Vol. 50 JANUARY 1974 No. 1 THE Pan-Pacific Entomologist PINTO — Courtship behavior in Linsleya compressicornis and its taxonomic significance (Coleoptera: Meloidae) SCOTT — Adult behavior and population biology of Poladryas minuta, and the relationship of the Texas and Colorado populations (Lepidoptera: Nymphalidae) HARADON — Vaejovis spicatus : A new scorpion from California (Scor- pionida: Vaejovida) LANGSTON — The maritime earwig in California (Dermaptera: Carcinophor- idae) CHEMSAK AND LINSLEY — Notes and descriptions of some lycid-like Neo- tropical lepturine Cerambycidae (Coleoptera) _ KOMPFNER — Larvae and pupae of some wrack dipterans on a California beach (Diptera: Coelopidae, Anthomyiidae, Sphaeroceridae) HORN — Observations on primary and secondary parasitoids of California oak worm, Phryganidia calif ornica , pupae (Lepidoptera: Dioptidae) DAILEY, PERRY, AND SPRENGER — Biology of three Callirhytis gall wasps from Pacific slope Erythrobalanus oaks (Hymenoptera: Cynipidae) ALCOCK — Observations on the behavior of Mallophora fautrix Osten Sacken (Diptera: Asilidae) STEINER — Unusual caterpillar-prey records and hunting behavior for a Podalonia digger wasp: Podalonia valida (Cresson) (Hymenoptera: Sphecidae) McDONALD, HEED, AND MIRANDA — The larval nutrition of Minettia flaveola and Phaonia parviceps and its significance to the Hawaiian leaf -breeding Drosophila (Diptera: Lauxaniidae, Muscidae, Dro- sophilidae) SCIENTIFIC NOTES 22, 52, 67, 72, 84, 85, 86, BOOK REVIEW BOOK NOTICE RECENT LITERATURE 27, PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY 1 9 23 28 35 44 53 60 68 73 78 87 83 27 34 90 SAN FRANCISCO, CALIFORNIA • 1974 Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES THE PAN-PACIFIC ENTOMOLOGIST EDITORIAL BOARD J. 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The Pacific Coast Entomological Society Officers for 1974 C. B. Philip, President Paul H. Arnaud, Jr., Treasurer H. V. Daly, President-elect Franklin Ennik, Secretary Statement of Ownership Title of Publication : The Pan-Pacific Entomologist. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher, Publisher and Owner : Pacific Coast Entomological Society, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118. Editor: Dr. John T. Doyen, Division of Entomology & Parasitology, University of California, Berkeley, California 94720. Managing Editor and Known Bondholders or other Security Holders : None. This issue mailed 15 July 1974 Second Class Postage Paid at San Francisco, California and additional offices. ALLEN PRESS, INC. LAWRENCE, KANSAS The Pan-Pacific Entomologist Vol. 50 January 1974 No. 1 Courtship Behavior in Linsleya compressicornis and its Taxonomic Significance 1 (Coleoptera: Meloidae) John D. Pinto Department of Entomology, University of California, Riverside 92502 The systematic position of the small, western North American genus Linsleya MacSwain has been the subject of recent controversy. Although members of this group were originally placed in Lytta on the basis of superficial adult resemblance (Horn, 1873), larval anatomy has now convinced most workers that they belong near Epicauta and Pleuro- pompha in the Epicautina (MacSwain, 1951, 1956; Selander, 1955, 1964a; Arnett, 1960; Werner, Enns and Parker, 1966). Yet, other authors (Kaszab, 1959; Gupta, 1965) continue to treat Linsleya as al- lied to Lytta. This study of courtship behavior was undertaken to pro- vide additional information bearing on this question. Courtship in Linsleya ( Linsleya ) convexa (LeConte) was studied previously but failed to indicate subtribal affinity (Selander and Pinto, 1967). The behavior of a second species, L. ( Linsleyina ) compressicornis (Horn), is described below. This species is of considerable interest since its courtship is remarkably similar to that of several species of Epicautina and unlike any behavior known in the Lyttina 2 . Adults utilized in this study consisted of 30 males and 28 females collected from their host plant, Menodora spinescens Gray (Oleaceae), 2 miles E of Big Pine, Inyo County, California, on 9 June 1973. Sexual isolation was effected 12 hrs after collection and was maintained when- ever observations were not in progress. All observations were made in the laboratory from 12-20 June 1973, at 26-30 °C. Specimens were maintained and observed in plastic cages IIV 2 cm long and 8 cm wide. 1 This study was supported by Grant GB-30907 from the National Science Foundation. 2 This paper follows Selander’s (1964b) classification in treating the Lyttina and Epicautina as subtribes within the Lyttini. Other authors (e.g. MacSwain, 1956) give these two taxa tribal status. The Pan-Pacific Entomologist 50: 1-8. January 1974 2 The Pan-Pacific Entomologist Ethological data are based on six hours of observation utilizing nu- merous randomly selected pairs, and include 200 feet of motion picture films. L. compressicornis has been divided into two subspecies by Selander (1955). This study is based on members of the nominate form. Courtship Behavior All components of courtship in L. compressicornis are performed by the male while positioned behind the female (posterior position). At no time does he mount her, or overtly contact her with his legs in any other way. For descriptive purposes courtship is divided into three phases, a preliminary phase, a display phase, and a genital phase. At the onset of the preliminary phase the male orients directly behind a quiescent female. His maxillary palpi are often extended and ir- regularly touch the female’s pygidium. The position and state of the male’s antennae vary. They may be held motionless and out to the side or, more frequently, with segments II-XI directed downward at an ap- proximate right angle to segment I and the anterior surface of each antenna facing the side of the female’s last abdominal segment. While in the latter position the antennae are often rapidly vibrated antero- posteriorly. Periods of vibration vary in duration from one to 20 sec and comprise, on the average, 50-60% of total courtship time. Antennal vibration is usually repeated several times before the start of the display phase. Due to decamping by the female, however, some bouts never progress beyond preliminary phase activity. Although antennal vibration is the dominant component of courtship in terms of time, its significance is not clear. Contact with the female apparently never occurs. Because of this and the lack of an obviously correlated female response, I have excluded this act from the display phase. Display in L. compressicornis consists of a single act, antennal press- ing (Fig. 1). This act is always preceded by antennal vibration. Prior to pressing the antennae are rotated so that their anterior surface is directed forward and, at the same time, they are moved slightly postero- medially to a position above the midline of the female’s abdomen. Tac- tual stimulation immediately follows and consists of the male pressing down on the last two abdominal tergites of the female one to five times in rapid succession. Both antennae are employed synchronously with the anterior surface of segments IV-XI making contact. A determination of the number of presses normally provided during each bout of display is complicated by female activity. Most females thwart courting males Vol. 50, No. 1, January 1974 3 Figs. 1-2. Courtship behavior in L. compressicornis. Fig. 1. Display phase. Male performing antennal pressing. Fig. 2. Genital phase. Male attempting gen- ital insertion. by either decamping, kicking, or abruptly elevating their abdomen at the onset of display. The majority of bouts, however, consist of three presses. Display was not common in this study. It occurred, on the average, only once during every four minutes of courtship. Display is invariably followed by a genital insertion attempt (genital phase) (Fig. 2). In attempting insertion the male maintains his pos- terior position, elevates his fore legs and, with genitalia partially ex- truded, curls the apex of his abdomen forward through his middle and 4 The Pan-Pacific Entomologist Figs. 3-4. Scanning electron micrographs of right antenna of L. compressicornis (anterior view). Fig. 3. Male (60x)- Fig. 4. Female (50x)- hind legs toward the gonopore of the female. If, as is usually the case, the female does not allow mating, he either returns to preliminary phase activity or ceases courting entirely. Genital insertion attempts only oc- cur after display. In my observations their duration varied from seven to 45 seconds. The display and genital phases appear to be tied behaviorally in this species since insertion is always attempted immediately after pressing. This is even true when the female decamps during display. In this case the male quickly pursues the female and upon continuing courtship sev- eral seconds later resumes with an insertion attempt. Situations where the female successfully eludes the male were not observed. Only a single mating was observed. This particular male inserted im- mediately after display, and within less than five seconds turned to his right and assumed the linear position without contacting the female with his fore legs. The duration of this mating was not recorded. Anatomical Correlates of Courtship Antennal modifications in the male of L. compressicornis are the most obvious correlates of courtship. As shown in Figs. 3 and 4, seg- ments III-XI of the male are more strongly broadened than those of the female, and their anterior surface is concave rather than convex. Ad- ditional modification is found in antennal surface structure. Most of the male’s segments have fewer setae on the anterior (contacting) surface, and segment XI is entirely denuded at its base. Examination of this surface at high magnification with the scanning electron microscope re- Vol. 50, No. 1 , January 1974 5 veals an abundance of minute cuticular pores. These are especially dense at the base of segment XI, the area of greatest density appearing as a small ellipse at 60X (Fig. 3). The function of these pores is unknown. Their association with chemical stimuli has been suggested for other meloines (Matthes, 1969; Pinto, 1973). Another modification in males of L. compressicornis is the highly developed fore tarsal pads. The pale, apically flared setae which com- prise the pads in this species are best developed on the first three seg- ments. In Linsleya convexa, similarly modified fore tarsal pads in the male are considered an adaptation for grasping the female while he is mounted on her (Selander and Pinto, 1967). In L. compressicornis , however, the male does not contact the female with his legs. Instead, the pads may aid the peripatetic males in maneuvering on the host plant surface during their persistent search for females and during courtship. Comparative Behavior and Discussion L. convexa is the only other species of Linsleya for which courtship data are available. This species shares few similarities with L. compres- sicornis besides traits typical of meloines in general 3 . In L. convexa the male courts entirely from a fully mounted position (dorsal position), and display consists of rubbing the dorsum of the female with his hind two pair of tarsi (Selander and Pinto, 1967). The only noteworthy similarity is that males of both species fail to manipulate the antennae of the female but stimulate her dorsum instead. Courtship in L. compressicornis is remarkably similar to that of Pleuropompha and certain species of Epicauta. In both species of Pleu- ropompha and in Epicauta ( Epicauta ) puncticollis Mannerheim (Pinto, 1973), and in E. ( Epicauta ) ruidosana Fall (Pinto, unpubl.) the male also courts entirely from an unmounted posterior position. The genital phase is performed in the same manner in all five species, and although the specifics of display vary, employment of the antennae by the male is characteristic of all but E. ruidosana. In all other nominate Epicauta studied, the genital phase and often the display phase as well are per- formed from a dorsally mounted position (Pinto, 1973). A dorsal mount, at least during part of courtship, is typical of Meloinae. Its com- plete absence has been reported only in the above-mentioned species. The act of antennal pressing in L. compressicornis is roughly similar to display in some other species of nominate Epicauta , namely E. par- dalis LeC., E. magnomaculata Martin, and E. ventralis Werner (Pinto, 3 Courtship traits typical to the Meloinae are discussed by Selander (1964b). 6 The Pan-Pacific Entomologist MS). In these three species the posteriorly positioned male rubs or presses the dorsum of the female with his antennae. The specifics of this act in these Epicauta are sufficiently different, however, as to preclude homology with antennal pressing. As in both species of Linsleya , dis- play in all nominate Epicauta that have been studied involves the stimu- lation of the female’s dorsum in some way (Pinto, 1973), and is never directed toward her antennae. Antennal manipulation is found in most other meloine taxa (Selander, 1964b; Pinto, 1972) including Epicauta subgenus Macrobasis (Selander and Mathieu, 1969). With respect to courtship then, L. compressicornis and certain species of nominate Epicauta are more similar to each other than to congeners. Unfortunately, we can not yet speculate knowledgeably as to whether posterior courtship, their shared pattern, represents the primitive or derived state in the Epicautina. However, it is clear that either posterior or dorsal courtship has been derived independently in the subtribe since the two patterns occur in both Linsleya and Epicauta. Such marked in- terspecific variation in the position of the courting male is surprising considering its constancy in other meloines. In other taxa (e.g. Meloe, Lytta , Eupompha , Pyrota ) dorsal mounting is typical, and the position assumed by the male varies only slightly between species. The adaptive significance of posterior versus dorsal courtship can not be profitably discussed without analyses of several aspects of reproductive behavior and related ecological factors. The similarities of behavior in L. compressicornis and certain Epicauta are marked. To the contrary, there are no important parallels to court- ship in any lyttine studied so far. In species of Lytta the male invariably courts from a fully mounted position, and display, when present, in- volves the manipulation of the female’s antennae (Selander, 1960; Mat- thes, 1972; Gerber and Church, 1973; Pinto, unpubl.). A similar pat- tern has been reported for the lyttine genus Lydus (Cros, 1912). Conclusion In spite of the failure of adult anatomy to clearly indicate relationship, both larval and ethological features can now be cited in support of plac- ing the five species of Linsleya in the Epicautina. The similarities of the first instar larvae of Linsleya, Epicauta and Pleuropompha are nu- merous (see MacSwain, 1956) and, in themselves, leave little doubt as to the affinity of these genera. Selander (1964a) has also pointed out that larvae of Linsleya sphaericollis (Say) appear to be parasitoids of grasshopper egg pods (as in Epicauta) rather than of bee nests (as in Vol. 50, No. 1, January 1974 7 Lytta and related genera). The behavioral data presented above are of importance since they clearly support these conclusions based on larval anatomy and bionomics, and, furthermore, suggest that Linsleya is most similar to the nominate subgenus of Epicauta. As mentioned above, Kaszab (1959) and Gupta (1965) continue to place Linsleya near Lytta 4 . Kaszab ’s rationale for this is not clear. Al- though his 1959 classification of the Meloidae is primarily based on wing venation, the wings of Linsleya are poor indicators of subtribal affinity as they are similar to those of both Epicauta and Lytta. Gupta’s reassignment of Linsleya was prompted by the absence of two structures in L. sphaericollis , V-shaped proventricular folds and well developed stomadaeal valves. Both traits are present in Epicauta and Pleuro- pompha ; other epicautine genera ( Psalydolytta , Denier ella and Anom- alonchus) were not studied. To reassign Linsleya solely on this basis is not convincing. If Linsleya is the most primitive genus of epicautine as both MacSwain (1951) and Selander (1955) believe, its lacking cer- tain specializations of the more derived genera should not be surprising. Acknowledgments Assistance by Messrs. Derham Giuliani and Gregory R. Ballmer is gratefully acknowledged. Figures 1 and 2 were prepared by Mr. Carl T. Conley. Identification of the host plant of L. compressicornis was kindly provided by Mr. Oscar F. Clarke (U. C. Riverside Herbarium). Literature Cited Arnett, R. H., Jr. 1963. The Beetles of the United States (a Manual for Iden- tification.) Wash. D.C., 1112 pp. Cros, A. 1912. Lydus algiricus L. Ses moeurs — Sa larve primaire. La Feuille Jeunes Natur., 42 : 78-86. Gerber, G. H. and N. S. Church. 1973. Courtship and copulation in Lytta nut- talli (Coleoptera: Meloidae). Can. Entomol., 105: 719-724. Gupta, A. P. 1965. The digestive and reproductive systems of the Meloidae (Coleoptera) and their significance in the classification of the family. Ann. Entomol. Soc. Amer., 58: 442-474. Horn, G. H. 1873. Revision of the species of several genera of Meloidae of the United States. Proc. Amer. Phil. Soc., 13: 88-117. Kaszab, Z. 1959. Phylogenetische Beziehungen des Fliigelgeaders der Meloiden (Coleoptera), nebst Beschreibung neuer Gattungen und Alien. Acta Zool. Acad. Sci. Hung., 5: 67-114. MacSwain, J. W. 1951. A new genus of Meloidae from North America. Pan- Pac. Entomol., 27: 58. 4 In a recent classificatory outline of the Meloidae, Kaszab (1969, Mem. Soc. Entomol. Ital., 48: 243) lists Linsleya as an epicautine. 8 The Pan-Pacific Entomologist MacSwain, J. W. 1956. A classification of the first instar larvae of the Meloidae (Coleoptera) . Univ. Calif. Publ. Entomol., 12: 1-182. Matthes, D. 1969. Die Fachelbalz von Cerocoma schafferi (L.) (Coleopt., Meloidae). Zool. Anz. Suppl. — Verhandlungen Zool. Ges., 33: 316-322. Matthes, D. 1972. Die Balz der Spanischen Fliege ( Lytta vesicaloria L.). Zool. Anz., Leipzig, 188: 441-447. Pinto, J. D. 1972. Comparative courtship behavior of Negalius, Phodaga and Cordylospasta , three closely related genera of blister beetles (Coleoptera: Meloidae). J. Kans. Entomol. Soc., 45: 459-476. Pinto, J. D. 1973. Sexual behavior in the genus Pleuropompha LeConte: A new mating display in blister beetles (Coleoptera: Meloidae). Can. Entomol., 105: 957-969. Selander, R. B. 1955. The blister beetle genus Linsleya (Coleoptera: Meloidae). Amer. Mus. Novitates, 1730: 1-30. Selander, R. B. 1964a. The systematic position of the genus Linsleya (Coleop- tera: Meloidae). Proc. Entomol. Soc. Wash., 66: 216. Selander, R. B. 1964b. Sexual behavior in blister beetles (Coleoptera: Meloi- dae) I. The genus Pyrota. Can. Entomol., 96: 1037-1082. Selander, R. B. 1960. Bionomics, systematics, and phylogeny of Lytta, a genus of blister beetles (Coleoptera, Meloidae). Univ. 111. Biol. Monogr., 28: 1-295. Selander, R. B. and J. M. Mathieu. 1969. Ecology, behavior ,and adult anatomy of the Albida Group of the genus Epicauta (Coleoptera, Meloidae). Univ. 111. Biol. Monogr., 41: 1-168. Selander, R. B. and J. D. Pinto. 1967. Sexual behavior in blister beetles (Cole- optera: Meloidae) II. Linsleya convexa. J. Kans. Entomol. Soc., 40: 396-412. Werner, F. G., W. R. Enns and F. H. Parker. 1966. The Meloidae of Arizona. Agr. Exp. Sta. Univ. Ariz. Tech. Bull., 175. Vol. 50, No. 1, January 1974 9 Adult Behavior and Population Biology of Poladryas minuta , and the Relationship of the Texas and Colorado Populations (Lepidoptera: Nymphalidae) James A. Scott Department of Entomology, University of California, Davis This paper is part of a comparative study of behavior and movements of eleven species of diurnal lepidoptera (Hesperiidae and Papilionoidea) , emphasizing the relationship between mate-locating behavior and move- ments (Scott, 1973a; 1973b). Included are studies of mate-locating be- havior by males, mating, movements and lifespan, oviposition, adult and larval foodplants, and basking behavior in Poladryas minuta Ed- wards. In addition I describe experiments in which the Texas sub- species ( minuta ) was hybridized and backcrossed in nature to the Colo- rado subspecies ( arachne Edwards), providing much of the behavioral information. The species was studied in 1969 on a treeless ridge just northwest of Cripple Creek, Teller County, Colorado, and in 1972 on Green Moun- tain, Jefferson County, Colorado. All times are 24-hour standard time. I thank Roy 0. Kendall, San Antonio, Texas, and Kilian Roever, Phoenix, Arizona, for helpful information, Jerry A. Powell, University of California, Berkeley, and Glenn R. Scott, Lakewood, Colorado, for reviewing parts of the manuscript. The University of California, Berke- ley, provided a grant for computer time. Methods Felt-tipped markers were used to give each individual a different number using the method of Ehrlich & Davidson (1960). Marks were placed on the upperside so that the number could usually be determined without capture. Individuals were marked and individually released at the site of capture. The following method of analysis of movements allows direct com- parison between sexes and between species, detection of change of move- ments with age, and separation of the velocity and distance aspects of movements. The capture points for each individual are plotted on sep- arate maps of the study site. The following statistics were determined for each individual: di, tj — distance in meters and time in days between capture i and cap- ture (i + 1) ; D — sum of all d’s of an individual; R — distance between The Pan-Pacific Entomologist 50: 9-22. January 1974 10 The Pan-Pacific Entomologist the two farthest capture points (range) ; T — time between first and last capture; V — overall velocity (D/T) ; Vj— velocity between successive cap- tures (di/tj). Means of the above statistics are computed for all individuals of each sex. Midpoint age is the age midway between two successive captures of an individual. Correlations between the above movement parameters and age determine whether movement changes with age. Population size, survival rates, and number of new insects per day were determined from mark-recapture data using the method of Jolly (1966). Mate-locating Behavior Most butterflies use one of two strategies to bring the sexes together for mating (Scott, 1973b) : in some species males wait at characteristic sites such as hilltops, gullies, or treetops, and locate females by investi- gating passing objects ( perching behavior ). In these species females fly to the perching sites after emergence. In other species males search for females by flying almost continuously ( patrolling behavior ) . P. minuta is the only known species of butterfly (and perhaps insect) which