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Volume 32 1978 Number 1
JOURNAL
of the
LEPIDOPTERISTS’ SOCIETY
Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS
7 April 1978
THE LEPIDOPTERISTS’ SOCIETY
EXECUTIVE COUNCIL
J. W. Trpen, President KENELM W. Putuip, Vice President I. F. B. Common, Ist Vice President JuLian P. DoNAHuE, Secretary Lionet Hiccrns, Vice President RONALD LEUSCHNER, Treasurer
Members at large:
F. S. CHEW R. A. ARNOLD J. F. EMMEL D. F. Harpwick E. D. CAsHATT R. R. GATRELLE J. B. ZrEcLER R. E. STANFORD A, P.\ Prats
The object of the Lepidopterists’ Society, which was formed in May, 1947 and formally constituted in December, 1950, is “to promote the science of lepidopterology in all its branches, . . . . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards these aims.
Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journal and the News of the Lepidopterists’ Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. There are four numbers in each volume of the Journal, scheduled for February, May, August and November, and six numbers of the News each year.
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Send remittances, payable to The Lepidopterists’ Society, and address changes to: Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A.
Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA
by Cyrit F. pos Passos
Price: Society members, $5.00 U.S.; non-members, $7.50 U.S. Paper covers, revisions of the Melitaeinae and Lycaenidae supplied separately.
Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni- versity of Louisville, Louisville, KY 40208, U.S.A.
The Lepidopterists’ Society is a non-profit, scientific organization. The known office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid at Lawrence, Kansas, U.S.A. 66044.
Cover illustration: Dasychira dorsipennata larva, dorsal and lateral views. From Fascicle 22.2, “Lymantriidae,” by Douglas C. Ferguson, in Moths of America North of Mexico. The drawing was done by E. R. Hodges, Scientific Illustrator, Department of Entomology, Smithsonian Institution. (Reproduced by permission of the author.)
JOURNAL OF
Tue LEpPIDOPTERISTS’ SOCIETY
Volume 32 1978 Number 1
Journal of the Lepidopterists’ Society 32(1), 1978, 1-2
ieee OF PARNASSIUS CLODIUS GALLATINUS (PAPILIONIDAE )
STEVE KOHLER
Montana Department of Natural Resources and Conservation, Division of Forestry, 2705 Spurgin Road, Missoula, Montana 59801
ABSTRACT. Types of Parnassius clodius gallatinus Stichel, 1907, were discov- ered in the remnants of the Elrod collection, at the University of Montana, Missoula. Proper labels have been attached to the specimens, and they have been placed in the collection of the American Museum of Natural History.
The description of Parnassius clodius gallatinus Stichel was based on a pair illustrated by Elrod (1906). The actual specimens were never seen by Stichel (1907).
While preparing a paper clarifying the nomenclature of Parnassius clodius Ménétriés subspecies found in the Rocky Mountains (Ferris, 1976), Cliff Ferris contacted me in an attempt to locate the types of gallatinus. A search through the remnants of the Elrod Collection, housed at the University of Montana, Missoula, led to the discovery of the two specimens matching El!lrod’s 1906 illustrations. They were figured on page 16 of “The Butterflies of Montana.”
At the suggestion of Ferris, the two specimens have been placed in the collection of the American Museum of Natural History. The follow- ing labels are affixed to the specimen pins:
Holotype Male: A white label partially machine printed in black ink and partially hand lettered in red ink which reads: Gallatin Co. Mont./ Eley. 6800/ Col. E. Koch/ 6-27 1900, and a red label hand lettered in black ink which reads: Holotype ¢/ Parnassius clodius/ gallatinus Stichel/ ex. Elrod Coll. Univ./ Mont. S. Kohler 1976.
Allotype Female: A white label partially machine printed in black ink and partially hand lettered in red ink which reads: Gallatin Co. Mont./ Elev. 6800/ Col. Cooley/ 6-27 1900, and a red label hand lettered in
bo
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
te
Fig. 1. Parnassius clodius gallatinus Stichel: a) holotype male, dorsal; b) same,
ventral; c) allotype female, dorsal; d) same, ventral. Photos approximately 7 natural size.
black ink which reads: Allotype ?/ Parnassius clodius/ gallatinus Stichel/ ex. Elrod Coll. Univ./ Mont. S. Kohler 1976.
The specimens are in good condition except for some minor dermestid damage to the abdomen of the female. An additional pair collected by Cooley at the same locality and on the same date was located in the collection of Montana State University, Bozeman, for Ferris by Dr. Nor- man L. Anderson. These are not designated as paratypes.
Because of the scarcity of copies of Elrod’s “The Butterflies of Mon- tana, the type specimens have been illustrated in Figure 1.
ACKNOWLEDGMENTS
I would like to thank Dr. James H. Lowe and the University of Mon- tana for allowing the type specimens to be placed at the American Museum of Natural History, and also Cliff Ferris for comments and assistance,
LITERATURE CITED
IeLnop, M. J. 1906. The butterflies of Montana. Univ. Montana Bull. 10: 1-174.
Fennis, C. D. 1976. A note on the subspecies of Parnassius clodius Ménétriés found in the Rocky Mountains of the United States (Papilionidae). J. Res. Lep. 15: 65-74.
SticHEL, H. 1907. Lepidoptera Rhopalocera Fam. Papilionidae Subfam. Parnas- siinae. Wytsman Gen. Ins. Fasc. 58: 1-60.
Journal of the Lepidopterists’ Society 32(1), 1978, 3-19
SPECIFICITY, GEOGRAPHIC DISTRIBUTIONS, AND FOODPLANT DIVERSITY IN FOUR CALLOPHRYS (MITOURA) (LYCAENIDAE)
Kurt JOHNSON
Department of Biology, City University of New York, City College, Convent Avenue and 138th Street, New York, New York 10031
ABSTRACT. The species C. siva, gryneus, hesseli, and turkingtoni are examined. Genitalic evidence of their non-conspecificity is provided along with discussion of particular localities of sympatry. Detailed distributional data are illustrated and a documented table of foodplant diversity included. C. siva and gryneus are oligoph- agous On numerous species of Juniperus (Cupressaceae) which replace each other geographically across the United States. C. hesseli is monophagous on Chamaecyparis thyoides (Cupressaceae); the foodplant of turkingtoni is unknown. Evidence indicates that all local populations are specific to one foodplant species.
Callophrys ( Witoura) nelsoni ( Boisduval), C. siva (Edwards), C. loki (Skinner), C. gryneus (Hiibner), and C. hesseli (Rawson & Ziegler), aside from taxonomic descriptions, have been subject to several biological and regional studies, but published works (Anderson, 1974; Johnson, 1972; Pease, 1963; Rawson et al., 1951; Remington & Pease, 1955) are very heterogeneous in content and comprehensiveness.
During the last four years I have been compiling data on their dis- tributions and larval foodplants as a base for taxonomic studies of the group. I have also been studying the genitalia of all Nearctic and Neotropical Callophrys (Mitoura) in detail (Johnson, 1976a). The pur- pose of this paper is to present detailed distributional data for three of these species (C. siva, C. gryneus, and C. hesseli), demonstrate that C. siva and C. gryneus are not conspecific, and summarize data on larval foodplants, a number of which are new to the literature. What bio- geographical data are known on the newly named C. turkingtoni Johnson (Johnson, 1976b) will also be presented. The specificity of C. siva and C. nelsoni involves several complex problems in the northwestern United States and will be treated in a separate paper (Johnson, 1977).
METHODS AND MATERIALS
Using the collection of the American Museum of Natural History as a basis, additional information on localities and possible local foodplants was gathered by correspondence and recorded county by county. Speci- mens or photographs were solicited in cases of peripheral or isolated populations, and available published records were included. The re- search aimed at definitive treatment on the species level only. Genitalic
+ JOURNAL OF THE LEPIDOPTERISTS SOCIETY
studies of males and females were performed in areas where C. siva and C. gryneus were reportedly sympatric. These genitalia were compared with those from many parts of the ranges of C. siva and C. gryneus, as well as with dissections of other congeners. The number of these speci- men dissections included: C. siva, 78; C. gryneus, 46; C. hesseli, 14; C. turkingtoni, 1; C. nelsoni, 83; C. rosneri, 46 (Johnson, 1976a); C. barryji, 19 (Johnson, 1976a); C. byrnei, 9 (Johnson, 1976a), and C. loki, 15. Geographic ranges were studied to discover areas of insect distribution not coinciding with present published foodplant knowledge, and efforts were then made to make the list for each species complete by identifica- tion of exact plants with which the adults were associated by perching behavior (Johnson & Borgo, 1976) or on which oviposition or larvae were observed. Full documentation of each of these methods is given in the foodplant table (Table 1) since a degree of fallibility has been demon- strated in each (Brower, 1958; Downey & Dunn, 1965). An ongoing effort to compile foodplant specimens at one institution was initiated, and plants collected thus far are cited in the table. Since the perching behavior of these insects limits general flight patterns to the vicinity of the foodplant, and since data not only in this study but another (John- sonn, in prep. ) indicate that C. siva and C. gryneus are exclusive Juniperus- feeders, some useful evidence on larval foodplants in areas where only one juniper species was regionally present could be culled from identifi- cation of the plants at the locality indicated on the specimen labels. The list of plants established as the only Juniperus species present in a
region (R) or at a locality (L), source of butterfly data (B), source of plant data (P) is:
C. siva: Juniperus deppeana Steud., (LL) 10 mi. NW Pine Springs, Culberson Co., Texas, (B) R. O. Kendall, (P) Herbarium, University of Texas, Austin; J. deppeana, (1.) 5 mi. W of McDonald Observatory, Jeff Davis Co., Texas (Bk. 'O: Kendall, (P) Herbarium, University of Texas, Austin. Juniperus occidentalis oc- cidentalis Hook. x J. osteosperma Torr. (Little), (R) Washoe Co., Nevada (Reno and vicinity westward), Ormsby and Douglas cos., (B) P. Herlan, (P) Vasek, 1966. Juniperus monosperma (Engelm.) Sarg., (LL) Sycamore Canyon, NW of Nogales, Santa Cruz Co., Arizona, (B) Share and Clark (American Museum of Natural History (AMNH)), (P) Herbarium, Arizona State University, Tempe. Juniperus pinchotii Sudw., (R) Reeves Co., Texas, (B) D. Stallings and M. R. Turner, (P) Adams, 1972, R. P. Adams, pers. comm. C. gryneus: Juniperus virginiana L., (R) Cass Co., Texas, (B) R. O. Kendall, (P) Adams & Turner, 1970, R. P. Adams, pers. comm. Juniperus ashei Buchholz, (R) McLennan Co., Texas, (B) R. O. Kendall, (P) Adams, 1972; Adams & Turner, 1970; R. P. Adams, pers. comm. Juniperus pinchotii Sudw., (R) Pecos Co., Texas, (B) R. O. Kendall, (P) Adams, 1972, R. P. Adams, pers. comm. Juniperus deppeana Steud., (L) Huejotitlan, Chihuahua, Mexico, (B) AMNH, (P) Little, 1971. J. deppeana, (i) Baboquivari Mountains, S of Baboquivari Peak, Pima iGo Arizona,” (ByenieaD: Gunder (AMNH), (P) Herbarium, Arizona State University, Tempe. Juniperus virginiana L. * J. horizontalis Moe nch., (1) Lynxville, along Mississippi River,
VoLUME 32, NUMBER 1
Aste I.
Larval foodplants established by the identification of exact plants.
Foodplant taxa and specimens
Butterfly taxa and specimens
Callophrys (Mitoura) siva
Juniperus scopulorum Sarg."
Plants, PL: Van Haverbeke, N-2; hybrid index at site—63 + 8% J. scopulorum; SL: Johnson, 1972.”
Plant specimens, K. Johnson #2 (Smiley Canyon = VanH. N-2; #1 (Chadron), Royal Ontario Museum, Toronto
(ROM ).°
Juniperus scopulorum var. columnaris Fasset
Plants, PL: Van Haverbeke ND-S, I: T. McCabe, (H) North Dakota State Univ., Fargo; SL: Van Haverbeke, 1968. Plant specimens, Van Haverbeke ND-8, Univ. of Nebraska (UN).
Juniperus scopulorum Sarg. X J. virgini- ana L.
Plants, PL: Van Haverbeke N-7;
hybrid index at site—48 + 6% J. scopulorum; Range of use of hybrids (Johnson, 1972 )—70 + 4% to 36 + 4% J. scopulorum.
Plant specimens, Van Haverbeke N-7, UN; K. Johnson #3 (Sizer, Keith Co. ) ROM.
Juniperus virginiana L.
Plants, 17 mi S of PL: Van Haverbeke N-4; hybrid index at site—36 + 4%
J. scopulorum; SL: Johnson, 1972. Plant specimens, Van Haverbeke N-4, UN; K. Johnson #4 (locality as above ) ROM.
Prostrate morph of J. scopulorum Sarg. x J. virginiana L. x J. horizontalis Moench.
s. siva; Dawes Co., Nebraska (Smiley Canyon ), W of Fort Robinson; Catholic Cemetery, Chadron.*
Butterthes, PE (ACS@: B)*: K. Johnson (AMNH).°
s. siva; Slope Co., North Dakota (Amidon, along burning coal vein). Butterflies, PC: T. McCabe.
s. siva; Garden Co., Nebraska (bluffs above N. Platte River, nr. Lewellen). Butterflies, PC (AC): L. Running, AMNH.
s. siva; Rock Co., Nebraska (Long Pine Rec. Area).
Butterflies, PC (AC, LC, B): K. Johnson, L. Running, AMNH.
s. siva; Saskatchewan, Canada (Val Marie, near Rosefield along Frenchman River ).
pm 1 Taxon of foodplant used (according to nomenclature of USDA (1953) and Little (1971)).
2Source of plant data:
PL — butterflies were specifically collected at a_ particular locality
studied by Cupressaceae taxonomists; their designation of the site is noted along with the date of their study. “Hybrid index” refers to these studies’ calculation of the degree of hybridity in plants at these areas. Plant identifications are noted as “7”: I, exact substrate plant identified pen Ee —Rerbarum at ==»), and! location of voucher specimens; I,, plant identified
by data sent to ______ by
; I,, foodplant established in original description of butterfly,
citation given; I,, plant identified from specimens sent to ——_——_; I,, substrate plant established
by matching herbarium specimens with butterfly data and establishing that no other species co- occurs, herbarium cited. An “*” following this category (‘‘Plants’”’) means this foodplant usage
is well
3 The label number and place of deposition of plant specimens collected in this study An “* means collection in progress at time of this writing.
known; SL = other literature which supports this identification.
>?
4Taxon of butterfly concerned (as designated in annotated list), with state and exact locality.
5 Source of butterfly data: PC = “personally collected by
Letters in parentheses
following mean: AC, adults commonly observed perching; AI, adults perching but not commonly observed; LC, larvae collected; LO, larvae observed; B, behavior studied in detail; O, oviposition
observed.
6 Location of specimens if not aforementioned person (AMNH =the American Museum of Natural History, New York). M =museum specimens were used as the source of data; ver.
means verified by
, and method. TL = type locality of the insect.
6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TAsrE
Continued
Foodplant taxa and specimens
Plants, Schurtz (1971) indicates this area would be included in his tri- parental swarm. Van Haverbeke (pers. comm.) supports this evaluation; I: (H)
Univ. Saskatchewan, Regina; I: D. F. Van
Haverbeke (data from R. Hooper, K. Johnson); SL: Little, 1971; Van Haverbeke, 1968; Fassett, 1945.
Plant specimens, (H) Univ. Sask. (Sask. Prairie Park); K. Johnson #5* ( Hooper ) ROM.
Juniperus osteosperma ( Torr.) Little Plants, I;: D. F. Van Haverbeke; SL: Emmel & Emmel, 1973; Johnson, 1977; Little, 1971.
Plant specimens, K. Johnson #15 (Running, locality as above) AMNH.
Juniperus californica Carr. Plants +, I: D. F. Van Haverbeke; SL:
Comstock, 1927; Emmel & Emmel, 1973.
Plant specimens, K. Johnson #16 (Leone, locality as above) AMNH.
Juniperus occidentalis occidentalis Hook. Plants, I: D. F. Van Haverbeke; SL: Johnson, 1977, Little, 1971.
Plant specimens, K. Johnson #17 (Buckingham, locality as above).
Juniperus occidentalis australis Vasek Plants, I: John H. Lane; SL: Vasek, 1966.
Butterfly taxa and specimens
Butterflies, PC (AC): R. Hooper (ver. K. Johnson, photo ).
s. ssp.; White Pine Co., Nevada (nr. McGill Junction).
Butterflies, PC (AC): L. Running, AMNH.
s. juniperaria; Los Angeles Co., Cali- fornia (Mint Canyon).
Butterflies, PC (AC): M. Leone, AMNH.
s. ssp.; Jefferson Co., Oregon (nr. Warm Springs, W on road to Twin Buttes ). Butterflies, PC (AC): F. Buckingham, AMNH.
s. ssp.; Tulare Co., California (vic. Kennedy Meadows ), San Bernardino Co., California (Big Bear Lake). Butterflies, PC (AC): John H. Lane.
Callophrys (Mitoura) gryneus
Juniperus virginiana L.
Plants, I: K. Johnson; SL: Klots, 1951, Little, 1971.
Plant specimens, K. Johnson #21 (locality as above).
Juniperus silicicola (Small) Bailey Plants, I: F. D. Fee, (H) Univ. Florida, Gainesville; L: (H) Univ. Gainesville; SL: Klots, 1951; Little, 1971.
Plant specimens, K. Johnson #10 (St. Augustine locality, Univ. Florida) ROM.
Juniperus scopulorum Sarg. * J. virgini- ana LL.
Plants, PL: Van Haverbeke M-1; hybrid index near site—27 + 4% J. scopulorum; Range of use of hybrids (Johnson,
g. gryneus; Ulster Co., New York ( West Park, Holy Cross Publications ). Butterflies, PC (AI): K. Johnson, AMNH.
g. sweadneri; St. Johns Co., Florida (along Ocean Rt. A1A, St. Augustine ). Butterflies, PC (AC): F. D. Fee.
g. gryneus; Jackson Co., Missouri (general ). Butterflies, PC (AC): J. R. Heitzman.
VoLUME 32, NuMBER 1
TABLE l.
Continued
Foodplant taxa and specimens
Butterfly taxa and specimens
1972 )—38 + 4% J. scopulorum to 27 + 4% J. scopulorum.
Plant specimens, K. Johnson #11 (Heitzman, Independence ) ROM.
Juniperus ashei Buchholz
Plants, I: J. R. Heitzman; SL: Little, 1971.
Plant specimens, K. Johnson #12* (Heitzman, ?) ROM.
Juniperus pinchotii Sudw. Plants [two examples], L: R. P. Adams (Scott, Roever); SL: Adams, 1972.
I: R. O. Kendall; SL: Little, 1971; Adams & Turner, 1970.
Juniperus virginiana L. x J. horizontalis Moench.
Plants, I: (by reason of Schurtz, 1971) D. F. Van Haverbeke; SL: Schurtz, 1971; Little, 1971.
Plant specimens, K. Johnson #13 (locality as above).
g. ssp.; Barry Co., Missouri (Eagle Rock ), also McDonald Co.; Washington and Carroll cos., Arkansas.
Butterflies, PC (AC): J. R. Heitzman.
g. castalis [two examples]; Armstrong Co., Texas (just below N rim of Palo Duro Canyon, 15-16 mi. S Claude).
Butterflies, PC (AC): M. Toliver,
H. A. Freeman, J. M. Burns, K. Roever (R. O. Kendall); J. Scott.
Bexar Co., Texas (Reo Seco Road, off U.S. Hwy. 281 N of San Antonio). Butterflies, PC (AC, O): R. O. Kendall.
g. gryneus; Dane Co., Wisconsin (10 mi. W of Madison). Butterflies, PC (AC): W. Sieker.
Callophrys ( Mitoura) hesseli
Chaemaecyparis thyoides (L.) B.S.P. Plants*, I: S. Hessel, G. W. Rawson,
J. B. Ziegler; I,: Rawson et al., 1952; Rawson & Ziegler, 1950 (therein det. by I. M. Johnston, Harvard Univ. ).
hesseli; Ocean Co., New Jersey (Lakehurst, TL).
Butteniess 2G. ( ACO. LEG) S. Hessel, G. W. Rawson, J. B. Ziegler.
Lacrosse Co., Wisconsin and 5 mi. W of Sauk City, Sauk Co., Wisconsin, (B) F. Amold and W. E. Sieker, (P) Ross & Duncan, 1949; Schurtz, 1971; D. F. Van Haverbeke, pers. comm. C. turkingtoni: Juniperus flaccida Schlecht., (R) 10 mi. E of Namiquipa, Chihauhua, Mexico, (B) W. Gertsch and M. Cazier (AMNH), (P) Little, 1971; Herbarium, University of Mexico, Mexico City.
RESULTS Genitalia of C. gryneus and C. siva Genitalia of males and females were studied in three regions where these species were reportedly sympatric (Davis Mountain, Texas; Guada-
lupe Mountains, New Mexico and Texas; and Baboquivari Mountains, Arizona) and found to be easily separable. However, some traditionally
8 JoURNAL OF THE LEPIDOPTERISTS SOCIETY
Figs. 1-9. Female genitalia of selected Nearctic Callophrys (Mitoura) spp. C. siva siva: 1, topotypical; 2, heavily sclerotized areas of lamellae and eighth sternite; 3, showing tufts of “hair” ailowing diagnosis by naked eye. C. gryneus: 4, topotypical. C. hesseli: 5, topotypical. Genital plates of sympatric species near Alpine, Texas: 6, C. gryneus and 7-9, C. siva.
used wing-pattern characters for distinguishing these species (ventral secondaries: post basal spots or pattern of mesial band) were shown to be less reliable (also noted in Johnson, 1976a, 1977). The diagnostic
genitalic characters are as follows:
Females (Figs. 1-9). C. gryneus (Figs. 4, 6): ductus bursa longer and not “club-ended” as on siva; lamellae tapering caudad from antrum, not shouldered as on siva, lamellae postvaginalis nearly as long as broad; juncture of lamellae and eighth abdominal sternite not heavily sclerotized or connected,
C. siva (Figs. 1-3, 7-9). Ductus bursa shorter than gryneus and “club-ended”; lamellae distinctly shouldered, lamella postvaginalis much broader than long. Junc- ture of lamellae and eighth abdominal sternite heavily sclerotized, in area between
VOLUME 32, NUMBER 1 9
Figs. 10-12. Male genitalia of selected Nearctic Callophrys (Mitoura) spp., lateral and posterior views with tip of aedeagus (right) and falces (left): 10, C. gryneus castalis, topotypical; 11, C. siva siva, topotypical; and 12, C. hesseli, topo- typical.
1. postvaginalis and 1. antevaginalis forming bulkly ridges and convolutions at their juncture, these binding lamellae tightly with eighth abdominal sternite and con- taining many spines.
[C. hesseli (Fig. 5). Easily recognized by unique shape of the lamellae and broad cephalad tapering from the antrum (figured for reference). C. turkingtoni: female unknown. ]
Males (Figs. 10-12). C. gryneus (Fig. 10). Valvae, lateral shape: only barely concave between dorsal and ventral articulation with vinculum; valvae, caudad saccus (dorsal or ventral view): rounded and indented, vaguely shouldered caudad. Saccus: long and broad.
C. siva (Fig. 11). Valvae, lateral shape: deeply concave and rounded between dorsal and ventral articulation with vinculum; valvae, caudad saccus (dorsal and ventral view): parabolic and unindented, no shouldering caudad. Saccus: short and much less broad than gryneus.
[C. hesseli (Fig. 12). Lateral shape of valvae less broad, quite concave between articulations with vinculum, and much longer caudad; valvae caudad saccus broadly round, indented, and extremely shouldered caudad (figured for reference). C. turkingtoni (Johnson, 1976a), easily recognized by extremely long caudad extension of valvae and by heavily sclerotized and spiny area of valvae, caudad saccus.]
JouRNAL OF THE LEPIDOPTERISTS SOCIETY
10
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VoLUME 32, NuMBER 1 ai
& — J. virginiana =
v 6) gs J. flaccida ZN J. californica
= J. pinchotii
poles occidentalis ee communis
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Sympatric C. (M.) siva and alc. (M.) gryneus, diagnosed by genitalia. @c.
Mexican populations assigned to species as indicated.
Fig. 14. Nearctic distributions of Callophrys (Mitoura) siva and its larval food- plant Juniperus spp., and known range of C. (M.) turkingtoni. Plant distributions adapted from Little, 1971. Distribution of Juniperus horizontalis shown only as it exceeds J. scopulorum northward; hybrid swarms of Juniperus spp. illustrated in
Pig, 1d.
