antonio Canyon, III-25-32,
1 3 (T. Craig, CAS). Doubtful or erroneous records: Brawley, II-18-25, 1 6 3 9
(Fortener, "E. A. Dodge Collection", CAS). "Near Brawly", V-6-25, 1 9 (Wehner,
CAS). Laguna Mts., "Sep. 18 25", 1 a 1 9 ("J. P. Strohbeen Collection", CAS):
[below this questionable date is "Brauley", which may explain the "Brawley"
Records for P. sonorensis form comstocki
CALIFORNIA. Los Angeles Co.: Mouth of Fish Canyon, II-25-30, 1 6
(Ingham, in Gunder, 1930). Fish Canyon, III-20-37, 1 6 (C. H. Ingham, AM).
Mint Canyon, 111-20-21, 1 6 (Malcolm, in Gunder, 1930). Mouth of San Gabriel
Canyon, III-9-30, 1 6 (Friday, in Gunder, 1930); III-4-33, 4 9; III-7-33, 1 8; II-18-34,
1 6; II-28-34, 1 9; III-8-34, 1 1 9; II-16-35,1 3 1 9; III-1-35, 1 ; III-12-35, 2 ; III-7-36,
1 3; III-15-36,1 8; III-28-36, 1 9; III-7-37, 1 5 (all C. Henne, CH; note, Henne estimates
that the ratio of comstocki to typical sonorensis here is 1 to 200). San Gabriel River,
Duarte, III-15-22, holotype (AMNH). Ventura Co.: Derrydale Canyon, 3600',
Los Padres Nat. For., IV-19-58, 1 3 (C. Henne, CH).
Records for P. sonorensis ab. sonoralba
CALIFORNIA. Los Angeles Co.: mouth of Fish Cyn., Duarte, III-12-35,
1 9; III-20-37, 1 9 (C. Henne, CH). San Gabriel Wash, III-5-24, 1 8 (J. A. Comstock,
in Gunder, 1930).
P. sonorensis in central California is found in association with Dudleya
cymosa (Lem.) Britt. & Rose. D. cymosa ssp. cymosa is used at North Fork American
River E. of Auburn, Placer Co. (det. R. Moran), where a larva collected V-13-70
emerged VIII-30-70 (9). Adults were reared from larvae on this ssp. at Briceburg,
Mariposa Co. (Shields #68, det. D. Breedlove), and adults were taken in association
with it at Skelton Creek Canyon E. of Jerseydale, Mariposa Co. (Shields #71,
det. D. Breedlove). Langston took adults in association with it at 3 mi. NE Tuolumne
City, Tuolumne Co. Adults have been collected around D. cymosa ssp. setchellii
(Jeps.) Moran at 5 mi. SW Paicines, San Benito Co.; Alum Rock Park, Santa Clara
Co.; and Del Puerto Canyon, Stanislaus Co. (Langston records). D cymosa ssp.
minor (Rose) Moran is the host at Partington Creek, 200-300', ca. 9 road mi. S.
of Big Sur, Monterey Co.; 3rd instar on V-10-69 emerged II-7-70 (J. F. Emmel).
Also eggs collected on this Dudleya V-14-68 produced 9 adults that emerged III-
25 to IV-3-69 (C. Henne). This subspecies is also found in at least 4 southern Calif-
nornia sonorensis localities: Arroyo Seco Cyn., San Gabriel Mts.; San Gabriel
Cyn.; Eaton Cyn., San Gabriel Mts.; and Millards Canyon, L. A. Co. R. Wells and
I searched D. cymosa ssp. gigantea (Rose) Moran colonies in the vicinity of
Mokelumne River, Sutter Creek, and Plymouth on II-22-70 without success.