utilizes completely different strategies of mate-location depending on the time of day. In the morning from about 0700 to about 1215 and espe- cially from 0800 to 1130, male P. m. arachne perch on hilltops and high points on ridges. In the afternoon until about 1600, males patrol near flowers, usually on hillsides or flats. Male P. m. minuta in Texas also perch in the morning on ridgetops and patrol in the afternoon on hill- sides. In the morning, perching males alight on bare spots or rocks and dart out at passing fluttering objects (usually other male P. minuta , and often males of Hesperia comma L., another perching, hilltopping spe- cies). Between investigative flights, males often visit flowers or fly short distances to adjacent perches. If they encounter a female during these short flights courtship ensues, resulting in the small percentage of morning copulations initiated by flying males. If a male investigates an adjacent perched male, an “encounter” usually results in which both fly vertically near each other for about 3 m, then separate and return to the ground or repeat the vertical flight. Passing males may be pur- sued briefly. Males usually return to the same or a nearby perch after investigating a passing object, but sometimes move to another spot on the ridge. In the afternoon, the behavior of males changes radically: males patrol rapidly near flowers, briefly pursuing male P. minuta or other butterflies and courting females. Vol. 50, No. 1, January 1974 11 Experiments with females dangling from a fishing pole showed that movement of the wings is necessary to attract a perched male. When the female fluttered, the male approached and then courted if the fe- male was dropped to the ground. Usually the female hung motionless, and was ignored by males, which would sometimes pass by the motion- less female to investigate a more distant much larger black Papilio but- terfly ( minuta is mottled orange). Flight is not an absolute require- ment to attract the male, since in two instances courtship commenced when a female was carefully placed next to a perched male without dis- turbing him. Mating Virgin females were released before perched males for analysis of courtship, and many courtships of native individuals were also observed. To obtain pure minuta matings, the male was released first, then the female was released near the perched male. Thirty-two completed court- ships in five male-female combinations (three pure combinations: mi- nuta $ X minuta $ (1), minuta nympha Edwards S X m. nympha 2 (1), arachne $ X arachne 2 (3) ; two hybrid combinations: arachne $ X minuta 2 (12), arachne $ X ( arachne S X minuta 2) 2 (15)) and many abortive courtships were observed. Courtship in these five combinations was identical, including all the behavioral elements de- scribed below. In the simplest form of courtship the female alights (usually within 3 m of the male’s perch), the male alights behind her and bends his abdomen laterally (11 right, 7 left in successful attempts) to copulate. Four additional components may be present. Male hovering consists of the male rapidly beating his wings at small amplitude a few cm above and downwind of the female for a few seconds. The male then alights behind the female. If the female flies, crawls vigorously away, or flutters during courtship, the male may hover before alighting be- hind her again. Male fluttering is similar to hovering except that the male is on the ground so the wings are moved more slowly, above the horizontal, and with greater amplitude while the body remains sta- tionary. Male fluttering occurred rarely (only 4 successful copulations) and only after alighting while the male was behind the female prior to attempted copulation. Male nudging consists of the male holding the wings about 40° above horizontal, the antennae directed backward, and pushing his head under a spread hind wing of a female, evidently positioning him alongside the female so that his laterally curved ab- 12 The Pan-Pacific Entomologist domen is in position for copulation. If the female’s wings are nearly closed the male immediately crawls alongside and attempts coupling. The male usually keeps his wings about 40-60° above horizontal even when it would appear advantageous to raise them to get closer to the female. Male nudging was observed in many successful courtships, especially early in courtship when the female’s wings were spread. Fe- males usually raised their wings prior to coupling but several males managed to couple by nudging under the female’s spread wings. Fe- male fluttering consists of females holding the wings about 40° above horizontal and fluttering the wings slightly. It occurs while the male is on the ground behind the female. Successful courtship last from 2 seconds to 2 minutes after both in- dividuals alight, but typically requires about 10-15 sec. Unreceptive virgin females discourage males in a variety of ways. They may crawl away, turn, fly a short distance, or flutter their wings (female fluttering) ; they rarely perform the stereotyped rejection dance of mated females (performed early in three successful courtships; see below). Occasionally they may lift the abdomen about 30° above hori- zontal so that the male cannot couple, and they usually keep the wings spread. Males often overcome moderate female unreceptivity by per- sistent hovering, nudging, and attempted coupling. The male crawls or flies behind the female, often hovering briefly when the female flies or crawls rapidly away, or sometimes he hovers after she flutters her wings. After many copulation attempts (up to 20-30 ), the female may become quiescent, raising her wings nearly to the vertical, and moving her abdomen to the horizontal or slightly below so that the male can couple. One female whose wings remained spread raised her abdomen slightly to nearly horizontal so that the nudging male could couple. The main causes of courtship termination by virgin females are: 1) the female eludes the male by crawling or flying so that the male can- not relocate her; 2) after courting a persistently unreceptive female the male flies away. Table 1 shows that male hovering and fluttering are associated with crawling, flying, or fluttering in unreceptive virgin females. In only five courtships was male hovering or fluttering not associated with these female responses. The function of hovering and fluttering seems to be to inform the female of the identity of the male and thereby to make her more receptive, but the male often hovered before landing and before the female could respond, suggesting that male hovering has be- come somewhat ritualized. Vol. 50, No. 1, January 1974 13 Table 1. Association of some of the behavioral elements of courtship in Poladryas minuta subspecies, based on 32 successful courtships with virgin females. Females less than two hours old indicated by (y) ; two days old, (o) ; other females were one day old. See text for descriptions of male and female behavioral elements. Male Behavior Hover- ing & Flut- tering Other* Total Hovering Flut- tering Female Flying 3 ( ly) 1 4 Response Fluttering 2 2 Crawling 2 ( ly ) 1 1 4 Flying and Fluttering 1 1 Flying and Crawling 3 (ly) 1 4 Fluttering and Crawling 5 (3y,lo) l(o) 6 None 4(ly,lo) 1 6(2o) 11 Total 20 1 3 8 32 * Includes nudging or absence of preliminary courtship. Females mated at ages of one hour to two days. Table 1 shows that young females were somewhat less receptive than older females, and that males hover slightly less frequently over older females. Males be- gin courting on the day of emergence, as one P. m. minuta male perched and chased 10 objects on a hilltop in a half hour period two hours after emergence, and attempted to copulate with a female. Another P. m. minuta male mated one day after emergence. Mated, unreceptive females have a rather stereotyped rejection dance. The female, when pursued by a male, slowly flies vertically about three m, then rapidly returns to the ground. The male frequently cannot follow her through the downward part of this flight. The dance is re- peated if the male follows her. This dance is almost identical to the vertical encounter between two perched males, suggesting that the visual similarity may function in discouraging courting males. The female rejection dance occurs in all four taxa studied, and in P. m. nympha in southern Arizona (Fred Thorne, written communication). Many of the elements of behavior during courtship seem to depend on visual cues. The perched male is attracted only to moving objects. 14 The Pan-Pacific Entomologist The rejection dance of mated females visually resembles the vertical encounter between adjacent perching males. Males sometimes hover over red Castilleja flowers possibly because the colors of the flower and butterfly upperside are similar ( Castilleja was not used as a nectar source). Perching males often follow other males which have a slow, linear flight similar to that of females, and often land beside very young released males and occasionally hover and try to copulate. One male grasped a newly emerged male for several minutes, but such homosexual behavior is uncommon. Behavior during copulation. The male usually remains motionless during copulation but the female often opens and closes her wings a few times after coupling, then usually remains motionless. Upon ter- mination, the male resumes mate-locating behavior, while the female remains motionless for several minutes. Duration of copulation depends on the history of prior mating of the male. If the male had terminated mating within the previous hour, copulation lasted 9 hr. and 10 hr. 41 min. (N = 2). If the male had mated the previous day, copulation lasted an average of 54 minutes (20-82, N = 8) . If the male had mated more than 1 day before or had no known history of mating, copulation lasted an average of 26 min. (18-49, N = 21). Some of the latter males may have mated pre- viously so that duration without recent mating of the male may be sig- nificantly less than 26 minutes. Number of matings. Males may mate at least five times. One native arachne male mated 5 times in 4 days, and 4 other arachne males mated at least 3 times each. Immediately after mating, males resume perching and courting, and two males mated only 11 and 15 minutes after ter- minating a previous copulation, but as noted above this resulted in a prolonged mating. Females usually mate only once, very rarely twice. Dissections of 51 field collected females indicated 14 virgins, 36 mated once (one spermatophore) , and only one mated twice ( a fresh female with two abnormally small spermatophores) . The male deposits a clear, solid plug in the female genital orifice. The plug is visible ex- ternally, and may act as a mechanical barrier to further mating. Time and location of mating. Courting and mating occur through- out the day. Six spontaneous matings were observed in the morning hours, and 29 experimental copulations were induced between 0732 to 1211 by releasing females in front of perching males. Many courtships by perching males were observed from 0756 to 1147, and many court- ships by patrolling males were observed from 1300 to 1530. The mating Vol. 50, No. 1, January 1974 15 behavior of males and females remains constant during the day, unlike mate-locating behavior of males, and all types of rejection behavior of females and courtship behavior of males were observed throughout the day. Virgin females were found at all hours, even late in the day, suggesting that some females do not mate until the second day of adult life. In the morning, all observed matings and courtships except one oc- curred on the ridgetops. A single mated pair on the hillside was found 100 m east of a Cripple Creek ridgetop indicating that courtship prob- ably occurred on the hillside. Two female P. m. minuta two days old left a hilltop after release but later returned and mated there. The per- centage of virgin females on ridgetops at Cripple Creek before 1200 was 35% (9 of 26) but only 8% (2 of 24) 20 m or more from a ridgetop. These figures suggest that until females fly to the ridgetops they remain near the sites of emergence, which are mainly on hillsides near Pen- stemon. After mating, females leave the ridgetops. In the afternoon, courtships occur mainly on hillsides and flat areas near the morning perching sites, where virgin and mated females often alight on flowers to feed. Both sexes usually emerge from pupae before 1100, males an average of a day earlier than females. In the afternoon males apparently ac- tively search for females which emerge that day, then wait on ridges the next morning for the females which did not mate on the day of emergence. Some females may immediately fly to ridgetops, but as shown above females are more receptive a day or more after emergence. Movements In 1969 males were marked and recaptured for 10 days along the top of a ridge at Cripple Creek, mainly in the morning (Fig. 1, Table 2). Males flew along the entire ridgetop, moving a distance (D) up to 470 m. The moderately high density with resultant numerous chases between males may have contributed to the high movement on the ridge, but it is more probable that the movement occurred because the ridge contained suitable perching spots and flowers along most of it length. At the Bear Creek study area described below, the hilltops and perch- ing sites were discontinuous and movement was less. Males are not territorial, as they wandered over the entire ridge and perched wherever suitable sites occurred. In 1970, 20 males were marked and released at Bear Creek, Chaffee County, Colorado. Six of these were recaptured. Over a two-day pe- 16 The Pan-Pacific Entomologist Fig. 1. Maps of Cripple Creek (A) and Bear Creek (B) sites. Contour interval 3 meters (A) and 6 meters (B). Numbered hilltops correspond to foci of activity discussed in text. riod only one of these males stayed where marked. Two males moved considerable distances from a hilltop to a ridge and vice versa. Three males moved from hilltop 30 (Fig. 1) in the morning to flowers on the hillside in the afternoon: one of these, observed on hilltop 30 in the morning of two successive days, appeared on the hillside visiting flowers on the afternoon of the next day, and returned to hilltop 30 the follow- ing morning. Another male remained on hilltop 30 in the morning and the hillside to the west in the afternoon for three days, then was found on hilltop 33 on the sixth day. Still another male resided on hilltop 30 during the mornings of three days in a 4-day span, and moved to the hillside west of hilltop 30 on two afternoons during this time. The last two examples suggest that males return to the same hilltop and hillside areas repeatedly, but this is probably because hilltop 30 and the hill- Vol. 50, No. 1, January 1974 17 Table 2. Movement data for Poladryas minuta arachne at Cripple Creek in 1969. T = time between first and last capture; t i? dj = time, and distance respectively, between capture i and capture (i + 1) ; R = distance between the two farthest capture points ; D = sum of all d’s of an individual; V = D/T; Vi = dj/h; * = p ^ 0.01 (other correlations p > 0.05). Significance of correlations was determined by tests of zero correlation. Movement Parameter X N Parameter Movement X N T (days) 3.66 68 di (meters) 56 151 ti (days) 1.65 151 V (meters/day) 42 68 R (meters) 93 68 Vi (meters/day) 35 151 D (meters) 125 68 r (ds, ts) = .270* r (di, age) r = .003 r (ti, age) = .184 Partial correlation (di, ti) , age constant - .275* r (vi, age) = -0.006 Partial correlation (di, age), ti constant rz -0.049 side to the west are the most favorable (the most prominent hilltop, and the nearest hillside with many flowers) sites in the immediate vicinity of the Bear Creek locality. Effect of copulation and age on movements. Males which copu- lated on Green Mountain were marked and their movements were stud- ied. The results suggest that 1) males do not need to have experience at a site to copulate there, and 2) copulation does not cause the male to remain at that spot. For example, two arachne males and a minuta male which were caught elsewhere on Green Mountain or laboratory reared were released in the morning on a small ridgetop where they immedi- ately perched and mated within 5 to 15 min. of release. Other arachne and minuta males perched immediately upon release, showing that, in the morning at least, males perch and mate in suitable topographic sites regardless of “territorial” experience. That males do not remain in areas of previous copulations is shown by their frequent movements. For example, one male mated twice on hill A on one day, flew elsewhere the next day, moved to hill B the third day, mated on hill A the sixth day, then spent the remainder of the sixth day on hill B. Another male mated on hills, A, A, and B on three successive days. Two males mated on hills B, A, and A on successive days; one of these courted on hill A the fourth day. Only one male mated many times on one hill (5 times in 4 days) . 18 The Pan-Pacific Entomologist Table 3. Population parameters of Poladryas minuta arachne males at Cripple Creek, 1969, estimated from multiple recapture data using the stochastic model of Jolly (1966). N = total population size; Phi = probability of survival from each sampling period to the next; B = number of new animals joining the population during that sampling period ; SE = standard error. Day N ± 1.96 SE Phi ± 1.96 SE B ± 1.96 SE August 12 — — .727 .342 — ■ — 14 154.7 120.4 .847 .381 118.9 163.9 15 249.1 158.6 .953 .408 7.8 160.4 16 245.1 131.7 .637 .314 132.7 119.8 17 287.1 140.8 .693 .441 146.9 199.9 18 300.0 228.5 .813 .494 63.4 188.5 19 273.0 151.1 .923 .481 81.9 136.1 20 267.4 127.9 — — — ■ — Correlations of movement parameters with age for males (Table 2) show positive correlations of dj with time between recaptures. Time between captures (tj increases slightly but not significantly with age, which may indicate less active flight, and hence smaller probability of capture, with age. Male Population Parameters A 10 day mark-recapture study of males at the Cripple Creek site indicated a population size of approximately 200, with an initial in- crease then a decrease (Table 3). An average of about 80 individuals per day joined the population. Average survival rates and expected life- span were .794 (4.3 days) using method 1, and .817 (5.0 days) using method 2 of Scott (1973a). Probably a small portion of the population emigrated from the area, so that the survival rate should be increased slightly. The potential lifespan is longer than this, of course. The two longest lived males survived at least nine days and nine others seven to eight days. Longer times might have been recorded if the mark-recapture study had been longer. Feeding and Oviposition Individuals of both sexes and all ages frequently visit flowers, espe- cially during the hottest part of the day. Males feed on flowers occasion- ally during the morning perching period, but more frequently in after- Vol. 50, No. 1, January 1974 19 noon. Both sexes visit yellow flowers most frequently, but also visit flowers of other colors. At Cripple Creek 84 individuals were observed on Heterotheca villosa (Pursh) Shinners and 16 were observed on other yellow flowers, while only 19 visits were recorded to white, orange, or blue flowers. Proportions were similar at other localities. Oviposition occurs as the female slowly flutters among small Pen- stemon plants which are usually without inflorescences (individuals have never been observed feeding on the flowers, which are blue, white, or red). Eventually she deposits a round cluster averaging 38 eggs (14— 87; N = 43 ) , usually on the lower surface of one of the lower leaves. Daily eggs counts of 16 laboratory reared P. m. minuta and Fi females mated on Green Mountain indicate a preoviposition period of 1-4 (usually 1-3) days. Most females deposited several clusters on the same day or over a period of several days, but several first oviposited on about the third day, then laid other clusters several days later. Larval foodplants. Larvae feed on various Penstemon species in different geographic areas, including P. alhidus Nutt, in Baylor County, Texas (P. m. minuta ; identified by Roy 0. Kendall), P. dasyphyllis Gray in Santa Cruz County, Arizona (P. m. nympha) , P. virgatus ari- zonicus Heller on Mt. Graham, Arizona (P. m. near arachne ; both Lil- ian Roever, written communication), and P. alpinus Torr. in Larimer County, Colorado (P. m. arachne ; Sperry & Sperry, 1932). Oviposition was on P. harbatus torreyi (Benth.) Keck in Boulder County, Colorado (P. 772. arachne ; Emmel et al. 1971), and on P. secundiflorus Benth. at Cripple Creek. Many unidentified Penstemon were fed to the larvae in the laboratory, with the following general results. The herbaceous or shrubby species with green, mostly hairless, delicate foliage were moderately or very acceptable to the larvae. A cultivated variety (“pin- nifolius”) with small needle-like leaves was moderately acceptable. The herbaceous species with thick leaves covered with a whitish or bluish bloom were completely unacceptable. P. m. minuta larvae readily ate the leaves of P. albidus , and ate lesser amounts of other Penstemon including P. barbatus. The Fi, backcross, and P. m. arachne larvae fed readily on most species tested (with the exception of the glaucous spe- cies) , including P. barbatus. The palatability of these Penstemon species is similar for grazing animals (Forest Service, 1937), suggesting that grazing may have caused the extinction of P. m. minuta over most of its former range in Texas (Kendall, 1971). First and second instars are gregarious feeders, but later instars be- come solitary due to mortality and dispersal. Developmental period 20 The Pan-Pacific Entomologist from egg to adult ranges from 46 to 57 days indoors at about 20° C (egg stage 8-10 days, pupal stage 11-14 days). There are 4-5 broods in Texas and 2—3 in Colorado, but emergence is asynchronous; distinct broods were vaguely defined at Cripple Creek and almost constant num- bers were observed on Green Mountain from early June to early Sep- tember. Diapause may occur in half-grown larvae. Hybridization Experiments Hybridization of these subspecies is significant not only in providing much of the behavioral data presented, but also because they have been considered to be distinct species (Bauer, 1961) . P. m. minuta was reared from larvae collected in Baylor County, Texas, and females were re- leased in front of wild perching males of P. m. arachne on Green Moun- tain and south of Glenwood Springs, Garfield County, Colorado. Some resultant Fi females were later mated to wild arachne males on Green Mountain and the resulting backcross larvae raised to adults. Sex ratio of Fi and backcross adults was approximately 1:1. Among 11 minuta 2 ( X