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However, if the frequency of copulations resulting in the transfer of two spermatophores in the laboratory is a good estimate of their frequency in the field (3/199 copulations observed in the laboratory), its quantitative effect should be small.
Spermatophores always leave recognizable remains within the corpus bursae of the female. This is not true in C. xami since in the laboratory it was not always possible to observe clear sper-matophore remains in very old females that had laid most of their eggs (pers. obs.). However, judging from wing wear, no female in this condition was sampled (see paragraph four above).
In conclusion, the possible violation of the first and the last assumptions, and the fact that some of the females may have mated again had they not been collected, results in an underestimation of the frequency of copulations in females; whereas the fact that some males transfer more than one spermatophore in one copulation results in an overestimation of the number of copulations. However, judging from the low frequency of "interrupted" copulations (4.4%), very worn females in the field (at least during the sampling period), and copulations resulting in the transfer of two spermatophores (1.5%), I conclude that spermatophore counts are a reasonably good estimate of female copulation frequency in C. xami.
Acknowledgments
I thank Gabriela Jimenez and Dr. Rogelio Macias for their valuable technical help, and Dr. J. M. Burns and an anonymous reviewer for their comments. This research was supported by a Consejo National de Ciencia y Tecnologia (Mexico) scholarship.
Literature Cited
Braby, M. F. 1996. Mating frequency in bush-brown butterflies (Nymphalidae: Satyrinae). J. Lepid. Soc. 50:80-86.
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Burns, J. M. 1968. Mating frequency in natural populations of skippers and butterflies as determined by spermatophore counts. Proc. Nat. Acad. Sci. U.S.A. 61:852-859.
Cordero, C. 1993. The courtship behavior of Callophrys xami (Lycaenidae). J. Res. Lepid. 32:99-106.
----------. 1998. Ecologia del Comportamiento Sexual de los Machos
de la Mariposa Callophrys xami, con Algunas Consideraciones Acerca de la Evolution del Semen de Insectos. Doctoral Thesis, UACPyP/CCH, UNAM, Mexico.
Drummond III, B. A. 1984. Multiple mating and sperm competition in Lepidoptera, pp. 291-370. In R. L. Smith, (cd.), Sperm competition and the evolution of animal mating systems. Academic Press, New York.
Eberhard, W. G. 1985. Sexual selection and animal genitalia. Harvard University Press, Cambridge, U.S.A.
----------. 1996. Female control. Sexual selection by cryptic female
choice. Princeton University Press, Princeton, U.S.A.
Lederhouse, R. C, M. P. Ayres&J. M.Scriber. 1989. Evaluation of spermatophore counts in studying mating systems of Lepidoptera. J. Lepid. Soc. 43:93-101.
Soberon, J., C. Cordero, B. Benrey, P. Parlange, C. Garcia-Saez & G. Berges. 1988. Patterns of oviposition by Sandia xami (Lepidoptera, Lycaenidae) in relation to food plant apparency. Ecol. Entomol. 13:71-79.
Carlos Cordero. Institute de Ecologia, Universidad Nacional Autonoma de Mexico, Apdo. Post. 70-275, C.P. 04510 D.E, and Cen-tro de Investigaciones Fisiologicas, Universidad, Autonoma de Tlax-cala, Apdo. Post. 262, C.P. 90070 Tlaxcala, Tlaxcala, MEXICO (Address for correspondence)
Received for publication 5 April 1999; revised and accepted, 16 December 1999.
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journal of the Lepidopterists' Society 53(4), 1999, 170-172
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ADDITIONAL NOTES ON PROSERPINUS CLARKIAE AND ARCTONOTUS LUCIDUS (SPHINGIDAE) LIFE HISTORIES FROM THE PACIFIC COAST OF NORTH AMERICA
Additional key words: Onagraceae, Rubiaceae, Gayophytum, Galium, Clarkia breweri, Clarkia modesta, Camissonia.
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Host associations for Proserpinus clarkiae (Boisduval) and Arctonotus lucidus (Boisduval) have recently been documented. Proserpinus clarkiae was found using Clarkia unguiculata (Lindley) in nature (Osborne 1995). Here, I compare results of my life history work on P. clarkiae with other results (Hardy 1959) on this species. The life history of A. lucidus is also known (Comstock & Henne 1942). However, the first natural host associations for A. lucidus were made by photographs and collections from Clarkia species in California, and are presented here along with observations on captive rearing of this moth. The immature stages of these related sphingid species have been confused in the field by some, possibly due to their sympatry, common use of Clarkia hosts, and superficial resemblance. Thus, I will also discuss morphological differences among these and other sympatric Clarkia feeding sphingids.