12 JouRNAL OF THE LEPIDOPTERISTS SOCIETY
ae re ew were ee ene pent es tors an
North Dakota
r South Dakota
Wyoming Nebraska
Mig. 15. Bi-parental and tri-parental swarms of divergence in Nearctic Juniperus: top, localities indicated by Van Haverbeke (1968) as “hybrid” J. virginiana x J. scopulorum; center, parental areas of J. horizontalis indicating possible centers for
VoLUME 32, NuMBER 1 13
Geographic Distributions and Available Larval Foodplants
Figs. 13 and 14 show the Nearctic distributions of species of Callophrys (Mitoura) in relation to the ranges of available or established larval foodplants. Fig. 15 shows areas of Juniperus ranges that have been botanically demonstrated as “hybrid swarms” (Van Haverbeke, 1968; Schurtz, 1971).
Summary of Data and Current Taxonomic Usages
The following is a review of the current common usage of trinomens in each group with a summary that includes distribution, foodplant(s) as established in this paper, general comments on the phenotype, and notes on the particular significance of each population. Where populations are under study by other lepidopterists, and especially where they are plan- ning to assign new names, | have called these subspecies “ssp.” and in- cluded the appropriate investigator's name in brackets.
Annotated List Callophrys (Mitoura) siva
C. (M.) siva siva. Type locality: Fort Wingate, McKinley Co., New Mexico. Distribution: Workers have named populations distinct from this taxon only on the West Coast, although others are undoubtedly present. Phenotype: There are two general morphs, based on ground color of the ventral secondaries. Great Basin populations (Fig. 14, squares) are brown beneath, whereas others (Fig. 14, plain black circles) are green. Populations of green-browns and mixed greens and browns occur in western Utah (Fig. 14, overlapping square and circles). The brown morph, which Peter Herlan, H. K. Clench, and I have investigated (Johnson, in prep.), is separately treated below. Foodplants: Many western Juniperus species (see Table 1) replace each other geographically. “Hybrids” (see Summary and Con- clusions) of J. virginiana x J. scopulorum, J. virginiana x J. horizontalis, and J. virginiana X J. scopulorum x J. horizontalis (possibly also J. occidentalis x J. osteo- sperma) occur in western Nevada, adjacent California, and eastern Oregon. All populations of Callophrys siva are on erect trees except for one local population (Val Marie, near Rosefield, Saskatchewan, along Frenchman River) on prostrate plants.
C. (M.) siva ssp. [Johnson, in prep.]. The research of Herlan, Clench, and Johnson involves naming this Great Basin population. Distribution: southern Nevada northward to Idaho; brown eastward to Salt Lake City; brown westward to southeast Oregon; broad interface with green morph C. (M.) siva siva in western Utah (e.g., Eureka, Dividend, Provo, western Millard Co.). Foodplants: Herlan reported J. osteosperma, but probably J. osteosperma x J. occidentalis near Reno, Nevada (Vasek, 1966). Apparently not J. scopulorum where it is available, al-
<<
“hybrid” status with J. virginiana; bottom, localities indicated by Schurtz (1970) as “hybrid” J. virginiana <x J. scopulorum xX _ J. horizontalis.
14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
though Idaho occurrences of C. (M.) siva may be lacking due to inadequate sampling.
C. (M.) siva juniperaria (Comstock). Type locality: Los Angeles Co., California. Distribution: western San Bernardino, northern Los Angeles, eastern Kern, and southern and northern Inyo cos.; transition with next subspecies in northern Ventura and northeastem Santa Barbara cos. Foodplant: Juniperus californica throughout its range, but in one locality (areas between Phelan and the San Gabriel Mountains, close to the Los Angeles, San Bernardino county lines) it is known to perch also on J. osteosperma (J. Lane, pers. comm. ).
C. (M.) siva mansfieldi (Tilden). Type locality: Simmler, San Luis Obispo Co., California. Distribution: southern California—western Kern and _ eastern San Luis Obispo cos. This is somewhat northward of the preceding subspecies. This taxon is ill-defined from that immediately below. Phenotype: deep green morph. Foodplant: J. californica.
C. (M.) siva ssp. [Lane, A]. Distribution: north of the preceding entity in the south inner coastal range of California (northern San Luis Obispo, southwestern Fresno, San Benito, eastern Santa Clara, and western Contra Costa cos.). Pheno- type: brown morph. Foodplant: J. californica.
C. (M.) siva ssp. [Johnson, in prep.]. The identity cf reputed C. (M.) siva specimens from eastern Oregon and Washington to western Idaho, and their rela- tion to the name C. (M.) nelsoni, has been resolved by detailed genitalic studies (Johnson, in prep.). Populations in nearly all of Oregon east of the Cascades to extreme southwest Idaho and extreme southeastern Washington are C. siva. Pheno- type: burgundy-brown morph. Foodplants: Oregon, Washington—J. occidentalis; Idaho—J. osteosperma. The taxon in press by Johnson includes only the J. occi- dentalis utilizers; those on J. osteosperma represent northward range of another subspecies distributed throughout Nevada and reviewed above.
C. (M.) siva ssp. [Lane, B]. Distribution: High altitudes in the southern Sierra Mountains (e.g., Kennedy Meadows, Tulare Co., California) and San Bernardino Mountains (e.g., Big Bear Lake, San Bernardino Co., California). Phenotype: green morph. Foodplant: J. occidentalis australis.
Callophrys (Mitoura) gryneus
C. (M.) gryneus gryneus. Type locality: Rappahanock Co., Virginia. Distribution: eastern and central North America in scattered populations wherever J. virginiana occurs. Phenotype: green morph. Foodplants: J. virginiana, but not observed feeding on sympatric J. communis, prostrate morphs of J. horizontalis x J. virginiana, or prostrate morphs of J. horizontalis northward; however, apparently utilizes erect morphs of J. horizontalis > J. virginiana (see Summary and Conclusions). Particular note: sometimes collected on nectar sources with C. (M.) hesseli, but foodplants are segregated by habitat in nature and not interchangeable.
C. (M.) gryneus sweadneri (Chermock). Type locality: St. Augustine, St. John’s Co., Florida. Distribution: Florida, perhaps southern Georgia, and north along the Atlantic Coast where J. silicicola occurs. Phenotype: green morph. Foodplant: J. silicicola.
C. (M.) gryneus castalis (Edwards). Type locality: McLennan Co., Texas. Distribution: mainly Texas, but also Chihuahua, Mexico, and areas west of the Mississippi River “gap” in juniper ranges; in addition, used by some workers as a form name within eastern United States populations. Phenotype: green morph. Foodplants: J. virginiana, J. ashei, and J. pinchotii, replacing each other westward. J. deppeana and possibly J. flaccida in Mexico.
C. (M.) gryneus ssp. [Johnson, in prep.]. Distribution: the Baboquivari Moun- tains eastward into Cochise Co., Arizona, and possibly southward in disjunct ranges of J. deppeana. Phenotype: green morph. Foodplant: J. deppeana suspected.
VoLUME 32, NuMBER 1 15
Callophrys (Mitoura) hesseli
C. (M.) hesseli. Distribution: see Fig. 13. Phenotype: green morph. Food- plant: Chamaecyparis thyoides.
Callophrys (Mitoura) turkingtoni
C. (M.) turkingtoni, a single specimen known from Namiquipa, Chihuahua, Mexico, in habitat of J. flaccida. Phenotype: brown morph.
SUMMARY AND CONCLUSIONS
Interspecific relations. Studies of C. siva and C. gryneus at several sympatric localities (21 specimens from the Baboquivari Mountains, Pima Co., Arizona; Cochise County (general), Arizona; Guadalupe Mountains, Eddy and Otero cos., New Mexico, Culberson Co., Texas; and Alpine, Brewster Co., Texas) confirmed that they are separable by genitalia of the males and especially the females (Johnson, in prep.). Since town, county, or mountain range is the only data available on some of these specimens, the extent of their microallopatry or microsympatry remains unknown. Biogeographic data suggest that the species may be altitudinally separated at some localities in Texas (C. siva on higher altitude J. deppeana, C. gryneus on lower altitude J. pinchotii), but it is likely that interspecific competition occurs at some locations. Sharing of nectar sources may occur, as reported in C. gryneus and C. hesseli (J. B. Ziegler, pers. comm.). These two species are generally segregated by the habitats of their foodplants. The female genitalia of C. hesseli have not been previously figured in the literature and are included in Fig. 5.
Foodplant relations. C. siva and C. gryneus utilize a broad spectrum of related and equally acceptable Juniperus species, which replace or exceed each other in geographic distribution over the Nearctic Realm. There is evidence that every species of Juniperus in the Nearctic is utilized, with two exceptions: J. communis L. and J. horizontalis Moench. Van Haverbeke (1968) and especially Schurtz (1971) have shown that J. horizontalis is actually part of a broadly distributed “swarm of diver- gence’ which involves the parental stock to which the names J. virginiana, J. scopulorum, and J. horizontalis have been applied. Van Haverbeke (pers. comm.) prefers Schurtz’s interpretation that each of these merits species status but that they are tied by their evolutionary histories, J. virginiana being an eastward evolutionary manifestation of J. scopulorum and J. horizontalis being a northward evolutionary manifestation of this biparental parent stock. Thus, there is little chemical or morphological reason (unless it is the number of needles versus fleshy leaves) that would prevent use of J. horizontalis by these Callophrys (Mitoura) especially where it is sympatric with utilized J. virginiana or J. scopu-
16 JouRNAL OF THE LEPIDOPTERISTS SOCIETY
lorum. Johnson & Borgo (1976) have shown that the perching behavior of C. siva and C. gryneus is distinctly patterned and preferenced for heights. They postulate that the nature of this patterned perching be- havior selects against prostrate morphs and is at least a partial boundary on their usage as a larval foodplant (Johnson & Borgo, 1977). The im- portance of the number of needled leaves on both J. horizontalis and J. communis needs investigation since first instar larvae burrow into these to feed.
Knowledge of the local specificities of the two oligophagous species is quite incomplete, although preliminary evidence from several localities indicates that populations are specific to particular plant species. In Palo Duro Canyon (Randall and Armstrong cos., Texas) J. scopulorum, J. pinchotii, J. monosperma, and hybrids of the latter two occur (Adams, 1972, and pers. comm.). Field data from collectors of C. gryneus indicate that J. pinchotii is the only foodplant. However, verification is needed by someone who can test this hypothesis directly. Peter Herlan (pers. comm.) reports that the Great Basin brown morph of C. siva feeds ex- clusively on J. osteosperma. Perhaps this is true, but Vasek (1966) has suggested that this species introgresses with J. occidentalis westward, and the taxonomic relationships of C. siva in the northwest basin are now indicated as including two, largely disjunct subspecies, one feeding on J. occidentalis in central and eastern Oregon and the other on J. osteosperma in Nevada eastward to Utah. In Missouri and Arkansas, C. gryneus populations are located on J. ashei where it occurs as “islands” within the range of J. virginiana. Other C. gryneus populations are on J. virginiana. This is another location ideal for specificity studies, as are the areas of diversity of juniper species in Arizona and New Mexico. In California, John Lane reports (pers. comm.) C. siva juniperaria perching on both J. osteosperma and J. occidentalis in an area where J. occidentalis has been reported as the foodplant. Thus, foodplant relations in C. siva and C. gryneus mirror situations reported in Burns (1964), Downey (1966), and Downey & Dunn (1965). Local specificities are due to oviposition by the female on the plant species it fed on as a larva. Thus, according to the familiar “Hopkins’ Host Principle,” specificity is main- tained. However, it is obvious that alterations do occur through time and space (as the above authors also indicate), and this is why such species show catholicity when their foodplant usage is viewed as a whole. The mechanism of ovipositional specificity and the nature of chance alterations need further elucidation. Downey & Dunn (1965) suggest that the patterning of Hopkins’ Host Principle is not genetic but physio- logical and undergoes divergence, convergence, and parallelism through
VoLUME 32, NUMBER 1 17
time and space. The present study indicates that similar foodplants available as replacers offer opportunity for divergence, since nearly all barriers posed to these insects by replacer plants have been crossed. Similarly, the remarkable coincidence of distinguishable morphs or sub- species generally within the distribution of one or another foodplant or foodplant relative suggests that foodplant adaptations play an important role in subspeciation. Callophrys gryneus sweadneri inhabits the areas of J. silicicola, C. gryneus gryneus those of J. virginiana, and C. gryneus castalis those of the transition of the latter plant to the ranges of J. ashei and J. pinchotii. Relations in the C. siva complex, although trinomial knowledge is less complete, are equally distinctive. If one assumes monophagous C. hesseli evolved through adaptations of some populations of early C. gryneus stock to Chamaecyparis thyoides, a similar mechanism is imaginable, especially since C. thyoides and J. virginiana have under- gone a change in their degree of sympatry through time (M. Rosenzweig, pers. comm.) in which populations of C. thyoides are now somewhat disjunct, and those of C. hesseli apparently extremely so.
Laboratory foodplant experiments with these species have not been extensive, and such data is of limited use in drawing inferences about foodplant utilization or preference in nature (Downey & Dunn, 1965; Downey & Fuller, 1962). However, studies to date indicate that quite divergent Cupressaceae species are at least nutritionally adequate and otherwise edible by some of the Callophrys (Mitoura) species. There is a need to further clarify the reported acceptance of J. virginiana by larvae of C. hesseli.
Distributional relations. One comment on the distribution of these insects, with regard to the frequency of transplanted populations is ap- propriate at this time. Cupressaceae species are widely used both in agricultural and landscape planting, and a number of transplanted Callophrys (Mitoura) populations have been noted (Figs. 13 & 14). Taxonomists should be especially aware of this when studying the com- parative morphology of these butterflies. The occurrence of C. siva in planted forest well isolated in central Nebraska, where juniper is raised from Rocky Mountain stock, is an extreme example, as is the occurrence of this insect in a shelter belt along the Missouri River.
ACKNOWLEDGMENTS
Many lepidopterists and botanists contributed data for this study. I owe a special debt to John Lane, Oakley Shields, and Arthur M. Shapiro (Univ. of California, Davis) for discussing Callophrys (Mitoura) species with me, and to Frederick H. Rindge (American Museum of Natural
18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
History, New York) for providing the facilities of the museum and editing this paper. For general support I am grateful to the staff of the Museum of Natural History, University of Wisconsin, Stevens Point, especially Charles A. Long, Director, and Robert Freckmann, Curator of the Herbarium. Debt is owed John C. Downey (University of Northern Iowa, Cedar Falls ) who introduced me to this group and suggested topics for research. The following provided data for which I am very grateful: R. A. Anderson, J. F. Gates Clarke, Ernst J. Dornfield, J. Donald Eff, George Ehle, Scott Ellis, Frank D. Fee, Clifford D. Ferris, Mike Fisher, Richard Funk, Richard Guppy, Lucien Harris, J. Richard Heitzman, Peter Herlan, Sidney Hessel, Ronald Hooper, Roderick R. Irwin, Roy O. Kendall, H. L. King, Alexander B. Klots, Steve Kohler, Henry A. LeBeau, Bryant Mather, James R. Maudsley, Tim McCabe, Lee D. Miller, James Mori, John S. Nordin, Roger Pease, Richard Priestaf, Kilian Roever, Michael Rosenzweig, R. E. Sanford, James Scott, Ernest M. Shull, William Sieker, Michael J. Smith, J. Bolling Sullivan III, Fred Thorne, and J. W. Tilden.
I am indebted to the herbaria at Arizona State University (Tempe); Colorado University (Boulder); University of Florida (Gainesville); Uni- versity of Mexico (Mexico City); University of Saskatchewan (Regina); and University of Wisconsin (Stevens Point) and especially to Cupres- saceae specialists R. P. Adams (Colorado State Univ., Fort Collins) and David F. Van Haverbeke (University of Nebraska, Lincoln) for their
comments and aid.
LITERATURE CITED
ApaMs, R. P. 1972. Chaemosystematic and numerical studies of natural popula- tions of Juniperus pinchotii Sudw. Taxon 21: 407-427.
Apams, R. P. & B. L. Turner. 1970. Chaemosystematic and numerical studies of natural populations of Juniperus ashei Buch. Taxon 19: 728-751.
ANDERSON, R. A. 1974. Southern records of Mitoura hesseli (Lycaenidae). J. Lepid. Soc. 28: 161.
Brower, L. P. 1958. Larval foodplant specificity in butterflies of the Papilio glaucus group. Lepid. News 12: 103-114.
Brown, F. M., D. Err, & B. Rorcer. 1957. Colorado butterflies. Denver Mus. Natur. Hist., Denver, 368 p.
Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif. Publ. Ent. 27: 1-216.
Comstock, J. A. 1927. Butterflies of California. By author. 344 p.
Downey, J. C. 1966. Host-plant relations as data for butterfly classification. System. Zool. 11: 150-159.
Downey, J. C. & D. B. Dunn. 1965. Variation in the lycaenid butterfly Plebejus icarioides III. Additional data on foodplant specificity. Ecology 45: 172-178.
Downey, J. C. & W. C. Futter. 1962. Variation in Plebejus icarioides (Lycae- nidae ). I. Food-plant specificity. J. Lepid. Soc. 15: 34-42.
EMMEL, T. C. & J. F. Emmet. 1973. The butterflies of southern California. Nat. Hist. Mus. Los Angeles Co. Sci. Ser. 26: 1-148 p.
VOLUME 32, NUMBER 1 19
Fassett, N. C. 1945. Juniperus virginiana, J. horizontalis, and J. scopulorum— IV. Hybrid swarms of J. virginiana and J. horizontalis. Bull. Tor. Bot. Club 72: 379-384.
Jounson, K. 1972. Juniperus (Cupressaceae) speciation and the ranges and evolution of two Callophrys (Lycaenidae). J. Lepid. Soc. 26: 112-116.
Jounson, K. 1976a. Three new Nearctic species of Callophrys (Mitoura), with a diagnostis (sic) of all Nearctic consubgeners (Lepidoptera: Lycaenidae). Bull. Allyn. Mus. No. 38, 30 p.
1976b. A new species of Callophrys (Mitoura) from Mexico (Lepidop-
tera: Lycaenidae). Pan-Pacific Ent. 52: 60-62.
Callophrys (Mitoura) nelsoni (Boisduval) and siva (Edwards) in the northwestern United States, Lycaenidae. (Ms. in prep.).
Jounson, K. & P. M. Borco. 1976. Patterned perching behavior in two Callophrys (Mitoura), Lycaenidae. J. Lepid. Soc. 30: 169-183.
Perching behavior as a partial boundary of distribution and foodplant utilization in Callophrys (Mitoura) (Lepidoptera: Lycaenidae). (in review). Kxuors, A. B. 1951. A field guide to the butterflies of North America, east of the
Great Plains. Houghton Mifflin Co., Boston, xvi + 349 p.
LirtLe, J. B. 1971. Atlas of United States trees. U.S.D.A. Misc. Publ. No. 1146.
Pease, R. W. 1963. Extension of known range of Mitoura hesseli. J. Lepid. Shen a Romer a
Rawson, G. W., J. B. Zrecier, & S. A. Hesset. 1951. The immature states of Mitoura hesseli Rawson & Ziegler. Bull. Brooklyn Ent. Soc. 46: 123-130.
Remincton, C. L. & R. W. Pease. 1955. Studies in foodplant specificity. 1. The suitability of Swamp White Cedar for Mitoura gryneus (Lycaenidae). Lepid. News. 9: 4-6.
Ross, J. G. & R. E. Duncan. 1949. Cytological evidences of hybridization be- tween Juniperus virginiana and J. horizontalis. Bull. Tor. Bot. Club 76: 419- 429. -
ScHurtz, D. L. 1971. A tri-parental swarm of Juniperus L. Unpubl. PhD Thesis, University of Nebraska, Lincoln. 234 p.
U.S.D.A. 1953. Checklist of native and naturalized trees of the United States (including Alaska). U.S.D.A. Agric. Handb. No. 41. 472 p.
Van HaversBEKE, D. F. 1968. A population analysis of Juniperus in the Missouri River Basin. Univ. Nebr. Studies, No. 38. 82 p.
Vasex, F. C. 1966. Distribution and taxonomy of three western Juniperus. Brit- tonia 18: 350-372.
Journal of the Lepidopterists’ Society 32(1), 1978, 19
IMPORTANT NOTICE TO CONTRIBUTORS
Authors should submit an abstract for all articles to be published in the Journal beginning with Volume 32, issue No. 1. The abstract should summarize the im- portant contents and conclusions of the paper in concise and specific sentences. It should not exceed 1-3% of the paper's total length, and should indicate the ob- jectives, methods, and topics covered by the paper. References to literature, illustra- tions, and tables should be omitted from the abstract, since this should be com- pletely self-explanatory. These abstracts will be printed at the front of each paper, and also will be forwarded to Biological Abstracts for possible inclusion therein. Such abstracts should accompany articles only. They are not required for general notes. Authors who now have papers in press should forward such abstracts to the new editor at their earliest convenience.
Journal of the Lepidopterists’ Society 32(1), 1978, 20-36
FOODPLANT, HABITAT, AND RANGE OF CELASTRINA EBENINA (LYCAENIDAE)
WarRREN HERB WAGNER, JR. Department of Botany, The University of Michigan, Ann Arbor 48109
T. LAwRENCE MELLICHAMP Department of Biology, University of North Carolina, Charlotte 28233
ABSTRACT. The larval foodplant of the recently described Celastrina ebenina Clench is Aruncus dioicus (Walt.) Fernald (Rosaceae), the Goat’s-beard. Over 150 adults were raised from eggs and young larvae. The range of the butterfly coincides nicely with that of the plant, from Pennsylvania to North Carolina and Missouri. The habitat for plant and butterfly is moist, rich forest. The closely related C. pseudargiolus Boisduval & LeConte is vastly more abundant and ubiq- uitous than W. ebenina, and has a wide variety of larval foodplants. Larvae of C. ebenina differ in several respects from those of C. pseudargiolus, including color pattern and stellate processes. Also described and discussed are the plant and butterfly associates of C. ebenina, flower visitations of the adults, experiments on foodplant specificity, feeding characteristics of the larvae, broods, botany of the foodplant, and geographical distributions, including a number of new locality records. A guide for discovering new colonies of this rare eastern American butter- fly is provided.
Except for brief reports (Clench, 1972; Wagner & Showalter, 1976), little has been published on the biology of the poorly known Dusky Blue Butterfly, Celastrina ebenina Clench, of the eastern United States. This lycaenid is notable for several reasons. Interpreted for over a century as an aberration or form, it was not recognized as a distinct species until 1972. The colors of the upper surfaces of the males and females are peculiar for being the reverse of the usual situation among plebejine blues in that the males are dull, dark grayish-brown or blackish, while the females are mainly lustrous blue. The insect is regarded as especially rare and local, having been reported previously, usually as just one or a few individuals, from only 12 localities. Knowledge of its foodplant, behavior, habitat, and geographical distribution has been incomplete or lacking.
The present paper records the results of research in 1976. We now understand the ecology of C. ebenina far better than we did in the past, and we believe that we have an explanation for the geographical distri- bution and sporadic occurrence of the species. At the outset of this study, as botanists, we entertained the possibility that the peculiarities of oc- currence of C, ebenina might be due to specialized larval foodplant preference,
Because of earlier reports of the species there, the area chosen for our field investigations was in the Daniel Boone National Forest, in and
VoLUME 32, NUMBER 1 2]
around the Red River Gorge in Powell and Menifee cos., Kentucky, a few miles north of the town of Slade. We found that habitats suitable for C. ebenina occur extensively, though sporadically, throughout this area, and in some places the butterfly is common or abundant, though extremely localized, flying with the much more numerous and ubiquitous Common Blue or “Spring Azure,” C. pseudargiolus Boisduval & LeConte. The topography in the Red River Gorge is made up of steep, abrupt hills and valleys, ranging from 700’ altitude in the river and stream beds to over 1300’ at the tops of the highest hills. Commanding cliffs of light tan or whitish sandstone crop out at the crests of some of the hills, but in the valleys where C. ebenina flies, the most conspicuous rock is a loose, broken, dark-gray shale. The general area is mainly traversed by narrow dirt roads, especially along the larger streams, and there are only occasional two-lane hardtop roads.
Our search for suitable habitats to study was initiated on 17 April 1976. We drove from place to place along the country roads, stopping wherever we encountered roadside puddles or wet streamside flats. In such spots accumulations of butterflies were the rule when it was bright and sunny, especially between the hours of 0900 and 1400. Among the guests at these “puddle parties” we encountered rare males of C. ebenina, prac- tically always with at least several and usually many C. pseudargiolus, the latter being much more conspicuous in flight. In spite of our success in encountering specimens of C. ebenina here and there over an area of perhaps a dozen square miles, at no place were there more than a few individuals. Rare observations of females showed them almost always to be in flight, and following them gave no clue to where their eggs were laid.