arachne $ ) crosses, two produced no eggs, three infertile eggs, and six fertile eggs which produced adults. Among 14 Ft $ (X arachne $ ) crosses, six females failed to oviposit, while eight laid many com- pletely fertile eggs which produced many adults. Among 15 controls (arachne $ X arachne $ ) nine females failed to oviposit, eggs of two females were infertile, offspring of two females died as first or second instar larve (apparently due to disease), and two females produced many viable offspring. P. m. minuta and P. m. arachne are morpho- logically very similar. Adults differ slightly in details of wing, antennal and abdominal color pattern, and larvae differ in the ground color be- tween the scoli of the dorsum of late instars (red in minuta , largely white in arachne ). Mr. Jack Harry, Denver, Colorado, collected a large series near Guadeloupe Peak, Culberson County, Texas, which although more similar to minuta than arachne , is clearly intermediate in several wing pattern characteristics. In addition the two subspecies are very similar in foodplant preferences and in all aspects of courtship behavior, as shown previously. These similarities, together with the high degree of genetic compatibility, clearly demonstrate the conspecificity of these two taxa. Discussion The most unusual feature of behavior in P. minuta is the alteration of mate-locating behavior during the day. This alteration appears to be Vol. 50, No. 1, January 1974 21 well adapted to the search for flowers and virgin females. In the morn- ing, before the females emerging that day are able to fly to ridgetops, males wait for females, which are more active and receptive after their first day of adult life, to come to ridgetops. In afternoon, males search throughout the habitat near flowers, often on hillsides and flats, for virgin females which often feed on flowers at that time. In the morning, temperatures are generally lower than in afternoon; males bask at tem- peratures too low for prolonged flight. In afternoon higher temperature results in more flower feeding, and enables continuous searching flight. Summary Males perch on hilltops and ridgetops from 0700 to about 1215 to locate females, and dart out at passing fluttering objects. Females fly to ridgetops to mate. In the afternoon, males patrol on hillsides and flats near flowers in search of females. Courtship is quite variable, as several male and female behavioral elements are present or absent de- pending partly on the receptivity of the female, which in turn depends partly on age. Unreceptive virgin females have a variety of means of discouraging males, but unreceptive mated females use a stereotyped vertical flight which resembles the encounter between adjacent perching males, and visual stimuli are of general importance both before and during courtship. Males are not territorial. Movement is most pro- nounced along continuous ridges, decreasing in those habitats with isolated hilltops. Both sexes feed predominantly at yellow flowers. Fe- males deposit clusters averaging 38 eggs on various species of Pen- stemon , usually beginning 2-4 days after emergence. P. m. minuta from Texas was hybridized and backcrossed with P. m. arachne from Colo- rado by use of natural courtships. Lack of barriers to interbreeding, and the similarity of adult behavior, larval and adult morphology, and foodplants, confirm the conspecificity of these two allopatric subspecies. Literature Cited Bauer, D. L. 1961. p. 131 In-. Ehrlich, P. R., & A. H. Ehrlich. How to know the butterflies. Wm. C. Brown, Dubuque, la. Ehrlich, P. and S. Davidson. 1960. Techniques for capture-recapture studies of Lepidoptera populations. J. Lepid. Soc., 14: 227-230. Emmel, J., 0. Shields, and D. Breedlove. 1971. Larval foodplant records for North American Rhopalocera. Part 2. J. Res. Lepid., 9: 233-242. Forest Service, U. S. 1937. Range Plant Handbook. U. S. Government Printing Office, Washington. Jae, R. 1972. Natural interbreeding of close nymphalid groups. J. Lepid. Soc. 26: 28. 22 The Pan-Pacific Entomologist Jolly, G. 1966. Explicit estimates from capture-recapture data with both death and immigration — stochastic model. Biometrika, 52: 225-247. Kendall, R. O. 1971. Lepidoptera in the unpublished field notes of Howard George Lacey, naturalist (1856-1929). J. Lepid. Soc., 25: 29-44. Scott, J. A. 1973a. Population biology and adult behavior of Lycaena arota (Lyacaenidae) . J. Lepid. Soc., in press. Scott, J. A. 1973b. Mate-locating behavior of butterflies. Amer. Midi. Nat., in press. Scott, J. A. 1973c. Mating of butterflies. J. Res. Lepid., 11: 99-127. Sperry, G. H., and J. Sperry. 1932. Notes on the larva of Melitaea pola Bdv. Bull. So. Calif. Acad. Sci., 31: 8. SCIENTIFIC NOTE Notes on Hesperus arizonicus with description of the female (Coleoptera: Staphylinidae ). — Hesperus arizonicus was described by Moore in 1958 (Trans. San Diego Soc. Natur. Hist. 12: 311) on the basis of a single male specimen from Patagonia, Santa Cruz County, Arizona. A single female specimen in the collection of the University of California at Riverside is from Cave Creek Ranch, elevation 5000', Cochise County, Arizona, taken on August 1, 1965 at ultraviolet light by G. R. Ballmer. It differs from the male in numerous respects. Description of the female. Color largely ferruginous; third through tenth antennomeres dusky; elytra with yellow band across basal third hounded basally and apically by a narrow piceous band, apical third ferruginous; fifth abdominal segment piceous at apical two-thirds; sixth abdominal segment yellow. Head wider than long, slightly wider than pronotum, dorsal surface feebly microreticulate, with coarse punctures generally separated by less than their diameters except for small elongate impunctate central area anteriorly, and area of nearly coalescing punctures behind eyes. Pronotum subquadrate, slightly longer than wide, slightly narrowed behind, sides somewhat sinuate before posterior angles; surface feebly microreticulate, not impresesd, as coarsely punctured as head but more sparsely so, with punctures generally separated by their width, and with a central impunctate strip. Elytra quadrate, wider and longer than pronotum; humeri broadly rounded, sides straight and hardly divergent to narrowly rounded exterior apical angles; surface finely, densely, and rather roughly punctured. Abdomen narrower than elytra, tapering slightly to apex; dorsal and ventral surfaces with fine, longitudinal microstrigulation, about as finely punctured as elytra but more sparsely so and not as roughly ; last sternite evenly arcuate at apex. The most important structural difference between this female and the holotype male is in the unmodified apical margin of the sixth sternite, a condition typical of many Staphylinidae. There is also considerable difference in color, some in ground sculpture and in the fact that the surface of the pronotum is not impressed. These latter differences are probably within the range of individual variation for the species and not of sexual significance. — Ian Moore, University of California, Riverside, 92502. Vol. 50, No. 1, January 1974 23 Vaejovis spicatus: A New Scorpion from California (Scorpionida: Vaejovidae) Richard M. Haradon San Mateo, California Described below is a new vaejovid scorpion species, discovered in tbe Little San Bernardino Mountains of southern California. A remarkably developed subaculear proturberance on the telson of this species suggests the name. Vaejovis spicatus Haradon, new species (Figs. 1-7) Diagnosis. — Males unknown. Females of V. spicatus are distinguishable from all other known species of Vaejovis by the presence of an elongate subaculear “tooth” (Figs. 4, 5). Females somewhat similar to those of Vaejovis joshuaensis Soleglad (1972, p. 190), also from southern California, but differ as follows: (1) fixed pedipalp finger about equal to or slightly shorter (not distinctly longer) than manus; (2) six and seven interior lateral granules on fixed and movable pedipalp fingers respectively (do not vary on each finger from four to six) ; (3) meta- somal segment V width greater than (not about equal to) half its length, and greater (not less) than width of segment I; (4) bristles along dorsal keels of metasomal segments I-IV, 0, 1, 1, 2 (not 0, 0, 1, 1) ; (5) total adult length about 17 mm (not 24 mm) . Description of holotype (adult female). — Size. — Measurements given in Table 1. Coloration. — Ground color brownish-yellow, fingers slightly darker; no contrast- ing markings. Granules along carinae usually reddish-brown. Tip of aculeus red. Carapace. — Anterior margin concave, with slight median notch. Entire surface finely granular. Median furrows narrow, shallow; posterior laterals broad, very shallow; all other furrows and associated keels obsolete. At least two pairs of lateral eyes, third indefinite. Median ocular tubercle set in shallow depression, forward of center; posterior carapace length/carapace length = 0.65. Chelicerae. — Superior margin of fixed digit with basal bicusp, single adjacent tooth, and terminating in single tine; inferior margin smooth. Movable digit bi- furcate distally, superior tine about % length of inferior; superior margin with two small subapical teeth, adjacent larger tooth, and fourth small, basal; serrula along smooth inferior margin, not extending to apex. Long white hairs along in- ferior and interior basal surfaces of fixed digit. Single dorsal bristle just posterior to digital commissure. Pedipalps. — Sparsely hirsute. Interior keels of humerus and brachium distinct and granular; exterior dorsal keels weak, lightly granular. Intercarinal areas uni- formly finely granular. Chelae relatively slender; manus contour somewhat rounded; all keels weakly developed; digital smooth, inner secondary very lightly granular, interior marginal vestigial. Movable digit distinctly longer than manus; interior margins with closely set, pointed denticles in seven (movable digit) and Tile Pan-Pacific Entomologist 50: 23-27. January 1974 24 The Pan-Pacific Entomologist Figs. 1-5. Vaejovis spicatus Haradon, new species. Holotype female. 1. Right chela, trichobothria ventral-exterior surfaces. 2. Right brachium, trichobothria exterior surface. 3. Right chela, trichobothria dorsal surface. 4. Telson, lateral view. 5. Telson, ventral view. six (fixed digit) rows, each row marked distally by enlarged denticle and adjacent interior lateral granule; slight even space between closed fingers. Trichobothria: humerus, one each proximally on interior, dorsal and exterior surfaces; brachium, two ventral, one interior, two dorsal, 14 exterior (Fig. 2) ; chela, two on interior base of fixed digit, five on exterior surface of manus (Fig. 1), 19 on fixed digit and manus viewed doreally (Fig. 3) . W diking legs . — Generally smooth ; all keels very weak or obsolete. Conspicuous bristles sparse on all segments. Single row of short white bristles ventrally in line with unguicular spine along entire length of distal tarsomere. Genital plate . — Completely fused medially. Pectines . — Moderately hirsute; extending to trochanter of fourth walking leg; 11 teeth overlapping slightly in series; six oval to subcircular middle lamellae; fulcrae subcircular. Mesosoma . — Terga extremely finely granular; vestigial median keel on terga four to seven ; two pairs of weak lateral keels on seven. Sterna smooth to extremely finely granular; lateral keels on last sternum weak, granular; stigmata short, slit-like. Metasoma . — Segments I— II wider than long, III about as wide as long. Keels: dorsals serrate, terminating in enlarged conical spine; superior laterals I-IV ser- rate, with enlarged spine on I— III and fin-like process posteriorly on IV, low Vol. 50, No. 1, January 1974 25 Figs. 6-7. Vaejovis spicatus Haradon, new species. Holotype female. 6. Dor- sal view. 7. Ventral view. rounded granules on V ; laterals granular in posterior % on I— II, posterior % on III, absent on IV, weak with scattered rounded granules in anterior % on V. In- ferior laterals similar to corresponding inferior medians; I weakly crenulate; II crenulate; III crenulate to finely serrate; IV-V serrate. Inferior median keels I-IV set with 3, 3, 3, 3 bristles; dorsal keels with 0, 1, 1, 2 bristles. Telson . — Vesicle laterally swollen; surface extremely finely granular; 16 long slender hairs on ventral and lateral surfaces. Aculeus short, sharply curved, with elongate subaculear protuberance (Figs. 4, 5) . Variation . — The only other adult specimen, also a female, does not differ sig- nificantly from the description of the holotype. Three minute immature specimens, of undetermined instar, were characterized most obviously by differences in ana- tomical proportions, lack of pigmentation, and incomplete carinal development; the subaculear tooth was well developed. Material. — Holotype from Berdoo Canyon, 6.9 miles NE of junction with Dilllon Road, Little San Bernadino Mountains, Riverside County, California (elevation 3600 feet), 31 March 1972, R. M. Haradon and J. L. Marks. Four paratypes from Berdoo Canyon, 3.2 to 5 miles NE of Dillon Road (elevation 2000-2500 feet). 18 26 The Pan-Pacific Entomologist Table 1 . Measurements (in millimeters) of adult Vaejovis spicatus Haradon, new species. Holotype (female) Paratype (female) Total length 17.30 16.10 Carapace length 2.35 2.25 Anterior width 1.15 1.10 Width at/of ocular tubercle 1.65/0.25 1.60/0.25 Posterior width 2.00 1.85 Mesosoma length (sum) 6.10 5.65 Metasoma length (sum) 6.60 6.05 segment I length/width 0.90/1.15 0.80/1.10 segment II length/ width 1.05/1.15 0.95/1.10 segment III length/width 1.15/1.15 1.05/1.10 segment IV length/ width 1.35/1.25 1.25/1.20 segment V length/width 2.15/1.30 2.00/1.20 Telson length 2.25 2.15 Vesicle length/width 1.65/1.05 1.60/1.15 Vesicle depth 0.80 0.80 Aculeus length 0.60 0.55 Pedipalp humerus length/width 2.00/0.60 1.90/0.55 brachium length/width 2.30/0.70 2.15/0.65 manus length/width 1.85/1.00 1.80/1.00 chelal/fixed digit lengths 3.65/1.80 3.50/1.70 movable digit length 2.20 2.10 Pectinal teeth (left/right) 11/11 10/11 Middle lamellae (left/right) 6/6 6/6 Margin lengths, anterior/dentate 1.35/1.10 1.25/1.00 March, 9 April and 5 May 1972, and 24 February 1973, R. M. Haradon and J. L. Marks. The holotype has been deposited at the California Academy of Sciences, San Francisco. Remarks. — The type locality and only known habitat is a narrow, sparsely vegetated, desert canyon. All specimens of V. spicatus were found on rocky canyon walls, by ultraviolet light at night. Also at the type locality occur at least six other scorpion species: Hadrurus ari- zonensis Ewing, Paruroctonus vachoni Stahnke, Vaejovis confusus Stahnke, Vaejovis deserticola Williams, Vaejovis joshuaensis Soleglad, and Vejovis hirsuticauda Banks. Acknowledgments. — Stanley C. Williams, California State Univer- Vol. 50, No. 1, January 1974 27 sity, San Francisco, kindly reviewed the original manuscript. Photo- graphic credit is due R. H. Howard, Los Angeles. Joseph L. Marks, Santa Monica, generously contributed much time and effort in the field, and is properly credited as the co-discoverer of V. spicatus. Literature Cited Soleglad, M. E. 1972. Two new scorpions of the wupatkiensis group of the genus Vejovis. Wasmann J. Biol., 30: 179-195. BOOK NOTICE Eastern Forest Insects. Whiteford L. Baker. U.S. Department of Agriculture, Forest Service, Miscellaneous Publications no. 1174, 642 pp. 1972. $5.00. This completely revised version of an earlier edition (For. Ser. Misc. Publ. 657) contains concise but informative accounts of the life histories of the important in- sects destructive to forest trees in eastern North America. The text has been ex- tensively rewritten and many new photographs included. The cited literature in- cludes over 800 entries, nearly all more recent than 1940, and both the insects and their host plants are indexed, which should make this volume an extremely valuable successor to the earlier work. — Editor. RECENT LITERATURE Ants of Deep Canyon. George C. Wheeler and Jeanette Wheeler. Philip L. Boyd Deep Canyon Desert Research Center, University of California, Riverside. 162 pp. 1973. $3.95. In this paperback volume the Wheelers have produced a highly interesting sum- mary of the biology of desert ants as well as a useful taxonomic treatment of the genera which occur in the arid regions of North America. Discussions of the eco- logical characteristics of the North American hot deserts and some of the morpho- logical, physiological and behavioral adaptations of desert ants are followed by de- tailed taxonomic and ecological discussions of the species occurring in Deep Canyon. The text is profusely illustrated with photographs of the ants, their nests, and typ- ical habitats, and provided with a glossary of technical terms. — Editor. 28 The Pan-Pacific Entomologist The Maritime Earwig in California (Dermaptera: Carcinophoridae) Robert L. Langston 31 Windsor Ave., Kensington, Calif. 94708 During an extensive survey of earwigs in California, particular em- phasis was focused on the maritime or seaside earwig, Anisolabis mari- tima (Gene). My special interest was due to the lack of material in most California museums, and to the rather narrow ecological habitat of this species. Based on the classification of Popham (1965), the placement of An- isolabis is in the subfamily Carcinophorinae under the Labiodea. Within California, this is the only species of the genus found under natural conditions. It is morphologically distinct, and is rarely con- fused with any other earwig occurring in the state. A condensed synonymy is given below to indicate only the original references for each generic change, combined with the first records for the United States and also California. Anisolabis maritima (Gene) Forficula maritima Gene, 1832, Sagg. Monogr. Forfic. Indig., p. 9. [Nice, France; Genoa and Tuscany, Italy; along Mediterranean.] Forficula ( Labidura ) maritima, Fischer, 1853, Orth. Eur., p. 68, pi. 6, figs. 4, 4a-d. F orficesila maritima, Serville, 1853, Ins. Orth., p. 27. Anisolabis maritima, Fieber, 1853, Lotos, III, p. 257. [South Carolina.] (Generic assignment and first United States record.) Brachylabis maritima, Dohrn, 1864, Stettin. Ent. Zeit. 25, p. 293. Anisolabis maritima, Essig, 1922, Pomona College Jour. Ent. & Zool., p. 75. [La- guna Beach, California,] (First California record.) Diagnostic Features and Life History The adult maritime or seaside earwig measures 20-32 mm including forceps. It is wingless, with the body shiny black, or sometimes very dark brown. The uniformly dark colored antennae vary from 20 to 24 segments in the adults, with a lesser number in the juveniles. (One to several lighter segments are diagnostic for some other species of ear- wigs.) The legs are pale yellow (dark banding on the legs is character- istic of a smaller, closely related species commonly found in California) . The forceps of the male are curved or sickle-shaped, and asymmetrical, the right being shorter. The forceps of the female are basally parallel, becoming curved slightly inward near the tips. The Pan-Pacific Entomologist 50: 28-34. January 1974 Vol. 50, No. 1, January 1974 29 The immatures have the body shape, color and straight forceps of the mature females, but juveniles can be differentiated by the number of segments in the antennae; the number increases with each successive stage. Preserved tenerals may remain brown, as in some of the faded specimens in museum collections. Maritime earwigs appear to be primarily predaceous. In the labora- tory they have been observed by Fulton (1924) to catch and devour crickets, sand-fleas and smaller earwigs, using their forceps. Foraging occurs at night. In the daytime I found these earwigs under rocks, logs, driftwood and various other objects washed up along shores where there is salt or brackish water, but they quickly crawl into cracks and crevices when exposed. Because of the flexible abdomen and the strong pinch- ing ability, they are most easily collected with forceps. Adults and juveniles have been collected during all seasons of the year. They occur in definite colonies, and if a single individual is found, usually many (up to several dozen) can be collected in the im- mediate vicinity. In my experience, there was a predominance of fe- males at all collecting sites. Based upon seven years (1964-69, 73) of collecting in California, adult females and the larger juveniles (third, fourth and fifth instars) were found in the winter and early spring months. Depending upon the climate, availability of food, and other factors, the instars tend to over- lap in size, and even the adults have considerable size variability. Some of the dwarf adults may be just slightly over half the size of the ma- jority of individuals. In late spring, summer and autumn, males were found in the colonies along with females and all stages of juveniles. Eggs were found in deep cracks or burrows from early summer to autumn, but are probably pres- ent at other times. The female guards her egg clutch, which is probably deposited over a period of several days. In one instance, 84 eggs were found tended by a single female. In another case, 88 eggs guarded by two females were unearthed from a chamber between rocks and dirt. Females maintain the eggs in neat piles, turn them over, and keep them free of dirt. The eggs are glistening white, nearly spherical and slightly over 1 mm in diameter when deposited. As the embryos develop, the eggs elongate to nearly 2 mm and darken. In California, Anisolabis maritima is found only under very restricted ecological conditions. Most individuals have been collected near the high tide level at San Francisco Bay, San Pablo Bay and Carquinez Strait (Langston, 1967). Below this level the conditions appear to be 30 The Pan-Pacific Entomologist too wet, with the water often seeping into the depressions beneath rocks or logs. In this zone the predominant animal life consists of Isopoda (kelp sowbugs), and other Crustacea. Above the high tide level the substrate becomes increasingly drier. In this zone the predominant animal life consists of Isopoda (dooryard sowbugs, common pillbugs), with the commoner insects including silverfish, crickets, European ear- wigs, carabid and staphylinid beetles, and ants. Many of the areas where this earwig was found were very rocky with considerable drift and de- bris, hut very little noticeable plant life. It is probable that A. maritima moves up and down from the high tide level to feed on most of the ani- mal forms mentioned above, except probably the larger Crustacea. Caudel (1913) stated that A. maritima is found only along the sea- shore. In California it seems to prefer bays and inlets (Langston, 1967) . Although generally distributed along the Gulf and Atlantic coasts, the established colonies in California are mostly restricted (see Fig. 1), with