In presenting the biology of P. clarkiae (Osborne 1995), I repeated the assertion made by Hodges (1971) that its life history was unknown. Since that time, Dr. Frederick Rindge (American Museum of Natural History) has drawn my attention to a life history of P. clarkiae that predates both works. Larvae and a pupa reared from Vancouver Island (Hardy 1959) were described by Hardy (1959), and match the immatures of P. clarkiae from California. Hardy obtained seven ova by confining females over potted Galium aparine (Lewis & Szweykowski) (Rubiaceae). He reared at least one individual to pupation on that plant, but a field host was not given. The single fifth instar larva of P. clarkiae from Vancouver Island had the lateral dark blotches contiguous in an undulating line, a trait consistent
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with some (< 5%) of the California material I reared (most California larvae had oblique blotches disjunct) (Osborne 1995). This dark form may be typical of cool, wet, north coastal localities, where darker maculation may impart local selective advantages, or may be an artifact of captive rearing.
Dr. Robert Raguso, who studied sphingid pollination of Clarkia species in central California (see Raguso & Pichersky 1995, Raguso et al. 1996, Raguso & Light 1998), sent me several suspected Proserpinus larvae, a reared pupa, and a photograph (Fig. 1) of a fifth in-star larva in nature on Clarkia breweri (A. Gray) E. Greene. These specimens were all collected from C. breweri and Clarkia modesta (Jepson) at Del Puerto Canyon, Stanislaus Co., California in May, 1991. However, instead of P. clarkiae, all were determined (by KHO) to be Arctonotus lucidus, a closely related species from a monotypic genus. Early instar A. lucidus larvae may be separated from P. clarkiae by the presence of a black anal horn which is absent in P. clarkiae. Fifth instar A. lucidus lose the anal horn, but have dorsal and lateral markings of olive green (but briefly black just after molt [Comstock & Henne 1942]), not black or gray as in P. clarkiae. In addition, A. lucidus can be distinguished from P. clarkiae on the basis of dorsal, transverse intersegmental lines of tan or cream breaking the olive green field, and ventral whitish or gray. The ground color in fifth instar A. lucidus larvae is variable (Comstock & Henne 1942), ranging from black to olivacious green to light green, to pink (Comstock & Henne 1942; D. Rubinoff pers. comm.; K. H. Osborne unpubl. obs.).
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Volume 53, Number 4
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Fig. 1. Raguso.
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Fifth instar Arctonotus lucidus larva on Clarkia breweri at Del Puerto Canyon, Stanislaus Co. CA., May, 1991. Photograph by Robert
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Raguso's photograph of an A. lucidus larva on C. breweri, and collections of A. lucidus from C. breweri and C. modesta, represent the first natural host records for this moth. Raguso (pers. comm. 1995) has seen [these?] larvae on Clarkia gracilis sonomensis (Hitchc.) near Lake Berryessa, Napa Co., California. (These records must be considered as likely A. lucidus but could possibly be P. clarkiae.). Additionally, one wandering fifth instar A. lucidus (determined by KHO) was found by M. Lynn (pers. comm.) in May, 1997 in the immediate vicinity of abundant Camissonia bistorta (Nutt), Camissonia strigulosa (Fisck. & Meyer) Raven [^Oenothera contorta Munz], and Clarkia purjjurea (Curt.) Nels. & Macbr. at Lake Skinner, Riverside Co., California, suggesting these plants as possible hosts. The Cammissonia species are used as larval hosts by related sphingids in southern California, Euproserpinus phaeton Grote & Robinson using C. bistorta (Osborne 1995), Euproserpinus euterpe Hy. Edwards using C. strigulosa (Tuskes & Emmel 1981, K. H. Osborne unpubl. obs.), and Hyles lineata (L.) using both (K. H. Osborne unpubl. obs.). Clarkia breweri is restricted to central California from Alameda Co. south to Fresno Co. (Munz 1959) and C. modesta ranges through California from Tehema Co. south to Santa Barbara Co. (Hickman 1993). The wide range of A. lucidus (Holland 1903, Hodges 1971) from British Columbia at least as far south as San Diego Co., California (Brown & Donahue 1989; Osborne unpubl. obs.) indicates A. lucidus must use other host plant species.