Finally, at around 1030 hrs the following day, we came upon an excel- lent locality—a moist, steep, rocky, north-facing wooded slope, along a narrow dirt road along the Red River. In only 20 min, ca. three dozen C. ebenina of both sexes were observed, the females numbering about twice as many as the males. The latter were all more or less worn, dull in appearance, and flying along the edges of the road, occasionally alighting on wet muddy spots (Fig. 1, lower photo). Most of the females, however, were in fresh condition and were flitting around the vegetation on the slopes above and below the road. It was obvious that many of them were engaged in oviposition. They flew rapidly in an “exploratory” pattern, pausing often at a single species of plant, the Goat’s-beard, Aruncus dioicus (Walt.) Fernald. Occasionally the females landed on the abundant and conspicuous Wild Hydrangea, Hydrangea arborescens L., but usually only momentarily.
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
bo bo
Fig. 1. Celastrina ebenina habitat along Red River, Powell Co., Ky. Upper: View upstream showing extensive understory growth on forest slope. Butterflies visit Geranium flowers here. Lower: View downstream showing damp ruts in road where males congregate. (Photo by J. M. Beitel. )
VOLUME 32, NUMBER 1 23
Figs. 2-6. Celastrina ebenina adults and eggs: 2, female laying eggs on young Aruncus shoots (R. P. Carr); 3, freshly emerged male, showing blue scaling (T. L. Mellichamp):; 4, eggs on leaf and inflorescence primordia of Aruncus (cf. fig. 14) (T. L. Mellichamp): 5, unhatched eggs; 6, eggs (two of them hatched) showing wall pattern detail (R. P. Carr).
On Aruncus the butterflies alighted on very young, unfolding leaflets, and then walked around slowly, laying eggs (Fig. 2). After an individ- ual would fly away, we could easily find the eggs, mainly on the lower blade surfaces, on and between the main veins of the leaflets (Fig. 4). At this time of year the main axis of the plant is still embryonic, and the habit is very different from the mature habit with the inflorescence fully developed (cf. Fig. 13, full-grown plant, and Fig. 14, stage at time of oviposition). When freshly laid, the eggs showed a grayish blue color.
24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Sometimes several eggs are laid in the same spot, but usually they are laid separately (Figs. 4, 5, & 6). Egg-laying occurred over 1% hours of observation, and there was no sign of abatement after 1200 hrs when we left the site.
For careful observations, a total of 18 young cuttings like those in Fig. 14 were randomly collected. The two oldest leaves overtop the main shoot at this stage. The main shoot, with its very young leaves and inflorescence primordium, is only about one-fourth the length of the oldest leaf (the large, bipinnate leaf on the right side of the figure) and one-half the length of the next oldest leaf. The softest, most embryonic tissues are those of the primordial main shoot. We found a total of 133 eggs altogether on the collected shoots—35 on the oldest leaves, 26 on the next oldest, and 66 on the young main shoots. Thus, our evidence suggests that the butterflies prefer to lay eggs on the youngest tissues. The number of eggs averaged seven per cutting, but one had 18.
Naturally we wondered whether at least some of the eggs we found did not represent the closely related C. pseudargiolus, which flies in large numbers at this locality, a species noted for its polyphagy. There- fore, we decided a couple of weeks later to conduct an experiment de- signed to help us answer this question, as will be described below.
The plant community on this slope is a rich mixed-mesophytic forest. We recorded a total of 31 trees and shrubs and 56 herbs (including ferns and graminoids ) in the area where C. ebenina was ovipositing. Most of these plants are typical associates of Aruncus dioicus, and some of the more prominent ones will be enumerated later in our discussion of this plant.
The best place to find C. ebenina adults is in association with other mud-loving butterflies (C. pseudargiolus, Callophrys henrici, Erynnis spp., and Papilio spp.) in damp spots along dirt roads and gravelly, sandy, or muddy river flats. Practically all of the “mudding” individuals we observed were males, sometimes as much as a quarter of a mile from the foodplant, although usually much closer. On only two occasions did we find females landing on wet soil. We disturbed one of them several times, but each time it returned.
The only flower which seemed to attract C. ebenina at this locality was the Wild Geranium, Geranium maculatum. The showy rose-purple flower has a flat five-petalled corolla 3.0-3.5 cm across. Bearded nectaries occur between and at the bases of the petals. The butterflies walk over the top surfaces of the corolla and probe between the petal bases. Later
'A complete list of the associated plants at this locality will be sent upon request to readers.
VOLUME 32, NUMBER 1 25
Figs. 7-12. Celastrina ebenina immature stages: 7 & 8, instar 1 caterpillars (J. G. Bruce III); 9 & 10, instar 2 caterpillars (J. G. Bruce III); 11, mature caterpillars, showing pale, poorly contrasting pattern and characteristic leaf damage; 12, pupa attached to Aruncus leaf (R. P. Carr).
we discovered that the bulk of individuals obtain their nectar from underneath the flower! Both sexes flit from flower to flower, landing on the peduncle or on the underside of the perianth, then walking toward the sepal bases where they insert their proboscises. So positioned on the flowers, the butterflies are invisible from above.” We recorded all of the species of butterflies we found in association * Curious to see whether other butterflies behaved in the same manner, we discovered (in
Michigan in the middle of May) that Erynnis juvenalis displayed the same routine on Geranium flowers.
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
pecaaimnnanat e —.rrtrs——C
1O 6M.
(‘a
—s
13, habit drawing of fully grown Aruncus dioicus showing inflorescence (apex) and form y); 14, young shoot of Aruncus at time of oviposition by C. ebenina (see text) (T. L.
~, Vg , Vial of WEN Nia Fog a ; AS . RSS Cor ) A neil
Figs. 13-14. Celastrina ebenina foodplant: of compound leaves in mid-June (Del. J. G. Lac
Mellichamp ).
VoLUME 32, NuMBER 1 27
ee
= *\\
Fig. 15. Geographical distribution of Aruncus dioicus (stippling) and docu- mented localities for Celastrina ebenina (dots) (T. L. Mellichamp).
28 JouRNAL OF THE LEPIDOPTERISTS SOCIETY
with C. ebenina on 18 April. The following list is a compilation, since no one locality had all of the species (those marked with an asterisk oc- curred with C. ebenina on wet soil): Papilio glaucus,* P. troilus,* Battus philenor, Pieris virginiensis,* Colias eurytheme, C. philodice, Polygonia comma, P. interrogationis, Boloria bellona, Phyciodes tharos,* Callophrys henrici,* Celastrina pseudargiolus,* Erynnis brizo,* E. juvenalis,* Amblyscirtes samoset, A. aesculapius, and Epargyreus clarus. On several later visits to the study area by Loran D. Gibson, Amos H. Showalter, and ourselves, the following additional species were observed, most of them, however, well beyond the normal flight period of C. ebenina, which ended the first week of May: Limenitis astyanax, Speyeria cybele, Asterocampa celtis, Ewptychia hermes, E. cymela, Lethe ~ creola, L. anthedon, Autochton cellus, Thorybes pylades, and Poanes hobomok.
Experiment on Foodplant Specificity
The question we addressed ourselves to was whether C. pseudargiolus also shared the foodplant of C. ebenina. The former is abundant in the Red River Gorge area and is in constant association everywhere with C. ebenina. Although we did not observe C. pseudargiolus ovipositing upon Aruncus, this would not preclude the possibility that it occurs. The num- ber of Celastrina eggs we observed was so great that it seemed reasonable to assume, because of their morphological similarity, that perhaps some of them belonged to the abundant species, especially since C. pseu- dargiolus is noted for its varied bill of fare. Literature records show that no less than 10 families of flowering plants contain larval foodplants for this species, and we have discovered early stages of it on such different families as Caprifoliaceae and Cornaceae. It would not have surprised us, therefore, to find C. pseudargiolus utilizing Aruncus, especially since its larvae have been reported on Spiraea, another closely related genus in the same subfamily of Rosaceae.
On 2 May, accordingly, we revisited the Red River Gorge area and collected from seven to nine shoots with attached eggs and minute larvae from each of five colonies of the foodplant, all of them a mile or so separate from one another. These shoots were brought back to The Uni- versity of Michigan in Ann Arbor, and the eggs and young larvae were raised to adulthood. Extra foodplant shoots were kept in plastic bags and refrigerated to preserve them until they were needed, and additional foodplants were grown for further supplies. The cut bases of the shoots of Aruncus bearing eggs and larvae were inserted in jars of water in | x 1 x 2 ft glass aquaria covered with a plastic material to keep the
VoLUME 32, NUMBER 1 29
larvae from escaping. Some of the caterpillars drowned when they walked down the main stems into the water, others escaped and were lost, and some died as a result of cannibalism. Nevertheless, a total of 153 butterflies were raised to maturity by the end of the second week in June, 76 males and 77 females. Another eight butterflies came from larvae which escaped from their containers. The first caterpillars pupated on 16 May, 14 days after the field collection, and the first adults to emerge appeared on 23 May. This second brood apparently does not occur in nature (see below).
The results of this experiment were striking. Not only were none of the “checkery” caterpillars of C. pseudargiolus noticed among the paler, less contrasty larvae of C. ebenina (Figs. 7-9), but not a single one of the adults was C. pseudargiolus; all were C. ebenina. Furthermore, the 14 additional butterflies from our 18 April eggs were all C. ebenina. Also, the four individuals that emerged from dormant chrysalids the following winter were all C. ebenina. With C. pseudargiolus as abundant as it is in the localities where our collections were made, one would expect at least some evidence of its occurrence on Aruncus if it fed on that plant at all.
It is interesting to note that ca. one-third of the males display blue scales (Fig. 3), at least at the time of emergence. Although the original description by Clench (1972, p. 37) records the male upperside as “uniform blackish brown when fresh,’ blue scales are conspicuous in these individuals, especially on veins R, to R; and the discal portions of M;, Cu,, and Cu, on the fore wings, and scattered in the discal area of the hind wings. The occurrence of blue scaling on the males may con- stitute a regional difference. The ground color above of the males varies from blackish gray to pale slate gray. The females vary greatly in ground color (whitish blue to fairly intense blue) and the amount of dark mark- ing. There is no confusing any of these specimens, however, with any of the forms of C. pseudargiolus known to us.
To make preliminary comparisons of the caterpillars of C. ebenina and C. pseudargiolus, we obtained eggs and first instar larvae of the latter in two localities near Ann Arbor in Washtenaw Co., Michigan. They were readily found on the young inflorescences of Grey Dogwood, Cornus racemosa and Red Osier, C. stolonifera (Cornaceae), and Nanny- berry, Viburnum lentago (Caprifoliaceae); the dates of collection were 8 and 22 May. We maintained the larvae under the same conditions as those of C. ebenina but kept them in a separate room to avoid any op- portunity of possible escapees getting mixed up.
We did not make detailed comparisons of the larval morphology, but
30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
certain differences were obvious. The final instar caterpillars of C. ebenina (Fig. 11) are considerably more uniform in color than those of C. pseudargiolus. They are pale whitish blue-green, with three slightly contrasting longitudinal stripes of yellowish white, two lateral and one dorsal. Between the dorsal and each lateral stripe is a line of yellow— white dots, and similar dots and dashes are scattered over the body. The caterpillars of C. pseudargiolus, as is well known, are more variable in eround color, ranging from pinkish or pale bronze to yellowish green; the pattern is much more conspicuously blotched as a rule. Earlier larval stages of the two species (young instars of C. ebenina shown in Figs. 7-10) are very similar in their uniform pale green color.
In both species the mature larvae are velvety because of elaborate stellate processes, each with a narrow and pointed filament arising from the center. In C. ebenina the filaments are only slightly curved and stand nearly erect. In C. pseudargiolus the filaments are more strongly curved and tend to be arched over such that they are distally nearly parallel to the body surface. This difference applies mainly to the more abundant, smaller filaments; larger ones in both species are more alike, only slightly curved and nearly erect.
Another difference between the larvae of the Dusky and Common Blues involves the use of their respective foodplants. Although larvae of C. ebenina occasionaily feed upon the tiny embryonic floral primordia of Aruncus, the bulk of their feeding is upon the blade tissue between the major lateral veins of the leaflets. Their feeding produces charac- teristic elongated perforations in the blades; these persist when the leaflets have achieved their full size, and give valuable clues during late spring and summer for localities of the butterfly. Fig. 11 shows the characteristic perforations. The younger caterpillars were observed on the tops of the leaves more often than the older ones. Feeding between a pair of lateral veins usually begins near the midrib of the leaflet and progresses outward, but only rarely all the way to the margin and in- cluding it. Very large caterpillars in the last instar may eat major lateral veins.
In our experience the larvae of C. pseudargiolus feed primarily upon floral primordia. Their eggs and larvae are found in inflorescenses. We tried moving larvae to leaves, but always, after eating a small amount, they would return to their floral clusters where they continued feeding.
Larvae of C. pseudargiolus bite into the sides of the closed flower bud or the inferior ovary and eat the entire contents or leave certain parts
(e.g., petals). The body of the caterpillar sits motionless on the floral pedicel or the side of the bud, and the extensible head is projected into
VOLUME 32, NuMBER 1 31
the cavity in the young flower to feed. From certain angles the larva appears headless.
Broods
Previous field experience of our own and others suggested either that C. ebenina is univoltine (in the Red River Gorge, flying from the second week of April to the first week of May), or else, if it has more than one brood, that adults of any later broods are so similar to those of C. pseu- dargiolus that they have not been recognized as distinct. The second lab- oratory brood is like the first brood. As it turned out, the vast majority of our pupae (Fig. 12) emerged within a week or two of pupation. Of those that failed to emerge and were kept until the following winter, only a few produced butterflies.
The first adult from our 18 April field collections of C. ebenina eggs emerged on 11 May. The first from our 2 May collections of eggs and young larvae emerged on 23 May. Emergence continued until 29 May, when it was stopped by placing the cultures in a coldroom from this date until 4 June to keep more butterflies from appearing while we were out of town. After we returned them to normal temperatures, emergence resumed on 5 June and continued to 12 June. A grand total of 175 butter- flies in our laboratory cultures seemed to demonstrate that there is a second brood in the wild that follows closely upon the first, the second brood flying from the second week in May to the first or second week in June.
We therefore returned to the Red River Gorge to determine whether a second brood occurred in nature. The results of our survey were most unexpected. Loran D. Gibson visited there on 26 May and saw not a single C. ebenina, although many other butterfly species were seen (in litt., 2 June 1976). We then reconnoitered the area on 4 June, when the butterflies should have been at their peak abundance, if there is a second brood in nature. We saw no C. ebenina despite the fact that C. pseu- dargiolus was common as well as 16 other species of butterflies. On 9 June, Amos H. Showalter searched the area and reported that “C. pseu- dargiolus was common, but no ebenina” (in litt., 28 June).
After 13 June, the 28 pupae that remained in our cultures apparently went into dormancy, and no more butterflies emerged. At the end of June, therefore, we placed them in a coldroom at a temperature of 2°C and left them there until 22 December. We hope that by keeping them thus, in a set-up that we have used for “winterizing” fruits and seeds, we might avoid the destructive effects other workers have had with C. pseudargiolus, which involve either drying out or molding of the chrys-
32 JouRNAL OF THE LEPIDOPTERISTS SOCIETY
alids. On 1 January 1977, two females emerged, one of which failed to expand its wings. Another female emerged on 4 January, and still an- other female failed to escape from the pupal skin. All the remaining pupae appeared to have died, eight of them having moldy surfaces and the rest having an unnatural brown color. }
What can we conclude regarding the broods of C. ebenina? In the field we found no evidence for a second brood. If there is one, it must be in extremely low numbers, i.e., a small “partial” brood. Somehow the conditions of our laboratory cultures must have caused an abnormal eclosion without the customary prolonged dormancy period. Some dia- pause stimulus that effects C. ebenina must have been weak or missing under the conditions of our experiment, and thus only a small percentage of the chrysalids went into long-term dormancy.
Botany of the Foodplant
The colloquial name of the foodplant, Aruncus dioicus, may cause some confusion, since its name “Goat’s-beard” is applied also to the un- related Tragopogon pratensis L. in the Asteraceae, a naturalized weed from Europe. The generic name Aruncus comes from the Greek and means literally “goat’s beard.” It is a member of the Rosaceae and is a native eastern American plant of rich, mature forests. It is famous among United States’ botanists as an illustration of convergent evolution, because superticially A. dioicus resembles closely the “False Goat’s-beard,” Astilbe biternata, of the Saxifragaceae. So closely do these plants resemble each other that they are regularly confused, even in herbaria. Ecologically the two look-alikes occupy almost identical niches, and they are both unusual among members of the mesophytic forest association in being dioecious (male and female flowers being borne upon separate plants). They are pollinated not by wind, which is the usual situation in dioecious plants, but rather by insects, mainly small Hymenoptera. The geograph- ical range of Aruncus dioicus is shown in Fig. 10. The range of Astilbe biternata is much narrower, mainly in the mountains of North Carolina and adjacent parts of Virginia, W. Virginia, Kentucky, Tennessee, South Carolina, and Georgia. Aruncus overlaps it completely, so that students of C. ebenina must be warmed of the danger of confusing the “True” with the “False” Goat’s-beards in the area of their sympatry. Accord- ingly, we have prepared a comparison of the two in Table 1, the most obvious characters marked with asterisks. A line drawing of a mature specimen of Aruncus nearly 1’4 m tall is reproduced in Fig. 1. The stage of growth when the plant serves as larval food for C. ebenina is shown in Fig. 2, corresponding to only the two bottom leaves and the lower
VOLUME 32, NuMBER 1] Oo
TaBLE 1. Comparison of “True” and “False” Goat’s-beards.
Aruncus dioicus Astilbe biternatum *1. Stipules Absent Present 2. Terminal leaflet Unlobed 3-lobed 3. Leaf base Attenuate Cordate (heart-shaped ) *4. Veins per leaflet 8-18 pairs 8 or less pairs *5. Leaf and stem hairs Absent Abundant ( glandular ) 6. Marginal teeth Convex Acuminate 7. Sepals per flower 5 5) 8. Petals per flower 5B 0-5 9. Stamens per flower 15-20 8-10 *10. Carpels per flower 3-4 2
sixth of the drawing. The tissues upon which the caterpillars feed are soft, and the earliest instars feed upon the most embryonic parts. The plants grow rapidly and come into flower 5-7 weeks after the butterflies lays their eggs.
Both Aruncus and Astilbe have rather massive underground stems that produce large roots or root masses which hold the plants firmly in place on steep slopes (Fig. 3). Many buds are present at the ground level, and some of these may develop into shoots at the crown, producing clumps of as many as eight flowering shoots. Their spreading compound leaves fill in the space where they grow, presumably allowing little growth of other plants beneath them.
The Goat’s-beards are most typical of rich, mesic woods, partly shaded roadsides, and sloping sides of streams and rivers. The ancient habitat was probably on steep eroding slopes and stream banks in dark forested areas, but man has stimulated its spread by creating new habitats where roads have been cut through the mesic forest. In a uniformly shaded forest stand only 10% of the plants may flower, but when released from the effects of low light levels, as on road cuts at the forest edge or steep, eroding stream banks, the populations may display up to 100% flowering. The plants require, however, relatively cool, moist conditions, and they exist almost exclusively upon north-facing slopes. Aruncus is known to occur as high as 5500’ altitude in the mountains (Buncombe Co., N.C.), although the average occurrence throughout its range is considerably less than 2500”.
Would-be collectors of C. ebenina should seek the foregoing site con- ditions, with the following array of associated species (based upon studies of a number of Aruncus localities by Mellichamp 1976): woody plants—Acer rubrum, A. saccharum, Aesculus octandra, Betula lenta, Carpinus caroliniana, Cornus florida, Fagus grandifolia, Lindera benzoin,
34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Liriodendron tulipifera, and Tilia americana; herbs—Adiantum pedatum, Athyrium filix-femina, Botrychium virginianum, Carex plantaginea, Cimicifuga racemosa, Geranium maculatum, Impatiens capensis, La- portea canadensis, Tiarella cordifolia, and Trillium spp. The presence of a majority of these species, together with Wild Hydrangea, Hydrangea arborescens, given the topographical conditions cited, especially a north- facing slope, should lead to colonies of Aruncus and therefore C. ebenina.
Ranges of Plant and Butterfly
There is a remarkable correlation between the known localities where the butterfly has been found and the geographical distribution of Aruncus dioicus. Clench (1972) has already discussed doubtful records for C. ebenina, including New York City and southern Colorado. We should like to add to the list of doubtful records that of Blatchley in Wabash Co., Indiana (Clench, 1972, p. 41), unless an actual specimen is dis- covered. The “black male” he referred to could have been a melanistic male of C. pseudargiolus, a wind-blown stray of C. ebenina, or a mis- labeled specimen. Clench mentions that Edwards had vaguely attributed the species to Tennessee and Georgia, although Clench himself had seen no specimens in these areas. It now seems very likely, as Clench (1972) suggested, that C. ebenina may turn up in both of those states.
Aruncus grows far to the west of the localities cited in earlier studies of C. ebenina (Clench, 1972; Wagner & Showalter, 1976), the western- most documented records for which were all east of Cincinnati, Ohio, and Lexington, Kentucky. We can now report records in Illinois, Missouri, and Arkansas. Charles L. Remington took a fresh male south of Elsah, Jersey Co., Illinois, on 15 April 1942 (in litt., 21 Jan. 1977). He found additional males in St. Louis Co., Missouri, in the late 1930’s. Also, J. Richard Heitzman (in litt., 14 July 1976) has informed us that he ob- tained by exchange “a male and a female taken 4 May 1924 at Creve Coeur Lake near St. Louis, St. Louis Co. There is no collector’s name, but it should have been one of the active collectors of the day, probably Ernst Schwarz, E. P. Meiners, or H. I. O’Burme. AII were active at the time and collected often at Creve Coeur.” The dot shown in Fig. 10 for Arkansas is based upon a single male taken in Hickory Flat Hollow, Washington Co., on 2 April 1973 by Edward Gage. Gage writes (in litt., Dec. 1976): “This male particularly stood out as it was flying about a mud puddle with several C. argiolus and Everes comyntas. .. . I im- mediately assumed that it was a melanic form and quickly collected it. The site was in close proximity to a draw or shallow canyon. About 90% of the immediate surrounding area is deciduous hardwood. ... Can-
VoLUME 32, NUMBER 1 By5)
yons and woodland extend all around Beaver Lake from the collecting site. ”
CONCLUSIONS
Celastrina ebenina may have larval foodplants other than Aruncus dioicus. However, it should be noted that none of the unquestioned localities of this butterfly lies outside the known range of Aruncus. Furthermore, we now have good reason to believe that the abundant Common Blue, C. pseudargiolus, does not share the foodplant of the Dusky Blue, C. ebenina. Our experimental raising of eggs and young larvae on Aruncus from areas in which C. pseudargiolus is abundant revealed not a single specimen of that species. All were C. ebenina.
One reason for apparent rarity of C. ebenina in comparison with its near relative is that its geographical range is much more limited. Another is that it is probably monophagous rather than polyphagous. Its food- plant is confined to one habitat—north-facing, richly wooded, shaded slopes, so that the butterfly tends to be highly localized and colonial. The multiple foodplants of C. pseudargiolus occupy many habitats, and the butterfly is therefore practically ubiquitous.
Celastrina ebenina is probably often overlooked. The dull males in flight may suggest badly worn individuals of C. pseudargiolus. Celastrina ebenina tends to fly closer to the forest floor (a concomitant of its under- story foodplant?), whereas C. pseudargiolus has a slow, up-and-down flight reaching the shrub and lower tree layer. The bright reflecting blue of C. pseudargiolus males plus their tremulous flight pattern through the woods at heights of roughly 2-10’ make them especially visible. The females of C. ebenina resemble dull females of C. pseudargiolus, but they are even more localized than the males, rarely even visiting mud puddles, occurring rather in the herb layer of the forest in more or less checkered sunlight. We wonder how many collectors (including our- selves!) in the spring have overlooked C. ebenina while they focused instead on such critical genera as Erynnis and Callophrys as well as such widely advertised rarities as Pieris virginiensis and Erora laeta, both of which are now known to fly in association with C. ebenina.
If the conclusions of this research are correct, we predict that C. ebenina will be found not only in many new localities in the states from which it is already known, but, in addition, southern Indiana, eastern Tennessee, northern Georgia, western South Carolina, and western Mary- lan. To help achieve this, we propose the following formula:
1. Locate areas of rich mesophytic forest in rolling or mountainous country.
36 JoURNAL OF THE LEPIDOPTERISTS SOCIETY
2. Follow roads or streams and find north-facing, cool, shaded forest slopes with some erosion or disturbance.
3. Look for plant associations including such trees and shrubs as Hydrangea, Acer, Aesculus, Betula, Carpinus, Cornus, Fagus, Lindera, Liriodendron, Rhododendron, and Tilia plus the majority of herbs given above.
4, Explore for large colonies of “Goat’s-beard,” Aruncus dioicus, the larval foodplant, especially on north-facing roadsides and streamside slopes.