few collections from along the open ocean coasts. Absence of many records on the immediate coast is not due to poor collecting, as two other species of earwigs, Euborellia annulipes (Lucas) and Forficula auricularia Linnaeus, have been taken abundantly. There are many records of both species either on or just above the seashore in north- ern, central and southern California. The first published report on the occurrence of A. maritima in Cal- ifornia was by Essig (1922) , who found it at Laguna Beach and vicinity during June and July, 1921. In 1944, a single male was collected at Costa Mesa also in Orange County. It has been taken in quarantine by the California Department of Agriculture at six localities in southern California. Geographical Distribution Anisolabis maritima is worldwide in distribution. It has been re- corded from most of the continents and major islands, with the excep- tion of the Arctic and Antarctic regions. Specifically, it has been re- ported from France, Italy, along the Mediterranean (Hebard, 1917) ; Canary Islands, Formosa, Japan, New Zealand, Puerto Rico, Santo Do- mingo (Burr, 1910); and Venezuela (Rehn, 1949). Published North American records include British Columbia, Canada (Hebard, 1933; Heifer, 1963) ; and the Gulf and Atlantic coasts of the United States as far north as Maine (Hebard, 1917) . California distribution : — The detailed California records include the term “juv.” for immature specimens, which are similar to the females until the penultimate stage. In this stage and the adults, the forceps dif- Vol. 50, No. 1 , January 1974 31 Fig. 1 . California distribution of Anisolabis maritima. Circles denote established populations. Triangles denote quarantine records. fer markedly between the sexes. Therefore, the specimens are not sexed in the detailed collection records unless close to, or in the adult stage. Established records of specimens are believed to represent naturalized populations. These include localities where large numbers were taken 32 The Pan-Pacific Entomologist or small numbers were accumulated over a period of years. A single record may be considered established if it is within the over-all distri- bution where there are adjacent or contiguous records. Quarantine rec- ords indicate specimens taken at border or ports-of-entry stations by the California Department of Agriculture. Quarantine also includes the within-state or county records where specimens were intercepted. These border and within-state records usually show the point-of-origin, and/ or the host material. The following initials refer to the institutions where specimens are on deposit: BVC = Bureau of Vector Control, California Public Health Service, Berkeley; CAS = California Academy of Sciences, San Fran- cisco; CDA = California Department of Agriculture, Sacramento; CIS = California Insect Survey, University of California, Berkeley; CSH = California State University at Hayward; LACM = Los Angeles County Museum of Natural History, Los Angeles; SFS = San Francisco State University, San Francisco; UCD = University of California, Davis; and UCR = University of California, Riverside. Alameda County: Alameda, 1 juv., No date or collector (CDA) ; Albany, 2 $ 2, X-17-1965 (R. L. Langston— CIS) ; Berkeley, 6 $ $ , 24 2 2, XII-16-1949, 40 $ $ . XII-24-1949, shore of S. F. Bay (J. W. MacSwain — CIS) ; Emeryville, 2 $ $ , 20 $ $ , 20 juv., X-17-1965, 4 $ 2 , 4 juv., 1-15-1966, mud flats (R. L. L.— CAS, CDA, CIS, LACM, UCD, UCR) ; Oakland, VII-17-1957 (N. L. Jones— CDA). Contra Costa County: Point Molate, Richmond, 1 $ , 4 2 2, III-23-1962 (A. G. Raske — CIS) ,12 $ 2 , 19 juv., 11-13-1966, 24 $ $ , 26 2 2, V-19-1966, shore of S. F. Bay (R. L. Langston— CAS, CDA, CIS, CSH, LACM, SFS, UCD, UCR), 23 S $ , 41 2 2 , 20 juv., III-26-1966 (R. L. L. & P. A. Opler — CAS, CIS) ; Point Richmond, 4 $ $ , 12 2 2,8 juv., V-19-1966, 2 $ $, 10 2 2,6 juv., V-6-1967, shore of S. F. Bay (R. L. L. — CIS, LACM) ; Pinole Point, 3 $ $ , 12 2 2,6 juv., IX-29-1973 (R. L. L. — BVC, CAS) ; Point San Pablo, Richmond, 2 $ $ , 8 2 2, IX-16-1967, 11 S $, 8 2 2, HI-9-1968, 15 $ $ , 25 2 2 , 20 juv., IX-21-1973, shore of San Pablo Bay (R. L. L.— BVC, CAS, LACM) ; Port Costa, 1 mi. W., 2 <$ £, 15 2 2 , V-30-1964, 7 2 2, VI-27-1964, 1 $ , 1-17-1965, 8 2 2,8 juv., 11-27-1966, shore of Carquinez Strait (R. L. L. — CAS, CDA, CIS, LACM, UCD, UCR) . Los Angeles County: Los Angeles, IV-10-1940, Quarantine from San Francisco on dahlias (CDA) ; San Gabriel, III-3-1941, Taken in quarantine on chrysanthemum (CDA). Marin County: Belvedere, 1 2, V-13-1947 (E. Tams — CDA); China Camp, 2 2 2, XI-7-1965, shore of San Pablo Bay (R. L. Langston — CIS) ; Near Peacock Gap, San Rafael, 3 3^,5 2 2,3 juv., XI-7-1965, 6 $ S , 20 2 2 , 84 eggs, VI-2-1966 (R. L. L.— CAS, CDA, CIS, LACM, SFS, UCR) ; Sausalito, 1 $, 14 2 2 , 10 juv., XI-7-1965, shore of S. F. Bay (R. L. L. — CAS, CIS, LACM, UCR). Orange County: Costa Mesa, 1 $, III-6-1944, ex firewood (R. G. Bumgardner — CDA) ; Laguna Beach and vicinity, VI & VII-1921 (Essig, 1922 — first California record). San Diego County: Chula Vista, 1-18-1933, ex lawn (CDA) ; Del Mar, III-9-1932, Quarantine on Araucaria sp. (J. Adams — CDA) ; El Cajon, IV-10-1940, Quarantine from San Francisco on dahlia tubers (CDA) ; Escondido, VI-29-1934, Vol. 50, No. 1, January 1974 33 Quarantine from OREGON (M. England — CDA) . San Francisco County: “Frisco Bay, Cal.” 1 $, 4 9 9, XII-10-1949, under rock (“W. V. G.”— CIS) ; San Francisco, 1 $, IX-1935 (E. S. Ross — CAS). Solano County: Benecia, 5 $ $, 12 $ $ , VI-2-1966 (R. L. Langston — CAS, CIS) ; Benecia, 1 mi. W., 7 $ $ , 13 9 9,8 juv., 88 eggs, VI-2-1966, 13 $ $ , 32 9 9 , 27 juv., X-16-1966, shore of Car- quinez Strait (R. L. L. — CAS, CIS, CSH, SFS) ; Glen Cove, 1 $ , 1 9, VI-2-1966, 1 $, 4 9 9, VI-11-1967, 1 $, 4 9 9, VI-15-1969, shore of Carquinez Strait (R. L. L. — BVC, CIS, LACM) . Limited searches with negative results were made on parts of the im- mediate coast within the boundaries of the inset in Fig. 1. It is possible that severe wave action excludes the earwig from the Pacific Ocean sea- shore, in the western part of this area. It appears to be absent along the northern periphery of San Pablo Bay (Black Point, Vallejo, mouths of the Napa and Petaluma Rivers), possibly because the water is too fresh from the river outflows. It may also be absent from the west delta area east of Benecia or Port Costa (Martinez, Port Chicago, Antioch, Bethel Island, Brannan Island, Rio Vista) because the water may be too fresh, although seasonal increases in salinity occur depending on the tides and time of year. No examples were taken south of Alameda or San Francisco. Perhaps the water is too salty or polluted. However, two other earwigs ( Euborellia annulipes and Forficula auricularia ) oc- cur abundantly around the South Bay. Acknowledgments The help of Mr. George Buxton, California Department of Agriculture, Sacramento is greatly appreciated. Complete freedom in the use of his files, examination of specimens, and prompt notification of new records as they appeared in quarantine were especially helpful. Thanks are ex- tended to Dr. J. A. Powell of the California Insect Survey, Berkeley, for revisions and suggestions in relation to a more complete manuscript on the Dermaptera, from which much of the data for this paper was ex- tracted. I gratefully acknowledge the cooperation of Dr. P. H. Arnaud, Jr., for the study of the collection at the California Academy of Sciences, San Francisco, which is particularly rich in specimens from many parts of the world. Literature Cited Burr, M. 1910. The Dermaptera (earwigs) of the United States National Mu- seum. Proc. U.S. Nat. Mus., 38: 443^167. Caudell, A. N. 1913. Notes on Nearctic orthopterous insects. I. Nonsaltatorial forms. Proc. U.S. Nat. Mus., 44: 595-614. 34 The Pan-Pacific Entomologist Essig, E. 0. 1922. Insect notes from Laguna Beach, California. Pomona College J. of Entomol. & Zool., 14: 75-78. Fulton, B. B. 1924. Some habits of earwigs. Ann. Entomol. Soc. Amer., 17 : 357-367. Hebard, M. 1917. Notes on the earwigs (Dermaptera) of North America, north of the Mexican boundary. Entomol. News, 28: 311-323. Hebard, M. 1933. Dermaptera in collection of the California Academy of Sci- ence. Pan-Pac. Entomol., 9: 140-144. Helfer, J. R. 1963. How to know the grasshoppers, cockroaches and their allies. Pictured-key Nature Series. Wm. C. Brown Co., Dubuque, Iowa, v -p 354 pp. Langston, R. L. 1967. The maritime earwig, Anisolabis maritima (Gene). Pan- Pac. Entomol., 43: 88. Popham, E. J. 1965. A key to dermapteran subfamilies. The Entomologist, June, 1965: 126-136. Rehn, J. W. H. 1949. Catalogue of the Dermaptera of Venezuela. Bol. Entomol. Venezolana, 8: 1-9. RECENT LITERATURE The Butterflies of Southern California. Thomas C. Emmel and John F. Emmel. Natural History Museum of Los Angeles, Science Series, 26, 148 pp. 1973. $4.00. Amid the plethora of regional natural history manuals published recently this volume is distinguished by its uniform professional quality, combined with a pop- ular, easily read style. The introductory discussion includes a concise review of the seasonal and geographic distribution of southern California butterflies in terms of Merriam life zones. Brief but extremely interesting historical sketches of the more important lepidopterists of the area are followed by informative accounts of the species, including summaries of distributions and larval food plants and short descriptions of immature stages. The color plates of adults are excellently reproduced, and many larvae and pupae are illustrated in text figures. The host plant index and a pertinent list of references are useful features not always found in such faunal treatments.- — Editor. Vol. 50, No. 1, January 1974 35 Notes and Descriptions of Some Lycid-like Neotropical Lepturine Cerambycidae (Coleoptera) John A. Chemsak and E. G. Linsley University of California, Berkeley, California The genus Megachoriolaus was proposed by Linsley (1970) for a Mexican species with lycid-like coloration. At the same time a number of other species were transferred to this genus from Euryptera, all of which are probably also mimetic. In 1971 Linsley and Chemsak added two of Bates’ species and described one additional new species. Subse- quent collections have yielded a number of additional new species which are described below because of their possible significance in Neotropical mimetic complexes. During the study of the Euryptera-Megachoriolaus complex (Linsley and Chemsak, 1971), a paper by Fuchs (1956) was inadvertently over- looked. In this publication, three new species of Euryptera were de- scribed and other species previously included in the genus were char- acterized. Judging by the original description, E. unilaticollis Fuchs is probably assignable to Euryptera as defined by Linsley and Chemsak; E. rotundipennis Fuchs to Choriolaus Bates; and E. brasiliensis to Mi- miptera Linsley. The National Science Foundation through grant #GB-4944X is grate- fully acknowledged. Genus Megachoriolaus Linsley Megachoriolaus Linsley, 1970, Pan-Pac. Entomol., 46:128; Linsley and Chemsak, 1971, Arq. Zool., 21:3. Type species: M. chemsaki Linsley, by original designation. All of the known species of this genus fall into one of two color types, yellow and black or red and black. Since few of the mimicry models are known, the significance of these patterns remains to be determined. The yellow and black species are: chemsaki Linsley; breviceps Linsley; spiniferus Linsley; patricia (Bates) ; filicornis Linsley and Chemsak; and unicolor (Bates). The red and black species include: flammatus Linsley; ignitus (Schaeffer) ; sabinoensis (Knull) ; imitatrix Linsley; cruentus (Martin) ; texanus (Knull) ; and the four new species described below. The Pan-Pacific Entomologist 50: 35-43. January 1974 36 The Pan-Pacific Entomologist Megachoriolaus lineaticollis, new species Female: Form moderate sized, elytra slightly expanded posteriorly; elytra dark Nopal red (Ridgeway, 1912), pronotum red orange except for broad, median, longitudinal black band, vertex with a black band extending onto neck, appendages black, prosternum black at sides, mesosternum orangish on coxae and mesosternal process. Head with vertex impressed behind antennal tubercles, finely, confluently punctate, pubescence fine, dense, recumbent; front moderately elongate, ratio of distance between apices of antennal tubercles and line midway between them and apex of labrum, 1.0:1. 8; antennal tubercles shallow, very finely punctate to large glabrous median triangle, pubescence fine, rather dense, depressed; clypeus with a few irregular punctures at base; labrum with a few setiferous punctures; genae finely punctate; antennae extending to about middle of elytra, segments to base of fifth shining, clothed with coarse black hairs, outer segments opaque, poriferous areas absent, segments from fifth thickened, third shorter than first, fourth shorter than third, fifth longer than third but shorter than first. Pronotum densely, rather finely punctate except for vague linear area at center behind middle; pubescence dense, reddish, appressed; sides sinuate, posterior angles acutely produced over humeri; prosternum subglabrous, shallowly impressed before coxae; mesosternum with intercoxal process extending to level of coxae, abruptly declivous in front; metasternum very finely, shallowly punctate, finely pubescent. Elytra about 2.2 as long as width across humeri; punctures very fine, dense, separated by about one puncture diameter; pubescence dense, long, reddish, subdepressed; apices shallowly emarginate truncate, outer angles strongly dentiform. Legs with posterior tarsi shorter than tibiae, first segment as long as remainder of tarsus; femora slender, densely, shallowly punctate. Abdomen very densely punctate except at middle; last sternite subtruncate at apex. Length, 13 mm. Holotype female (United States National Museum) from Volcan de Chiriqui, 1000 M., Panama. (Friedr. Tippmann, Wien). This is the only known species of reddish Megachoriolaus with a black pronotal band. The rather fine dense punctation of the pronotum and the dense pubescence of the elytra also make this species distinctive. Megachoriolaus nigricollis, new species Male (?) : Form moderately robust, elytra subparallel, slightly tapering toward apices; integument black, elytra almost Nopal red (Ridgeway, 1912). Head with vertex vaguely impressed on each side of midline, punctures vague, pubescence dense, golden, appressed; front elongate, narrow, ratio of distance between apices of antennal tubercles and line between them and apex of labrum, 1.0:2. 3; antennal tubercles not prominent, minutely shallowly punctate, median glabrous triangle large; clypeus finely punctate at base, labrum with several small punctures at sides; genae very finely, sparsely punctate; antennae extending to about apical one-third of elytra, segments to fourth shining, remainder opaque, segments from fifth thick- ened, poriferous areas vague on outer segments, third segment slightly shorter than first, fourth shorter than third, fifth equal to third. Pronotum moderately densely punctate, punctures a little larger than those of elytra; pubescence red- orange, dense, depressed; sides sinuate; posterior angles acute, extending over hu- meri; prosternum finely punctate over apical %, pubescence very sparse; meso- sternum with intercoxal process broad, protruding slightly above level of coxae, Vol. 50, No. 1, January 1974 37 abruptly declivous in front; metasternum glabrous at middle, sides and margins very finely, densely punctate, pubescence sparse. Elytra a little more than twice as long as broad; punctures rather fine, well separated; pubescence moderately dense, suberect, reddish; apices slightly obliquely truncate, outer angle obtusely dentate. Legs with posterior tarsi about as long as tibiae, first segment as long as remainder together; front tibiae densely pubescent internally; femora moderately densely clothed with bronze pubescence externally. Abdomen very finely, densely punctate at sides, glabrous at middle, last sternite almost all punctate, apex very broadly emarginate, angle dentate. Length, 13-16 mm. Holotype male (?) (California Academy of Sciences) and six male (?) para- types from 30 miles NE Tehuantepec, Oaxaca, Mexico, 8 July, 1955 (Derham Giuliani) . No other known species of Megachoriolaus has the entire body except the elytra black. The type series shows no variation in this color pattern and all specimens are very similar. In addition to the black head and pronotum, the characteristics of the head, mesosternum and abdomen will separate M. nigricollis from other species. This species has no obvious secondary sexual characters. Megachoriolaus celestae, new species (Fig. 1) Male (?) : Form moderately robust, elytra subparallel before tapering at apices; elytra a scarlet Brazil red (Ridgeway, 1912), head and pronotum orangish, mouth- parts, antennae, eyes, legs and most of underside black, abdomen often partially pale. Head with vertex shallowly impressed on each side of midline, finely, con- fluently punctate, finely clothed with brownish depressed pubescence ; front elon- gate, ratio of distances between apices of antennal tubercles and line midway be- tween them and apex of labrum, 1.0:2. 3; antennal tubercles prominent, minutely opaquely punctate to large glabrous median triangle, pubescence very dark, brown- ish, depressed; clypeus rather coarsely, shallowly punctate over basal %, apex impunctate, labrum impunctate; genae minutely, vaguely punctate; antennae ex- tending to about apical one-third of elytra, basal segments to middle of fifth shining, rather sparsely clothed with coarse, suberect black setae, outer segments opaque, minutely densely punctate and pubescent, poriferous areas of outer seg- ments small, third segment slender, about as long as first, fourth slender, shorter than third, segments from apex of fifth very slightly thickened, fifth about as long as third, eleventh bluntly tapering at apex. Pronotum very densely, moderately coarsely punctate except for median impunctate line; pubescence dense, subde- pressed, colored similarly to elytra; sides sinuate, posterior angles acute, extending over humeri; prosternum impunctate, very sparsely clothed with very short pubes- cence; mesosternum very finely, densely punctate, moderately densely clothed with short, depressed, bronze pubescence, mesosternal process lying below surface of coxae, gradually arcuate anteriorly; metasternum finely densely punctate, moder- ately densely pubescent. Elytra about twice as long as broad ; punctures fine, dense, separated by about two puncture diameters; pubescence dense, rather short, sub- depressed, colored as surface; apices obliquely truncate, outer angles dentiform. Legs with posterior tarsi slightly shorter than tibiae, first segment longer than 38 The Pan-Pacific Entomologist Fig. 1. Megachoriolaus celestae Chemsak and Linsley. Vol. 50, No. 1 , January 1974 39 remainder of tarsus; femora finely densely punctate externally, densely clothed with short, dark and golden pubescence. Abdomen shining, finely densely punc- tate basally, less densely at middle toward apex; last sternite narrowly rounded at apex. Length, 13-15 mm. Holotype male (?) (California Academy of Sciences) from SE side of Cerro Tres Picos, Montane rain forest, 1524-1829 M., Chiapas, Mexico, 25 May 1972 (D. E. Breedlove) ; one male (?) paratype from N. slope of Cerro Bola, N. Cerro Tres Picos, 1524-2134 m., Chiapas, Mexico, 5 May 1972 (D. E. Breedlove). M. celestae is very distinctive by the unique scarlet red coloration of the elytra; none of the other reddish species of Megachoriolaus has this particular hue. The species also differs by the non-expanded elytra and form and color of the head. Of the other red-headed species, M. ignita (Schaeffer) has the pronotum more coarsely punctate; M. imi- tatrix Linsley has shorter antennae and sinuate-truncate elytral apices, and M. sabinoensis (Knull) has a much shorter front of the head. As in most species in this genus, there are no apparent secondary sex- ual differences. It is equally possible that both our specimens may be females or that both sexes are represented. The shape of the apex of the last abdominal sternite in M. celestae is quite unlike most other spe- cies which have the apex emarginate truncate and the angles dentate. We take pleasure in dedicating this beautiful species to Celeste Green in appreciation for the preparation of the illustrations which accompany this and many of our prior publications. Megachoriolaus gracilis, new species Male (?): Form moderate sized, slender, tapering posteriorly; tips of mouth- parts, eyes, antennae, tarsi, tibiae except bases of front pair, hind femora at apical one-half, middle femora at apical one-third, and front pair narrowly at apices, black, elytra approaching Nopal red (Ridgeway, 1912) in color, head, pronotum, underside and parts of femora orangish. Head with vertex impressed behind antennal tubercles, finely confluently punctate, pubescence very fine, golden; front moderately elongate, ratio of distance between apices of antennal tubercles and line midway between them and apex of labrum, 1.0 :2.2 ; antennal tubercles shallow, very finely, shallowly punctate to large glabrous median tri- angle, pubescence short, fine, subdepressed; clypeus with a few larger, shallow punctures irregularly dispersed; labrum with a few shallow setiferous punctures; genae almost impunctate; antennae almost as long as elytra, segments to middle of fifth shining, clothed with coarse black hairs at apices, outer segments opaque, poriferous areas obsolete, basal segments slender, segments from apex of fifth very slightly thickened, third a little longer than first, fourth equal to first, fifth equal to third. Pronotum moderately coarsely densely punctate except for midline be- hind middle; pubescence moderately dense, reddish, depressed; sides sinuate, pos- terior angles acute, extending over humeri; prosternum glabrous; mesosternum with intercoxal process tapering, lying below surface of coxae, very gradually arcu- ate anteriorly ; metasternum very finely, densely punctate, densely clothed with 40 The Pan-Pacific Entomologist short depressed pubescence. Elytra a little less than two and one-half times as long as broad; punctures fine, dense, separated by about one puncture diameter; pubescence dense, moderately long, depressed, colored as surface; apices obliquely rounded, outer angles dentiform. Legs with posterior tarsi about as long as tibiae, first segment longer than remainder of tarsus; femora slender, densely punctate. Abdomen shining, finely densely punctate at side; last sternite narrowly rounded at apex. Length, 11-13 mm. Holotype male (?) (California Academy of Sciences) from La Pacifica, 4 Km N.W. Canas, Guanacaste