Galium, suitable for P. clarkiae in captivity (Hardy 1959), was rejected by A. lucidus as were two unnamed Oenothera species (Corn-stock & Henne 1942). In captivity, A. lucidus larvae would accept leaves of C. breweri, C. modesta and Clarkia affinis (H. Lewis & M. Lewis) (Raguso pers. comm.), but were hesitant to accept Clarkia
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unguiculata (Lindley) and Fuchsia (K. H. Osborne unpubl. obs.; Raguso pers. comm.). Most prepupal A. lucidus larvae wandered and died without pupating in dry, sandy soil, and the one that did pupate, about 3 cm below ground in loose gravel, was attacked by mold and never emerged (K. H. Osborne unpubl. obs.; Raguso pers. comm.). Dan Rubinoff (pers. comm.) reported success getting A. lucidus to pupate on moist potting soil and when reared by Comstock & Henne (1942) larvae pupated as deep in the soil as possible (in cages).
Hyles lineata is common on Clarkia (C. unguiculata at Gates Canyon, Solano Co, CA [unpublished records] and C. breweri, at Del Puerto Canyon [Raguso]) when A. lucidus and P. clarkiae may be present. Hyles lineata is easily distinguished from larvae of A. lucidus and P. clarkiae by its prominent orange or yellow anal horn in all larval stages and by distinctive (but variable) longitudinal markings (Hodges 1971).
ACKNOWLEDGMENTS
I thank Dr. Frederick Rindge for informing me of prior life history work on P. clarkiae, Dr. Robert Raguso who sent me immatures, photographs and notes, and reviewed this manuscript. Daniel Rubinoff also reviewed the manuscript and made helpful suggestions. Finally, my deepest gratitude goes to an anonymous reviewer who scoured the manuscript, making many insightful comments and improvements.
Literature Cited
Brown, J. W. & J. P. Donahue. 1989. The Sphingidae (Lepi-doptera) of Baja California, Mexico. J. Lepid. Soc. 43:184-209.
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Comstock, J. A. & C. Henne. 1942. The early stages oiArctonotus lucidus Bdv. (Lepidopt.). Bull. South. Calif. Acad. Aci. 41:167-171.
Hardy, G. E. 1959. Notes on the life histories of four moths from southern Vancouver Island. Entomol. Soc. British Columbia, Proc. 56(4):49-52.
Hickman, J. C. (ed). 1993. The Jepson manual: higher plants of California. University of California Press, Berkeley, Ca.
HODGES, R. W. 1971. Moths of North America, north of Mexico. Fascicle 21 (Sphingidae). 170 pp.
Holland, W. J. 1903. The moth book. Doubleday, Page and Co., New York. 479 pp.
Osborne, K. H. 1995. Biology of Proserpinus clarkiae (Sphingidae). J. Lepid. Soc. 49(l):72-79.
Munz, P. A. 1959. A California flora. University of California Press, Berkeley. 1681 pp.
Raguso, R. A. & E. Pichersky. 1995. Floral volatiles from Clarkia breweri and C. concinna (Onagraceae)—-recent evolution of floral scent and moth pollination. Plant Systematics and Evol. 194(l-2):55-67.
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Raguso, R. A., D. M. Light & E. Pichersky. 1996. Electroan-tennogram responses of Hyles lineata (Sphingidae, Lepi-doptera) to volatile compounds from Clarkia breweri (Onagraceae) and other moth-pollinated flowers. J. Chem. Ecol. 22(10):1735-1766.
Raguso, R. A. & D. M. Light. 1998. Electroantennogram responses of male Sphinx perelegans hawkmoths to floral and 'green-leaf volatiles' Entomologia Experimentalis et Applicata. 86(3):287-293.
Tuskes, P. M. & J. F. Emmel. 1981. The life history and behavior of Euproserpinus euterpe (Sphingidae). J. Lepid. Soc. 35:27-33.
Kendall H. OSBORNE. 24292 Lysanda Drive, Mission Viejo, California, USA 92691
Received for publication 3 June 1999; revised and accepted 6 February 2000.
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