5. Visit the area in April and early May in search of C. ebenina— males on muddy spots, females around the foodplant, and both sexes on Geranium flowers (careful!—they may be underneath the petals).
6. Or, if the weather is cloudy or rainy, search the young shoots of Aruncus for greenish blue, rough-surfaced eggs, and (or) pale green caterpillars, the latter evidenced by narrow perforations in the soft leaf tissue between the veins of young leaflets.
To sum up, our evidence thus far indicates that Celastrina ebenina is a “specialist,” not a “generalist.” When compared with C. pseudargiolus, it has a narrow range (not broad), one foodplant (not many), a single brood (not several), and an essentially uniform morphology (not many varieties and forms ).
ACKNOWLEDGMENTS
We thank the following persons for their help in this project: J. M. Beitel, J. G. Bruce III, Robert P. Carr, D. J. Harvey, Loran D. Gibson, J. Richard Heitzman, Janice Glimn Lacy, Amos H. Showalter, Florence S. Wagner, and K. S. Walter. We are especially grateful to Harry K. Clench for his encouragement and critical advice.
LITERATURE CITED
CLeENCH, Harry K. 1972. Celastrina ebenina, a new species of Lycaenidae (Lepidoptera) from the eastern United States. Ann. Carnegie Mus. 44: 33-44.
Me.iicuamp, T. L. 1976. A comparative study of Aruncus (Rosaceae) and Astilbe (Saxifragaceae ), and the problem of their relationships. Doctoral Thesis. The University of Michigan, Ann Arbor.
Wacner, W. H., Jr. & AMos H. SHowatrer. 1976 Ecological notes on Celastrina ebenina (Lycaenidae). J. Lepid. Soc. 30: 310-312.
Journal of the Lepidopterists’ Society 32(1), 1978, 37-48
STUDIES ON RESTINGA BUTTERFLIES. II. NOTES ON THE POPULATION STRUCTURE OF MENANDER FELSINA (RIODINIDAE)
Curtis J. CALLAGHAN IBM do Brasil Ltda., P.O. Box 1830, Rio de Janeirv, Brazil
ABSTRACT. The objective of the study was to describe various aspects of the adult behaviour and population dynamics of the riodinid butterfly Menander felsina (Hew). The conclusions were based on four years of field observations and a marking-recapture study conducted over a period of 15 weeks.
The population was characterized by low intensive and extensive frequencies (rarity in numbers and space respectively), characteristics shared by many other forest riodinid species. M. felsinad maintained constant population levels of about 19 individuals over the 15 week marking-recapture period, due to 1) longevity above that of most holarctic lycaenids, 2) low egg laying frequency both in time and space, and 3) male territoriality, which results in older males doing most of the mating. Limited adult and larval food sources were discounted as an explanation.
The population was found to be distributed in small groups or colonies near food plant localities. The reasons for this were the low extensive distribution of foodplants coupled with high female vagility. Depending upon conditions at each foodplant locality, such as predation and exposure to the elements, each colony could become extinct, only to be reestablished by another wandering female.
Studies on the ecology of neotropical butterflies to date have been concerned for the most part with the larger species such as heliconiines (Crane, 1955, 1957; Turner, 1971; Ehrlich & Gilbert, 1973), nymphalines (Benson & Emmel, 1973; Young, 1972), and papilionids (Cook e¢ al., 1971). Smaller butterflies such as riodinids have received only super- ficial treatment, mainly consisting of scattered notes on observed habits and population density. Early writers such as Bates (1864) and Seitz (1913) made reference to the rarity of individuals and the great num- bers of species that characterize neotropical riodinids. This has been confirmed statistically by Ebert (1969), who used the terms low ex- tensive and intensive frequency to denote the rarity of populations and individuals, respectively. However, there has been no attempt to explain the mechanism(s) involved in maintaining populations in spite of such low intensive and extensive frequencies. The opportunity to study this phenomenon was provided to me by the discovery of a reasonably large colony of Menander felsina (Hew.) near Rio de Janeiro, Brazil. Whereas the ecology of this butterfly resembles closely that of other forest- dwelling but less common species, it also provides the basis for con- clusions of a general nature applicable to other members of this group.
The purpose of this paper is to describe the adult behavior of M. felsina in connection with feeding, male patrolling, and mating. The
38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
results of a marking-recapture program are presented with an analysis and discussion of the population parameters.
METHODS
The observations that form the basis of this study were gathered over a period of more than 4 years at Pedra de Itauna, an area near the coast west of Rio de Janeiro, Brazil. The vegetation is typical of the restinga (coastal dune community) described in detail elsewhere (Cal- laghan, 1977). The area for the mark-recapture study was a trail some 125 m in length just inside the south (seaward) side of the low woods surrounding the Itauna Rock. Because of the density of the brush, little collecting was possible elsewhere in the forest. The mark-recap- ture project took place over a 14-week period, from 18 July-15 November 1973. Marking was done every weekend except for five near the end, which were lost as a result of inclement weather. An average of ca. 5 h collecting (900-1400 hrs) per day was devoted to the experiment on study days. Individuals, once captured, received a predetermined mark that indicated the day and were immediately released at the same spot. Occasionally, a butterfly would show signs of shock after marking by fluttering to the ground, in which case it was killed and eliminated from the experiment. For analyzing the data, the Fisher and Ford method (Fisher & Ford, 1947) was used because of the relatively small sample sizes.
RESULTS
General Behavior: Feeding, Mating, and Patrolling
Menander felsina inhabit exclusively the low woods and nearby flats on the seaward side of the forested areas. They prefer open habitats such as trails and small clearings in the woods, shunning completely the deep shade found in the higher forests. Yet, they are one of the few butterflies that fly on cloudy days, albeit in smaller numbers than on sunny ones. Both sexes can be found sunning in the early morning hours with wings outspread on the upper surfaces of leaves 2-3 m above the ground. When disturbed, they fly off with a rapid, jerky flight, circling a few times before settling, wings outspread on the undersides of leaves not far from where they departed. After a few minutes they sometimes return to the same spot. When resting, they appear to thermoregulate, raising the wings to sharper angles depending on the intensity of the sun’s rays.
Group feeding takes place in the morning hours from 800-1100, and to a lesser degree in the late afternoon. The M. felsina will visit prac-
VOLUME 32, NUMBER 1 39
tically any plant that happens to be in bloom in the habitat, often becoming so “engrossed” in feeding that they can be removed with forceps. Males and females feed together, the latter being found com- monly only at this time.
Starting ca. 1230 hours, males begin taking up positions on the edges of trails and clearings to await females. They sit motionless for long periods on the upper or undersurfaces of leaves from 0.2—2 m above the ground with their abdomens slightly raised. Males seldom leave their perches to investigate other species of butterflies. The only ones in- vestigated were small white pierids (Eurema sp.), which to some extent resemble M. felsina in flight. Larger butterflies and skippers are com- pletely ignored, which indicates that sight plays a role in the recogni- tion of rivals or mates. This activity continues until ca. 1500. This behavior is in contrast to the habits of lycaenids (Powell, 1968; Scott, 1974) and skippers (MacNeill, 1964). In these cases the males would investigate any object flying past them, including small rocks (MacNeill, 1964).
To determine the extent to which spacing occurs among males, a section of trail some 20 m long was observed on three occasions. In the middle of this area is an opening in the woods that faces outward toward the knee-high vegetation of the flats. Here, an older, slightly worn male took up a station at 1255. From time to time he would take off, flying around in an area some 5 X 3 m and perching for a few minutes on various plants within this area. At 1326, two fresh males moved into the study area, taking up positions to the right and left of the older male. When one of these flew near the older male, the latter rose up, flew around in circles with the other for a few seconds, then returned to his original spot, the fresh male alighting a few yards out- side the area of the older male. At 1405, a female alighted on the outward side of the older male’s area. He immediately flew over to the female, who took off, and followed her out onto the flats. A half hour later, he had not returned, nor had other females appeared in the areas of the fresh males. The next day at the same time, the same older male was again observed in the same area.
On another occasion only fresh males were observed. They showed less exclusiveness with regard to patrolling areas than the older males. Another male passing nearby would sometimes be engaged in a circular mutual chase, but both individuals would then settle down on leaves sometimes only a meter apart. Once a male left an area, another would often move in. During the observation periods, no females were seen entering the fresh males’ areas.
40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TasLe 1. Basic capture-recapture data on a population of Menander felsina. The capture or recapture of each individual butterfly is denoted by 1.
Aug. Sept. Oct. Nov. 18 1 9 17 24 a 14 20 15 1 iL it il 1 1 1 1 if il if 1 il 1 Il Il 1 il 1 1 1 i 1 Il il i 1 I 1 Ht 1 il iL 1 Jl 1 1 iH 1 1 it it 1 1 Il il il Hi it il 1 1 1 a I i il i 1 1 1 1 Ju 1 1 1 1 1 1 1 IL 1 1 1 1 iL 1 1 1
Tatal Malaise, 122 kL none 17 ile 19 7
bo i bo iN)
I conclude that M. felsina males do engage in spacing, the older individuals defending their chosen areas more vigorously than the younger, more inexperienced males. That the older male was successful in meeting a female suggests that they may choose and defend the preferred “rendezvous” areas.
VoLUME 32, NUMBER 1 Al
TABLE 2. Type A data trellis derived from data in Table 1. Units under “Date of Marking” refer to marks and not animals.
Date of Marking
Date Captured Released 18 1 9 7) 24 ve 14 200 t5 18 Aug. 22, 22,
1 Sept. 14 14 2
9 Sept. 15 15 2 4 17 Sept. i ley i! 4 8 24 Sept. 15 15 ip I.
7 Oct. 19 19 1 2; 14 Oct. ba ha 1 20 Oct. 24 24 5 3 15 Nov. 22, 22, 2 5
1 Low captures due to inclement weather.
On one occasion, a complete courting sequence was observed. About 1416, a female alighted on a leaf, wings outspread ca. 1 m from a perched male, who immediately flew around her several times, then alighted and, with wings moving slowly up and down, walked to a face to face position. There they remained for ca. 30 sec. Then, the female walked around to the underside of the leaf, the male followed, and copulation was initiated. This lasted for ca. 22 min. On another occasion, a pair discovered in copula under a leaf remained so for 8 min before breaking off. Sexual activity continues until late in life. On 15 November 1973, a male that had been marked three weeks previously was found in copula with a freshly emerged female. Finally, mating appears to be done by older territorial males since, of the three cases observed, two involved males that had been previously marked.
POPULATION SIZE AND MorTALITY
Tables 1 and 2 show the basic marking and recapture data gathered during the study, after Sheppard & Bishop (1973). It is instructive to note that few individuals were captured more than once, which indi- cates that collecting was not very efficient over time. Low captures on 14 October were due to rainy weather. The survival rate (Fisher & Ford, 1947) was 0.45 per week or ca. 0.91 per day. The average life span was 1.82 weeks by the formula E = il , which as-
1 — survival rate sumes that all deaths occur just before sampling. Potential survival is up to 5 weeks in the field, as confirmed by a recaptured female. Ob- served field survival for males is up to 4 weeks. When the Lincoln index modified by Bailey (1952) is applied to the study data, the weekly
42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TABLE 3. Population composition over time of new and recaptured butterflies, considering all animals captured and recaptured before and after a determined date to constitute part of the population even though they were not captured on that date.
Date Total New % Recaptures % 18 Aug. DD, 22, 100 —- - 1 Sept. 17 WP, WL 5 29 9 Sept. 20 9 45 We 55 17 Sept. 19 4 DAML 116) 79 24 Sept. 16 13 81 3 19 7 Oct. 23 19 82 A 18 14 Oct. 11 6 54 5 46 20 Oct. DB) 16 64 9 36 15 Nov. 22, Ike 68 Tf ol Average 19.4 10.4 = WA! = Standard Error +22.6% +49.2% _ +42.4% ~
estimates of total population size show great variation as a result of the large differences in recaptures from one sampling to the next.
These fluctuations were felt to be more due to inaccessibility of individuals because of the thick brush on either side of the trail than to changes in the population level. Therefore, to make the capture data more realistic a third table was mounted assuming that those animals captured in week one and recaptured in week three, for example, formed part of the population in week two even though they were not captured at that time. In Table 1 we see that on 18 August, 22 butterflies were marked and released of which five were later recaptured: two on 1 September, two others on the 9th, and one on 19 September. On the Ist, at least three butterflies were not captured.
Therefore, rather than a total of 14 for that date, we have 17 which we know formed part of the population at that time: 12 new plus 2 recaptured plus 3 which were recaptured at later dates. The numbers thus derived were entered in Table 3. The procedure was conducted for each capture period during the study.
This results in considerably smoother total capture figures, which show that if allowance is made for inefficiency in collecting, the esti- mates of numbers of individuals in the study area was quite stable over the 14-week period (Table 3) with a mean of 19.4 individuals and a standard error of 4.4 or + 22.6%. More consistent results can be ob- tained by eliminating data for 14 October. When efforts were made to capture all the M. felsina in the study area—22 July 1973 and 6 June 1974—21 and 22 individuals were recorded, respectively, which again suggests that the number of M. felsina in the study area remains stable over long periods with a low number of individuals.
VoLUME 32, NUMBER 1 43
Sex ratio data gathered during the study were unsatisfactory as a result of the similar appearance and behavior of males and females, except during oviposition and territorial displays. Because of the delicate nature of these butterflies, they had to be kept in the net during marking and afterwards immediately released. This prevented detailed examina- tion. Females, however, were a small minority of all captured. On the two occasions referred to above, 19 males were captured each time with 2 and 3 females, respectively. The reason is that the females entered the study area sporadically only for mating and feeding, thus not being as accessible as the males. The number of females caught under these circumstances is not truly representative of the female population; thus, for the purpose of this study, males and females were considered together.
DIscUSSION AND CONCLUSIONS
As noted above, tropical butterfly populations, especially those of theclids and riodinids, are characterized by low intensive and extensive frequencies. The data from the marking-recapture program and other observations on the M. felsina population permits a number of sugges- tions as to why this is so.
‘Low Intensive Frequency
Low intensive frequency means that an animal is represented by a small number of individuals in a given population. As shown by the recapture data, the number of M. felsina frequenting the study area remained low and fairly constant for long periods. Other students have made similar observations on neotropical butterfly populations. Benson & Emmel (1973) observed a roost of Marpesia berania (Hewitson) in Costa Rica that maintained constant equilibrium of population size for more than 3 months because of constant rates of recruitment and mortality. Ehrlich & Gilbert (1973) recorded a similar structure for Heliconius ethilla Godt in Trinidad over a period of 2 years, as did Young (1972) for Siproeta epaphus (Latreille) in Costa Rica. Why do these populations have an equilibrium level and what is the mechanism that enables them to maintain it? Ehrlich & Gilbert (1973) explain the constant population level in H. ethilla through the constant recruitment of new adults over time which equals mortality. This was thought to be due to unvarying predator pressure on the immatures and not be- cause of a lack of foodplant, which was quite common in the habitat. The other factor was limited adult nectar resources, which controlled egg production, thereby regulating the production of immatures. Young
44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
(1972) in his study of S. epaphus likewise found that egg and adult numbers remained constant throughout the study period. As a mecha- nism, he suggested low fecundity and low adult vagility as well as the sheltered nature of the forest understory community. Both studies dis- count outmigration as a factor.
My observations on M. felsina allow the following conclusions to be made. First, neither foodplant nor nectar availability appeared to be a limiting factor. The larval damage was always small compared with the amount of foodplant available, showing little pressure on foodplant resources. The adult’s preference for nearly anything that is in bloom and its willingness to travel good distances eliminates lack of adult nectar sources as a possibility. The factors that appear to be most significant are low egg-laying frequency, increased longevity, and male spacing.
Low egg-laying frequency was observed on several occasions. The maximum number of eggs seen laid in any one afternoon by the same female was four, these being laid singly and widely dispersed on the foodplant (Callaghan, 1977). The results of this were shown in the low numbers of widely separated larvae in all instars that could be found on the same foodplant. Pupation and emergence were likewise staggered, meaning a fairly even flow of adults over time. Since the larvae were attended and protected by ants (Callaghan, 1977), preda- tion was kept down to a minimum and thus did not appear to be a significant factor as it was in the case of H. ethilla (Ehrlich & Gilbert, 1973). For M. felsina, then, the low frequency of oviposition assures the low intensive frequency of its populations. Why egg production should be low could not be determined during the course of the present study.
Another factor is longevity. The 1.82 weeks observed for M. felsina is high when compared with holarctic butterflies, but low with respect to those tropical nymphalines that have been studied. Scott (1974) reported an average life span of 4.2 days of Lycaena arota Bois with a potential of 8 days. Labine (1968), Turner (1971), and Cook e¢ al. (1971) reported the life expectancy after marking for 6 holarctic species in the field as from 2.8-12.1 days. Powell (1968) gave a maximum of 16 days for Incisalia iroides (Boisduval). On the other hand, Benson & Emmel (1973) demonstrated that the neotropical nymphaline Marpesia berania has an average longevity of 43.9 days with a potential of at least 157. Heliconids are especially long lived. Benson (1972) reported Heliconius erato petiverana Doubleday in Costa Rica with average observed longevities of 52 days and an individual alive 6 months after
VOLUME 32, NUMBER 1 45
marking. Turner (1971) and Ehrlich & Gilbert (1973) reported similar results. Greater longevity ensures low intensive frequency and is of considerable selective value in the tropics since it enables the wide- spread dispersal of eggs in both space and time, which may diminish parasitism and ensure a larger number of offspring reaching maturity (Benson & Emmel, 1973). Also, having individuals in all stages of development would permit survival of a natural disaster that might eliminate one stage but not the others. The causes of adult mortality of M. felsina are not precisely known, since no actual predation or other forms of natural mortality were observed during the study. However, potential predation exists in the form of lizards, spiders, ants, and birds. Rapid flight and hiding beneath leaves are two methods used by adult M. felsina and many other riodinids to avoid predation. On the other hand, its fairly sedentary habits might mean greater predator pressure and lower survival rate for M. felsina than for heliconines and nympha- lines, which enjoy mimicry, distastefulness, and/or strong flight. Lower survival would be more likely true for males because of their conspicu- ous behavior.
Finally, male spacing seems to select for longevity, since older males have been observed to be more aggressive and fixed to their habitual areas and thus are easily able to drive off younger, more inexperienced newcomers. The result is that females appearing at the rendezvous area will be more likely to mate with strong, long-lived males. This is turn assures longer-lived offspring, which can effectively distribute their eggs widely in space and time. These three factors combined, then, might provide for the perpetuation of the low intensive frequencies observed in the M. felsina population. Although to date other forest riodinids have not been studied in such detail, I suspect that their low intensive frequencies may be explained on much the same basis.
Low Extensive Frequencies
Low extensive frequency means that populations are found rarely within a given faunistic region. This is the case with M. felsina as well as other forest riodinids. The key factor here in the case of M. felsina is foodplant distribution. Individuals of Norantea brasiliensis Choisy (Marcgraviaceae), the M. felsina foodplant, are widely and sparingly distributed on the restinga, but this does not prevent their being visited by ovipositing females.
There is a tendency on the part of the newly emerging butterflies to establish a colony at a suitable locality near the site of their foodplant. This was observed on several occasions, the most notable of which was
46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
when two males were seen engaging in patrolling behavior on a small clump of bushes near a foodplant some 500 m from the main colony. These same two individuals were there one week later, a little bit worn but still recognizable. Before the next visit, they had disappeared and no others had taken their place. On the nearby foodplant, larvae were discovered in various instars. I did not observe any other case of actual colonization occurring on that particular clump of bushes. Undoubtedly, this attempt failed as a result of the rather harsh, unsheltered conditions in that particular section of the restinga. Females, then, are the col- onizers for M. felsina, traveling considerable distances many times in search of foodplants on which to lay their eggs.
This phenomenon has been recorded indirectly with other riodinid butterflies. On numerous occasions a single butterfly will be encoun- tered at a particular place and time, whereas subsequent visits under comparable conditions will fail to turn up additional specimens. In the same woods as the M. felsina habitat, a single male Calydna lusca (Geyer) was captured on 18 July 1973. On 7 July 1972, a male and female Nymphidium lisimon attenuatum Stichel were taken and a single female Leucochimona philemon (Cramer) was taken on 7 July 1973. Before and since, 46 collecting days over 4 years have failed to reveal additional examples from this small wood. The best explanation is that the females are very vagile, always on the move searching for new foodplant localities. Similar conclusions have been reached by other students of tropical riodinids. Ebert (1969) mentions that little species such as riodinids “migrate continuously within a great area of favorable biotypes. . ...”
Finally, the question arises concerning the barriers that female rio- dinids will cross in their movement. In the case of M. felsina, open flat areas do not appear to provide a serious obstacle, although this might be expected because of their preference for a low forest habitat. Ob- servations on deep forest riodinids are few but significant. On 19 January 1975, a lone female Nymphidium mantus (Cramer) was ob- served passing through dry secondary shrub near Linhares, Espirito Santo, Brazil, an area very different from its normal habitat in the deep forest near the edges of swamps. Water does not appear to be a sig- nificant barrier since many riodinid populations on either side of large rivers such as the Amazon are virtually indistinguishable (Callaghan, in prep. ).
SUMMARY
The results of the marking-recapture and observations of adult be- havior allow a number of conclusions to be drawn with respect to the
VOLUME 32, NuMBER l AT
population structure of the neotropical riodinid butterfly M. felsina. It was found that this butterfly exhibits the structure common to most forest butterflies, that of low intensive and extensive frequencies. The reasons for the former are low egg-laying frequency, longevity, and male spacing. The latter was explained by a combination of high female vagility and low intensive and extensive foodplant distribution, which lead to the establishment of new colonies by females at widely dispersed foodplant localities. Depending on conditions at these localities, such as parasitism and exposure to the weather, the colony may become extinct only later to be reestablished by another wandering female. It is sug- gested that similar population structures for other neotropical forest butterflies, particularly riodinids, may be explained on this same basis.
ACKNOWLEDGMENTS
I wish to thank Drs. Woodruff Benson and Keith Brown for making many helpful comments on the manuscript and the former especially for his helpful discussion and encouragement during the initial phases of the field work. To Dr. Alfredo de Rego Barros my thanks for use of the facilities of the Museu Nacional do Rio de Janeiro.
LITERATURE CITED
BartEy, N. T. J. 1952. Improvements in the interpretation of recapture data. J. Animal Ecol. 21: 120-127.
Bates, H. W. 1864. The naturalist on the River Amazons. University of Cali- fornia Press (1962 reprint of the second edition, John Murray, London). 465 p.
Benson, W. W. 1972. Natural selection for Miillerian mimicry in the Heliconius erato in Costa Rica. Science 176: 936-939.
Benson, W. W. & T. C. Emmet. 1973. Demography of gregariously roosting populations of the nymphaline butterfly Marpesia berania in Costa Rica. Ecology 54: 326-335.
CaLLAGHAN, C. J. 1977. Studies on restinga butterflies. I. The life cycle and immature biology of Meander felsina (Riodinidae). J. Lepid. Soc. 31: 173-182.
Cook, L. M., K. Frank, & L. P. Brower. 1971. Experiments on the demography of tropical butterflies. I. Survival rate and density in two species of Parides. Biotropica 3: 17-20.
Crane, J. 1955. Imaginal behavior of a Trinidad butterfly Heliconius erato hydara Hewitson, with special reference to the social use of color. Zoologica 40: 167-196.
1957. Imaginal behavior in butterflies of the family Heliconiidae: Chang- ing social patterns and irrelevant actions. Zoologica 42: 135-146.
Esert, H. 1969. On the frequency of butterflies in eastern Brazil, with a list of the butterfly fauna of Pocos de Caldas, Minas Gerais. J. Lepid. Soc. 23: Suppl. 3, 48 p.
EnruicH, P. R. & L. E. Gimperr. 1973. Population structure and dynamics of the tropical butterfly Heliconius ethilla. Biotropica 5: 69-82.
48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fiser, R. A. & E. B. Forp. 1947. The spread of a gene in natural conditions in a colony of the moth Panaxia dominula (L.). Heredity 1: 143-174. LasineE, P. A. 1968. The population biology of Euphydryas editha. VIII. Ovi- position and its relation to patterns of oviposition in other butterflies. Evolu-
tion 22: 799-805.
MacNeri, C. D. 1964. The skippers of the genus Hesperia in western North America with special reference to California (Lepidoptera: Hesperiidae). Univ. California Publ. Ent. 35: 230 p.
PowEL., J. A. 1968. A study of area occupation and mating behavior in Incisalia iroides (Lepidoptera: Lycaenidae). J. N. Y. Ent. Soc. 76: 47-57.
Scott, J. A. 1974. Population biology and adult behavior of Lycaena arota (Lycaenidae). J. Lepid. Soc. 28: 64-72.