Dudleya abramsii Rose ssp. abramsii is not known to be a host but is found
in the Sierra Juarez, B. C., and on Monument Pk., Laguna Mts., where sonorensis
D. lanceolata (Nutt.) Britt. & Rose is reported to be the sonorensis foodplant
in San Gabriel Canyon (Martin, 1951; Hovanitz, 1967). Lanceolata occurs also at
the following sonorensis localities: Mint Canyon, Arroyo Seco, Fish Canyon,
Point Loma, and San Diego (ex herbaria records).
D. saxosa (Jones) Britt. & Rose ssp. aloides (Rose) Moran is found at Sentenac
Canyon and vicinity Scissors Crossing, San Diego Co. where sonorensis flies.
F. T. Thorne found 20 sonorensis ova on the flat-leaves near Scissors Crossing on
D. stolonifera Moran occurs only at Aliso and Laguna Canyons, Orange Co.;
sonorensis flies at the former, so this may be a possible foodplant.
D. edulis (Nutt.) Moran may be used by sonorensis where it flies at Torrey
Pines, La Jolla, and Sweetwater Dam in San Diego Co.
D. ingens Rose occurs at two known sonorensis localities in Baja Calif.:
San Telmo and Hamilton Ranch. However, D. attenuata is also found at San
Telmo (data from Moran, 1951a).
Langston (1963) found no adult sonorensis on Dudleya farinosa (Lindl.)
Britt. & Rose and D. caespitosa (Haw.) Britt. & Rose in central coastal California
based on several seasons of spring collecting. In San Diego Co. I have not seen
sonorensis at D. pulverulenta (Nutt.) Britt. & Rose locals at the proper time. Re-
ports in the literature of Sedum as a foodplant appear erroneous.
Following Moran's (1951a) classification, Dudleya (Crassulaceae;
Echeverioideae) has 38 species in western North America grouped as follows:
I. Subgenus Veradudleya
A. Section Stylophyllum--9 species
B. Section Eududleya-25 species
II. Subgenus Hasseanthus--4 species
This is a difficult genus with many unsolved problems, especially in section
Eududleya which is not easily separated into species and which has a high
degree of polyploidy (39% of the species). Dudleya reaches its greatest development
on the coasts and islands of California and Baja, and extends inland to southern
Nevada and central Arizona, with the San Diego coastal region the center of
distribution (above data from Moran, 1951a).
When the known and suspected sonorensis foodplants are superimposed
on Moran's classification, an interesting pattern emerges. Nearly all the species
used fall within the first 10 primitive members of Eududleya:
1. stolonifera--possible host
2. guadalupensis--Guadalupe Island, B. C. only
4. farinosa--Langston (1963) says it is not used, at least in part of its range
5. caespitosa-Langston (1963) says it is not used, at least in part of its range.
6. greenei--San Miguel, Santa Rosa, Santa Cruz, and Santa Catalina Islands
9. linearis--San Benito Islands, B. C. only
10. abramsii--possible host
The only other Eududleya used, ingens, ranked no. 18 of 25, may or may not be
a foodplant (D. attenuata of the Stylophyllum being the other candidate). In section
Stylophyllum, only edulis, ranked as the most advanced member, is perhaps
used, although attenuata, ranked close to edulis, may also be used. D. densiflora
(Rose) Moran, known only from Fish Canyon and San Gabriel Canyon (where
sonorensis flies) is apparently closely allied to edulis. No Hasseanthus are known
to be hosts.
Moran states that it is uncertain if the prototype was Stylophyllum or
Eududleya. The sonorensis foodplant relationships suggest that it was an ad-
vanced Stylophyllum which gave rise to Eududleya. A hybrid between edulis X
stolonifera is known; stolonifera "seems intermediate in some respects between
the subgenera Eududleya and Stylophyllum" (Moran, 1949).