Province, Costa Rica, 30 May 1972 (P. A. Opler) ; three male (?) paratypes with same data. The moderately slender, tapering body form of this species makes it easily recognizable. The coloration of the body is also distinctive. In this respect there is no variation within the type series. Eurylemma, new genus Form moderately robust; elytra expanded posteriorly. Head with front short, deeply impressed transversely; palpi unequal, apical segments cylindrical; eyes small, finely faceted, deeply emarginate, upper lobes small, widely separated above ; tempora prominent, elongate, parallel; antennae short, segments moderately thick- ened, basal segments with coarse setae at apices, poriferous areas vague, scape robust, longer than third segment, fourth shorter than third, fifth equal to first. Pronotum rather flattened, sides sinuate; hind angles acute, not extending over humeri; apex about as broad as head, basal margin broadly lobed; disk impressed at sides; prosternum narrow, deeply impressed at apex, intercoxal process slender, expanded at apex, coxal cavities partially open behind, coxae rather small; meso- sternum with intercoxal process lying below surface of coxae. Elytra with broad, elevated lines; apices broadly rounded. Legs slender, short; posterior tarsi shorter than tibiae, first segment a little shorter than remainder of tarsus together. Ab- domen normally segmented. Type species: Eurylemma auricollis, new species. This genus is apparently related to the group of genera presently re- ferred to as the Euryptera-Megachoriolaus complex. It superficially re- sembles Euryptera but differs by the more slender antennae, longer tempora, broader front of the head, non-elevated mesosternal process, and the absence of dentate angles at the apex of the last abdominal ster- nite. The shorter front of the head, prominent tempora, shape of the pronotum, and costate elytra will distinguish Eurylemma from Mega- choriolaus. A single lycid-like species is presently known. Eurylemma auricollis, new species (Fig. 2) Female: Form moderate sized, depressed, elytra expanding posteriorly; integu- ment black with a bluish overcast, pronotum broadly yellow down sides, pubescence dark, golden on head and sides of pronotum. Head with front rather short, antennal tubercles short, finely punctate; clypeus with a few scattered punctures, lahrum Yol. 50, No. 1, January 1974 41 Fig. 2. Eurylemma auricollis Chemsak and Linsley 42 The Pan-Pacific Entomologist almost impunctate; vertex broad, minutely punctate, each side with a band of dense, golden, appressed pubescence extending from antennal tubercles to neck; antennae extending to about middle of elytra, segments slightly thickened, segments to base of fifth shining with a metallic caste, remainder opaque. Pronotum broader than long; sides yellowish, densely clothed with golden recumbent pubescence; disk impressed at sides near base; punctures fine, contiguous, middle with a glab- rous longitudinal line; pubescence at middle short, dark, depressed; prosternum vaguely punctate, pubescence sparse; meso- and metasternum very finely, separ- ately punctate, moderately clothed with fine subdepressed pubescence. Elytra about three times as long as basal width, broadly expanding toward apices; each elytron rather strongly tricostate and with a feeble costa toward lateral margin; punctures minute, dense; pubescence short, depressed; apices broadly rounded. Legs slender, short, finely pubescent. Abdomen shining, almost glabrous; last sternite truncate at apex. Length 17 mm. Holotype female (California Academy of Sciences) from Cerro Huitepec, W. of San Cristobal de las Casas, 2591 M., Chiapas, Mexico, 23 May, 1972 (D. E. Breedlove) . This species has a very distinctive appearance as a result of the dark integument with a bluish caste and the golden recumbent pubescence of the head and pronotum. Euryptera albosterna, new species Female: Form moderate sized, elytra strongly expanding posteriorly; color black, head with parts of front and vertex yellow orange, pronotum with two broad red bands down sides extending over humeri, humeri red, red band extending acutely back to about basal %, underside of head, middle of prosternum and mesosternum yellow; pubescence black on dark areas, bright red and appressed on red bands of pronotum and elytra. Head moderately elongate, ratio of distance between apices of antennal tubercles and a line midway between them and apex of labrum, 1.0:1.75; front moderately impressed, indistinctly punctate, median tri- angle vague, clypeus sparsely, moderately coarsely punctate, labrum sparsely, mod- erately coarsely punctate, pubescence fine, subdepressed; vertex plane, median line becoming a carina to neck, punctures vague, pubescence short, subdepressed; genae finely, sparsely punctate; palpi very unequal, last segment of maxillary very elongate; antennae rather stout, extending to about basal % of elytra, basal seg- ments shining, clothed with coarse black setae, segments from sixth opaque, mi- nutely pubescent, scape conical, longer than third segment, third longer than fourth, fifth longer than third. Pronotum minutely punctate; pubescence long, appressed, colored as background; sides sinuate, hind angles acutely produced over humeri; base lobed at middle; prostemum glabrous, not impressed, intercoxal pro- cess fairly broad, rounded; mesosternum with intercoxal process broad, extending above coxae, abruptly declivous in front; metasternum minutely punctate, moder- ately densely pubescent. Elytra about 2% times as long as basal width, broadly expanding toward apices; punctures minute, dense; pubescence dense, suberect, depressed along humeral red area, sides fringed; each side of suture at basal % with a strong costa terminating in a sharp spine at apical inner angle; epipleurae strongly margined; apices broadly emarginate, outer angles dentate. Legs short, slender; posterior tarsi shorter than tibiae, first segment shorter than remainder Vol. 50, No. 1, January 1974 43 of tarsus. Abdomen minutely, densely punctate, segments narrowly glabrous pos- teriorly; last segment sinuate at apex, outer angles spinose, Length, 12 mm. Holotype female (California Academy of Sciences) from Turrialba, Costa Rica, 31 January 1973 (V. M. Kirk) . This species shares most of the generic characteristics of Euryptera latipennis Serville, and the only relative differences of E. albosterna are the wider prosternal process and reduced costae of the elytra. E. albosterna may be readily recognized among the Central American eurypteroid lepturines by the appressed and suberect pubescence of the elytra, the distinctive red and black dorsal coloration, and by the sutural costae and spines of the elytra. We wish to thank V. M. Kirk for making this specimen available for study. Genus Mimiptera Linsley Mimiptera Linsley, 1961, Jour. New York Entomol. Soc., 69:139. Type species: Euryptera fulvella Bates, by original designation. Specimens of this genus are extremely rare in collections. Since the genus was described only one additional example has been seen, from Hacienda Comelco, 24 Km. NW Canas, Inter-Am. Hwy., Guanacaste, Costa Rica, on flowers of Casearia, 16 November, 1971 (E. R. Heithaus). Based on the coloration, it would be regarded as M. fulvella (Bates) but structurally it is identical with M. costaricensis (Melzer). As in other mimetic species, such as those of the genus Ely- troleptus (Chemsak and Linsley, 1965), M. fulvella apparently has more than one lycid-like color form, that is with the elytra transversely black banded or wholly yellow. M. costaricensis is therefore considered synon- ymous with M. fulvella. Literature Cited Chemsak, John A. and E. G. Linsley. 1965. A revised key to the species of Elytroleptus with notes on variation and geographical distribution. Pan- Pacific Entomol., 41: 193-199. Fuchs, E. 1956. 2. Beitrag zur kenntnis der neotropischen Cerambyciden. En- tomol. Art. Mus. G. Frey, 7: 567-576. Linsley, E. G. 1961. The North and Central American species of Euryptera and a related new genus. Jour. New York Entomol. Soc., 69: 131-141, 1 fig. Linsley, E. G. 1970. New genera and species in the lepturine complex related to Euryptera and Choriolaus. Pan-Pacific Entomol., 46: 128-141, 4 figs. Linsley, E. G. and John A. Chemsak. 1971. An attempt to clarify the generic status of some Neotropical species currently assigned to Euryptera, Chontalia and Ophistomis. Arq. Zool., S. Paulo, 21: 1-40. Ridgeway, R. 1912. Color Standards and Color Nomenclature. 43 pp., 53 pis. 44 The Pan-Pacific Entomologist Larvae and Pupae of Some Wrack Dipterans on a California Beach (Diptera: Coelopidae, Anthomyiidae, Sphaeroceridae) Helen Kompfner Hopkins Marine Station of Stanford University Introduction The kelp (wrack) flies, Coelopa vanduzeei Cresson (1914), Fucellia costalis Stein (1910), Fucellia rufitibia Stein (1910), and Leptocera johnsoni Spuler (1925) are common on California beaches (Cole, 1969). The life histories of such related European species as Fucellia maritima and Coelopa frigida have been studied (Egglishaw, 1960a and b), but little is known about the immature forms of the Pacific coast species. Preliminary observations of fly larvae found living in wrack suggested that a study of distribution and biology of larvae on the beach would be valuable in further understanding how different species of flies relate to the wrack, as a food source and habitat. This investiga- tion was carried out April— June, 1972, on two pocket beaches in Mon- terey Bay, California: (1) the west beach of Mussel Point, Pacific Grove, and (2) the Point Lucas beach, Pacific Grove. My studies of larvae were accompanied by a study of the activity patterns and be- havior of adult wrack dipterans by another investigator (Hyatt, 1972) at Mussel Point. Characteristics of Larvae and Pupae (Fig. 1 and 2) Larvae were collected in the field and raised on samples of damp wrack kept in the laboratory at room temperature. Initial identifications were made on the basis of emerged adults. Samples of these adult flies were examined and identified by Dr. Paul Arnaud of the California Academy of Sciences, and George C. Steyskal of the U.S. Dept, of Agriculture. Coelopa vanduzeei. — Third instar larva. Length 9.0-12.0 mm, great- est width and height 1.5 mm at caudal end, body narrowing anteriorly, second segment 0.5 mm wide. Color white, with grey ventral spines on abdominal segments. Caudal plates yellow with black borders, sur- rounded by pale hairs; 3 spiracles. Anus four-lobed, located ventro- posteriorly on segment 12. Anterior prothoracic spiracles bi-lobed, each The Pan-Pacific Entomologist 50: 44-52. January 1974 Vol. 50, No. 1, January 1974 45 Fig. 1 . Larval structures of wrack dipterans discussed in text. A. Coelopa vanduzeei B. Fucellia costalis C. Fucellia rufitibia D. Leptocera johnsoni. (1) left lateral view of mouthparts (2) left lateral view of entire third instar larva (3) caudal view, showing caudal plates and posterior spiracles. lobe marginally scalloped to form 10 petaldike processes. Antennae smooth, finely pointed. Puparium. Length 3. 5-6.0 mm; greatest width at segment 8, about half this ; greatest height at segment 8, approximately one third length ; ventral surface and dorsal surface of anterior three segments flattened; dorsum arched; lateral edges of each segment indented; color yellow- brown to brown-black. Fucellia costalis . — Third instar larva. Length 9.0-12.5 mm; body widest at caudal end, 1.5 mm, tapering to 0.5 mm at anterior end of second segment. Color white, or off-white. Caudal plates white with three amber spiracles, no caudal plate hairs. Anterior prothoracic spi- racles each with a single cluster of 8-10 lobes. Antennae blunt-tipped with annular ridges. Puparium. Length 6.0-7.0 mm; greatest width and height at seg- ments 7 and 8, 3.0 mm. Shape fusiform, circular in cross-section. Color dark reddish brown, segments 1-11 with wrinkled surface, segment 12 matte. 46 The Pan-Pacific Entomologist 1mm D Fig. 2. Puparia, dorsal views. A) Coelopa vanduzeei , B) Fucellia costalis, C) Fucellia rufitibia D) Leptocera johnsoni. Fucellia rufitibia . — Third instar larva. Length 8.5—10.5 mm, greatest width 1.2 mm, at caudal end, tapering to 0.3 mm at anterior end of seg- ment 2. Color white, grey at twelfth segment. Caudal plates white with yellow or red spiracles, no caudal plate hairs. Anterior prothoracic spi- racles each one-lobed, with a single cluster of 7-11 processes. Antennae blunt-tipped, with annular ridges. Puparium. Length 4.0-5. 0 mm, greatest width and height at segments 7 and 8, 1.5 mm. Shape fusiform, circular in cross-section. Color red- dish brown, surface texture smooth, shiny. Leptocera johnsoni . — Third instar larva. Length 2. 0-3. 5 mm. Color white, with black spines on abdominal ridges at anterior end of segments 6-12. Caudal plates on caudal stalks, plates black surrounded by long, black hydrophobic hairs in tufts of 4 or 5. Prothoracic spiracles with 5 slender radiating marginal processes. Puparium. Length 2. 0-2. 5 mm, width and height greatest at seg- ments 5-11, 0.8 and 0.5 respectively, dorso-ventrally flattened especially segments 1-2 and 12. Color yellow brown, segments 1-4 often translu- cent. Segment 12 has two caudal stalks. Field Studies Distribution of fly larvae was studied on beaches at Mussel Point and Point Lucas. Samples were taken at high, mid and low sections of the beach. Since most larvae were found in wrack banks (piles greater than 15 cm in depth, as defined by Backhand, 1945), vertical distribu- tion within the banks was also studied. Samples of wrack from various elevations on the beach, and samples taken at depths of 10, 20, and Vol. 50, No. 1, January 1974 47 w < G G > Q o 111 o < I- UJ o oc 111 Q. Fig. 3. Distribution of larvae on beach in relation to tidal height. 30 cm within wrack banks, were collected in plastic bags and returned to the laboratory. Larvae were extracted from the wrack by shaking the samples vigorously with water, then capturing the smaller larvae with a fine aquarium net. The wrack sample was then carefully examined for any remaining larger larvae. Fig. 3 shows the distribution of fly larvae from wrack banks stranded at various elevations above mean lower low water. The majority of wrack deposits at Mussel Point centered around the +3.0 tidal level and consisted of mixed red and brown algae and the surf grass Phyl- lospadix. The major deposits at Point Lucas consisted of banks of the brown alga Macrocystis concentrated on the upper beach at tidal heights of +5.0 to 6.5 feet. Coelopa vanduzeei larvae were found exclusively in lower beach wrack banks. Fucellia costalis larvae were found almost exclusively in the upper beach banks. F. rufitibia larvae were found at all levels, but predominately on the mid and upper beach. L. johnsoni larvae also occurred widely on the beach but were more evenly dis- tributed. The occurrence of larvae in wrack corresponds well with the distribution of the adult flies found on the same beach in studies by Hyatt (1972). Vertical distribution of fly larvae within wrack banks is shown in Fig. 4A, B. In banks on the lower beach Coelopa vanduzeei predomi- 48 The Pan-Pacific Entomologist nated in the top 10 cm, and L. johnsoni at the lowest levels. A small number of F. rufitibia were found, largely at intermediate levels. F. costalis was absent. Observation of a lower beach bank before and after complete immersion during a high tide showed that numbers of C. vanduzeei present did not change ; numbers of L. johnsoni diminished, however, suggesting a loss accompanying wave action. In the upper beach bank (Fig. 4B) Fucellia costalis larvae predominated, and F. rufitibia outnumbered L. johnsoni at the lowest levels. C. vanduzeei was absent. High beach wrack banks tended to dry out, and as this happened, larger larvae were observed moving down in wrack over time. Observations on Feeding, Behavior, and Temporal Variation Larvae appeared to eat the algae on which they were found. Frag- ments of Macrocystis were found in the guts of all species. Larvae were never observed on very fresh wrack; apparently some decomposition must occur before the algae are eaten. Nematodes were abundant where fly larvae were found and may be involved in initial algal breakdown, as suggested by Egglishaw (1960b). Wrack consisting of the kelp Macrocystis seemed to be preferred above other algae by all larvae collected from natural wrack. When wrack was experimentally sorted out into separate piles containing only brown algae, only reds, only Phyllospadix, or a mixture of all types, and placed on the beach for several days, larvae were only found on brown and mixed wrack. Larvae placed on red algae and Phyllo- spadix died within a day in the lab, while those on Macrocystis flour- ished. Those kept without food died within two days, and larvae kept without moisture died overnight. Of the four species observed, the larvae of C. vanduzeei and L. john- soni were the most active and moved most rapidly. Over the 5 week period of the study some temporal variation was observed in the numbers of C. vanduzeei larvae. In the last two weeks of April none were found on the beach at Mussel Point. By May 2, they were found in small numbers and in the week of May 8-15, occurred in multitudes in buried wrack on the lower beach there and further south along the coast at Pebble Beach and Carmel. The latter period coincided with neap tides, and formation of major wrack deposits on the beaches. By May 17 no more larvae were found and the wrack was also gone from the beach. Fucellia costalis , F. rufitibia , and Leptocera johnsoni were found in fairly constant numbers, but never as abundantly as Vol. 50, No. 1, January 1974 49 C. vanduzeei F ruf itibia L. johnson i F. costal i s 13% 13% 74 / 86 % 14 1 59 % 27 % 1 4 % N=15 N=7 N^5t 377o . 62% 1 % N=73 N=61 17% 52% 31% N=142 10 20 30 1 0 20 30 A B WRACK HEIGHT IN CENTIMETERS Fig. 4. A) Vertical distribution of larvae in lower beach wrack bank. B) Ver- tical distribution of larvae in upper beach wrack bank. Coelopa vanduzeei at its peak, over the time interval noted. Some adult L. johnsoni were observed mating in all wrack deposits observed over 5 weeks. Laboratory Rearing of Larvae Attempts to obtain eggs and subsequent larval stages in the laboratory from captive populations of flies maintained on wrack in large plastic bags were not successful. However, larvae of C. vanduzeei , F. costalis , F. rufitibia , and L. johnsoni were reared successfully on Macrocystis stipes (examined first for foreign larvae or eggs) in Petri dishes at room temperature. Cultures were wetted and new Macrocystis added every two days. Pupae were removed, placed in corked vials, and checked daily for hatching. This method of rearing does not eliminate the possibility that some larave may have fed on each other. Some species of Fucellia are known to be predaceous as larvae, i.e. F. rejecta Aldrich. The picture of development obtained, while incomplete, provides the beginning of a timetable for the life history. Data available are shown in Fig. 5. Duration of the third instar was shortest in Coelopa vanduzeei , longest in Fucellia costalis. The former lives lowest in the intertidal zone, the latter highest on the beach. Duration of pupal period showed considerable variation among species and within C. vanduzeei. For the latter, there may be some connection between intertidal height on the beach and length of pupal period. Although later development took 50 The Pan-Pacific Entomologist unknown N Larva second I | Pupa i nstar Fig. 5. Developmental period of lab reared larvae. place in the laboratory, the 12 individuals of C. vanduzeei which hatched after a 4 day pupal period were collected from buried wrack at the +3.0 foot tide level, while the individual which spent 9 days as a pupa was taken from an upper beach wrack bank. L. johnsoni had a pupal period of 6-7 days, F. rufitiba and F. costalis 12—15 days. F. costalis larvae may remain in the third instar for considerable periods; one batch of 10 remained in this stage for 25 days before pupation. Discussion The flies appear to use different strategies in exploiting the available food and habitat resources for larvae on the beach. C. vanduzeei is found almost exclusively in lower beach wrack which is wetted at least once a day and seldom remains on the beach longer than two weeks. The main pressure here is for quick larval development and a short pupal period. The caudal plate hairs allow the larva to breathe from air bubbles trapped in the wrack during periods of tidal immersion. Immature F. costalis and R. rufitibia, found mainly on the mid and upper beach levels, are able to use wrack for longer periods. Caudal hairs are lacking. The danger of being washed out to sea is less than that of dessication. The pupal period reflects this, being relatively long. L. johnsoni larvae are found in large numbers and are distributed both high and low on the beach. Their caudal plate hairs allow them to re- spire when submerged in wrack beds low on the beach, but their small size makes them vulnerable both to being swept out of low immersed wrack, and to desiccation in higher beach wrack. Perhaps this is why Vol. 50, No. 1 , January 1974 51 L. johnsoni tends to occur well below the surface of wrack banks. A comparison of mean tidal height of distribution and mean pupal period for each species suggests that species found higher on the beach may have longer pupal periods, although the variation within each species is large enough to eliminate meaningful statistical comparison with present data. Summary Larvae and pupae of Coelopa vanduzeei , Fucellia costalis, Fucellia rufitibia, and Leptocera johnsoni were studied on two California beaches. Third instar larvae and puparia are described. Studies of distribution of larvae on the beach showed C. vanduzeei occupies lower beach wrack banks and F. costalis the upper beach banks. F. rufitibia larvae, while widespread, occur more in mid and upper beach wrack, while those of L. johnsoni are found at all levels. Larvae were observed to prefer the kelp Macrocystis both as a habitat and food source. C. vanduzeei larvae were observed in large numbers only during a two week period coin- ciding with a major wrack deposit. F. costalis, F. rufitibia, and L. john- soni were found in constant numbers over 5 weeks. Duration of pupal periods in the laboratory were: C. vanduzeei, 4-9 days; L. johnsoni, 6-1 days; F. costalis and F. rufitibia 12-15 days. Average duration of the pupal period appears to increase with increasing tidal height of the larval habitat. Acknowledgments I would like to thank Dr. Paul Arnaud of California Academy of Sci- ences and George C. Steykskal of the U.S. Department of Agriculture for identifying the adult flies of this study, and Joel Hyatt, for many helpful discussions about flies and larvae on the beach. I am especially indebted to Dr. Donald P. Abbott of Hopkins Marine Station, for his unfailing patience and encouragement with this study and the editing of this manuscript. Dr. I. A. Abbott also provided much appreciated encouragement during this study. Literature Cited Backlund, H. O. 1945. Wrack fauna of Sweden and Finland. Ecology and chorology. Opuscula Entomol. Suppl., 5: 1-238, Lund, Sweden. Cole, F. R. 1969. The flies of Western North America. University of California Press, Berkeley. Egglishaw, H. 1960a. The life history of Fucellia maritima (Haliday) (Diptera, Muscid.) Entomologist, 93: 225-231. 