Seitz, A. 1913. Macrolepidoptera of the World. Vol. 5. The American Rhopaloc- era. Alfred Keren Verlag, Stuttgart. 1143 p.
SHEPPARD, P. M. & J. A. BisHop. 1973 (1974). The study of populations of Lepidoptera by capture-recapture methods. J. Res. Lepid. 12: 135-144. Turner, J. R. G. 1971. Experiments on the demography of tropical butterflies. II. Longevity and home-range behaviour in Heliconius erato. Biotropica 3:
21-31.
Younc, A. M. 1972. The ecology and ethology of the tropical nymphaline but- terfly, Victorina ephaphus. I. Life cycle and natural history. J. Lepid. Soc. 26: 155-170.
Journal of the Lepidopterists’ Society 32(1), 1978, 48
THE INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE
The draft third edition of the International Code of Zoological Nomenclature is now available for comment by zoologists. Copies may be obtained (price £2.50 surface mail, £5.00 air mail) from the Publications Officer, International Trust for Zoological Nomenclature, c/o British Museum (Natural History ), Cromwell Road, London SW7 5BD, U.K. Comments should be sent as soon as possible, and in any case before 30 November 1978, to the Secretary, International Commission on Zoological Nomenclature, at the above address.
A paper explaining the major changes proposed by the Commission’s Editorial Committee to the existing Code has been published in the Bulletin of Zoological Nomenclature, vol. 34, part 3. Copies may be obtained (price 50p) from the same address as copies of the draft Code.
Journal of the Lepidopterists’ Society 32(1), 1978, 49-54
AN ANALYSIS OF THE HELIOTHIDINE TYPES (NOCTUIDAE) OF HERMAN STRECKER WITH LECTOTYPE DESIGNATIONS
D. F. HAarpwIick
Biosystematics Research Institute, Ottawa, Canada
ABSTRACT. The authenticity of the nominal type specimens of species of Heliothidinae described by Herman Strecker is evaluated. A number of the nom- inal type specimens are judged to be spurious. Lectotypes for the following Strecker species are selected: Schinia approximata, Schinia dolosa, Heliothis fastidiosa, Schinia labe, Schinia lora, and Schinia pyraloides.
In anticipation of future revisionary work on the Heliothidinae I took the opportunity in October of 1976 to examine Herman Strecker’s type material belonging to this subfamily at the Field Museum of Natural History in Chicago. Strecker’s species names have always presented a problem to Lepidopterists, firstly because his original descriptions were often very brief, and secondly because he evidently had the habit of augmenting or replacing his original type series. Thus, although Heliothis regia was described from a single specimen, there are now six specimens each labelled in his hand as “type” of regia.
In the earlier years of his career, Strecker evidently had no type con- cept, or at least a very nebulous one. As a result, many of the specimens on which he based his early original descriptions must have been either destroyed or misplaced. In later years, however, with increasing aware- ness of the value of type specimens, Strecker presumably tried to rectify his earlier laxity by labelling specimens other than the “originals” as his types.
If such substitutions can be demonstrated, then obviously the spurious types have no status under the “Rules.” Nevertheless such pseudotypes do have value in indicating Strecker’s concept of his species in, the maturity of his later years, and should be considered in any subsequent neotype selection procedures if these are found to be necessary. At the present time the Strecker Collection is housed as a separate entity within the collections of the Field Museum of Natural History, and Strecker’s arrangement of species and his hand-printed labels have been retained. In my discussion of type specimens which follows, the species names are arranged alphabetically.
Rhododipsa aden Strecker
Strecker, 1898, p. 11. The original description of aden was based on a single specimen. The male labelled as “original type” in the Strecker collection matches the original descrip-
50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
tion well and is evidently the one on which Strecker based his description. It is labelled as follows: “Col.”; “384”; “S. Aden, Orig. Type’. There is a major piece of the left hind wing broken off and the anal angle of the right front wing is missing. A genitalic slide (no. FM Hel 1) has been prepared from the holotype.
Schinia approximata Strecker
Strecker, 1898, p. 10.
This species was described on the basis of three specimens collected by Boll near Dallas, Texas. The three females labelled as “original types” match the orig- inal description well and are assumed to be authentic. Because of its superior condition the specimen numbered 76 is hereby selected as lectotype; it is labelled as follows: “76”, “S. approximata, 374, Orig. Types”. A genitalic slide (no. FM Hel 2) has been prepared from the lectotype.
Schinia ar Strecker
Strecker, 1898, p. 10.
The single male in the Strecker Collection labelled as “original type” matches the original description well, and is evidently the one on which the name was based. The specimen is labelled as follows: “371”; “S. ar, 371, Orig. Type”. A genitalic slide (no. FM Hel 3) has been prepared from the holotype which is in excellent condition.
Schinia dolosa Strecker
Strecker, 1898, p. 9.
The original description was based on two specimens taken near San Antonio by Boll. The two males in the Strecker Collection labelled as “original types” are evidently authentic. I hereby select the slightly larger specimen as lectotype; it is labelled as follows: “tex”; “S. Dolosa, Orig. Type”. A genitalic slide (no. FM Hel 3) has been prepared from the specimen.
Heliothis fastidiosa Strecker
Strecker, 1876, p. 121.
The original description of fastidiosa was based upon two specimens collected by Boll in Texas. The two males in the Strecker Collection match the original description well and undoubtedly represent the specimens on which it was based. I hereby select the smaller specimen bearing the individual “31” label as lectotype. The specimen is labelled as follows: “31”; “S. Fastidiosa, 31, Orig. Types”. A genitalic slide (no. FM Hel 5) has been prepared from the lectotype.
Heliothis gloriosa Strecker Strecker, 1877, p. 132.
A single specimen in the Strecker Collection is labelled as “original type” and this is evidently the one on which the original description was based. The specimen, a female, is in excellent condition except for lacking a portion of the right antenna. It expands 1%,” and is labelled as follows: “18”; “S. gloriosa Orig. Type”. A genitalic slide (no. FM Hel 6) has been prepared from the holotype.
Schinia hanga Strecker Strecker, 1898, p. 9. The species was described on the basis of one specimen collected by Boll at Dallas, Texas. The male in the Strecker Collection labelled as “original type” is
VOLUME 32, NUMBER l 5li
evidently this specimen. It expands 144” and is labelled as follows: “70”; “393”; “S. Hanga, Orig. Type”. A genitalic slide (no. FM Hel 7) has been prepared from the holotype, which is in excellent condition.
Heliothis imperspicua Strecker
Strecker, 1876, p. 122.
The original description of imperspicua was based upon a single specimen, bear- ing the number 53, which was collected in Texas by Boll. There are two specimens in the Strecker Collection each labelled as “original type” but these are evidently both spurious. One specimen is labelled as having been collected in Colorado; the other is without a locality label and bears the number “49”. Neither specimen differs from the rather generalized original description in any striking detail. The true type of imperspicua must be presumed lost.
Heliothis inclara Strecker
Strecker, 1876, p. 122.
The original description was evidently based upon a single specimen collected by Boll in Texas, which was numbered 46. There are now two specimens in the Strecker Collection labelled as “original types”. One of these bears the number 78 and is considerably smaller than the specimen cited in the original description. The other specimen is without collection number but corresponds well with the original description and may be the true type.
Schinia labe Strecker
Strecker, 1898, p. 10.
The original description of labe was based upon two specimens collected at Dallas, Texas by Boll. The two specimens in the Strecker Collection labelled as “original types” are apparently these. I hereby select the smallest of the two, which bears a separate “372” label, as lectotype. The lectotype is a male ex- panding slightly less than 34” and is labelled as follows: “372”; “S. Labe, 372, Orig. Types’. A genitalic slide (FM Hel 10) has been prepared from the specimen.
Heliothis lanul Strecker
Strecker, 1877, p. 132.
There is a single male in the Strecker Collection labelled as “original type” and this is evidently the specimen on which the original description was based. There is no locality data indicated in the original description nor on the specimen. The holotype is labeled as follows: “85”; “S. Lanul, 85., Orig. Type”. A genitalic slide (no. FM Hel 11) has been prepared from the type.
Schinia lora Strecker
Strecker, 1898, p. 10.
The original description of lora was based on three specimens, two from Boll collected near Dallas, Texas, and one from Heiligbrodt at Bastrop, Texas. Only two specimens in the Strecker Collection are labelled as “original types” and these are apparently authentic. There is another, unlabelled specimen in the collection which may represent the third specimen of the type series. Of the two specimens labelled as “original types” I hereby select the specimen with the separate “73” label as lectotype. The lectotype is a male in generally good condition which bears the following labels “73”; “373”; “S. Lora, 373, Orig. Types”. A genitalic slide (no. FM Hel 12) has been prepared from the lectotype.
52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Schinia neglecta Strecker
Strecker, 1898, p. 10.
The single specimen labelled as “original type” in the Strecker Collection matches the original description well and is evidently the one on which the name neglecta was based. According to the original description the holotype was collected at Loveland, Colorado. The specimen is a female, expands 1”, and bears the following labels: “Col.”; “377”; “S. Neglecta 377., Orig. Type”. A genitalic slide (no. FM Hel 14) has been prepared from the holotype.
Heliothis nubila Strecker
Strecker, 1876, p. 122.
Strecker’s original description of nubila was evidently based on a single specimen taken in Texas by Boll (number 48). There are two specimens in the Strecker Collection labelled as “original types”. Neither of these matches the original de- scription very well, there being no red shading on the underside of the wings, and both are numbered “72” rather than “48”. I consider these specimens to be spurious; the three types must be presumed lost.
Schinia obscurata Strecker
Strecker, 1898, p. 10.
The single specimen labelled as original type of obscurata in the Strecker Collec- tion is obviously the one on which the original description was based. The holotype is in good condition except for having a notch in the left forewing. The female specimen is labelled as follows: “St. Vincent, Pa.”; “378”; “S. Obscurata, 378, Orig. Type’. A genitalic slide (no. FM Hel 15) has been prepared from the specimen.
Schinia pyraloides Strecker
Strecker, 1898, p. 9.
The four specimens on which the original description of pyraloides was based are in the Strecker Collection and labelled as “original types”. The type series was taken at Glenwood Springs, Colorado by Bruce. I hereby select the male specimen with the individual “Col”. label as lectotype. The specimen is in generally good condition except for lacking most of the left antenna and having a slit in the right hind wing. The lectotype is labelled as follows: “Col.”; “S. Pyraloides, Orig. Type, Colorado”. A genitalic slide (no. FM Hel 16) has been prepared from the specimen.
Heliothis regia Strecker
Strecker, 1876, p. 121.
So far as can be determined from the original description, the name regia was based upon a single specimen. There are, however, six specimens in the Strecker Collection labelled as “type”. According to the original description the holotype was taken in Texas by Boll but none of the six nominal “types” bears a locality label. I have compared each of the specimens with the original description and one female matches it very well, and I construe this to be the true type. It is labelled as follows: “S. Regia, Type”. A genitalic slide (no. FM Hel 17) has been prepared from the holotype.
Heliothis rubiginosa Strecker Strecker, 1876, p. 122.
Strecker’s description of rubiginosa was evidently based upon a single specimen taken in Texas by Boll; there are now six specimens, each labelled as “Type” in the
VOLUME 32, NUMBER 1 53
Strecker Collection. None of these, however, bear the “50” label mentioned in the original description. One of them, a male, matches the original description well and may represent the true type but the evidence is insufficient to make a definitive judgement.
Heliothis siren Strecker
Strecker, 1876, p. 122.
Strecker’s original description of siren was evidently based on a single specimen collected by Boll in Texas. There are now two specimens in the Strecker Collection labelled as “original type” but neither of these bears the “45” label mentioned in the original description. One of the two is without number label and the other bears an “80” label. The unnumbered specimen matches the original description well and may represent the true type but there is no way of establishing this with certainty.
Heliothis spectanda Strecker
Strecker, 1876, p. 122.
The original description of spectanda was based upon a single specimen taken in Texas by Boll. When the Strecker Collection was examined, no specimen labelled as type of spectanda was found. There was, however, a specimen in the series of Heliothis virescens bearing the number “52” which was cited in the original de- scription as belonging to the type. The specimen matches the original description very well and I construe it to be the holotype. A genitalic slide (no. FM Hel 20) has been prepared from the specimen which is a female.
Heliothis sulmala Strecker
Strecker, 1878, p. 1862.
The original description of sulmala was based upon a single male taken at Pagosa Springs (Colorado). Strecker evidently mislaid the specimen because it was found in 1939 in a drawer of miscellaneous moths; it lacks Strecker’s charac- teristic type label. The specimen matches the original description very well, how- ever, and I construe it to be the holotype. The specimen is labelled as follows: “Heliothis Sulmala Streck., Pagosa Springs Col., (Orig. Type), McCauley, Found (1939) in a drawer with misc. moths.” A genitalic slide (no. FM Hel 21) has been prepared from the holotype.
Schinia tanena Strecker
Strecker, 1898, p. 10.
Strecker described tanena on the basis of a single specimen taken at Bastrop, Texas by Heiligbrodt. There is a single male in the Strecker Collection labelled as “type” and there is no reason to doubt its authenticity. The specimen is labelled as follows: “tex”; “380”; “S Tanena, 380, Orig. Type”. A genitalic slide (no. FM Hel 22) has been prepared from the holotype.
Schinia ultima Strecker
Strecker, 1876, p. 122.
The original description of ultima was evidently based upon a single specimen taken in Texas by Boll. There are two specimens labelled as “original type” in the Strecker Collection, but both specimens bear the number “71” rather than the “49” indicated in the original description and both specimens differ from the original description in several details. I do not consider either specimen to represent the holotype, and the latter must be presumed lost.
54 JoURNAL OF THE LEPIDOPTERISTS SOCIETY
ACKNOWLEDGMENTS
I appreciate the cordial co-operation of Dr. Rupert Wenzel, Chairman of the Department of Zoology, and Dr. Eric Smith of the Division of Insects, during my visit to the Field Museum of Natural History.
LITERATURE CITED
SrrRECKER, H. 1876. Lepidoptera, Rhopaloceres and Heteroceres indigenous and exotic. Number 13. Reading, Pa. P. 109-124. 1877. Id. Number 14. P. 125-134. . 1878. Lepidoptera in Annual report of the engineers for 1878. Appendix SS. Washington, D.C. P. 1750-1865. . 1898. Lepidoptera, Rhopaloceres and Heteroceres, indigenous and exotic. Supplement 1. Reading, Pa. P. 1-12, pl. 1, 2.
Journal of the Lepidopterists’ Society 32(1), 1978, 54
Letter to the editor:
A Comment on Monarchs and a “Tragedy of the Commons” in Science
When the paper by Urquhart & Urquhart appeared in this journal (1976, Vol. 30: 153-158), I sat down and wrote a letter criticizing the editorial policy of allowing an observation to be published without providing sufficient information to allow verification by other biologists working with Lepidoptera.
While I shared the fear that publication of the exact locale of the Mexican roost would possibly endanger it, I felt that the authors should have at the very least volunteered to disclose the site to responsible qualified scientists researching monarch biology.
Subsequent events have made me regret not sending in my original comment. Incredibly, a scientist of international reputation, Lincoln Brower, was denied the locality information by Professor Urquhart. I do not consider such secrecy to be in the spirit of modern science, nor necessary in this particular instance.
Anyone familiar with Brower’s body of work on the monarch would not question his scientific stature. Anyone who has seen his environmental film on the Connecti- cut River System cannot doubt his sensitivity to ecological problems.
We all respect the effort that Professor Urquhart has put into studying monarch migration. That does not, however, give him territorial rights over monarch roosting areas or free him from the scientific pesponsthality of allowing other scientists to verify his results.
Much of the controversy and ill will which apparently has followed L. Brower’s independent visit to the Mexican monarch roosting area might have been avoided had the study of the monarch proceeded as unselfish science rather than a race for glory in glossy magazines.
In the future I would hope that this journal will insist that authors be willing to disclose their study sites to responsible colleagues.
LAWRENCE E. GILBERT Department of Zoology University of Texas Austin, Texas 78712
Journal of the Lepidopterists’ Society 32(1), 1978, 55-56
ATOPOTHOURES A. BLANCHARD: A SYNONYM OF GOYA RAGONOT (PYRALIDAE)
A. BLANCHARD P.O. Box 20304, Houston, Texas 77025
ABSTRACT. Atopothoures ovaliger A. Blanchard becomes Gaya ovaliger (A. Blanchard ), close to, but different from Goya stictella Hampson.
Karan and Jay Shaffer, my wife and I went collecting, 17-24 May 1977, at the Welder Wildlife Foundation Refuge, near Sinton, Texas. Dr. Shaffer made a special effort to collect Peoriines and was well satisfied with the results of this trip. On their way back home the Shaffers spent an afternoon with us at Houston, so that he could examine my collection of Peoriines. This is when he discovered that what I had un- fortunately described as Atopothoures ovaliger (Blanchard, 1975) should have gone under the genus Goya Ragonot.
The male genitalia of G. ovaliger are extremely close to those of G. stictella Hampson which is not too uncommon in Texas, but the two
Figs. 1-5. Goya: 1-4, stictella: 1, male, Welder Wildlife Refuge, Sinton, San Patricio Co., Texas, 30 June 1975 (U.S.N.M.); 2, male genitalia of same (slide A.B. 3828); 3, male genitalia of another male, same location, same date, (slide A.B. 3827; 4, same enlarged to show gnathos. 5, ovaliger, El] Rancho Cima, Hays & Comal cos., Texas, 29 August 1975, slide A.B. 3826 enlarged to show gnathos.
56 JoURNAL OF THE LEPIDOPTERISTS SOCIETY
species are definitely distinct. The habitus of G. ovaliger (Blanchard, 1975, Figs. 1-4) is quite different from that of G. stictella (this paper, Fig. 1 and Shaffer, 1968, Fig. 23). The differences between their male genitalia are not so obvious. Fig. 2 shows the genitalia of G. stictella prepared in the conventional manner. In Fig. 3 they are prepared in the manner favored by Shaffer (1968, page 3). Figs. 4 (stictella) and 5 (ovaliger) show the enlarged gnathos and the webs or ribs which sup- port its apical process from beneath; this is where the most obvious difference between the two species is to be found. Dr. Shaffer, who had the opportunity to look at many more specimens than I had, gave me the following information: “These ribs are provided in ovaliger with a double row of teeth (two or three to six in each row). In stictella the number of teeth per row varies from zero to two. Counting is com- plicated by the fact that in both species the size of the teeth varies from large and well developed ones to tiny, barely discernible nubbins.”
LITERATURE CITED
Biancuarp, A. 1975. A new phycitine genus and species (Pyraloidea). J. Lepid. Soc. 29: 95-97.
SHAFFER, J. C. 1968. A revision of the Peoriinae and Anerastiinae (Auctorum) of America north of Mexico. Bull. U.S. Nat. Mus. 280, 124 p.
Journal of the Lepidopterists’ Society 32(1), 1978, 56
PROTECTIVE BEHAVIOR IN AMPLYPTERUS GANNASCUS (SPHINGIDAE)
During August 1974, I spent about two weeks collecting Lepidoptera on the grounds of the Inter-American Institute of Agricultural Sciences, approximately 45 km SE of San Jose, Costa Rica (near the town of Turrialba). On two separate occasions I witnessed an interesting behavioral response in the sphingid Amplypterus gannascus (Stoll), which was common in the area. A few gannascus would some- times remain resting high up on the whitewashed walls of the Institute buildings until about 1000, having been attracted to these sites by the lights on the buildings the night before. In two cases it was possible to touch individuals by means of tossing a multi-segmented net about twelve feet long at them. The individuals responded to being touched by releasing their grip and sailing slowly to the ground in a slow spiralling descent, with their wings held rigidly in a swept-back V position. Once on the ground the moths remained passive in spite of being nudged, and only attempted to escape after being seized by hand. The appearance of this behavior was strikingly similar to the appearance of a dead leaf wafting to the ground from a tree, and would seem to be a behavioral adaptation to escape predators by imitating an unappetizing plant fragment.
Jerr Ross, Department of Museum Science, Texas Tech University, Lubbock, Texas 79403.
Journal of the Lepidopterists’ Society 32(1), 1978, 57-58
OENEIS ALBERTA (SATYRIDAE) IN MONTANA
Oeneis alberta Elwes has been taken to date in widely scattered colonies from Alberta, Manitoba and Saskatchewan (alberta), Colorado (oslari Skinner), Arizona (daura (Strecker) and New Mexico (capulinensis Brown). On 19 and 20 May 1976, a series of 21 males and 6 females of alberta was taken by the author from the high grasslands in the Little Snowy Mountains of central Montana, Fergus and Golden Valley counties. This is a new state record for the species in Montana. Habitat of the Golden Valley County colony is pictured in Fig. 1. Elevation is approximately 6500 feet. Specimens from the colony are shown in Fig. 2. The Montana colonies represent the nominate subspecies.
Additional colonies of alberta will probably be discovered in Montana as suitable habitat in areas east of the Continental Divide is explored at the proper time of year. Colonies should also be expected to occur in Wyoming and Utah, though none have been found thus far.
STEVE KoHLER, Montana Department of Natural Resources and Conservation, Division of Forestry, 2705 Spurgin Road, Missoula, Montana 59801.
Fig. 1. Habitat of Oeneis alberta in the Little Snowy Mts., Golden Valley Co., Mont.
58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Mig. 2. Oeneis alberta from the Little Snowy Mts., Golden Valley Co., Mont. (a & c) males, dorsal; (b & d) same, ventral; (e & g) females, dorsal; (f & h) same, ventral. All photos natural size.
Journal of the Lepidopterists’ Society 32(1), 1978, 59-60
NEW OR INTERESTING LEPIDOPTERA RECORDS FROM WESTERN TEXAS
The Panhandle and South Plains areas of Texas have probably received less close attention from lepidopterists than other areas of the state because of the dearth of resident collectors and the greater number of interesting species in other parts of Texas. In view of the relative lack of information on Panhandle-Plains species, it seems worthwhile to publish certain significant records from my collection and the Texas Tech University collection at this time. I report herein one new Texas record (Pieridae) and additional records of species not usually associated with these areas of Texas.
PIERIDAE
Kricogonia lyside (Godart) has previously been reported from the Panhandle- Plains area only in October (Kendall and Freeman 1963, The Butterflies and Skippers of Texas: A Tentative List, Sinton, Texas, 6 p.). The Texas Tech University col- lection contains two males of this species from Lubbock, Texas (Lubbock Co.), both of which are in very good condition. One was collected on 12 July 1970 by D. W. Kiser, and the other on 24 September 1967 by U. Barber. In addition to these specimens, I collected one male and two females in fair condition at Palo Duro Canyon State Park, Randall Co., Texas, on 6 May 1977.
Phoebis agarithe maxima (Neumoegen). Although this species was not cited in Kendall and Freeman’s checklist as having been recorded from the Panhandle-Plains region of Texas, the Texas Tech collection contains two males, in fair condition, from Lubbock, Texas (Lubbock Co.). One was collected on 7 July 1967 by “E J W,” and the other on 16 September 1970 by P. M. Allen. I observed numer- ous males and females of agarithe maxima in Lubbock, Texas throughout September of 1976, and captured a single worn male on 19 September 1976.
Pieris napi (Linnaeus). The Texas Tech collection contains a single perfect male specimen of an undetermined subspecies collected on 17 August 1970, at Canyon, Texas (Randall Co.) by Walt Fournier, a former Tech graduate student. As far as can be determined, this record is a new one for the state of Texas.
LYCAENIDAE
Lycaeides melissa melissa (Edwards). I collected a single perfect male of this species on 31 August 1975 at the Buffalo Springs Lake Recreation Area (4 mi. E Lubbock, Lubbock Co., Texas). This record tends to support the contention by Rickard and Vernon (1975, J. Lepid. Soc.: 150) ihat this heretofore rarely reported species has probably just been overlooked in the past.
NYMPHALIDAE
Chlosyne janais (Drury). A single female of this common neotropical species was collected by me at my residence in Lubbock, Texas, on 11 June 1977. The specimen has badly torn hindwings but is in fair condition otherwise. This species has not previously been reported from the Panhandle-Plains region of Texas.
SATURNIIDAE
Hemileuca hera hera (Warris). Although Douglas C. Ferguson states that hera is “widespread in the West but not known from Texas” (1972, Bombycoidea- Saturniidae in part, p. 106. in R. B. Dominick, et al., The Moths of America north of Mexico, Fascicle 20.2B), the Texas Tech collection contains a single male in very good condition, collected on 16 September 1969 in Dickens, Texas (Dickens Co.) by M. Hughes.
Callosamia promethea (Drury). The Texas Tech collection contains one female,
60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
in fair condition, collected on 2 September 1973 at Junction, Texas (Kimble Co.) by Tech graduate student Sandy M. Benbow. According to Ferguson (p. 235), promethea has not been cited previously as occurring west of Tyler, San Jacinto, and Montgomery counties in eastern Texas. Thus this record suggests a possible range extension of several hundred miles into the Edwards Plateau area. Texas Tech University students and faculty collect annually at a field campus in Junction, so it should be possible to determine if promethea is more than just a stray in the area.