The diploid cymosa appears to intergrade with the polyploid caespitosa
and lanceolata and perhaps with farinosa and abramsii. It is possible that
cymosa ssp. setchellii and abramsii ssp. murina are connected in the inner South
Coast Range, and it is also possible that cymosa ssp. minor intergrades with
abramsii ssp. abramsii in the San Bernardino and San Jacinto Mts. D. saxosa
ssp. aloides morphologically connects with lanceolata (above data from Moran,
Clausen, Moran, & Uhl (1945) note possible connecting link species be-
tween Hasseanthus and Stylophyllum (chromosomal resemblance among
Hasseanthus, D. pumila, D. viscida); H. elongatus is near Dudleya in appearance
and in the condition of the basal leaves. None of these are known sonorensis
foodplants. Wild hybrids are known between Dudleya edulis X Hasseanthus
blochmaniae (Moran, 1951b).
The rock substrate was sampled from two localities where sonorensis
flies abundantly on D. cymosa ssp. cymosa. At Murderer's Bar, 4 mi. NE Auburn,
Placer Co., the Dudleya grows on metavolcanic green schist; and at Skelton
Creek Canyon, E. of Jerseydale, Mariposa Co., it grows on slopes of slates
(=metatuff), greenstone (metavolcanics), and metaquartzite or siliceous metatuff
(samples det. by Dr. E. M. Moores, UCD Geology Dept.). In Del Puerto Canyon,
22 mi. W. of Patterson, Stanislaus Co., D. cymosa ssp. setchellii grows on rocky
serpentine slopes where sonorensis flies.
I have had no success with sonorensis in the foothills W. and NW of
Sacramento. D. cymosa ssp. is fairly common in Thompson Canyon, Vaca Mts.,
Yolo Co., where it grows on Upper Cretaceous sedimentary rocks of shales,
sandstones, siltstones, and conglomerate lenses. No sign of sonorensis there
during weekly Feb.-March collecting over a three-year period. In Rumsey Canyon,
3 mi. W. Rumsey, Yolo Co., on exposed S.-facing wall of canyon, on III-13-71,
much D. cymosa ssp. but no sonorensis (leg. R. Wells). At an ideal habitat on
serpentine outcrops W. of Leesville, Lake-Colusa Co. line, no sign of sonorensis
on III-15-73, D. cymosa ssp. common.
Fig. 1. SEM (Cambridge) micrograph of a P. sonorensis egg surface from
a top view, ca. 460 X; egg collected at Chili Bar, 4 mi. N. Placerville, El Dorado
Co., Calif., II-21-70.
Fig. 2. Same, close-up of micropyle region, ca. 1600 X.
LIFE HISTORY NOTES
A detailed life-history description of sonorensis is given by Comstock &
Coolidge (1930), and the egg is figured in Comstock (1927).
Confined females can lay at least 100 ova. A female from National City,
San Diego Co., captured III-3-52, produced 100 ova; one confined Y from Bennington
Monument, San Diego Co., taken II-19-33, laid 27 ova Feb. 20, 39 ova Feb. 21, 24
ova Feb. 22, hatched Feb. 25 & 26 (90 ova); same locality, III-4-33, 6 99 kept alive
laid over 200 ova March 6 and 50 ova March 7 (F. T. Thorne records).
Along an abandoned road between slope above Murderer's Bar and junction
of Middle & North Fk. American River, NE of Auburn, Placer Co., II-14-71, leg.
O. Shields & R. Wells. Counted sonorensis eggs on Dudleya cymosa plant clumps
(up to 4-7 plants per clump; those plants more than one foot away were counted
as another clump) at serpentine rock outcrops along steep 450 angle roadcut:
(1) Location #1, SW exposure, receives afternoon sun. On W. Side, 36 plant
clumps, 10 having eggs (28%); on E. side, 30 plant clumps, 2 with eggs (7%).
(2) Location #2, S. exposure, receives morning and afternoon sun. On W. side,
32 plant clumps, 13 with eggs (41%); on direct S. side, 6 plant clumps, 5 with eggs
(83%); on E. side, 19 plant clumps, 2 with eggs (11%).
(3) Location #3, direct S. exposure, 8 clumps, 6 with eggs (75%).