52 The Pan-Pacific Entomologist Egglishaw, H. 1960b. Studies of the family Coelopidae (Dipter) . Trans. R. Entomol. Soc., 112: 109-140. Hyatt, Joel. 1972. Behavior of wrack dipterans, F. rufitibia (Anthomyiidae) and C. vanduzeei (Coelopidae) on a California beach. Unpub. MS on file at Hopkins Marine Station Library. SCIENTIFIC NOTE The Occurrence of Trichocorixa reticulata in the Gulf of California (Hemiptera: Corixidae). — During the course of ground observations in sup- port of the NASA Earth Resources Satellite Program 1 , plankton tow samples were taken in the delta region of the Colorado River at the extreme northern end of the Gulf of California. One of these tows (Plankton Station D-4 at N. end of Isla Montague in the Colorado River Delta, 31° 49.9' N. latitude, 114° 48.5'W. longi- tude, 16 December 1972, J. R. Hendrickson) included a female of Trichocorixa reticulata (Guerin-Meneville) . While the occurrence of this species in salt water has been noted previously (see Williams 1944, Proc. Haw. Ent. Soc., 12: 186-196, and Sailer 1946, Proc. Haw. Ent. Soc., 12 : 617-620) , it has not been recorded from ocean waters. Wil- liams found it in ponds more saline than sea water, accounting for its success in colonizing salt water habitats. The locality where the specimen was collected is one of strong tidal currents produced by semidiurnal tides which can vary by as much as 9 meters between extreme high and extreme low. The water carries a very heavy silt load (standard Secchi disk visibility range is 2-15 cm depth) ; salinity has been measured as low as 37 and as high as 44 parts per thousand, depending upon tidal state and season (world normal salinity is about 35 parts per thousand, normal for the northern Gulf of California is about 37 parts per thousand) . Accompanying the specimen were the normal components of a marine plankton tow (copepods, arrow worms, jellyfish, marine fish juveniles) . The specimen was swimming actively in the con- centrated plankton sample when first noted. — J. T. Polhemus, Martin Marietta Aerospace, Denver, Colorado 80201 and J. R. Hendrickson, Department of Bio- logical Sciences, University of Arizona, Tucson 85721. 1 NASA Contract NAS5-21777, Identification No. UN603. This work was also supported by NSF Grant GB34675. Vol. 50, No. 1, January 1974 53 Observations on Primary and Secondary Parasitoids of California Oakworm, Phryganidia calif ornica. Pupae (Lepidoptera: Dioptidae) David J. Horn 1 Department of Entomology, The Ohio State University, Columbus 43210 Introduction The California oakworm, Phryganidia californica Packard, periodi- cally defoliates evergreen and deciduous oaks ( Quercus ) in coastal Cal- ifornia. In the San Francisco Bay region, irruptions occur at 4- to 7- year intervals and last 2-3 years each, followed by spectacular declines in numbers. The oakworm normally has two generations a year. Larvae overwinter on evergreen oaks, particularly coast live oak, Quercus agri- folia Nee, and feed on new foliage in April and May. Pupation occurs as a chrysalid on branches and trunks of oaks, and in surrounding vegeta- tion, during June. The weak-flying adults emerge in a few weeks, mate, and oviposit on both evergreen and deciduous oaks, especially Q. lobata Nee, valley oak. Eggs hatch, and larvae feed in July and August, to pu- pate in September and emerge as adults in October and November. These oviposit on evergreen oaks, eggs hatch, and the larvae may feed but are usually inactive until the onset of warmer weather and new foliage in late March and April (Harville, 1955, and my observations). Early workers (Kellogg and Jack, 1895; Doane, 1912) attributed oakworm population declines to parasitization because dense infestations usually yielded high percentages of parasitized oakworm pupae. Har- ville (1955), following exhaustive field study in which pupal parasitism never exceeded 75%, concluded that population declines were caused by a variety of factors, particularly cold winter weather killing over- wintering larvae. Among the pupal parasitoids, Harville found Itoplectis behrensii (Cresson) most commonly. It is probably an obligate parasitoid of oakworm pupae (though Muesebeck et al., 1951, cite its presence also in Recurvaria milleri Busck) . It is bivoltine, like its host, and the long- lived adults can be collected throughout the year (Townes and Townes, 1960). Harville (1955) observed that I. behrensii females hunt tac- tilely on oak branches and trunks, “stumbling” onto potential hosts, and being unable to distinguish previously parasitized hosts. He found that 1 Formerly Department of Biological Science, California State University, Hayward. The Pan-Pacific Entomologist 50: 53-59. January 1974 54 The Pan-Pacific Entomologist Table 1. Population density, parasitization, and survival of Phryga- nidia californica pupae. Hayward, Alameda Co., Calif. May-July. Year Pupae/ft 2 Sample size Total parasitization Moth emergence 1969 1 23.4 270 110 (41%) 100 (37%) 1970 30.3 478 339 (71%) 56 (12%) 1971 .01 15 2 (13%) 12 (80%) 1972 0.4 35 2 (6%) 26 (74%) 1 Sampling commenced 20 June. Parasitization probably lower, and moth emergence greater, for whole season, than indicated value. /. behrensii in turn was parasitized by 3 secondary parasitoids: Gelis tenellus (Say), Mastrus aciculatus (Provancher) , and Dibrachys cavus (Walker) ; the total incidence of parasitism by all three on I. behrensii was less than 1% in eight study sites though D. cavus was sometimes locally more abundant. Harville also found Brachymeria ovata (Say) as a primary pupal parasitoid of the oakworm. It has no secondary parasitoids. B. ovata is facultative, having been recorded from over 100 species of lepidopterous pupae (Burks, 1960), and Harville sug- gested that it might survive on alternate hosts, possible species of Mala- cosoma , during times of oakworm scarcity. My preliminary observations of oakworm and parasitoid populations near Hayward, California in 1969 revealed more extensive secondary parasitization of 1. behrensii than that Harville reported. This suggested that the interrelationships between oakworm populations and those of the several parasitoid species warranted further study. Methods I selected a wooded hillside adjacent to the Ecological Field Station, California State University, Hayward. The vegetation of the study area, described in detail by Cogswell (1966) is a broadleaf evergreen wood- land, dominated by coast live oak and California laurel ( Umbellularia californica ) . No insecticides have ever been used here. In 1969-70, but not in 1971—72, the spring generation of oakworms severely defoli- ated oaks in the study area. On each visit to the study plot in 1969-70 and 1972, I counted fresh oakworm pupae in each of 10 1ft 2 areas of tree-trunk undersurface chosen randomly from a group of 5 trees. Pupae were so few in 1971 that I obtained no meaningful estimate of density. I determined the extent of parasitism by collecting a sample of 50- Vol. 50, No. 1, January 1974 55 Table 2. Emergence of parasitoid species from Phryganidia cali- fornica pupae. Hayward, Alameda Co., Calif. June-July 1969, May- July 1970. Year Total parasitoids /. behrensii B. ovata G. tenellus M. aciculatus Bathy- thrix D. cavus 1 1969 110 32 9 21 1 5 42 1 1970 339 107 19 66 6 69 72 1 1 Number of hosts parasitized, not number of emerging parasitoids. D. cavus is gregarious, with 3-10 parasitoids/host. 100 pupae randomly from lower, accessible, oak limbs and trunks, and kept pupae individually in 4-oz jars until moths or parasitoids had emerged. I repeated the procedure every 5 days from the onset of pu- pation until no new pupae appeared. Pupae from which nothing emerged were visually inspected, then dissected. While collecting pupae, I spent several hours observing the activities of I. behrensii and B. ovata. Wlien I encountered pupae of other Lepidoptera, I saved these for possible parasitoid emergence. These were primarily Hemerocampa vetusta Bois- duval (Liparidae), Hemihylaea edwardsi Packard (Arctiidae), Mala- cosoma calif ornicum (Packard), and M. constrictum (Edwards) (Lasio- campidae) . Results Six species of parasitoids emerged from field-collected oakworm pu- pae; 5 were those Harville (1955) found; the sixth, Bathythrix sp., a secondary parasitoid of I. behrensii , represents a new host record. Table 1 shows trends in oakworm numbers, pupal parasitization, and moth emergence during 1969-72. Oakworm density and parasitization increased from 1969 to 1970, then the oakworm population declined dramatically. Fewer moths were produced in 1971-72 than in 1969-70, but the percentage produced was much greater. Table 2 shows relative proportions of the several parasitoid species in 1969-70. Parasitoids of I. behrensii were much more abundant in 1970 than in 1969. In 1971-72 I found only two parasitized oakworm pupae each year; all yielded I. behrensii. No secondary parasitoids or B. ovata were found in oakworm pupae in 1971 or 1972. I reared B. ovata from one pupa of Hemerocampa vetusta and one of Hemihylaea edwardsii. The latter is a new host record. From 35 pupae of Malacosoma spp. in 1972, I reared two G. tenellus but no B. ovata. Fig. 1 illustrates seasonal trends in the incidence of parasitization 56 The Pan-Pacific Entomologist Fig. 1. Trends in oakworm pupal density and parasitization at Hayward, Ala- meda County, California, 1970. Vol. 50, No. 1, January 1974 57 in 1970, the year for which I obtained most data. Parasitization by B. ovata, though never exceeding 10%, was greatest on earlier oakworm pupae, with a small second generation in early July. Percent parasitiza- tion by 7. behrensii increased from 23% in May to over 90% in July. Over the entire sampling period from May to July, oakworm pupal mortality was 88%, as follows: B. ovata 4%, 7. behrensii 67%, and un- known 17%. Some of the unknown mortality may have been due to fungal infections and some may have resulted from oviposition by para- sitoids or feeding by predaceous Hemiptera (Horn, 1973) . Occasionally, I encountered desiccated parasitoid larvae within oakworm pupae. These are included as unknown mortality because I could not determine the species of parasitoid involved. The incidence of secondary parasitization of I. behrensii increased from 0 in May to 100% in late June and July. G. tenellus , D. cavus , and Bathythrix sp. were equally abundant and showed no seasonal separation, while Mastrus was much less common (Table 2). Together, secondary parasitoids killed 69% of Itoplectis present in the samples over the sea- son. A few B. ovata may have been parasitized in early July. I observed searching behavior of 10 I. behrensii females in 1970 for periods up to 45 minutes each. My observations agreed with those of Harville (1955), that 7. behrensii searches very rapidly and efficiently, running over the branches and trunk, and ovipositing readily in pre- viously parasitized hosts. In every case, females turned away when en- countering their own trail, indicating the presence of a trail odor (Price, 1970). Two B. ovata females that I observed moved about more slowly than 7. behrensii and oviposited in nearly all oakworm pupae encount- ered. One moved directly from an oakworm pupa to oviposit in a co- coon of 77. vetusta. Superparasitism apparently occurs readily in each species. Discussion Conclusions based upon observation of a local population through only a portion of a cycle are tentative at best, but the increase in para- sitization immediately preceding the oakworm population decline sug- gests that 7. behrensii may be partially responsible for oakworm pop- ulation declines. Of interest are the interactions among the parasitoids and their use of a fluctuating resource. 7. behrensii, having no alternate hosts, must disperse readily and search quickly, as its probability of survival is low, especially in years when oakworm pupae are scarce. It averages twice the size of B. ovata, has much longer wings, and adults are relatively 58 The Pan-Pacific Entomologist long-lived. Other parasitoid species display these characteristics in un- stable environments (Price, 1972). B. ovata, being facultative, moves to alternate hosts, at least to Hemerocampa vetusta and Hemihylaea ed- wardsii, when oakworms are scarce. It is smaller and stockier than I. behrensii and is thus capable of searching a wider variety of habitats, including ground litter and bark crevices, as well as the surfaces of branches and trunks. This befits a parasitoid with over 100 recorded hosts. Malacosoma may not be as likely a reservoir of alternate hosts as Harville (1955) suggested. I did not find B. ovata in a small sample (N = 35) of Malacosoma pupae, nor did Langston (1957) discover it in a large survey of parasitoids of California Malacosoma, though Stehr and Cook (1968) recovered it from Malacosoma elsewhere. I. behrensii thus has some attributes of an r-strategist whereas B. ovata, relatively, is a K-strategist (Force, 1972) ; populations of the former fluctuate in number along with their fluctuating resource, while numbers of the latter probably remain relatively constant due to the supply of alternate hosts. B. ovata occurs earlier and later than the bulk of I. behrensii in oakworm pupae. This suggests that I. behrensii may be the superior competitor when both species parasitize a single host pupa. In the laboratory I attempted to induce females of each spe- cies to oviposit in each of 20 oakworm pupae, from which I recovered 14 I. behrensii and 5 moths. However, I was not certain whether that B. ovata actually oviposited, so the results are inconclusive. The facultative secondary parasitoids, at least Gelis tenellus and Di- brachys cavus, also move to alternate hosts including parasitoids of Malacosoma (Langston 1957) when /. behrensii is rare. Mastrus aciculatus has been recorded from several hosts, mostly Tenthredinidae (Bobb 1965, Price, 1970), as have species of Bathythrix (Muesebeck et al. 1951). There was no seasonal separation of their occurrence on I. behrensii, but they do differ in size and this may affect competition among them. More must be known of their relative abundance on other hosts to ascertain their competitive interactions, which may be rapidly changing. For instance, Bathythrix occurred commonly in my 1970 field samples yet was not present in any of Harville’s study populations. It may have recently broadened its host preference to include I. behrensii. The total incidence of parasitization on I. behrensii by all 4 secondary parasitoids (52% in 1969 and 69% in 1970) was far greater than the 1% reported by Harville (1955). This suggests that significant local changes may occur in the parasitoid complex of the oakworm, with po- tentially important consequences for its population dynamics. Vol. 50, No. 1, January 1974 59 Acknowledgments The work was partially supported by a grant from the Research Foundation, California State University, Hayward. C. E. Dasch assisted with identification of parasitoids. R. S. Horn helped search for host pupae and she and J. P. Harville read drafts of the manuscript. The insatiable appetites of C. A. and R. B. Horn kept me abundantly sup- plied with 4-oz jars. Literature Cited Bobb, M. L. 1965. Insect parasite and predator studies in a declining sawfly population. J. Econ Entomol., 58: 925-926. Burks, B. D. 1960. A revision of the genus Brachymeria Westwood in America North of Mexico (Hymenoptera: Chalcidae) . Trans. Amer. Entomol. Soc., 86: 225-273. Cogswell, H. L. 1966. Thirtieth breeding bird census 40. Broadleaf evergreen forest with shrub-filled openings. Audubon Field Notes, 20: 637-643. Doane, R. W. 1912. Some insect pests of the California live oaks. J. Econ. En- tomol., 5: 346. Force, D. C. 1972. r- and K-strategists in endemic host-parasitoid communities. Bull. Entomol. Soc. Amer., 18: 135-137. Harville, J. P. 1955. Ecology and population dynamics of the California oak moth Phryganidia calif ornica Packard (Lepidoptera) Microentomol., 20: 83-166. Horn, D. J. 1973. Leptocoris rubrolineatus, an occasional predator of the Cali- fornia oakworm, Phryganidia calif ornica. Pan-Pac. Entomol., 49: 196. Kellogg, V. K. and F. J. Jack. 1895. The California phryganidian. Proc. Calif. Acad. Sci. (Ser. 2), 5: 562-570. Langston, R. L. 1957. A synopsis of hymenopterous parasites of Malacosoma in California. Univ. Calif. Publ. Entomol., 14: 1-50. Muesebeck, C. F. W., K. V. Krombein and H. K. Townes. 1951. Hymenoptera of America north of Mexico. Synoptic catalog. USDA Agr. Monog., 2: 1-1420. Price, P. W. 1970. Trail odors: recognition by insects parasitic on cocoons. Science, 170: 546-547. Price, P. W. 1971. Parasitoids utilizing the same host: adaptive nature of dif- ferences in size and form. Ecology, 53: 190-195. Stehr, F. W. and E. F. Cook. 1968. A revision of the genus Malacosoma Hubner in North America (Lepidoptera: Lasiocampidae) : systematics, biology, immatures, and parasites. U.S. Nat. Mus. Bull., 276: 1-321. Townes, H. and M. Townes. 1960. Ichneumon flies of America north of Mexico. 2. Subfamilies Ephialtinae, Xoridinae, Acaenitinae. U.S. Nat. Mus. Bull., 216: 1-676. 60 The Pan-Pacific Entomologist Biology of Three Callirhytis Gall Wasps from Pacific Slope Erythrobalanus Oaks (Hymenoptera: Cynipidae) D. Charles Dailey, Tim Perry, and Christine M. Sprenger Sierra College, Rocklin, California 95677 This paper presents life histories of two species of alternating unisex- ual and bisexual generation gallmaking cynipids, Callirhytis eldoradensis (Beutenmuller) (= Andricus eldoradensis Beutenmuller, 1913) and Callirhytis flora Weld $2 (1922a) ( = Callirhytis milleri Weld 9, 1922b) new synonym, and a description of the males of a third species, Callirhytis per foveata (Kinsey) $ 2 (1922). All three of these gallmaking cynipid species occur on all three species of Pacific slope oaks in the subgenus Erythrobalanus : Quercus agrifolia Nee, Quercus kelloggii Newberry, and Quercus wislizenii A. DeCandolle. Callirhytis eldoradensis (Beutenmuller), new combination Callirhytis eldoradensis emerges from integral acorn hull galls in aborted acorns (Fig. 1). The original description included only males and erroneously associated them with galls of Callirhytis milleri Weld. The bisexual generation female and correct gall were described by Weld (1922). This generation is unusual because the male is larger than the female and emergence is during late September and October rather than in the spring or early summer. The previously undescribed unisexual generation emerges in late spring or early summer from a spherical bud gall (Fig. 2). Biology. — During October 1970 a collection of C. eldoradensis was reared indoors and individuals of both sexes were confined together in bags on three native Q. wislizenii. On the following two days seven naturally emerged (not escaped) males were collected on the bags’ ex- teriors. This suggests the possibility of a female emitted pheromone. When the natural female emergence was beginning and bagged females began oviposition, three consecutive days of rain and cold weather inter- fered with observations and caused a high mortality of reared and pos- sibly naturally emerged insects. When the bags were removed in late March, 1971 the twigs contained green spherical bud galls about four mm in diameter with white basal hairs. The galls abort the apical meristem and are surrounded basally by the bud scales. Four of these galls figured as numbers 191—194 by The Pan-Pacific Entomologist 50: 60-67. January 1974 Vol. 50, No. 1, January 1974 61 Figs. 1-2. Callirhytis eldoradensis. Fig. 1. Bisexual generation gall, 5X- Fig. 2. Unisexual generation gall, 15X- Houard (1935) were erroneously identified as Andricus attractans Kin- sey, now in Callirhytis. They do not secrete honeydew as do C. at- tractans galls. They contain a succulent layer of nutritive tissue around the larval cell which provides a food and moisture reserve for the larva after the gall falls to the ground in April. Emergence is during the late spring or early summer, apparently the next year. One specimen reared at the U.S. National Museum emerged 27 April 1923 from galls collected May 1922. Another, reared indoors by the author, emerged prior to 15 June 1971 from galls collected April 1970. None of the adjacent limbs or trees, including the original source of insects, harbored the galls, probably due to both previous intensive collecting and inclement weather. Nor did any other unisexual generation galls occur on the isolated twigs. Morphology of the unisexual generation requires transfer of the species from its previous assignment in Eumayria Ashmead to Calli- rhytis Foerster group B, which includes almost exclusively species as- sociated with acorns. The following additional circumstantial evidence also substantiates this association of alternate generations. Bisexual generations usually emerge about two to three months later than the unisexual generation, which in this case suggests natural emergence and oviposition of the uni- sexual generation about May or June. By this time the trees are usually completing seasonal growth and though galls can be induced in some differentiated nearly mature plant tissue (Lyon, 1970), one of the few 62 The Pan-Pacific Entomologist plant tissues still growing sufficiently to be an optimum oviposition site for this unisexual generation female is the rapidly enlarging acorns. Callirhytis flora $ 2 , C. milleri £ , and Callirhytis carmelensis Weld $ are the only other gallmaking cynipids known to be associated with Pacific Slope Erythrobalanus acorn galls. The first two are alternate generations of the same species and thus neither could be an alternate generation of C. eldoradensis. It is highly unlikely that both generations of a cynipid would occur on a tissue so limited in availability as acorns, especially since a large brood of C. carmelensis would abort the acorns needed for a brood of C. eldoradensis. Also though acorns of both Q. kelloggii and Q. wislizenii require two seasons to mature and might re- ceive eggs during the first % of their time on the tree and sustain the bisexual generation of C. eldoradensis , Q. agrifolia matures acorns in only one season. Thus the typical winter emerging unisexual generations, such as C. carmelensis, cannot