ACKNOWLEDGMENTS
I wish to thank Roy O. Kendall of San Antonio, Texas, for reviewing this paper and confirming the identification of these specimens, and Dr. David E. Foster of the Texas Tech entomology faculty for allowing me to examine material in the Tech entomology collection.
Jerr Ross, Department of Entomology, Texas Tech Uniwersity, Lubbock, Texas 79403.
Journal of the Lepidopterists’ Society 32(1), 1978, 61-62
ERYNNIS BRIZO LACUSTRA AND HESPERIA COLUMBIA IN THE SIERRA NEVADA
Burns (1964, U. C. Publ. Entomol. 37: 1-214) reports no records for Erynnis brizo lacustra Wright for the Sierra Nevada, and MacNeill (1964, U. C. Publ. Entomol. 35: 1-221) lists no records for Hesperia columbia Scudder from there except one female in the AMNH from “Sier. Nev.” Both are indicator species of the coast range serpentine belts north of San Francisco. Until recently, serpentine outcrops have been little collected in the western foothills of the Sierra Nevada of east-central California. Table 1 (next page) lists the new distribution records there.
Sometimes the adults may fly a few miles from their serpentine areas to hilltop: e.g., both hilltop on Rocky Ridge, 1700-1900’, N. of Monticello Dam, Yolo Co., a non-serpentine area composed of Upper Cretaceous marine rocks of the Venado Formation. The nearest serpentine occurs in the extensive Mesozoic ultrabasic in- trusive rocks and the Franciscan Formation some 6 miles to the west. Similarly, Footman Ridge, Mariposa Co., is Paleozoic marine (also the area to the N & E), and to the south is Mesozoic granitic rocks, with no serpentine nearby. The nearest serpentine is found 5 mi. W. as Jurassic-Triassic metavolcanic rocks and 8 mi. SW near Mariposa as Mesozoic ultrabasic intrusive rocks. In the meadows, forests, and canyon immediately adjacent to Footman Ridge on the W & N, neither species has ever been collected.
On 15 May 1970, E. slope Walker Ridge along Brim Grade, c. 1800’, SW of Leesville, Colusa-Lake Co. line, I noticed a female lacustra ovipositing on the terminal growth of a Quercus durata Jepson bush growing on serpentine soil along a roadbank, at 1125. Bums (1964) says “the skipper invariably occurs in direct association with QO. durata, a serpentine obligate” (see Whittaker et al., 1954, Ecology 35: 258-288). However, in some areas, it may also use Quercus dumosa Nutt. which hybridizes with Q. durata and grows in strictly non-serpentine soils (see Forde & Faris, 1962, Evolution 16: 338-347 ).
Heretofore, these skippers were considered more coastal in their California distribution.
OaxkLEy SHIELDS, Department of Entomology, University of California, Davis, California 95616.
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JoURNAL OF THE LEPIDOPTERISTS SOCIETY
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Journal of the Lepidopterists’ Society 32(1), 1978, 63-64
BOOK REVIEWS
THE BUTTERFLIES AND Morus oF HAMPSHIRE AND THE ISLE OF WicHT (being an account of the whole of the Lepidoptera) by B. Goater. 1974. E. W. Classey, Ltd., Faringdon, Oxon, England. xiv + 439 pp. Price: £6.50, postpaid.
Perhaps nothing so authoritative as this book has been written on so small an areas fauna. The book is annotated with published records, manuscript notes and personal observations on all of the butterflies and moths of Hampshire and the Isle of Wight since the beginning of collecting there. Make no mistake, this book is an historical document, and as a record of what is (and was) in the County, and on the island, it is invaluable.
Goater has drawn records from many contemporary sources, and there are some prominent English entomological contributors to the list: such as D. W. Ffennell, John Heath, E. C. Pelham-Clinton, and the Baron C. G. M. de Worms, names well-known in English lepidopterology. This information provides a regional list unlike any we have seen and the coverage is complete through 1972.
Despite the lovely picture of Argynnis paphia on the cover, this is no “coffee table” book. It sticks strictly to business, and those looking for pretty pictures by which to identify British Lepidoptera should be forewarned to stay away from it. As a book of information (isn’t that what we really need? ), it is superb, and from it one can discover when, where and at what time any species of butterfly or moth has been captured in that area, and, if it has been reared, on what foodplant. I suspect that in this alone the book has fulfilled its purpose, and, additionally, it should stimulate the collector in the area to “fill in the blanks”.
The nomenclature used is standard perhaps only to the British, since it is derived from the Kloet and Hincks Checklist of British Insects, part 2, Lepidoptera, 1972. In this treatment, the Hesperioidea and Papilionoidea directly follow the Ptero- phoroidea and precede the Bombycoidea. To a North American rhopalocerist this arrangement will seem strange, even incomprehensible, since there are few other classifications that follow this one. Most schemes place the butterflies and skippers above the Noctuidae, the “top” family in Goater’s system. If you are interested in the butterflies, by the way, look on pp. 214-245. The sphingids may be found on pp. 307-312, and the saturiid (there is only one) on p. 248, while Catocala are on pp. 404-406. This gives a bit of a “road map” to the reader just trying the book for the first time (I confess to a great deal of initial confusion ).
As stated before, don’t buy this book on the basis of the pretty picture on its cover. Neither is this an identification manual. But if you are interested in a superb compendium of what is known about a limited fauna, by all means get the volume. It sets a fine standard, despite a few typographical errors not alluded to here (they happen to everyone! ), for future lists on small faunas.
Lee D. Miter, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580.
BUTTERFLIES OF West MALAYSIA AND SINGAPORE by W. A. Fleming. 1975. E. W. Classey Ltd., Farington, Oxon, England. Vol. 1: vii-x+64 pp., pls. 1-54; vol. 2: vii-x + 92 pp., pls. 55-90. Price: £19.50, postpaid.
This book, effective as it is, is something of an enigma. I find it impossible to rationalize making it in two volumes if the series is only to be sold as a whole, and not broken into separate volumes, if the buyer so desires. The text is identical in both volumes to page 15, so it is only in the plates and the parts following them that the two volumes differ. A little elementary arithmetic shows that if the volumes
64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
were combined into a single one, there would be a very manageable and useful single book of four prefatory pages, 144 text pages and 90 plates. Surely this would have been a better plan. Maybe others will have the problem I did—I would refer to “the” book to identify a Malaysian butterfly, and I almost inevitably selected the wrong volume. The only excuse I can think of for publishing this work in two volumes is economic, and perhaps £19.50 for a single book would put off some buyers, but with the price of books what it is today, I doubt it.
But enough of the complaining. There is a multitude of information in this book, even though the style is such that it takes some acclimation. The nomenclature is up-to-date and applied to the right insects. Species in which the illustrations are not enough for identification are characterized in the text, and in those instances where genitalic dissection is necessary for final determination, the fact is noted, even though the genitalia are not figured. If Fleming had included some biblio- graphic citations to these problem areas, and to the many included foodplants records, the book would have been more authoritative, and the space these refer- ences would have added could not have been that much.
The illustrations, however, are where the books truly excel. All of the photographs of specimens illustrate the salient points well and facilitate the identification of the insects in question. Al] of the specimens used are not perfect; some are downright tatty, such as the illustrated female of S12, Lethe europa malaya Corbet on Plate 24, but these were the best specimens available in collecions, and the photographs mercifully have not been “prettied up”. The color fidelity is very high, and at least most of the specimens are fresh, rather than century-old museum relics. Iden- tification of even the difficult Malaysian lycaenids is facilitated by them, though of course it is not made simple—no book could achieve that!
I particularly appreciated the accurate citation of the authors of various taxa, even though these names were not bracketed where appropriate. At long last, both of the Felders are cited as the authors of names proposed in the “Reise Novara’, not just a blanket “Felder”. This latter practice seems to have dated from “Seitz” where only Cajetan Felder was given credit for the descriptions in the work, even though the authors themselves cited “nobis” on every new name, rather than the singular “mihi”.
On balance, this is an exceilent book, the foregoing criticisms notwithstanding, and one that is remarkably free of typographical errors. The text portions are per- haps a bit too abbreviated, and authority is not given for many statements. I personally would have preferred a single volume about the size of Corbett and Pendlebury’s The Butterflies of the Malay Peninsula, but the present book accom- plishes some things that the earlier authors could not: Fleming has made the identification of Malaysian butterflies considerably easier than before. No more can be asked of any author! If your interests lie in the butterflies of southeastern Asia, by all means buy this book.
Lee D. Mitier, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580.
EDITORIAL STAFF OF THE JOURNAL Austin P. Puatr, Editor
Department of Biological Sciences University of Maryland Baltimore County, 5401 Wilkens Avenue Catonsville, Maryland 21228 U.S.A. Dovuctas C. Frercuson, Associate Editor THEODORE D. SARGENT, Associate Editor
NOTICE TO CONTRIBUTORS
Contributions to the Journal may deal with any aspect of the collection and study of Lepidoptera. Contributors should prepare manuscripts according to the following instructions.
Abstract: A brief abstract should precede the text of all articles.
Text: Manuscripts should be submitted in duplicate, and must be typewritten, entirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch paper. Titles should be explicit and descriptive of the article’s content, including the family name of the subject, but must be kept as short as possible. The first men- tion of a plant or animal in the text should include the full scientific name, with authors of zoological names. Insect measurements should be given in metric units; times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only where italics are intended. References to footnotes should be num- bered consecutively, and the footnotes typed on a separate sheet.
Literature Cited: References in the text of articles should be given as, Sheppard (1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically under the heading LrreRATuRE Crrep, in the following format:
SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 209 p.
196la. Some contributions to population genetics resulting from the study of the Lepidoptera. Adv. Genet. 10: 165-216.
In the case of general notes, references should be given in the text as, Sheppard (1961, Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. Roy. Entomol. Soc. London 1: 23-30).
Illustrations: All photographs and drawings should be mounted on stiff, white backing, arranged in the desired format, allowing (with particular regard to lettering ) for reduction to their final width (usually 4% inches). Illustrations larger than 8% x 11 inches are not acceptable and should be reduced photographically to that size or smaller. The author’s name, figure numbers as cited in the text, and an indication of the article’s title should be printed on the back of each mounted plate, Figures, both line drawings and halftones (photographs), should be numbered consecutively in Arabic numerals. The term “plate” should not be employed. Figure legends must be typewritten, double-spaced, on a separate sheet (not attached to the illustrations), headed ExpLANATION OF FicuRES, with a separate paragraph devoted to each page of illustrations.
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CONTENTS
Types OF PARNASSIUS CLODIUS GALLATINUS (PAPILIONIDAE). Steve
Bove oO Ses PS CON La aaa i 1 SPECIFICITY, GEOGRAPHIC DISTRIBUTIONS, AND FOODPLANT DIVERSITY IN Four CALLOPHRYS (MITOURA) (LYCAENIDAE). Kurt Johnson 3 FooppLANT, Hapirat, AND RANGE OF CELASTRINA EBENINA (LyY- CAENIDAE). Warren H. Wagner, Jr. and T. Lawrence Mellichamp 20 Strupies ON Restinca Butrerruises. IJ. NoTes ON THE POPULATION STRUCTURE OF MENANDER FELSINA (RiopINDAE). Curtis J. Callaghan, 2 37 AN ANALYSIS OF THE HELIOTHIDINE TyPEs (NocrumAE) OF HERMAN STRECKER WITH LEcToTyPE DesicNaATIoNs. D.F. Hardwick... 49 ATOPOTHOURES A. BLANCHARD: A SYNONYM OF GOYA RAGONOT (Pyratmar). A. Blanchard 0. ee 50 GENERAL NOTES Protective behavior in Amplypterus gannascus (Sphingidae). Jeff Robb 56 Oeneis alberta (Satyridae) in Montana. Steve Kohler __.. 57 New or interesting Lepidoptera records from Western Texas. Jeff Robb 59 Erynnis brizo lacustra and Hesperia columbia in the Sierra Nevada. Oakley Shields i000 ee 61 Noes) AND News 2220230 ies a 19, 48 Book (REVIEWS. i eT aa a 63
Volume 32 1978 Number 2
JOURNAL
of the
LEPIDOPTERISTS’ SOCIETY
Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS
19 July 1978
THE LEPIDOPTERISTS’ SOCIETY
EXECUTIVE COUNCIL
J. W. TuwpeNn, President KENELM W. Puut.ip, Vice President I. F. B. Common, Ist Vice President JuLtian P. DoNnAHUE, Secretary Lionet Hiccrs, Vice President RONALD LEUSCHNER, Treasurer
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Cover illustration: Dasychira dorsipennata larva, dorsal and lateral views. From Fascicle 22.2, “Lymantriidae,” by Douglas C. Ferguson, in Moths of America North of Mexico. The drawing was done by E. R. Hodges, Scientific Illustrator, Department of Entomology, Smithsonian Institution. (Reproduced by permission of the author. )
J OURNAL OF
Tue LeprpopreRists’ SOCIETY
Volume 32 1978 Number 2
Journal of the Lepidopterists’ Society 32(2), 1978, 65-74
STUDIES ON THE INTERACTIONS OF MORPHO PELEIDES (MORPHIDAE) WITH LEGUMINOSAE
ALLEN M. YouNG
Invertebrate Division, Milwaukee Public Museum Milwaukee, Wisconsin 53233
ABSTRACT. The butterfly Morpho peleides Kollar is a widespread species throughout tropical America, exploiting several wild genera and species of Leguminosae as larval foodplants. Field studies show that this species feeds on a broad spectrum of wild legumes on a regional basis. This interaction was explored in the laboratory by rearing caterpillars on peanut plants and alfalfa, cultivated legumes. The life cycle is completed successfully on these artificial foodplants, but feeding on alfalfa taken from an expressway led to mass mortality of caterpillars. Apparently the alfalfa was contaminated from some environmental source. In the native habitats of this butterfly, the Leguminosae are both diverse and numerous locally. This suggests that the mo- nophagous feeding habit provides sufficient ecological flexibility for exploiting different genera and species of the family. This is sufficient to maintain breeding populations of M. peleides in secondary habitats. Forest-dwelling species of Morpho are predicted to be experiencing different types of selection pressures favoring polyphagous feeding.
In the premontane tropical wet forest life zone (Tosi 1969) of north- eastern Costa Rica, a larval foodplant of the butterfly Morpho peleides Kollar (Lepidoptera: Morphidae) is the vine Machaerium aff. flori- bundum Benth. (Leguminosae). The vine and butterfly occur in stands of mixed primary and secondary tropical wet forest (Fig. 1). It is known that M. peleides utilizes several leguminous woody vines and trees as larval foodplants in Costa Rica (Young and Muyshondt 1973) and the species can be reared on commercially available peanut plants both in Costa Rica and Wisconsin (Young 1974). This present paper examines further the feeding habits of M. peleides larvae, using eggs obtained from a population in premontane tropical wet forest (rather than from montane forest, as in a previous study), and involves plants not studied previously (Young 1974). The results further support the assumption
66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Mig. 1. Above: mixed primary-secondary tropical wet forest habitat of the butterfly Morpho peleides at Finca El Tigre, near La Virgen, Heredia Province (Sarapiqui region), Costa Rica. At the spot shown here, M. peleides and M. granadensis are microsympatric. Below: Machaerium aff. floribundum, a forest leguminous vine which is a larval food plant of M. peleides (and probably M. granadensis) in the wild.
VOLUME 32, NUMBER 2 67
that larvae of M. peleides are monophagous feeders on many temperate and tropical genera of Leguminosae.
METHODS
A female of M. peleides was captured on bait of rotting bananas placed near the edge of a forest habitat (Fig. 1) on February 14, 1977 at “Finca El] Tigre,’ a farm adjacent to “Finca La Tirimbina,” a few km from La Virgen, Heredia Province (Sarapiqui region), Costa Rica (220 m elev.). She was placed in a clear plastic bag containing a fresh cutting of M. aff. floribundum, and within eight days had produced a total of 40 viable eggs. The female was then preserved, and the eggs were brought to Milwaukee, Wisconsin for rearing. The eggs began to hatch on February 25 in Costa Rica and by the time the morphos arrived in Wisconsin, they were all Ist instar. In Costa Rica these larvae were fed leaves of Dioclea wilsoni (Leguminosae) but they were switched to peanuts (Arachis hypogea L.—Leguminosae) upon arrival at the Milwaukee Public Museum. The larvae were kept on potted peanut plants placed in a covered glass tank in a laboratory. A growth light was kept over this rearing chamber. The rearing program in Wisconsin extended from March 3 through May 25, 1977 (the date of the late eclosion). Records were kept on body lengths and head capsule widths of all caterpillars. The sources of peanut plants used were (1) Olds Seeds from Madison, Wisconsin and (2) Crop Science Department of North Carolina State University (Raleigh). Near the end of the experiment (April 22), the foodplant was switched to alfalfa (Medicago sativa L—Leguminosae); at this time most of the larvae were in the late 4th instar. The alfalfa plants used were obtained from a farm in Waukesha County, Wisconsin. Later (May 2) the remaining 5th instar larvae (several had pupated) were fed alfalfa collected from the side of an expressway in downtown Milwaukee. Like the peanuts, the alfalfa plants were potted, but this time soil brought in with the plants from the field was used. One 4th instar caterpillar was offered a seedling of Erythrina crista-galli L. (Leguminosae) from Brazil. Records were kept on larval survival throughout the study. The adults obtained were kept for further examination.
RESULTS
Both young and older larvae of M. peleides ted successfully on peanut and alfalfa leaves in the laboratory, followed by normal eclosion (Fig. 2). In addition, at least the 4th and 5th instar larvae will feed on Erythrina. Although some caterpillars feed intermittently throughout the day, the
68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fig. 2. Right column: second and fourth instar catepillars of M. peleides on peanut leaves at the Milwaukee Public Museum; note eaten areas of leaf in first illustration. Leaf column: fifth instar caterpillar on peanut plant and freshly-enclosed adult clinging to empty pupa case (at Milwaukee Public Museum ).
VOLUME 32, NUMBER 2 69
MEAN AND RANGE OF BODY LENGTH (MM) OF CATERPILLARS
20
3 . . 16
14
tt
ae
MAR. 8 MAY 2 (DAY 1) SUCCESSIVE DAYS OF OBSERVATION (DAY 38)
Fig. 3. Growth and size (body length) patterns for Morpho caterpillars reared in the laboratory. The vertical lines give the range in body lengths.
greatest amount of feeding occurred in the late afternoon and early morning (e.g., 16:00-19:30 hrs/C.S.T.). Fourth and 5th instar larvae rested on the rims of the pots containing the peanut plants, and they would crawl up the plants to feed. Younger ones rested on leaves and shoots.
Survival both on peanuts and on “farm alfalfa” was 100%. However, larvae fed “expressway alfalfa” showed considerable mortality: between May 4 and May 12, the number of healthy caterpillars dropped from 32 to 13. Very shortly after being fed the expressway alfalfa, many died. Death was preceded by a drastic contraction of body length, and spasmic
70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TasLe 1. Head capsule size statistics for caterpillars of the tropical butterfly Morpho peleides.
No. Mean head capsule Range of head Instar measured width (x = S.D.) capsule widths (mm ) 1 17 1.50 + 0.08 1.4-1.5 2, 34 2.19 + 0.09 2.0—2.3 3 315} BA ae Vpllc 3.3-3.8 4 oll 4,91 +0.11 A4.7-5.1 5 Ia 6.09 + 0.45 5.2-6.7
Note—data obtained from molted head capsules at the end of each instar [distorted (crushed) head capsules are not included in the calculations]. All head capsules were collected within 24 hours after each molt.
waves of movements lengthwise. Afflicted caterpillars fell from the plants and wriggled on the bottom of the cage before dying. When attempts were made to replace them on the foodplant, they again lost hold and fell off. Their feces had a reddish-orange component that was quickly absorbed on paper toweling. A few died as prepupae.
Initial signs of the affliction included a larva remaining stationary on the plant, with the anterior half of the body hanging off to one side. Such a state lasted a few days before the larva fell off the plant and died. As a result of this sudden and epidemic-like mortality, only those indi- viduals that had already pupated by May 2 survived to emerge as adults. A total of 12 adults, all males, were obtained. Thus, the mortality was 100% among the 19 individuals that were still larvae on May 4.
Based on a sample of ten randomly chosen larvae, molting is not syn- chronous. The 5th instar is the longest, and despite changes in the — foodplant type, development proceeds without major accelerations or decelerations in the daily pattern of growth (Fig. 3). With the exception of the Ist instar, there is considerable variation in the body length and the magnitude of this variation is about the same for the four instars (Table 1). Excluding a small bias introduced by unequal sample sizes, the range of variation in head capsule width is very low for all instars (Table 1). Body length and head capsule width are used here as estimates of larval size. The total development time of 91 days is broken down as follows: (1) egg = 11 days; (2) caterpillar = 65 days; (3) pupa = 15 days. The right forewing length for the adult male butterflies ranged from 55 to 65 mm (N = 12), but the exclusion of two individuals reduced this range to 61-65 mm.
DIscUSSION
The data are useful for discussing (1) feeding behavior of the cater- pillars of M. peleides, and (2) apparent effects of environmental con-
VoLUME 32, NUMBER 2 fol
tamination of a foodplant of an exotic butterfly. The latter was an unexpected outcome of the study.
Morpho accepts peanuts and other legumes, not used as foodplants in the wild, as discussed previously (Young 1974). But to these records I add alfalfa and Erythrina as acceptable foodplants of M. peleides cater- pillars in the laboratory. Erythrina is native to the New World tropics (Bailey 1969), although it is not known if it is a natural foodplant of Morpho. Greenhouse cultures of this plant are usually infested with herbivorous insects, suggesting few effective defenses operative against such attacks. Alfalfa, a near relative of peanut, is commonly cultivated, and occurs as a weed species along roadsides; it is native to the Old World. As a weed species, alfalfa may possess few defenses against herbivores as energy allocation is likely to be directed toward high repro- ductive potential (Lewontin 1965). Thus, cultivars such as peanut and alfalfa, weeds such as alfalfa, and ornamentals, such as Erythrina, are examples of leguminous plant species with few defenses against her- bivores, perhaps making them ideal to serve as food for Morpho larvae. It is not known if Morpho will oviposit on these plants in the laboratory.
I observed previously that caterpillars of M. peleides are primarily “dawn-dusk” feeders in the wild (Young 1972a), and this is also true for laboratory cultures experiencing the Wisconsin dawn-dusk cycle (Young 1974; pers. obs.). Apparently the larvae are programmed with a rhythmicity for peak periods of feeding activity in both tropical and temperate situations. In the wild, 4th and 5th instar caterpillars rest on the trunks of the foodplant where they blend in with the background (Young 1972a), and this behavior explains why they rested on the rims of the pots containing the peanut plants.
Using a larger sample size, Young (1974) estimated the total develop- mental time for M. peleides on peanuts to be about 105 days, or about 14 days longer than the estimate obtained in the present study. Body length of 5th instars in the previous study, was about 73 mm as com- pared to 70-71 mm in the present study. Several factors may be relevant here: (1) the eggs in the two studies came from different regions. Thus selection pressures could have been different in terms of effects on development time; (2) differences in the foodplant as related to time of the year, and other factors. The discrepancy in the development time is in the length of the larval stage; the egg and pupal stages are the same (Young 1974; pers. obs.). Thus, differences in the foodplant may be involved. Switching to alfalfa (not done in the previous study ) might have accelerated development. To test this, eggs will have to be reared entirely on this plant in a future study. The transfer to alfalfa was done
—~l bo
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
in the fifth instar, the time of greatest food intake. In the previous study, older peanut plants were used, and it may well be that older plants have resistant properties more expressed than younger ones. There is some evidence that the defense systems of peanut plants change with age: in the present study I observed that Morpho larvae refuse to eat the first set of leaves of a peanut seedling, eating only shoot and leaf tissue above this point (S. Borkin and A. Young, pers. obs.).
The observed high level of mortality among 5th instar larvae on alfalfa plants taken from the downtown expressway may have been due to a contamination of these particular plants by an industrial or automobile residue. An exotic insect such as Morpho, when exposed to a con- taminated foodplant, may be expected to encounter such mortality. Since only male adults were reared I presume that all the females died as Sth instar larvae, since M. peleides has a sex ratio of unity (Young 1972b, 1973; Young and Muyshondt 1973). Female larval development generally takes longer than that of males. The observed inability of afflicted larvae to grasp the foodplant and feed was very likely due to the contaminant affecting the nervous system. It is not likely that a strain difference affecting feeding ability by morphos exists between the expressway alfalfa and farm alfalfa, since larvae did eat the former and until the time of obvious signs of illness, their feeding behavior appeared normal.