The range was 1-9 eggs per plant clump having eggs (average = 2.8). Eggs on leaf
upper surface = 66; eggs on leaf under surface = 41. II-17-73, Murderer's Bar,
only 2 eggs, no adults seen (one egg was collapsed, other held its shape but was
perhaps infertile?). The Dudleya were not yet large size, mostly medium sized.
On II-23-73, Murderer's Bar, leg. OS & S. Sims, found 8 unhatched eggs, played
on leaf uppersides toward upper part of leaf near the margin, and on center of
small leaves at center of rosette. II-21-70, Chili Bar, 4 mi. N. Placerville, El Dorado
Co., leg. OS & R. Wells, found ca. 25 eggs, mostly on 6-8 plants close together on
exposed rocky outcrop, mostly on basal half of leaves of young plants. Dudleya
plants further up the river where partial shade had no eggs or adults. On II-22-
70, 12 more eggs. On II-27-70, most of the eggs had emerged and larvae appeared
sizable if still 1st instar. They eat out the side of the egg and burrow into the
Dudleya leaf in close proximity to the egg site, generally. Larvae were feeding
head down and burrowed deep within the leaf; several found crawling over the
Observations of larvae in the wild:
(1) 3 mi. NE of Auburn, Placer Co., V-13-69, three 5th instars in D. cymosa
leaves along road; feeding evidence of earlier larvae in many plants.
(2) 4 mi. N. Placerville, So. Fk. American River, El Dorado Co., V-4-69, leg.
OS & R. F. Denno. Found about 3 last instar, 4-6 3rd-4th (?) instar larvae in
Dudleya leaves on a small slate knob by river. Larvae red, mine inside of
leaves, located by disturbed leaves and frass by entrance holes; all were inside
(3) Vicinity of Briceburg, Mariposa Co., V-6-69, leg. OS & N. La Due. Along old
RR track line on N. side of river. Numerous larvae (ca. 50) found mining leaves
of D. cymosa. Plants in exposed places had mined leaves but no larvae, other
plants in less exposed places had ca. 2nd-5th instar larvae. Larvae were
sometimes found on the outside of the leaves. Some larvae also found on
road cut along W. side of road ca. /2-/4 mi. S. of Briceburg.
Observations of adult behavior:
(1) Murderer's Bar, II-13-71, leg. OS & R. Wells. 58 5S 12 99 sonorensis taken,
8& flitting about tops of a weed (perhaps in search of freshly emerged 99)
and flying over areas where Dudleya cymosa grew, along roadcut banKs
at various rock outcroppings and S.-facing slope of monolithic limestone
rock above the Bar. 99 noted flying low, slowly over Dudleya and alighting
on Dudleya. Most adults taken between 11:00 am-1:00 pm PST, in wind pro-
tected areas behind rocks. Warm, dry day, clear in morning, 65-75, probably
up to 800 in protected areas near rocks, cooler and high overcast later in the
afternoon (2 pm on). A small swift lizard noted following a 9 sonorensis
that flew close to rocks over Dudleya's, almost got her. II-12-72, no sonorensis
on a warm day; Dudleya mostly small, few nearly full-grown. Nearly all Jan-
uary was cold. Warm weather the next week, and 65 adults (10 99) taken on
II-19 & 20-72, also a few eggs. By May 6th, the Dudleya's were fast drying
out and most in bloom; much evidence of feeding in dried-out leaves but no
(2) Chili Bar, 4 mi. N. Placerville, II-21-70, leg. OS & R. Wells. 16 6' 3 99
mostly freshly emerged, taken between 1:00-2:30 pm PST, all showed "flutter
flight" in a small outcrop of slate with Dudleya, exposed directly to sun.
Several 3( also perched on vegetation and rocks.
(3) 18 road mi. W. Patterson, Del Puerto Canyon, cal. 1700' Stanislaus Co.,
III-7-70, 1 9 worn sonorensis "sunning" near Dudleya plants on slope.