be part of the life cycle of C. eldoradensis because on Q. agrifolia they would have no acorns available in which to oviposit. There are no other known spring or early summer emerging unisexual generations associated with acorns which might potentially be considered as the alternate generation of C. eldoradensis. Systematics. — Female. Head: nearly massive, broadly triangular in front view, widest at antennal socket level, slightly narrower than thorax. Occiput darker, elevated about % width of ocellus above head contour. Cheeks widened behind eyes. Malar groove absent, malar area pubescent with fan-shaped striae radiating from clypeus corners. Antenna 13 segmented, last segment twice the length of the preceding. Thorax: mesoscutum sparsely pubescent, with fine transverse rugosity. Notaulices weakly complete, distinct only posteriorly. Anterior parallel lines and parapsidal lines extending nearly % length of mesoscutum. Median groove polished, Vs length of mesoscutum. Scutellum coriaceous centrally, reticulate laterally; foveae reticulate medially, polished smooth laterally. Mesopleuron ridged longi- tudinally. Tarsal claws simple. Wings: veins pale yellow-brown; faintly clouded, surface with short pubescence, margin bare. Radial cell open, aerolet % length of open cubital cell. Abdomen; first tergite polished smooth with dorsally inter- rupted pubescent ring at base, succeeding tergites polished micropunctate, three tergites visible dorsally. Ventral spine 12 times as long as wide, sparsely pubescent, forming 90° angle with ventral valves, equal to length of entire hind tarsus. Color: amber, abdomen darker. Type Deposition . — Plesiotype female in U.S. National Museum. Callirhytis flora Weld Callirhytis flora, which was described from females (though Weld also reared males) that emerged from an integral hard midrib leaf gall (Fig. 3) during May and June, is the alternate bisexual generation of Callirhytis milleri, new synonym which emerges between late Feb- Vol. 50, No. 1, January 1974 63 ruary and mid March from lumpy aborted acorn cotyledons (Fig. 4). Unisexual generation galls have been illustrated by Beutenmuller (1913) and Weld (1922b). The former erroneously associated them with Cal- lirhytis eldoradensis. Bisexual generation galls are also figured by Weld (1922a, 1957). Both generations belong to Weld’s group B, which in- cludes primarily unisexual generations from hard acorn galls. C. flora and C. milleri are hereby synonymized, with Callirhytis flora Weld having priority. Biology. — On 20 March 1969 numerous unisexual generation females placed on Q. wislizenii with rapidly elongating buds readily oviposited in the midrib of the lower surface of young leaves. Bisexual generations usually take two or three months to develop, and the only known bi- sexual group B Callirhytis emerging from a leaf gall on California Erythrohalanus oaks during May to June is C. flora. The long ovipositor of C. flora (9.3 X head width) caused Weld to believe it oviposited in acorns. Such an ovipositor would only be nec- essary for ovipositing deep in plant tissue, such as the center of a ma- turing acorn. The senior author collected ten bisexual generation fe- males of C. flora while they were ovipositing in acorns of Q. agrifolia and in second year acorns of Q. wislizenii during May and June 1967 to 1969. These acorns drop to the ground in the fall and the unisexual generation larvae remain in the galls at least IVz years before pupation. Weld (1922b) has recorded a larval period lasting 2% years. C. flora and C. milleri key out to the same couplet (Weld, 1922b). This structural similarity of alternate generations is paralleled by that of Callirhytis pomiformis (Bassett) (Lyon, 1959). The only other known Pacific Slope gallmaking cynipids reared from Erythrohalanus acorn galls are C. eldoradensis and Callirhytis carmelensis Weld. Neither of these could result from the oviposition of C. flora because of the fol- lowing reasons. C. eldoradensis from acorns is also a bisexual gen- eration, and the alternate unisexual generation emerges from a spherical bud gall. Acorn pip galls of C. carmelensis develop during March and April and drop to the ground about May. The acorns of Q. kelloggii and Q. wislizenii , which require two years to mature, could receive eggs nearly any time during the first or second years and produce galls of C. carmelensis. However, Q. agrifolia produce mature acorns between February and October of the same year. Since the bisexual generation of C. flora emerges in June, any Q. agri folia acorns in which they might oviposit would drop to the ground that autumn and not remain on the tree until the following spring. C. flora bisexual generation individuals 64 The Pan-Pacific Entomologist Figs. 3-4. Callirhytis flora. Fig. 3. Bisexual generation gall, 2.5 X- Fig. 4. Unisexual generation gall, 2.5 X- do not live through the winter until the following spring to be able to oviposit in the newly forming crop of acorns. C. carmelensis could therefore not be the alternate generation of C. flora. Systematics. — Male. Head: massive, not widened behind eyes, malar space grooved; antennae 15 segmented, third excavated dorsally, last equal to preceding. Thorax: scutum coriaceous, coarse posteriorly, notaulices complete, anterior par- allel lines distinct; foveae shallow, ridged, weakly separated; scutellum rugose, smoothest centrally; mesopleuron smooth dorsally and ventrally, coriaceous med- ially; wings pubescent, margins ciliate, veins pale beyond second crossvein. Ab- domen: smooth, bare, brown, lightest dorsally, not pedicellate. Length: 1.7 mm. Color: frons, genae, antennae, and legs yellow-brown, remainder red-brown. A unisexual generation male sport which emerged 22 January 1969 differs from bisexual generation males by its larger size (2.8 mm) ; uniformly brown wing- veins; last antennal segment slightly longer than preceding; scutum finely rugose anteriorly, coarser posteriorly; polished coriaceous abdominal tergites. Type Deposition . — Plesiotype bisexual generation male and unisexual generation male sport in U.S. National Museum. Plesioparatype males in the Weld collection in the possession of Robert J. Lyon, Los Angeles City College (3), California Academy of Sciences (10), senior author’s collection (60), and U. S. National Museum (5) . Vol. 50, No. 1, January 1974 65 Figs. 5-6. Callirhytis perfoveata, bisexual generation galls. Fig. 5. Position of galls on leaf, 2X- Fig. 6. Detail of gall structure, 7X- Distribution. — Probably found throughout the range of the host plants. Parasite Biology. — Euderus craw fori Peck (Hymenoptera: Eulophi- dae) adults were dissected from larval cells of bisexual generation C. flora which contained partly consumed adult gallmakers indicating the parasites did not attack the gallmakers until the latter completed pu- pation. Callirhytis perfoveata (Kinsey) When Kinsey described the rapidly developing spring cynipid C. perfoveata from six females (1922) he stated, “such species of cynipids are usually bisexual, so the male may yet be discovered for the species.” This has been confirmed with the rearing of males from Quercus wis- lizenii. Biology. — The green, nearly spherical, succulent, monothalamous in- tegral leaf galls (Figs. 5 & 6) were noted by the senior author in the new growth of Q. wislizenii at Folsom Lake, Placer Co., California, 19 March, 1967. They were fully developed and contained mature larvae. On 23 March some contained white pupae. Galls collected then and maintained indoors 65-75° F yielded two females on 1 April. Galls still on the trees showed no evidence of natural emergence. From galls collected 15 April both males and females emerged from 20 to 29 April (indoors). By May over half of the galls examined on the trees had emergence holes. Sprenger collected C. perfoveata from Q. wislizenii in Rocklin, Placer Co., California on 13 and 14 April, 1973. Insects 66 The Pan-Pacific Entomologist emerged 13 to 27 April while being maintained indoors. Neither the oviposition site of the females nor the alternate unisexual generation is known. Galls have also been seen at Lemoncove, Tulare Co., and Clear- lake, Lake Co., California. Systematics. — Female. Similar to Kinsey’s 2 description, but with the following variation: proximal antennal segments yellow-hrown darkening to deep brown distally; cheeks and malar space reddish-brown to black; mandibles yellow-brown to reddish-brown; median groove sometimes indistinct. Using maximum width of the head as a base, the mesonotum length ratio is 1.3, antennae 2.3, wings 3.7. Length of 48 specimens, 2.1-2.6 mm. Average 2.4 mm. Male. Head coriaceous; from above transverse, as broad as thorax, not broad- ened behind eyes; in anterior aspect, interocular space twice as wide as high, malar space 0.2 eye height without groove; antennae 15 segmented. Mesonotum cori- aceous, notaulices complete, broader behind, median groove less distinct than in females; scutellum rugose, smoother medially, fovea smooth and polished, separated by ridge; mesopleuron nearly bare, more striate ventrally. Wing hyaline, margin ciliated, veins brown, radial cell open, aerolet % wing length. Claws simple. Abdomen non-pedicellate, shorter than head plus thorax, tergite II smooth. Using maximum head width as base, mesonotum length is 1.4, antennae 3.3, wing 4.1. Length of 27 specimens, 1.8-2.4 mm, average 2.1 mm. Body, head, and antennae black, abdomen dark brown, lightest dorsolaterally on tergite two. Type Deposition. — The series contains 27 males and 48 females. Plesioallotype male in U.S. National Museum. Plesiotype insects of both sexes and freeze dried galls have been deposited at the U.S. National Museum, California Academy of Sciences, Weld collection in the possession of Robert J. Lyon of Los Angeles City College, University of California at Davis, and in the authors’ collections. Acknowledgments The authors are grateful to Mr. Robert J. Lyon of Los Angeles City College for the use of Weld’s collection and field notes, to Dr. B. D. Burks of the U.S. Natl. Museum for identifying the chalcid parasites, and to Linda Campbell and Barbara Young for the illustrations. Literature Cited Beutenmuller, W. 1913. The North American acorn galls with descriptions of new species. Bull. Brooklyn Entomol. Soc., 8: 101-105. Houard, C. 1935. Les collections cecidologiques du laboratoire d’Entomologie du Museum d’Histoire naturelle de Paris: galles des Etats-Unis. Marcellia, 28: 162-163. Kinsey, A. C. 1922. New Pacific coast cynipidae. Amer. Mus. Nat. Hist. Bull., 46: 287-288. Lyon, R. J. 1959. An alternating sexual generation in the gall wasp Callirhytis pomiformis (Ashm.). Bull. So. Calif. Acad. Sei., 58: 33-37. Lyon, R. J. 1970. Heterogony in Callirhytis serricornis (Kinsey) . Proc. Entomol. Soc. Wash., 72: 176-178. Vol. 50, No. 1, January 1974 67 Weld, L. H. 1922a. Notes on cynipid wasps, with descriptions of new North American species. Proc. U.S. Natl. Mus., 61: 20. Weld, L. H. 1922b. Notes on American gallflies of the family cynipidae pro- ducing galls on acorns, with descriptions of new species. Proc. U.S. Natl. Mus., 61: 1-32. Weld, L. H. 1957. Cynipid galls of the pacific slope. Privately Printed, Ann Arbor, Mich., 64 pp. SCIENTIFIC NOTE Recovery of Anax junius from a glacier in the Sierra Nevada (Odonata: Aeschnidae). — On August 16, 1972, while traversing the Lyell Glacier in Yosemite National Park with Charles Dickson, ranger-naturalist at Tuolumne Meadows, I discovered several specimens of a dragonfly, Anax junius (Drury), frozen into the ice at an altitude of 12,200 feet. The winter of 1971-72 had been unusually mild, resulting in one of the lightest snowpaclcs of this century. As a consequence, many of the Sierran glaciers retreated, and surfaces of ice deep within them were ex- posed. These particular dragonflies were undoubtedly swept onto Mount Lyell by upslope winds to perish on the glacier, where they have remained embedded for uncounted years. R. W. Garrison of the Dept, of Entomological Sciences at the University of Cal- ifornia, Berkeley, who confirmed the identification, also noted that one of the females was teneral and was therefore probably trapped on the glacier during her first week of life, possibly on her maiden flight. Anax junius is a very widespread species, occurring throughout North and Cen- tral America. During previous summers in the high Sierra, I have taken numerous specimens in Dana Meadows at Tioga Pass, where it undoubtedly breeds in sub- alpine lakes there. — R. P. Papp, Department oj Entomological Sciences, University of California, Berkeley 94720. 68 The Pan-Pacific Entomologist Observations on the Behavior of Mallophora fautrix Osten Sacken (Diptera: Asilidae) John Alcock Department of Zoology, Arizona State University, Tempe 85281 Mallophora fautrix subspecies bromleyi Curran (Cole and Pritchard, 1964) is a robust asilid with a close mimetic resemblance to the bumble- bee Bombus sonorus Say (see Plate 53, Linsley 1960). Linsley has pro- vided a brief account of the ethology of this species (under the name M. bromleyi ) with emphasis on the prey taken. The purpose of this paper is to offer additional information, largely on the mating and oviposition behavior of the fly. My observations were made during the first two weeks of August 1973 at a site near Cave Creek, approximately 2 km east of Portal, Arizona and 1 km north on the road to San Simon. The flies were abundant in an area of dense vegetation watered by a small irrigation ditch; the predominant plant was Cortyza coulteri Gray although scattered cockleburrs, Xanthium saccharatum Wallroth, and other desert vegetation were also present (see Linsley and Cazier, 1972, for a description of the locality and its flora). Mating behavior. — Asilid activity at the site began at about 0900- 0930 Mountain Standard time and continued at a high level until about 1230. The morning was apparently devoted primarily to reproductive behavior in as much as I never saw a fly with prey during this time despite the presence of many individuals. Males flew from one perch to another with brief intervals of rest. In the course of their flights they frequently hovered with the hindlegs widespread near conspicuous burrs, large seed heads, and perched individuals of the same species. They then darted at these objects sometimes striking a plant part re- peatedly before flying away. They also pursued conspecific individuals in the air in a noisy, bobbing “roller-coaster” flight which often carried them 3-4 m high. Although the onset of copulation was not observed, I did see five attempted matings (1020-1245) and four coupled pairs (1000-1205). Males struck and attempted to grasp perched or egg-laying females. If successful they would fall to the ground holding their would-be mate firmly. They then probed with their abdomen in an effort to achieve genital coupling. In the cases of attempted matings that I watched the male was often oriented with respect to the female in ways that made The Pan-Pacific Entomologist 50: 68-72. January 1974 Vol. 50, No. 1, January 1974 69 Fig. 1. Two positions assumed by copulating Mallophora fautrix. copulation impossible (for example, one male held the female by her abdomen while probing her head and thorax) and sooner or later the the pair broke apart. Mating pairs (Fig. 1) were typically observed clinging to the vegetation in the tail-to-tail position, while disturbed pairs flew from one area to another with one fly trailing behind the other. BEE 0830 1000 1200 1400 Fig. 2. The flight periods of the bumblebee B. sonorus and its mimic, Malloph- ora fautrix, at the Portal site. Data collected on 10 August (0830-1100 Mountain Standard Time) and 12 August (1100-1400) 1973. Black boxes indicate that at least one flying bee or asilid was seen during a five min. interval within five m. of the observer. On 13 August observations between 1400-1700 produced just three sightings of a bumblebee and one of the asilid. 70 The Pan-Pacific Entomologist Fig. 3. A female Mallophora fautrix in the act of producing an egg case. Note that the case is constructed of thin coils of white oothecal material. Flight activity. — The period of considerable flight activity coincided closely with the time when the fly’s abundant model, B. sonorus, was also active in the area (Fig. 2) . Whether the synchronous flight periods reflect a common thermal constraint on energetic flight for both species or whether selection has favored flies which time their conspicuous aerial reproductive activities to match the flight period of the model could not be determined. Perhaps both factors are responsible for the daily pattern of activity of M. fautrix. Egg-laying. — Females laid eggs between 1035-1245 (N = 10) . As Linsley surmised (1960), this species places its egg cluster at the tip of dried weed stems (especially Conyza coulteri ) at a height of 1-2 m above the ground. Females first alighted head-up on a suitable site and then moved about probing with their abdomen, apparently searching Vol. 50, No. 1, January 1974 71 for an open area on which to place the egg case. Having located such a site, the females then curled their abdomen and began to exude a thin coil of white foam (Fig. 3). By moving their abdominal tip back and forth in a semi-circle the insects created a half-cylinder with a cavity which was filled with masses of eggs at short intervals. Egg cases ranged in size from 0.5-2. 0 cm; females were sometimes interrupted in the midst of egg-laying by a male (three observations). Discussion Lavigne and Holland (1969) have summarized the information avail- able on the courtship and oviposition behavior of the Asilidae (see also Lavigne, 1970a, b; Rogers and Lavigne, 1972, for additional details). Although complex courtship displays have evolved in a few genera of asilids, most members of this family omit this component of reproduc- tive behavior. Mallophora fautrix appears to belong with the majority in this regard although, as noted, males will hover near certain objects {e.g. burrs) with their hind legs extended. This behavior could con- ceivably provide a female with a courtship signal. In addition, the func- tion of the oscillating flights may also be related to courtship although this remains speculative. The oviposition behavior observed for this species is unusual for the Asilidae. Apparently only species of Mallophora and Mallophorina produce chalky egg cases and place them on open stems. However, a small number of asilids, including some species of the closely allied genus Protomachus , oviposit on vegetation, generally in some protected site, such as the florets of a grass or the space between a grass stem and leaf (Lavigne and Holland, 1959). Acknowledgments Mr. Joe Wilcox identified specimens of the fly for me. Dr. Mont A. Cazier directed me to the relevant literature and identified the plants named in the paper. This study was made possible by a faculty Grant- in-Aid from Arizona State University. Literature Cited Cole, F. R. and A. E. Pritchard. 1964. The genus Mallophora and related asilid genera in North America (Diptera: Asilidae). Univ. Calif. Pubs. Entomol., 36: 43-93. Lavigne, R. J. 1970a. Courtship and predatory behavior of Cyrtopogon auratus and C. glarealis (Diptera: Asilidae). J. Kans. Entomol. Soc., 43: 163- 171. 72 The Pan-Pacific Entomologist Lavigne, R. J. 1970b. Courtship and predation behavior of Heteropogon macu- linervis (Diptera: Asilidae) . J. Kans. Entomol. Soc., 43: 270-273. Lavigne, R. J. and F. R. Holland. 1969. Comparative behavior of eleven species of Wyoming robber flies (Diptera: Asilidae). Univ. Wyo. Agr. Exp. Sta., Sci. Monog., 18: 1—61. Linsley, E. G. 1960. Ethology of some bee- and wasp-killing robber flies of southeastern Arizona and western New Mexico (Diptera: Asilidae). Univ. Calif. Pubs. Entomol., 16: 357-392. Linsley, E. G. and M. A. Cazier. 1972. Diurnal and seasonal behavior patterns among adults of Protoxaea gloriosa (Hymenoptera, Oxaeidae) . Amer. Mus. Novit, 2509: 1-25. Rogers, L. E. and R. J. Lavigne. 