The data indicate that feeding flexibility of M. peleides caterpillars is considerable in the sense that it allows this monophagous tropical her- bivore to exploit a broad range of genera and species locally. Foodplant records for M. peleides from Costa Rica and El Salvador indicate that many different wild Leguminosae are used, and allied South American species exhibit similar behavioral flexibility (Otero 1971; pers. comm. ). Secondary forest habitats in the wetter regions of Central America locally support a wealth of leguminous vine, shrub, and tree species, many of which are used by M. peleides. It is therefore not surprising that cater- pillars will feed successfully on allied legumes not used as natural food- plants, including cultivated forms such as peanuts and alfalfa. On a per unit area basis, secondary habitats in the tropics support large patches of Mucuna, Dioclea, Machaerium, Inga, etc. and many patches may occur locally. Thus, in terms of larval foodplants, the environment is very certain or predictable creating selection pressures favoring eco- logical specialization such as monophagy. As foodplant patches increase in number and size in an area, monophagy is considered an optimal feeding strategy for a herbivore (Levins and MacArthur 1969).
Some species of Morpho that live in the canopy of primary tropical forest deposit eggs on different families of trees and woody vines (Otero,
VoLUME 32, NUMBER 2 73
pers. comm.) and these species may be polyphagous. Miller (1968) lists several Morpho foodplant families, although the degree to which each species oviposits on more than one family has not been determined. Polyphagy in Morpho is adaptive in habitats where individuals of each foodplant species are greatly dispersed over large areas, making it ener- getically difficult to exploit a single family of foodplants. In forest hab- itats where each foodplant species is greatly dispersed over large areas, the environment is less certain in terms of a female butterfly locating suc- cessfully an individual of that particular plant. As more plants are added to the local foodplant niche, the environment becomes more certain; the incorporation of additional local foodplants implies the evolution of polyphagy, since member genera and species of individual plant families in tropical forests are greatly dispersed over large areas. Thus, although M. peleides and its near allies such as M. achilles may exhibit considerable feeding flexibility within the Leguminosae, they are monophagous species; polyphagous species of Morpho are expected to occur in primary forests. These include such likely candidates as M. amathonte, theseus, grana- densis, and cypris in the Central American rain forests.
ACKNOWLEDGMENTS
This research would not have been possible without the financial support of the Friends of the Museum (of the Milwaukee Public Museum) and James R. Neidhoefer. Susan Borkin and Joan Jass ( Mil- waukee Public Museum) conducted the rearing studies and assisted with taking measurements. Dr. J. Robert Hunter allowed me to work at Finca El Tigre and Dr. Ridgway Satterthwaite of the Associated Colleges of the Midwest provided logistical assistance in various ways. Luis D. Gomez of the Museo Nacional de Costa Rica provided a field vehicle. Janice Mahlberg of the Milwaukee Public Museum provided photo- graphic assistance. The assistance of Dr. Martyn Dibben and Neil Luebke with growing peanut plants at the museum is greatly appreciated. I also thank Joe Sugg, head of the North Carolina Peanut Council and Dr. Harry Cobel (Crop Science, North Carolina State University) for arranging peanut plants to be sent to me when the museum supply was defoliated. I thank Cheryl Castelli for typing the manuscript. To all of these people I am very grateful.
LITERATURE CITED
Baitey, L. H. 1969. Manual of cultivated plants. MacMillan Co., Toronto.
Levins, R. & R. MacArruur. 1969. An hypothesis to explain the incidence of monophagy. Ecology 50: 910-911.
74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Lewontin, R. C. 1965. Selection for colonizing ability. In The Genetics of Colo-
nizing Species (H. G. Baker and G. Stebbins, eds.). Academic Press, New York, 2 11-94.
Rien. L. D. 1968. The higher classification, phylogeny and zoogeography of the Satyridae (Lepidoptera). Mem. Amer. Entomol. Soc., No. 24, 174 pp.
Orero, L. S. 1971. Instrucoes para criacao da borboleta “Capitao-do-mato” (Morpho achillaena) e outras especies do genero Morpho (“Azul-seda’, “Boia’, “Azulao-branco”, “Praia-grande”). Inst. Brasileiro Desenvolv. Florestal, Rio de Janeiro, 27 p.
Tost, J. A., Jr. 1969. Mapa ecologico. Republica de Costa Rica. Centro Cientifico Tropical, San Jose, Costa Rica.
Younc, A. M. 1972a. Adaptive strategies of feeding and predator-avoidance in the larvae of the neotropical butterfly Morpho peleides limpida (Lepidoptera: Morphidae). J. New York Entomol. Soc. 80: 66-82.
. 1972b. Community ecology of some tropical rain forest butterflies. Amer.
Midl. Nat. 87: 146—157.
. 1973. The comparative ethology and ecology of several species of Morpho
butterflies in Costa Rica. Studies Neotrop. Fauna 8: 17-50.
1974. The rearing of Morpho peleides (Morphinae) on peanuts. J. Lepid. Soc. 28: 90-99.
Younc, A. M. & A. MuysHonpt. 1973. The biology of Morpho peleides in Central America. Carib. J. Sci. 13: 1-49.
Journal of the Lepidopterists’ Society 32(2), 1978, 75-85
NOTES AND DESCRIPTIONS OF EUPTYCHIINI (LEPIDOPTERA: SATYRIDAE) FROM THE MEXICAN REGION
Ler D. MILLER Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580
ABSTRACT. Several species of Euptychiini (Lepidoptera: Satyridae) are dis- cussed and/or described. Described as new: Taygetis mermeria griseomarginata (Guerrero, Mexico), Splendeuptychia kendalli (Tamaulipas, Mexico), and Cyllopsis wellingi (Cayo dist., British Honduras). The previously unknown female of Cyllopsis dospassosi L. Miller is described and figured.
Some years ago Mr. Roy O. Kendall of San Antonio, Texas sent me a strange euptychiine satyrid from northern Mexico for identification. It was apparent that the specimen was a representative of a new species in the genus Splendeuptychia Forster (1964), a group hitherto known from no further north than Panama. Since the Panamanian species, S. salvini (Butler), was unrepresented in the Allyn Museum collection, I compared the Mexican insect with the colored figures of salvini given by Butler (1866) and by Godman and Salvin (1880 [1879-1901]). Many dis- crepancies between the two insects became obvious, and a hurried call to Mr. Gordon B. Small, Jr. of Balboa, Canal Zone resulted in his sending two males of S. salvini that confirmed the superficial differences between it and the Mexican butterfly as well as genitalic ones.
Once Mr. Kendall and his wife had managed to rear the Mexican Splendeuptychia they needed a name on which to base the paper that follows. Accordingly, I am taking this opportunity to describe the new Splendeuptychia and some other euptychiines from the Mexican region. Additional data are given on species that have come to my attention since the publication of portions of my revision of the tribe. Some of the new species described herein are members of genera not covered in the revision yet, but it is felt that publication of these parts may be so far in the future that other workers could benefit by having the names proposed at this time.
Taygetis mermeria griseomarginata L. Miller, new subspecies Figs. 1-5
Male: Head, thorax and abdomen clothed with dark brown dorsal and tan to reddish-tan ventral hairs. Palpi reddish-tan, somewhat darker laterad. Antennae dark brown dorsad, reddish-brown checkered with tan ventrad. Legs clothed with brown hairs laterad, reddish-tan ones on inner portions of segments.
Wings with acutely falcate forewing apices as in T. mermeria excavata Butler (1868). Upper surfaces of wings dark, rich brown, unmarked except for broad (4-8 mm) grayish overscaling along margins of all wings and more narrowly and
76 JouRNAL OF THE LEPIDOPTERISTS SOCIETY
1.0 mm
Figs. 1-5. Taygetis mermeria griseomarginata, n. ssp. 1-2, Holotype ¢ upper (1) and under (2) surfaces; MEXICO: GUERRERO: Acahuizotla (Allyn Mus. photos 101476-7/8); LFW (length of forewing) 47.0 mm. 3-4, Paratype 9 upper (3) and under (4) surfaces; MEXICO: COLIMA: Comala (Allyn Mus. photos 101476-9/10); LFW 55.2 mm. 5, @ genitalia of Holotype; slide M-2732 (Lee D. Miller).
VoLUME 32, NuMBER 2 Wale
less prominently along forewing costa. Under surfaces of all wings mottled in vari- ous shades of brown, reddish-brown grayish-tan or ochreous (highly variable indi- vidually) with forewing mesial bands poorly developed and only the extradiscal bands of the hindwings well developed (usually delimited by some gray-green scaling distad of the dark brown bands themselves); ocelli of both wings varying from very well developed to obsolescent. Fringes of all wings gray above, tan to reddish-brown below.
6 genitalia similar to those of other Mexican specimens (excavata) with some- what stubbier valvae than those of South American representatives.
Length of forewing of Holotype ¢ 47.0 mm, those of the 21 ¢ Paratypes rang- ing from 44.6 to 53.3 mm, averaging 50.35 mm.
Female: Similar in appearance to the ¢, differing chiefly in the paler coloration both dorsally and ventrally and by the presence of a poorly defined transcellular band of the hindwings beneath that is not shown by the ¢.
Lengths of the forewings of the seven 2 Paratypes range from 54.1 to 58.0 mm, averaging 55.95 mm.
Described from 29 specimens, 22 males and seven females, from the western slope of the Sierra Madre Occidental, Mexico.
Holotype ¢@: Mexico: Guerrero: Acahuizotla, ix.1957 (T. Escalante); @ genitalia slide M-2732 (Lee D. Miller).
Paratypes: all Mexico. GuERRERO: same locality as Holotype, 14 viii.1957, 5 4 ix.1957, 16 x.1957, 12 viii.1958, 12 iii.1958 (all T. Escalante); Tierra Colorado, 126 42 viii-ix.1971 (all A. Diaz Frances). Nayarir: vic. Compostela, 14 Posen bo Kiots). Comma: Colima, 14 11.1.1968; Comala, 14 31.x.1967, 12 14.41.1968 (all R. Wind).
Disposition of type-series: Holotype ¢, 17é¢ and seven @ Paratypes in Allyn Museum of Entomology; single ¢ Paratypes will be placed in the American Museum of Natural History, the National Museum of Natural History, Carnegie Museum and the British Museum (Natural History ).
The name of this subspecies refers to the broadly gray dusted margins of both wings on the upper surfaces. This situation is only hinted at in specimens of T. m. excavata in which the maximum development of this marginal gray scaling is about 1-1.5 mm on the forewing and virtually absent on the hindwing. In the present subspecies this gray marginal scaling is most prominent on the hindwing, but the forewing scaling is more extensive than on any excavata specimen.
This gray-margined subspecies is apparently restricted to the western slopes of the Sierra Madre Occidental from at least Nayarit to Guerrero. While I have not seen material from all of the states in this area, I feel confident that griseomarginata will be found in Jalisco, Michoacan and possibly southernmost Sinaloa. A single specimen in the Allyn Museum collection from Chiapas (Tuxtla Gutierrez, 13.viii.1961, leg. “M. S.”) that was part of the Jae collection is referable to griseomarginata, but since all material from Oaxaca and Chiapas that I have seen has been referable to only excavata, I have excluded this Chiapas specimen from the type-series. It may have been mislabelled, or it may represent a genetic “throwback”, but it certainly is not typical of Chiapas-Oaxaca before me. All of the specimens I have seen from the Nayarit to Guerrero
78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
range have been referable to griseomarginata, and the presence of a single specimen from outside this range should not be taken as “proof” that the subspecies does not exist.
Splendeuptychia salvini (Butler), 1866 Figs. 6-8
Euptychia salvini Butler, 1866: 498 (type locality: Lion Hill, [Canal Zone], “Panama” ).
Forster (1964: 128 ff.) erected the genus Splendeuptychia for 23 Neo- tropical species, most of which are restricted to South America. The species included in the present genus are among the loveliest of the Euptychiini, and their pattern is unmistakable. Only S. salvini has thus far been reported from Central America, and it is restricted to the Canal Zone and adjacent Panama, possibly as far south as the Darien. S. salvini seems to be a rare butterfly, at least in collections. I suspect that this appearance of rarity is real, since the insect is one of the more spectacular Satyridae and should not be overlooked by even a casual collector.
Two males were obtained from Gordon B. Small, Jr. for examination and comparison with the Mexican species described below. The differ- ences are cited under the new species, but it suffices to say here that the two are not conspecific.
Splendeuptychia kendalli L. Miller, new species Figs. 9-13
Male: Head, thorax and abdomen clothed with gray-brown dorsal and ochreous- tan ventral hairs. Palpi pale gray laterad and dark gray ventrad and dorsad. An- tennae brown dorsad, reddish-brown ventrad; tip of club slightly darker. Legs clothed with gray hairs, but those of tarsi tan.
Wings above dull brown with three dark brown marginal lines separated by tan; otherwise unmarked, but the markings of the under surface showing through vaguely on this surface. Forewings below gray-brown in proximal half, tan in distal half; two thickened rust-brown lines, one across cell, the other just outside cell dividing the gray-brown from the tan ground color; three almost straight dark brown marginal lines; between the marginal lines and the distal band is a row of black-edged silver spots from M,—M:. to Cu:—2A, the whole spothand surrounded by a thin brown ring. Hindwings below with gray-brown proximal and tan distal ground color; thickened rust-brown bands of forewing continued on hindwing; three thin, dark brown marginal lines following the slightly crenulate wing outline; a mesial to submarginal yellow patch from Rs—M; to Cu.-2A encompassing silvered spots in the interspaces, those in Rs—M:, M.-M2 and Cu,-Cu: with well defined black irides; a subsidiary black line between the marginal lines and the yellow patch from Cu, to the tornus; along M, and Cu; are two black submarginal patches. Fringes pale gray above, tan below.
genitalia as figured, differing from those of S. salvini (Fig. 8) in many re- spects, especially the shorter gnathos arms and the simpler valvae.
Length of forewing of Holotype ¢ 17.8 mm, those of the 374 Paratypes rang- ing from 16.7 to 18.8 mm, averaging 17.67 mm.
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0.5 mm
Figs. 6-8. Splendeuptychia salvini (Butler). 6-7, ¢ upper (6) and under (7) surfaces; PANAMA: PANAMA: Bayano, nr. Pina (Allyn Mus. photos 010677— 11/12); LFW 15.9 mm; G. B. Small, Jr. collection. 8, 4 genitalia of same speci- men; slide M-3418 (Lee D. Miller).
Female: Very similar to the ¢, differing chiefly in size, slightly paler coloration and the more extensive yellow patches of the hindwing under surface.
Lengths of forewings of the 36 2 Paratypes range from 17.3 to 21.0 mm, averag- ing 18.84 mm.
Described from 74 specimens, 38 males and 36 females, from the Mexican states of Tamaulipas and San Luis Potosi.
Holotype ¢: Mexico: Tamauuipas: Gonzalez Ranch, nr. Los Kikos, ex ovum on Bambusa aculeata, emerged 9.i.1975 (R. O. and C. A. Kendall); chromosome specimen no. 3A-32-M; ¢ genitalia slide M-3651 (Lee D. Miller).
Paratypes: all Mexico. TamMau.ipas: same locality as Holotype, 2¢ 19 Momeni ovat LOV3., 194 M39 x 1973, 26 29 11974, Lea wl974, 36 49 xi.1974, 26 12 xii.1974, 16 792 i.1975, reared from Bambusa aculeata (all col- lected or reared by R. O. and C. A. Kendall or W. W. McGuire). San Luis Porost: Ciudad Valles, 29 vii.1970, 14 vii.1972, 36 39 vii.1973 (all collected by H. A. Freeman); El Naranjo, 3¢ 1@ ii.1976 (all collected by R. O. Kendall); Tama- zunchale, 1@ vii.1951 (T. Escalante).
10) JoURNAL OF THE LEPIDOPTERISTS SOCIETY
0.5 mm ho
Figs. 9-13. Splendeuptychia kendalli, n. sp. 9-10, Holotype ¢ upper (9) and under (10) surfaces; MEXICO: TAMAULIPAS: Gonzalez Ranch, nr. Los Kikos (Allyn Mus. photos 010677-16/17); LFW 17.8 mm. 11-12, Paratype 2 upper | (11) and under (12) surfaces; same locality as Holotype (Allyn Mus. photos
010677-14/15); LFW 19.6 mm. 13, ¢ genitalia of Holotype; slide M-3651 (Lee D. Miller).
Disposition of type-series: Holotype 2, 10¢ and 109 Paratypes in the collec- tion of the Allyn Museum of Entomology; nine ¢@ and 10 2 Paratypes returned to R. O. Kendall; 194 and 1592 Paratypes returned to W. W. McGuire. These series will be divided later among other museum collections.
I take great pleasure in naming this distinctive Mexican satyrid for
VoLUME 32, NUMBER 2 81
Mr. Roy O. Kendall who reared the Holotype and several other examples in the type-series. His work on the life histories of various Mexican, as well as Texan, butterflies has been of the greatest value to lepidop- terology and promises even more future benefits to the science.
The Tamazunchale specimen came from the Escalante collection and bore a cryptic determination label in an unknown hand identifying the specimen as S. salvini. I had previously discounted salvini as the name for the Mexican butterfly, and the receipt of true salvini confirmed my previous analysis.
The genitalia are somewhat aberrant for members of Splendeuptychia (Forster, 1964: figs. 161-164), especially in regard to the aborted enathos.
In addition to the genitalic dissimilarities between kendalli and salvini, the former may be distinguished by the following superficial characters: 1) the ground color of kendalli is browner, both dorsad and ventrad; 2) the marginal lines on the upper surface are better developed in kendalli; 3) the ventral forewing of salvini bears four dark bands proximad of the silver spotband, whereas in kendalli the basal of these is missing al- together and the distal band is merely a thin line forming part of the ring around the silvered spots; 4) the ventral hindwing of kendalli also lacks the basalmost band that is prominent in salvini; 5) the yellow patch of the ventral hindwing is more extensive in kendalli, whereas in salvini this patch is poorly developed to absent posteriad of vein Cu,; and 6) the silver spotband of the ventral forewing which extends posteriad as far as 2A in kendalli reaches no further posteriad than Cu, in salvini.
This species is apparently restricted to the mesic environments found in a few places in the eastern foothills of the Sierra Madre Oriental. Thus far the butterfly has been found in a very few localities from Tamazunchale north to Tamaulipas where colonies of B. aculeata grow. What we know about the bionomics of S. kendalli is given in a following paper (Kendall, 1978). Obviously, the insect is multivoltine.
Members of Splendeuptychia are almost uniformly rare. I suspect this is a real occurrence, since they are much more attractive than are most Euptychiini. Perhaps the relative abundance of S. kendalli and its as- sociation with Bambusa will make possible the discovery of greater num- bers of other species of Splendeuptychia. The association of this genus with bamboo is further confirmed by S. S. Nicolay who brought me several specimens of an as yet undetermined Splendeuptychia that he took in a bamboo thicket in eastern Ecuador.
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Figs. 14-15. Cyllopsis dospassosi L. Miller, @ upper (14) and under (15) surfaces; MEXICO: SAN LUIS POTOSI: El Salto Falls (Allyn Mus. photos 010677-2/3); LFW 17.7 mm; R. O. Kendall collection.
Cyllopsis dospassosi L. Miller, 1969 Figs. 14-15 Cyllopsis dospassosi L. Miller, 1969 (“1968”): 53; 1974: 84-86 (type locality:
52 mi. E Ciudad Victoria, Tamaulipas, Mexico ).
The type of this species remained unique until Mr. and Mrs. Kendall collected one at El] Salto Falls, San Luis Potosi on 16.i.1975. This speci- men is the second known example of dospassosi and, fortunately, is the first female. It is quite comparable to the male, but the ground color of the upper side is slightly darker, and that of the under surface is some- what less olivaceous. Nevertheless, the maculation of the under surface is comparable to that of the male with the addition of an ochreous outer element to the extradiscal band of the hindwing. The “gray patch” en- closing the ocelli of the ventral hindwing is obscure, as in the male. The length of the forewing is 17.7 mm. I have not done a genitalic dissection of this specimen since the female genitalia are not diagnostic in Cyllopsis (L. Miller, 1974: 4).
The Kendalls’ specimen of this species (which is in their collection) extends the known range of C. dospassosi from the Sierra de Tamaulipas to the dry eastern flanks of the Sierra Madre Oriental, presumably of Tamaulipas, as well as San Luis Potosi. The range of C. dospassosi may be much wider than previously thought, and its rarity in collections may be attributable to the usual lack of collecting of the smaller Euptychiini.
Cyllopsis wellingi L. Miller, new species Figs. 16-20
Male: Superficially like C. nayarit (R. Chermock), but differing in the follow- ing particulars: somewhat larger, approaching size of C. pephredo (Godman);
VOLUME 32, NUMBER 2 83
0.5 mm. ne |
Figs. 16-20. Cyllopsis wellingi, n. sp. 16-17, Holotype ¢ upper (16) and under (17) surfaces; BRITISH HONDURAS (BELIZE): Cayo District: Pine Ridge, Thousand Foot Falls (Allyn Mus. photos 021777-1/2); LFW 17.3 mm. 18-19, Paratype 2 upper (18) and under (19) surfaces; same locality as Holotype; (Allyn Mus. photos 021677-1/2); LFW 18.5 mm. 20, @ genitalia of Holotype; slide M-3667 (Lee D. Miller).
84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
under surface bands on both wings redder than in either species; transcellular bands of both wings below less well developed than in nayarit; ochreous markings of hind- wing below much more extensive than in nayarit (these are only hinted at in peph- redo); and ocelli not edged inwardly with ochreous in gray patch area, as in nayarit.
4 genitalia as figured, not at all resembling those of pephredo. The genitalia do bear some resemblance to those of C. pseudopephredo (R. Chermock) (L. Miller, 1974: fig. 141), a species that is otherwise quite distinct from wellingi and other members of the pephredo subgroup in its lack of an androconial patch. The valvae of the present species are somewhat broader than those of pseudopephredo, but the characteristic inwardly directed teeth (which usually appear as dorsally diverted ones) are very similar.
Length of forewing of Holotype ¢ 17.3 mm, those of the 11 ¢ Paratypes rang- ing from 17.0 to 18.2 mm, averaging 17.48 mm.
Female: Differs from the @ of C. nayarit in much the same manner as does the ¢, with the additional characteristic of a reddish flush on the upper surface of some specimens that is not shown in other members of the pephredo subgroup.
Lengths of forewings of the eight 2 Paratypes range from 17.9 to 19.1 mm, averaging 18.45 mm.
Described from 20 specimens, 12 males and eight females, from British Honduras ( Belize ).
Holotype ¢: British Honpvuras: Cayo District: Pine Ridge, Thousand Foot Falls, 650 m, 2.ix.1976 (E. C. Welling M); @ genitalia slide M-3667 (Lee D. Miller ).
Paratypes: all same locality as Holotype, 2—3.ix.1976 (E. C. Welling M.), 11é 82 (males all determined genitalically ).
Disposition of type-series: Entire type-series placed in Allyn Museum of Ento- mology, but the series may be subdivided later.
It is with great pleasure that I name this little satyrid for Sr. Eduardo C. Welling M. of Mérida, Yucatan, Mexico. He has consistently been available to collect specimens of Euptychiini for me, and has often placed undescribed and unexpected species at our disposal for systematic work.
This species is something of a puzzle. One male of thirteen in front of me was a specimen of C. pephredo, but all of the others were the new insect, as demonstrated by genitalic dissections. I have no idea why the single pephredo was intermingled with wellingi at the type locality of the latter, and this single specimen represents the first record of pephredo from British Honduras. These two species now bring the total number of Cyllopsis from that country to three (C. gemma freemani {Stallings and Turner] also occurs there).
ACKNOWLEDGMENTS
I am most grateful to Messrs. Roy O. Kendall, E. C. Welling M., Gordon B. Small, Jr. and Alberto Diaz F. and Drs. W. W. McGuire and Tarsicio Escalante for providing the material on which this is based. Mr. A. C. Allyn took the photographs used to illustrate the paper. Mr. Allyn and my wife and colleague, Jacqueline, read and suggested upon the paper. To all of these individuals I owe a great debt of gratitude.
VoLUME 32, NUMBER 2 85
LITERATURE CITED
Butter, A. G. 1866. A monograph of the genus Euptychia, a numerous race of butterflies belonging to the family Satyridae; with descriptions of sixty species new to science, and notes on their affinities, &c. Proc. Zool. Soc. London, [1866]: 458-504; ill.
1868. Catalogue of the diumal Lepidoptera of the family Satyridae in the collection of the British Museum. London, Trustees British Mus.: vi + 211 pp.; ill.
Forster, W. 1964. Beitrage zur Kenntnis der Insecktenfauna Boliviens. XIX. Lepidoptera III. Satyridae. Ver6dff. Zool. Staatssamml. Miinchen, 8: 51-188; ill.