(4) Briceburg, III-11-72, 1 9 oviposited on leaf underside of D. cymosa at 12:15
pm PST, after alighting on plant, walking down leaf, flexing hind wings back
and forth, then walking underneath, in rather jerky motions.
(5) At Skelton Creek Canyon E. of Jerseydale, Mariposa Co., in 1970, J. F.
Emmel & I noted several 8C adults that rocked from side to side while alighted,
with wings partway open (4-5X), = "clicking" motion.
(6) Sentenac Canyon and Two Mile Hill, San Diego Co., II-14-65, R. Breedlove
took 16 36 2 99, temperature 550 F., the only butterflies on wing at this cool
temp. (Thorne records).
Fig. 3 P. sonorensis larva, ca. 4th instar, entering a new leaf of Dudleya
cymosa to feed; note also the sonorensis egg; Middle Fork American River, 4 mi.
NE Auburn, Placer Co., Calif., March 1971 (courtesy J. Mori).
(7) Scissors Crossing, San Diego Co., III-4-56, 6 &3 3 99, so windy they were
almost unable to fly (F. T. Thorne).
I have seen 6' in "patrol" flights up and down the canyons at side draw in
Sentenac Canyon and vicinity of Scissors Crossing, and at La Pressa Canyon,
SE of Dictionary Hill, in San Diego Co. Adults fly over open rocky slopes where
Dudleya is growing at Jerseydale and Del Puerto Canyon.
Adults feeding on flowers appears to be rarely observed. One fresh 5
on "sweet alyssum flowers", Lobularia maritima, a European escape, at La
Tuna Cyn., Los Angeles Co., III-5-48 (W. H. Evans, AM). At Chili Bar,
4 mi. N. Placerville, some adults were seen on Lomatium sp., II-18 and III-15-
71 (leg. R. Wells). At Briceburg, III-11-72, several fresh S5 were noted hovering
around a damp seep.
I would like to thank the following people for loaning material or providing
records for the sonorensis and speciosa studies (* = in litt.): (AM) Allyn Museum
of Entomology, Lee D. Miller; (DB) David L. Bauer; (JB) *James Brock; (CAS)
Calif. Acad. Sci., Paul Arnaud; (CIS) Calif. Insect Survey, Jerry Powell; (CNC)
*Canadian National Collection, R. Langston (in litt.); (SD) *Stanley K. Dvorak;
(JG) *John S. Garth; Carll Goodpasture; (GG) Glenn A. Gorelick; (CH) *Chris
Henne; (KH) Keith Hughes; (JJ) *J. W. Johnson; (NL) Noel La Due; (JL) John Lane;
(RL) *Robert L. Langston; (LACM) Los Angeles County Museum, Julian Donahue;
(JM) James Mori; (MCZ) Museum of Comparative Zoology, John Burns; (NSM)
Nevada State Museum, Peter Herlan; (PO) Paul A. Opler; (RP) *Richard Priestaf;
(HR) *Herman Real; (SB) Santa Barbara Mus. Nat. Hist., Nelson Baker; (CS)
*Charles A. Sekerman; Steven Sims; (WS) *William L. Swisher; (FT) *Fred T.
Thorne; (UCD) Univ. Calif. Davis Coll., R. O. Schuster; (UI) Univ. Idaho Coll.,
D. E. Foster & D. P. Levin; (UW) Univ. Wisconsin Coll, Lutz J. Bayer; (EW)
Fig. 4. P. sonorensis male sunning on a weed, Middle Fork American River,
4 mi. NE Auburn, March 1971 (courtesy J. Mori).
*Erich Walter; (RW) Ralph Wells; and (YU) Yale Univ. Coll., Charles L. Remington.
I am deeply indebted to John F. Emmel who first encouraged me to undertake the
Philotes project and who has assisted extensively in the field work. Paddy Mc
Henry kindly provided some original descriptions. To all I am grateful.
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