Gopman, F. D. & O. Satvin. 1879-1901. Biolcgia Centrali-Americana. Insecta. Lepidoptera-Rhopalocera. London: 2 vols. + 1 vol. of ill.
KENDALL, R. O. 1978. Larval foodplant, life history notes and temporal distribu- tion for Splendeuptychia kendalli (Satyridae) from Mexico. J. Lepid. Soc., 32: 86-87.
Minter, L. D. 1969. On Mexican Satyridae, with description of a new species. J. Res. Lepid., 7 (“1968”): 51-55; ill.
. 1974. Revision of the Euptychiini (Satyridae). 2. Cyllopsis R. Felder.
Bull. Allyn Mus., (20): 98 pp.; ill.
Journal of the Lepidopterists’ Society 32(2), 1978, 86—87
LARVAL FOODPLANT, LIFE HISTORY NOTES AND TEMPORAL DISTRIBUTION FOR SPLENDEUPTYCHIA KENDALLI (SATYRIDAE) FROM MEXICO!
Roy O. KENDALL? Route 4, Box 104-EB, San Antonio, Texas 78228
ABSTRACT. Larval foodplant, Bambusa aculeata, Gramineae, rearing notes, ecologic and temporal distribution at its northern distributional limit, are recorded for Splendeuptychia kendalli Miller.
Field-collected adults of the satyrid, Splendeuptychia kendalli Miller, were found at 2 locations: 1) along the Rio Sabinas at Rancho Pico de Oro near Ciudad Mante, Tamaulipas, and 2) along the Rio Salto at El Naranjo, San Luis Potosi. This species is closely associated with its larval foodplant, Bambusa aculeata (Ruprecht) Hitchcock, Gramineae. It is doubtless that this insect will be found at other locations along water courses where its larval foodplant grows. Adults were taken in January, February, July, October, November, and December over a 4- year period. The areas were not visited during the other months. It is therefore unknown whether this multivoltine species is continuous brooded,; it may have a reproductive diapause.
Rearing. At Rancho Pico de Oro, 21 December 1972, I observed a 2 deposit a single egg on a juvenile leaf of B. aculeata. The ? was not captured, but the egg was recovered and preserved.
Again at this location on 22 January 1974, two females were collected and kept for egg production. Between 23 January and 2 February, 51 eggs were deposited in confinement on B. aculeata. Most of the eggs were deposited by 1 female which died 2 February. The other female was killed at this time, and both adults were preserved in alcohol to- gether with 6 eggs. The remaining eggs hatched between 28 January and 7 February. Larval losses were rather high resulting from an in- adequate supply of fresh food. Attempts to keep the plants fresh in a refrigerator were only moderately successful. Earlier, several specimens of the foodplant were transplanted to the Los Arcos Courts gardens at Ciudad Mante, our field headquarters, but they did not survive. In an attempt to circumvent a 60-mile trip every few days for larval food, the larvae were offered bermuda grass, Cynodon dactylon (L.) Pers. The
' Contribution No, 380, Bureau of Entomology, Division of Plant Industry, Florida Department of Agriculture and Consumer Services, Gainesville 32602. * Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, Florida
Department of Agriculture and Consumer Services.
VoLUME 32, NUMBER 2 87
larvae ate the bermuda grass, and it was thought a laboratory solution had been found for rearing this species. However, the larvae soon began to die. The bermuda grass may have been toxic to the larvae, but the lack of proper nourishment in the grass was suspect. On 6 April 1974 all remaining larvae (24) were preserved.
Once again a 2 collected 23 November 1974 from this location de- posited 27 eggs between 24 and 29 November and died 3 December 1974. These eggs hatched between 28 November-l11 December, and a maximum effort was made to rear them. Numerous trips were made to the collection site for fresh bamboo. Even so, there were several larval casualties attributed to rapid desiccation of the cut bamboo. Eleven larvae pupated between 27 December 1974 and 12 January 1975. Adults emerged (2 6, 72) between 7 and 23 January 1975. Two larvae and 2 pupae (one deformed ) were preserved.
Field-Collected Adults. In addition to the above, other field collections include: Rancho Pico de Oro, 21 December 1972 (24,12), 9 January 1974 (32 ), 22 Janu- ary 1974 (146), 22 February 1974 (14), 23 November 1974 (44, 19), 4 December 1974 (14), 6 December 1974 (39 ), and 8 January 1975 (1¢), all leg. Roy O. and C. A. Kendall. At the same location, 27 December 1972 (124), 18 July Poaee ee 20nfuly 1973 (2° ), 22 October 1973 (176, 82), 25 October 1973 (36, 22) all leg. W. W. McGuire. At E] Naranjo, 13 February 1976 (1@), 14 February 1976 (224), and 29 February 1976 (14) all leg. Roy O. and C. A. Kendall.
ACKNOWLEDGMENTS
Mrs. Kendall and I wish to thank Sr. and Sra. Carlos Gonzales for permission to conduct field research at their rancho, and for their warm hospitality. To Sr. and Sra. Fernando Reyes Bugarin and their family we are most grateful for the comfortable field headquarters provided and for the use of their botanical gardens in our research.
Journal of the Lepidopterists’ Society 32(2), 1978, 88-96
NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF VANESSA ANNABELLA (NYMPHALIDAE)
THomMas E. Dimock? 111 Stevens Circle, Ventura, California 93003
ABSTRACT. Observations on the life history of Vanessa annabella (Field) show the early stages to be quite variable: the eggs in rib structure, and the later larval stages in color pattern and behavior. Immature and adult behavioral charac- teristics are similar to those of other Vanessa. V. annabella is usually present throughout the year in coastal southern California.
Vanessa annabella (Field), the West Coast Lady, is a common and familiar butterfly in western North America. Because it can usually be found throughout the year in coastal southern California, opportunities to study its life history are almost always present. However, there are few published records available and none has included photographs of the complete life cycle. Of published reports, Dyar (1889) gave one of the more complete written accounts; Huguenin (1921) made some general observations on the life cycle and natural history; and Coolidge (1925) described the egg in detail and listed the larval foodplants. More recently Emmel & Emmel (1973) illustrated paintings of a light form of the last instar larva and the pupa and gave brief descriptive notes.
Specimens used for the present descriptions of the life cycle stages were collected as freshly laid ova by following an ovipositing female at the type locality in Ventura, California (Dimock, 1972). The leaves on which these eggs were laid were placed in plastic containers 11 cm square by 4 cm deep. Humidity was maintained by dampened tissue paper placed on the container bottom. The containers were kept indoors in a room temperature which varied from 17 to 25°C. Photographs and measurements were made of each stage. Other specimens were reared upon cut stalks of nettle placed in water so that leaf shelter construction and other activities could be observed. Afternoon sunshine provided direct and ambient light.
Full descriptions of the adults are given by Field (1971); thus, the following adult descriptions are limited to those characteristics which help distinguish V. annabella from related North American species.
Life Cycle Stages
_ Ege (lig. 1). Barrel-shaped, light green, with 10 to 14 transparent vertical ribs. Measurements (Coolidge, 1925): 0.72 mm tall, 0.52 mm wide, tapering to 0.30 mm at base and 0.26 mm at top. Duration 4 days.
‘Museum Associate in Entomology, Natural History Museum of Los Angeles County, 900 Ex- position Blyd., Los Angeles, California 90007. ;
VoLUME 32, NUMBER 2 89
Figs. 1-12. Vanessa annabella (Field): (1) egg, ca. 0.6 mm wide; (2) first instar larva, 3 mm; (3) second instar larva, ca. 4 mm: (4) third instar larva, ca. 9 mm; (5) fourth instar larva, ca. 13 mm; fifth instar larvae, all ca. 30 mm: (6) dark morph, black and yellow, (7) intermediate morph, orange and gray, (8) rusty orange morph, (9) light morph, gray; (10) head, fifth instar, light tan morph; (11) head, fifth instar, dark morph; (12) prepupa.
First instar larva (Fig. 2). Head shiny black, setae and thoracic legs black. Ground color grayish brown after feeding for 2 days. When mature, body with vague brownish mottling. Segments A-2, A-4, and A-6 with a pair of light yellow spots between subdorsal and supralateral setae. Grows to 3 mm in 5 days.
Second instar larva (Fig. 3). Head shiny black. Ground color mottled dark brown. Short branched spines black except for middorsal spines on A-4 and A-6 and subdorsal spines on A-2, A-4, and A-6, which are yellow. A narrow pair of vague yellow lines divided by a narrow middorsal line of dark ground color running from about T-1 to A-8. Grows to 4.5 mm in ca. 3 days.
Third instar larva (Fig. 4). Head shiny black with black setae arising from black chalazae. Ground color usually black, but may begin to lighten as in lighter morphs. Spines black except for subdorsal spines on A-2, A-4, A-6, and usually
90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Wj
ee. Z iY Wy, » ty
Fig. 13. Vanessa annabella (Field): fifth instar larval nest on Urtica holosericea Nutt.
middorsal spines on A-4 and A-6, which are yellow with black tips and yellow bases confluent with dorsal double yellow lines. These paired lines, variable in expression, separated by a middorsal line of ground color, are interrupted by ground color at bases of non-yellow spines. Grows to ca. 9 mm in ca. 3 days.
Fourth instar larva (Fig. 5). Head black with bronze hightlights. Head cap- sule width 1.5 mm. Ground color and markings nearly as variable as in fifth instar (see following description). Grows to ca. 14 mm in ca. 3 days.
Fifth instar larva (Figs. 6-9). Head blackish or brownish black (Fig. 11) with bronze highlights, or less often with vertical whitish tan stripes in light morphs (Fig. 10). Capsule width 2 mm. Extremely variable in ground color and mark- ings. Ground color varies from black to greenish white or grayish white, including various browns and tans. Light markings present in fourth instar here vary from dark rusty reds and oranges to yellow or various browns and tans. Extent of mark- ings and lateral line varies independently of color; these tend to disappear alto- gether in morphs of whitish ground color. Lateral line may be absent in any morph. tusty orange spots may appear between subdorsal and supralateral spine bases, varying in extent from absence to confluence with other pattern elements. Spines branched, variable from black in dark morphs to whitish in light morphs, or dark anteriorly and light posteriorly in intermediate morphs. Arrangement of spines: middorsal row on segments A-1 to A-8, subdorsal rows on T-2 to A-8, supra- lateral rows on T-2 to A-10, and lateral rows on A-1 to A-8. Body shape thickest at midabdominal segments. Grows to ca. 25-30 mm in ca. 5 days.
Prepupa (Fig. 12). Light markings darken somewhat. Larva becomes slightly shorter and thicker. Duration 1 day.
Pupa (Figs. 14-16). Head without projections. Mesothorax with a_ raised middorsal point and a pair of subdorsal points. Metathorax with two large sub-
VOLUME 32, NUMBER 2 91
Figs. 14-20. Vanessa annabella (Field): (14) pupa, dorsal view, 19 mm; (15) pupa, lateral view; (16) pupa, ventral view; (17) adult male, dorsal view, 47—mm expanse; (18) adult male, ventral view; (19) adult female, dorsal view, 49-mm expanse; (20) adult female, ventral view.
dorsal white spots, raised anteriorly. Abdomen with two small subdorsal white spots on A-1 bordering those on metathorax. A-2 to A-7 each with one very small middorsal point and a pair of more prominent subdorsal points. Color variable from overall tan to mottled dark browns, sometimes with a greenish golden cast.
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A spiracular line, darker brown than ground color, is variable in expression. Mea- surements (average of five specimens): length 19 mm; width 7 mm; depths: thorax 6.25 mm, saddle 5.25 mm, abdomen 7.25 mm. Duration 8-11 days.
Adults (Figs. 17-20). Sexual dimorphism subtle, females having a more rounded hindwing than males, especially at M:, Cu, and Cuz. Color pattern same in both sexes: tawny orange with black markings, white subapical forewing spots, and blue pupilled hindwing ocelli. On upperside, forewing cell crossed completely by a black bar. Forewing costal bar, between cell end and apex, orange. Forewing apex pointed, not rounded, at M,, with marginal tawny spot in interspace Rs. Hindwing with four blue pupilled submarginal ocelli in interspaces M:, M2, Ms, and Cu, and often a small solid black ocellus in interspace Rs. Hindwing under- sides mottled principally in various buffs, tans, browns, and grays, with a whitish triangle in interspace M» at cell end. Expanse averages between 40 and 48 mm, females often larger than males.
Total developmental time for this species is ca. 30 to 36 days.
Natural History
Vanessa annabella uses a variety of foodplants in the families Urti- caceae and Malvaceae. Native foodplants most frequently used in southern California are Urtica holosericea Nutt. (Urticaceae), Sida spe- cies, Sidalcea malvaeflora (DC.) Gray, and Sphaeralcea ambigua Gray (Malvaceae). Introduced plants include Malva species, especially M. parviflora L., and Althaea rosea (L.) Cav. (Malvaceae). John F. Em- mel, M.D. (pers. comm.) also reports the use of Urtica urens L. ( Urticaceae ).
The eggs are laid singly, usually on the uppersides of the leaves. On nettles (Urtica) the eggs are often attached to the sides of the stinging spines.
The hatching larva eats away the top and adjacent walls of the egg and crawls to a suitable place on the leaf uppersurface to construct a shelter. This consists of fine silk webbing tied across a leaf midrib, petiole, or small wrinkle on the leaf margin. The young larva lives under this webbing as it feeds on the leaf and places its frass into the webbing, creating a protective camouflage. In the second instar the larva may enlarge the old nest or construct a new one nearby. When the larva is at the growing tip of a nettle stalk, the nest may incorporate two or more of the tiny new leaves. By the third instar the larva is capable of folding a larger area of the leaf or constructing a deeper nest at the petiole. In the fourth instar the entire leaf may be folded together (on Urtica) or closed about the top edges (on Malwa). Frass is allowed to fall out of the nest but often accumulates in piles in the nest bottom. The fifth instar larval nest is usually larger and may incorporate neigh- boring leaves and stems (Fig. 13). Sometimes leaves of nearby plants which are not foodplants are also tied into the nest even though they are
VoLUME 32, NUMBER 2 93
not eaten. On plants with small leaves, such as young Malwa, the larva may tie together many leaves before a nest enclosure is completed. On Urtica holosericea, when a single leaf is used, larvae of V. annabella usually construct nests on the uppersides of the leaves, either by folding over one edge and securing it to the leaf surface or by tying both edges together to form an enclosure. The petiole or nearby midribs may or may not be partially cut to cause the leaf to hang vertically. Less frequently, the larvae will fold the leaf edges underneath so that the undersurface forms the nest interior.
Pupation sites are on either the foodplant or nearby objects. When the foodplant is used, a leaf chamber is constructed with firm webbing and the larva suspends itself from the chamber ceiling. Larvae in other loca- tions may secure together any nearby objects to approximate an en- closure or may simply pupate in exposed places, such as from twigs or branches. Pupae often react to disturbances by wiggling laterally.
Emerging adults hang from the pupal shell or adjacent perch to ex- pand their wings. A reddish brown meconium is ejected and the adult is ready for flight in an hour.
Adults of V. annabella may be encoutered in any life zone from sea level to alpine areas where open sunny places are preferred. Both sexes visit flowers. In the afternoon males tend to congregate on hilltops or other exposed places such as forest openings, glades, meadows, and streamside slopes, especially when patches of dry, bare earth are avail- able for sunning. Many man-made situations are particularly favorable: windbreaks of trees, orchard rows, trails, firebreaks, garden paths, and paved sidewalks and driveways. At these locations, when not occupied in sunning, males will chase after each other and the other vanessid butterflies Vanessa atalanta rubria (Fruhstorfer), V. cardui (L.), and V. virginiensis (Drury), along with unrelated butterflies which con- gregate in the same places. They often bravely chase larger insects and birds and in general will investigate anything that flies through their established area, including falling leaves and objects thrown overhead. These activities ultimately bring the males into contact and subsequent courtship and mating with females, but between these encounters the males spend a great deal of time and energy simply chasing each other. From observations made on hilltops in the vicinity of Ventura, California, during November 1976 when all four Vanessa were present, it was noted that any one species will chase the same or any other species, and two or more individuals may join in the chase. The butterflies may chase each other to a height of ca. 20 m or more before breaking chase and quickly gliding down to land once again on the ground with backs to the
94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
sun and wings spread. The butterflies’ wings frequently come into con- tact during these encounters, but without damaging effects, and the re- sulting noise can be heard nearby.
Females, although not congregating in the manner of the males, are likely to be found anywhere, feeding, seeking foodplants, or ovipositing, including hilltop localities when the foodplants or nectar sources occur there also.
The flight of V. annabella is composed of glides with the wings held horizontally, interrupted frequently by several fluttering beats. Chasing is mostly vigorous fluttering, and the return dives are composed of gliding and braking.
Diapause was not investigated, but if it does occur in this species it is almost certainly during the adult stage, as it is in the other vanessid- nymphalinid butterflies. If adults of V. annabella are unable to survive prolonged or severe frosts, the species probably reinvades the greater part of its northern and eastern range from the milder southwestern areas where breeding is continuous throughout the year.
Vanessa annabella is easily attracted to suburban gardens by planting Althaea rosea (Hollyhock) or encouraging Malva parviflora (Cheese- weed) to become established. It is an easy butterfly to raise in captivity, even under poor conditions. Larvae collected in the wild on one food- plant (for example, Urtica holosericea) can be switched to other foodplants (Malva, Althaea) when the former is less easily obtained. The larvae are very often parasitized by tachinid flies, which emerge from the mature butterfly larvae or pupae as mature maggots.
DiIscussIon
Coolidge (1925) noted that the egg ribs of V. annabella varied in number from 11 to 13, with 11 the most common number. My ob- servations confirm this, but eggs with 10 and 14 ribs were found during the present study. This is in partial disagreement with Field (1971), who stated that the genus Cynthia (in which annabella was placed) had from 14 to 19 egg ribs, and Clench (in Howe, 1975), who gave 14 or 15 as the number of egg ribs in the subgenus Cynthia.
The larvae of V. annabella are extremely variable, as are the larvae of V. atalanta rubria and V. cardui, and this variability not only makes a written description difficult but compounds the task of providing re- liable characteristics with which the three species can be separated. In general, the descriptions given for the larvae in this article can be com- pared with those for V. a. rubria and V. cardui in subsequent articles. However, fifth instar larvae of V. annabella can always be distinguished
VoLUME 32, NUMBER 2 95
by their smaller head capsule width of 2 mm; in V. a. rubria and V. cardui the fifth instar larval head capsule is nearly 3 mm. Once V. anna- bella is identified, V. a. rubria is distinguished by the numerous white cephalic chalazae, which in V. cardui are black.
From observations made on V. annabella larvae collected in various locations and larvae reared under controlled conditions, it was noted that the variations in ground color were at least partly due to environ- mental conditions. The light, grayish-white morphs were more fre- quently encountered on plants exposed to full sunshine, whereas the darker morphs were found mostly on plants in secluded, shaded areas. Darker morphs also resulted when larvae were reared under crowded conditions.
Because Malva species are especially successful in disturbed areas and are abundantly available throughout the year as foodplants, V. annabella has probably become much more common since the introduction of these weeds from Europe. This is the situation in the coastal southern California lowlands, where a favorable climate prevails and V. annabella can be found in every month of the year.
In his revision of the Vanessa butterflies, Field (1971) resurrected the genus Cynthia for the carye, cardui, and virginiensis species groups. Clench (in Howe, 1975) reunites all the species in Vanessa, treating Cynthia as a subgenus. Emmel & Emmel (1973) used the cases of hybridization between V. a. rubria and Cynthia annabella to demonstrate the “close genetic relationship and probable generic identity” of the two species. With no disrespect to the fine work of Field, I favor a treatment similar to that of Clench, with reservations on the precise placement of annabella. There are 10 known cases of hybridization between V. a. rubria and V. annabella: one specimen reported by Edwards (1877), one by Grinnell (1918), one by Gunder (1930), three by Dimock (1973), one by Emmel & Emmel (1973), one specimen collected by Kirby in the collection of the Natural History Museum of Los Angeles County, and two specimens raised by Henne and Ingham in the Peabody Museum collection. Mr. William D. Field (pers. comm.) discovered upon dis- section the partially crippled specimen designated as “Hybrid #3” in Dimock (1973) to be a female, not a male as erroneously reported. In my opinion these occurrences support, at least, the arrangement of Clench and the generic identity suggested by Emmel & Emmel. Biolog- ically, the examples of hybridization may also demonstrate the presence of an as yet incomplete reproductive isolatory mechanism caused by a recent invasion of V. annabella from South America or a recent invasion of V. a. rubria from Eurasia, or invasions by both species.
96 JoURNAL OF THE LEPIDOPTERISTS SOCIETY
ACKNOWLEDGMENTS
I wish to thank my father, M. W. Dimock, for printing the photographs which accompany this article; Mr. Julian P. Donahue of the Natural History Museum of Los Angeles County for access to the Museum col- lection; Mr. Christopher Henne of Pearblossom, California, for an account of his hybrids; Mr. William D. Field of the Smithsonian Institution, Washington, D.C., for information on the hybrids; Dr. John F. Emmel, Hemet, California, for reviewing the manuscript; and Mr. Alberto Muy- shondt of El Salvador for recommendations on photographic techniques and whose format for life history observations of El Salvador butterflies is the one I followed for this article.
LITERATURE CITED
Cootmncr, K. R. 1925. California butterfly notes, III. Bull. Brooklyn Ent. Soc. 20(3): 146-147.
Dimock, T. E. 1972. Type locality and habitat—Cynthia annabella. J. Res. on the Lepid. 10: 265-266.
1973. Three natural hybrids of Vanessa atalanta rubria x Cynthia annabella (Nymphalidae). J. Lepid. Soc. 27: 274-278.
Dyar, H. G. 1889. Preparatory stages of Pyrameis carye Hiibner. Canadian Ent. 21: 237-238.
Epwarps, H. 1877. Pacific Coast Lepidoptera, No. 22. Notes on some diurnal Lepidoptera, with descriptions of new varieties. Proc. Calif. Acad. of Scis. 7: 163-174.
EMMEL, T. C. & J. F. Emmeu. 1973. The butterflies of southern California. Nat. Hist. Mus. of Los Angeles County, Sci. Ser. 26: 1-148.
Fretp, W. D. 1971. Butterflies of the genus Vanessa and of the resurrected gen- era Bassaris and Cynthia (Lepidoptera: Nymphalidae). Smiths. Contribs. to Zool., Number 84: 1—105.
GRINNELL, Jr., F. 1918. Some variations in the genus Vanessa (Pyrameis). Psyche 25: 110-115, pl. 4.
Gunpver, J. D. 1930. Butterflies of Los Angeles County. Bull. Southern Calif. Acad. of Sci. 29: 39-95.
Howe, W. H., coordinating editor. 1975. The Butterflies of North America. Doubleday and Co., Inc., Garden City, L. I., New York. xiii + 633 p. + 97 pl.
HucuEnin, J. C. 1921. Life history of Pyrameis caryae in California (Lep., Rhop.). Ent. News 32: 216-217.
Journal of the Lepidopterists’ Society 32(2), 1978, 97-102
A NEW SPECIES OF HEMILEUCA FROM THE SOUTHWESTERN UNITED STATES (SATURNIIDAE)
Pau M. TusKEs
Department of Environmental Toxicology, University of California, Davis. Davis, California 95616
ABSTRACT. JHemileuca griffini Tuskes which occurs in southern Utah and northern Arizona was collected for the first time in 1974. The adult moth is a black and white day flying saturniid which is active during September and October. The larval hostplant is black brush, Coleogyne ramosissima. This species has a unique taxonomic position in that both the adult and larva exhibit morphological characters which are intermediate to the Pseudohazis and Hemileuca groups, thus, a continuum of characters exists between these two previously separated genera.
The genus Hemileuca consists of 23 described species, 16 of which have partial or complete distributional patterns north of Mexico. The moths within this genus are large to moderate in size, and exhibit a great deal of hostplant and habitat diversity. Adults are characterized by hav- ing the labial palpi fused to each other forming a small unsegmented bilobed structure; also, the male has bipectinate antennae. Members of Coloradia, the genus most closely related to Hemileuca, have labial palpi which are separate, and males have antennae which are quadripectinate.
The last Hemileuca described as a distinct species was chinatiensis (Tinkham), in 1943. The significance of H. chinatiensis as a species with genitalic characters intermediate between Pseudohazis and Hemi- leuca was overlooked by Tinkham; not until Ferguson (1971) was its taxonomic position made clear. Michener (1962) combined the genera Pseudohazis and Hemileuca on the basis of their morphological similarity, but made no mention of chinatiensis. Although Michener included four subgenera within Hemileuca Ferguson chose to abandon the subgeneric names and to consider them as species groups.
It is the purpose of this paper to describe a new species of Hemileuca collected for the first time in 1974, and to present additional mor- phological evidence to support the merger of Hemileuca and Pseu- dohazis. The new species