1972. Asilidae of the Pawnee National Grass- lands in northeastern Colorado. Univ. Wyo. Agr. Exp. Sta., Sci. Monog., 25: 1-35. SCIENTIFIC NOTE Lepturine Longhorns Collected on Chinquapin Bloom (Coleoptera: Cerambycidae) . — The forested regions of western Oregon are moderately rich in members of the cerambycid tribe Lepturini. Adults of most species of these anthophilous beetles appear in the spring and early summer and collectively visit the blossoms of a wide variety of herbaceous and woody plants. Periods of activity vary with altitudinal distribution but usually by late July many species and their host flowers have disappeared. However, there are some lepturines which fly throughout July and August in the middle elevations of the Cascades, and for these chinquapin appears to be a favored and important source of pollen. Giant chinquapin, Chrysolepsis chrysophylla (Dougl. ex Hood.) Hjelmquist, is widely distributed in the Cascade and Coast ranges and its showy catkins of creamy white, heavily fragrant flowers are commonly in evidence from July to early September. Eleven species of longhorns were collected on chinquapin blossoms as follows: Anoplodera amabilis LeC., A. canadensis Oliv., A. chrysocoma Kirby, A. crassi- cornis LeC., A. crassipes LeC., A. dehiscens LeC., A. dolorosa LeC., A. tibialis LeC., Leptura obliterata Hald., L. propinqua Bland, and Stenocorus flavolineatus (LeC.). Species diversity and abundance varied with locality except for A. cana- densis and S. flavolineatus which were represented by single specimens. Collection records include: % mile east of Suttle Lake Junction, Jefferson County, August 1, 1973; three miles east of Santiam Pass, Jefferson County, August 1, 1973 and; 13.2 miles south of Marion Forks, Linn County, August 4, 1973. A. crassicornis has been found abundantly on chinquapin on Doak Mountain, Klamath County, along with a few specimens of A. amabilis and A. dehiscens. — Richard L. Penrose and Richard L. Westcott, Oregon Department of Agriculture, Salem 97310. Vol. 50, No. 1, January 1974 73 Unusual Caterpillar-Prey Records and Hunting Behavior for a Podalonia Digger Wasp: Podalonia valida (Cresson) (Hymenoptera: Sphecidae) A. L. Steiner Department of Zoology, University of Alberta, Edmonton, Alberta, Canada Observations on Podalonia valida (Cresson) were made in late sum- mer and fall 1972, in an area immediately adjacent to the Chiricahua Mountains in extreme southeastern Arizona, about one mile west of the Chiricahua National Monument entrance, and about 35 miles south of Willcox, Cochise County. The study area, described in more detail elsewhere (Steiner, in prep- aration) , was situated in a riparian habitat along a temporary creek (Bonita Creek), in the Upper Sonoran/Transition Zones. This oasis-like area was surrounded by more arid desert scrub and grassland. Within the study area, a lush herbaceous vegetation, tall trees, and a shrubby understory were present. Four female Podalonia valida were observed in the study area be- tween September 10 and October 9, 1972. Detailed prey records were made on only one individual. This individual paralyzed and stored in her nests, large, very hairy caterpillars of the genus Estigmene (Arcti- idae), exclusively. In seven observed cases, all prey were large hairy caterpillars ranging from a black color, with rufous-reddish latero- ventral parts, to brownish-yellowish. All caterpillars were probably Estigmene acraea (Drury), or very closely related forms. Predatory Behavior of Caterpillar Hunting Wasps Lepidopterous larvae of various species, shapes, sizes, colors and habitats are preyed upon by digger wasps of the genera Ammophila and Podalonia (and some other genera including some solitary Vespidae). The majority of these wasps prey on naked caterpillars only (see for instance Murray, 1940; Muesebeck et al ., 1951; Evans, 1959; Powell, 1964; Krombein, Burks et al., 1967, for Nearctic species, and Berland, 1925; Roth, 1928; Tsuneki, 1968, for Palearctic species). A few rec- ords of odd prey species are available, however. Thus according to Ap- pleton (in Murray, 1940), Podalonia occidentalis Murray preys on the tent-caterpillar. Ammophila pilosa pilosa (Fernald) (= azteca Cam- eron) was found to store a Lycaenid larva, about 14 mm length, densely covered with short secondary setae, and also a Pterophorid larva (about The Pan-Pacific Entomologist 50: 73-77. January 1974 74 The Pan-Pacific Entomologist 15.5 mm length), with numerous tufts of elongate secondary setae; these caterpillars have rather concealed feeding habits (Powell, 1964) . Neither Lycaenids nor Pterophorids seem to have been reported as prey of other North American Ammophila (Powell, 1964). According to Ferton (1901, 1914), the Palearctic Podalonia hirsuta (Scopoli) (var. mervensis Radoszkowsky : “Ammophila ebenina” Costa) takes naked prey such as Agrotis , Cucullia chamomillae, but also hairy caterpillars, for instance Epinephele jurtina (Berland, 1925). A number of Podalonia species of known biology prey on Agrotidae- Noctuidae (cutworms), sheltered underground during daytime, e.g., Podalonia robusta (Cresson) : Evans, 1963; luctuosa and/or communis (Smith) : Hicks, 1931, 1932, for Nearctic species — hirsuta (Scopoli) : Fabre, 1879, 1882; affinis (Kirby) : Marchal, 1892; tydei (Le Guillou) : Fabre, 1879; Picard, 1903; atrocyanea Eversmann ( = chalybea Kohl), microhirta Kohl and/or caucasica Morawitz: Tsuneki, 1968, for Pale- arctic species). The wasps excavate these nocturnal prey from their underground shelters. Podalonia valida , in contrast, hunts diurnal lepidopterous larvae in the open, usually on dense herbaceous or low brushy vegetation such as white horsemint ( Monarda pectinata ) , Helianthus (Compositae) , and goldweed ( Verbesina encelioides) . In the area considered, these lepi- dopterous larvae were found to move from one plant species to another quite extensively. Whether this was related to polyphagous feeding be- havior or to escape behavior, in response to disturbances by hunting wasps, was not determined, but captive specimens readily accepted some plants of the sunflower family. Podalonia valida was capable of follow- ing with great accuracy the fresh scent trails left by the caterpillars, while moving on the ground from one plant to another. The wasp proceeded head down, tapping her antennae on the ground, especially where the caterpillar had stopped and rested for some time (mostly at the bases of plants), or had made a sudden and sharp change in direction. This ground level search behavior often led the wasp to the base of one plant, where a concentrated search would be undertaken, the excitation of the wasp increasing steadily in the process. There is little doubt that the wasp was picking up more and more fresh olfactory evidence of a re- cent occupancy of the site by the prey. Often, the wasp would then climb up the plant and investigate it thoroughly. At other times the wasp would fly directly onto the plants, without first searching the ground. Discovery of the caterpillar did not necessarily follow, however, even if the wasp came very close to the prey. Such failure to capture the prey Vol. 50, No. 1, January 1974 75 Fig. 1 . Podalonia valida carrying a large, paralyzed lepidopterous larva ( Estig - mene acraea ) (Arctiidae), using her mandibles and fore legs to hold the prey. Locomotion, with two pairs of legs only, is assisted by wing beats. was particularly frequent at early stages of hunting phases. This sug- gests that “motivational” deficiencies, rather than deficiencies in the ability to detect or locate the prey, were then involved. Such incomplete behavior is very frequent at early stages of the different phases of the nesting cycle in digger wasps (Steiner, 1962). These abortive activities are particularly conspicuous during the nest-digging phase, resulting in one or several incomplete burrows abandoned at once. Such trun- cated behavior also occurs at early stages during other phases of the nesting cycle, including the hunting and prey-stinging stages (Steiner, 1962), and several aborted pursuits, attacks or stinging of prey by Podalonia valida were recorded in this study. In these cases the wasp did not make any further attempt at pursuing the escaping prey. At such early stages, in contrast to later stages of the hunting phase, the wasp would not usually accept a suitable prey, even if dropped right in front of her. Such apparent unwillingness to capture a detected prey has also been reported from other wasps, including other Podalonia species. For example observations made on Podalonia luctuosa (and/ or communis ) (Newcomer, 1930; Hicks, 1931, 1932, in Murray, 1940), which preys on cutworms (Noctuidae), concealed beneath the soil by day, revealed abortive termination of digging behavior during hunting. No evidence of hunting by sight was found in P. valida , except per- haps at very close range. Little information on prey stinging was ob- 76 The Pan-Pacific Entomologist tained, since this occurred mainly in very dense vegetation and was ac- complished within a very short period of time. The heavy, cumbersome prey is straddled and carried over the ground in typical Podalonia fashion, using the mandibles and also the front legs, which embrace the larva (Fig. 1). Wing beats provide some additional propulsive force. A more detailed account of the biology and behavior of this Podalonia species will be presented elsewhere. Acknowledgments I wish to express my gratitude to the Directors, staff and guests of the Southwestern Research Station, Portal, Arizona; of The American Mu- seum of Natural History (New York) ; and of the Chiricahua National Monument, Arizona, for their hospitality, facilities provided, help and useful suggestions at various stages of this project. The wasp specimens were identified by R. M. Bohart, University of California, Davis, and the prey specimens by D. M. Weisman, U.S.D.A., Agriculture Research Center, Beltsville, Maryland (R. H. Foote, Chief Systematic Entomology Eaboratory). Some of the plants were identified by S. Mouat. Special thanks are due to B. and E. Erickson and L. Riggs, for kindly giving me free access to their property and ranch. This work was supported in part by NRC grant A-3499. Literature Cited Berland, L. 1925. Faune de France — Hymenopteres vespiformes. I (Sphegidae, Pompilidae, Scoliidae, Sapygidae, Mutillidae) . Lechevalier, Paris. 364 pp. Evans, H. E. 1959. Observations on the nesting behavior of digger wasps of the genus Ammophila. Amer. Midi. Nat., 62: 449-473. Evans, H. E. 1963. Notes on the prey and nesting behavior of some solitary wasps of Jackson Hole, Wyoming. Entomol. News, 74(9): 233-239. Fabre, J. H. 1879. Souvenirs entomologiques. Paris. Fabre, J. H. 1882. Nouveaux souvenirs entomologiques. Paris. Ferton, C. 1901. Notes detachees sur l’instinct des Hymenopteres melliferes et ravisseurs. Ann. Soc. entomol. Fr., 70: 83-148. Ferton, C. 1914. Notes detachees..., 8e serie. Ann. Soc. entomol. Fr., 83: 81- 118. Hicks, C. H. 1931a. On the digger wasp, Podalonia luctuosa (Smith). Pan- Pacif. Entomol., 8: 49-51. Hicks, C. H. 19316. The hunt and capture of the prey of a digger wasp. Bull. South. Calif. Acad. Sci., 30: 75-82. Hicks, C. H. 1932. Notes on the prey and inquilines of Podalonia violaceipennis form luctuosa (F. Smith). Psyche, 39: 150-154. Krombein, K. V., B. D. Burks et al. 1967. Hymenoptera of America North of Vol. 50, No. 1, January 1974 77 Mexico. Second supplement. U.S. Department of Agriculture, Washing- ton, 584 pp. Marchal, P. 1892. Notes sur la vie et les moeurs des Insectes. Observations sur Y Ammophila affinis Kirby. Arch. Zool. exper., 23-36. Muesebeck, C. F. W., K. V. Krombein, H. K. Townes et al. 1951. Hymenop- tera of America north of Mexico — Synoptic Catalog. U.S. Dept. Agric. (Agriculture Monograph No. 2), Washington. 1420 pp. Murray, W. D. 1940. Podalonia (Hym. : Sphecidae) of North and Central America. Entomol. Amer. (n.s.), 20: 1-82. Newcomer, E. J. 1930. Notes on the habits of a digger wasp and its inquiline flies. Ann. Entomol. Soc. Amer., 23: 552-563. Picard, F. 1903. Moeurs de YAmmophila tydei Guill. Feuille des jeunes natural- istes, 34: 15-17. Powell, J. A. 1964. Additions to the knowledge of the nesting behavior of North American Ammophila (Hym.: Sphecidae). Jour. Kans. Entomol. Soc., 37: 240-258. Roth, P. 1928. Les Ammophiles de l’Afrique du nord. Ann. Soc. entomol. Fr., 97: 153-240. Steiner, A. L. 1962. Etude du comportement predateur d’un Hymenoptere Sphegien: Liris nigra V.d.L. ( — Notogonia pompiliformis Panz.). Ann. Sci. Nat. Zool. Biol. Anim. Ser. 12, 4: 1-126. Tsuneki, K. 1968. The biology of Ammophila in East Asia (Hym., Sphecidae). Etizenia, 33: 1-64. 78 The Pan-Pacific Entomologist The Larval Nutrition of Minettia jlaveola and Phaonia parviceps and its Significance to the Hawaiian Leaf -breeding Drosophila (Diptera: Lauxaniidae, Muscidae, Drosophilidae) J. F. McDonald, W. B. Heed 1 and Mary Miranda Departments of Genetics and Food Sciences and Technology, University of California, Davis, California 95616 The purpose of the present report is to point out the similarities and differences in the larval ecology and nutrition of several species in the two Dipteran families, Lauxaniidae and Drosophilidae. The habit of mining in decaying detached leaves by representatives of these two fam- ilies presents an unusual opportunity to study the dynamics of host plant selection and related phenomena such as competition and nutrition. The leaf-breeding habit in the Drosophilidae is known only in the Hawaiian Islands and is universal in the genus Antopocerus and three related groups of species within the genus Drosophila (Heed 1968). Species in other groups and in the genus Scaptomyza also breed in fermenting leaves and the habit appears to have evolved independently several times. In total there are approximately 80 to 100 leaf-breeding species endemic to the Islands. Holarctic members of the Lauxaniidae have long been known to occupy this niche (Hering, 1951; Oldroyd, 1964). However until the present report no one has compared larval nutrition in the drosophilids and the lauxaniids. The overall similarity of the rotting leaf niche available for the two groups is so striking that a de- tailed account of recent observations and tests is given below. The breeding biology of the Hawaiian leaf-breeders has been com- pared previously (Heed 1971) to the unusual biology of Rhagoletis (Tephritidae) , as observed by Bush (1969). Results and Discussion On October 10, 1972, eggs and first-instar Dipterous larvae were dis- covered in a redwood forest by one of us (WBH) at Prairie Creek Camp- ground, north of Orick, Humboldt County, California, on the surface of fermenting leaves littering the ground beneath a California Bay tree ( Umhellularia californica (H. & A.) Nutt.). The majority of eggs and larvae were acalyptrate but 1 predatory larva and an egg of a species of Muscidae were also noted. The leaves were stored on moist sand in 1 Permanent address : Department of Biological Sciences, University of Arizona, Tucson, Arizona 85721. The Pan-Pacific Entomologist 50: 78-82. January 1974 Vol. 50, No. 1, January 1974 79 jars at 68° F at U. C. Davis, and adults were first noticed on 24 Jan- uary, 1973. They were identified as Minettia flaveola (Coquillet) (Laux- aniidae) . Many puparia and puparial cases were found within the leaves, but only 3 puparia were washed from the sand. On 28 December, 1972, another collection was made at the same tree and large third-instar larvae were observed mining within the leaves. On 21 June, 1973, no larvae were found and the leaves were mostly dry. Three other collection localities produced fermenting Bay leaves that contained larvae: (1) along the Carmel River at Carmel Valley, Mon- terey County, 25 November, 1972; (2) McDonald Ranch at Pope Val- ley, Napa County, 22 January and 5 February, 1973; and (3) 5 miles west of Lake Berryessa on State Route 128, Napa County, 1 February, 1973. In addition to M. flaveola, the Napa County samples produced another lauxaniid which proved to be a new species of Homoneura. Predaceous larvae were observed in all of the collecting sites, and one was seen to attack and consume a lauxaniid larva in the latter McDonald Ranch sample. The adult of this larva was reared and identified as Phaonia parviceps Malloch (Muscidae). The larva of P. keilini, has been reported to prey on mosquito larvae (Oldroyd 1964). The Prairie Creek collection in October was assayed for yeasts. Sam- ples were taken from the surface of 3 leaves (2 of which had tiny larvae on the surface) and innoculated on acid-malt- agar plates in the field by direct plating and by previous dilution in 2 ml. sterile water (Table I). Five of the species were represented by less than 10 colonies each in all the plates. Rhodotorula minuta var. minuta was abundant (1000 col- onies) on 1 leaf and Torulopsis fujisanensis was common on 2 leaves (100 colonies each). Most of the species are considered to be wide- spread and all are nonfermentative. T . fujisanensis, however, has not been reported from the Pacific Northwest (Phaff et al. 1972). In ad- dition to the yeasts, bacteria and mold colonies were abundant on the original plates. The collection from Carmel Valley was assayed for yeasts from the somewhat fluid interior of the fermenting leaves. Five leaves, 4 with a large larva in each, were sampled by direct plating and by the addition of sterile water in the leaf. All slants were made on acid-malt- agar plates. Only 28 yeast colonies appeared among many mold colonies in 9 plates. Twenty-four of the colonies originated from the leaf with no larvae (Table 1). Two leaves produced 2 colonies each and the two remaining leaves produced no yeast at all. The significance of these observations and tests lies in the similarities 80 The Pan-Pacific Entomologist Table 1. Yeasts isolated from fermenting leaves of the California Bay Tree. Species Total no. colonies California localities Yeast niche Cryptococcus albidus (Saito) Skinner var. diffluens (Zach) Phaff et Fell 4 Prairie Creek Surface Cryptococcus laurentii var. laurentii (Kufferath) Skinner 1 (3 leaves) If II Leucosporidium scottii Fell, Statzell, Hunter et Phaff 5 If II Rhodotorula minuta var. minuta (Saito) Harrison 1000 1 fl If Torulopsis fujisanensis Soneda 230 2 1 1 II Candida moscorum di Menna 9 If II Candida buffoni (Ramirez) Van Uden et Buckley 2 II II Pink colonies (lost) 43 s II 11 Cryptococcus albidus var. diffluens 1 Carmel Valley Interior Cryptococcus infirms-miniatus (Okumuki) Phaff et Fell 26 4 (5 leaves) If 11 Debaryomyces hansenii (Zopf) Lodder et Kreger-van Rij 1 If II 1 1 leaf. 2 2 leaves. 3 Probable identification : Cryptococcus infirms-miniatus. i 24 colonies from one leaf, no larvae; 2 colonies from feces of single larva in a second leaf. and differences they exhibit with the identical larval leaf niche in the Hawaiian Archipelago which is occupied by a large number of endemic species in the genus Drosophila (Drosophilidae) (Heed, 1968; Kircher, 1969; Kircher and Heed, 1970; Carson et al., 1970). There are 4 major similarities between the 2 regions. (1) Both the coastal region in north- ern California and the windward slopes in the Hawaiian Islands are belts of equable temperatures and high humidity for a major portion of the year. This is important for the constancy of the resource (ever- green foliage) and the slow soft-rot decay of leaves by mold and bac- teria. (2) The fresh leaf of the Bay tree has a familiar aromatic fra- grance when crushed. When the leaf is fermenting on the ground its scent is prevalent in a somewhat modified form. All of the leaves utilized by Drosophila larvae in Hawaii such as Cheirodendron, T etraplasandra , Vol. 50, No. 1, January 1974 81 Ilex, Clermontia, Pelea, Myoporum , Myrsine and others have their own characteristic aroma produced by the breakdown products and their essential oils. These distinctive scents are believed to play a major role in the very high host plant specificity observed in the Hawaiian leaf breeders and probably are also important to the lauxaniids discussed here. Mr. Ray Miller (personal communication) relates that 7 species in 5 genera of the Lauxaniidae have been reported in rotting leaves. Dr. B. A. Foote, Kent State Univ., and R. Miller have reared species of the genera H omoneura, Lyciella and Minettia from decaying leaves of sugar maple, sassafras, alder, elm, and wild black cherry in deciduous forests in Ohio (personal communication). (3) The nutrition of the Hawaiian leaf-breeding larvae is known to be based chiefly on bacteria and broken down plant parts for the reason that no yeast of any conse- quence has been isolated from the interior of the leaves (Robertson et al. 1968). Also the leaf-breeding species are notoriously difficult to raise on laboratory media containing yeasts. However, a variety of yeasts have been isolated from the rotting leaf exterior in Hawaii (H. J. Phaff and M. W. Miller, personal communication). The observations in the present report are similar to the extent that the interior of the rotting leaves significantly lack the variety and density of the yeasts found on the leaf surfaces. Hering (1951) considers that the lauxaniids which are found in rotting leaves are modified leaf miners that had a sa- prophagous origin. (4) Both fly-substrate systems in California and Hawaii support larval predators in the family Muscidae. In Hawaii the genus is Lispocephala (Heed 1968) . One of the major differences between the lauxaniids and the drosoph- ilids is that the former oviposit on the leaf surface and the first-instar larva presumably develops there for a while before entering the leaf. Up to this time, the major nutritional source could come from yeasts. The Hawaiian Drosophila insert the eggs into the mesophyll of the leaf with only the filaments emerging (Kambysellis and Heed 1971). Another difference is that pupation takes place within the leaf for Mi- nettia and Homoneura but in Drosophila it occurs several inches below the soil surface. Finally, Drosophila apparently do not use the leaf niche in North America (Carson 1971). Presumably the absence of native lauxaniids in Hawaii (Hardy personal communication) may have been a contributing factor for the occupation of this niche by Drosophila. Acknowledgments We wish to express our appreciation to the following investigators for the identification of specimens in the Lauxaniidae and Muscidae: 82 The Pan-Pacific Entomologist Marius Wasbauer, California Department of Agriculture; Ray Miller, Iowa State University; R. H. Foote and R. J. Gagne, U. S. Department of Agriculture. Professor Herman J. Phaff verified the identification of the yeasts. The second author wishes to express his appreciation to Drs. Herman J. Phaff, Martin W. Miller, Herman T. Spieth and Th. Dobzhansky for the use of their laboratories during his sabbatical leave at U. C. Davis. This study is supported in part by NSF Grant GB-28953X1 to W. B. H. and H. W. Kircher and in part by NIH Trainee Grant STOIGM00701-13 to J. F. M. Literature Cited Bush, G. L. 1969. Sympatric host race formation and speciation in frugivorous flies of the genus Rhagoletis (Diptera, Tephritidae) . Evolution, 23: 237- 251. Carson, H. L. 1971. The ecology of Drosophila breeding sites. Univ. Hawaii Harold L. Lyon Arboretum Lecture, 2: 1-28. Carson, H. L., D. E. Hardy, H. T. Spieth, and W. S. Stone. 1970. The evolu- tionary biology of the Hawaiian Drosophilidae. In Essays in Evolution and Genetics in Honor of Theodosius Dobzhansky, M. K. Hecht, W. C. Steere, eds. Appleton-Century-Crofts, New York, pp. 437-543. Heed, W. B. 1968. Ecology of the Hawaiian Drosophilidae. In Studies in Genetics IV. M. R. Wheeler, ed. Univ. Texas Publ., 6818, pp. 387-420. Heed, W. B. 1971. Host plant specificity and speciation in Hawaiian Drosophila. Taxon, 20: 115-121. Hering, M. E. 1951. Biology of the Leaf Miners. N. V. Drukkerij Hooiberg, EPE, Netherlands. 420 pp. Kambysellis, M. P., and W. B. Heed. 1971. Studies of oogenesis in natural populations of Drosophilidae. I. Relation of ovarian development and ecological habitats of the Hawaiian species. Am. Natur., 105: 31-49. Kircher, H. W. 1969. Sterols in the leaves of the Cheirodendron guadichaudii tree and their relationship to Hawaiian Drosophila ecology, J. Insect. Physiol., 13: 1869-1874. Kircher, H. W., and W. B. Heed. 1970. Phytochemistry and host plant speci- ficity in Drosophila. In Recent Advances in Phytochemistry III, C. Steelink and V. C. Runeckles, eds. Appleton-Century-Crofts, New York, pp. 191-209. Oldroyd, H. 1964. The Natural History of Flies. W. W. Norton & Co., Inc., New York. 324 pp. Phaff, H. J., M. W. Miller, M. Yoneyama, and M. Soneda. 1972. A compara- tive study of the yeast florae associated with trees on the Japanese Is- lands and on the West Coast of North America. Proc. IV IFS: Ferment. Technol. Today: 759-774. Robertson, F. W., M. Shook, G. Takei, and M. Gaines. 1968. Observations on the biology and nutrition of Drosophila disticha Hardy, an indigenous Hawaiian species. In Studies in Genetics IV. M. R. Wheeler, ed. Univ. Texas Publ. 6818, pp. 279-299. Vol. 50, No. 1, January 1974 83 BOOK REVIEW The Bark and Ambrosia Beetles of California (Coleoptera: Scolytidae and Platypodidae) . D. E. Bright, Jr. and R. W. Stark. 1973 Bull. Calif. Insect Survey, 16: 1-169. University of California Press. $8.50. Curtis G. Benjamin, in an article in Science (1973, 183(4122) : 282-284) has lucidly examined the economic afflictions of scientific monographs. Since 1957, per page prices have risen from 2.50 to 6.10, an increase of 144 percent. Simul- taneously, average sales of such works declined from about 5000 to 3000 copies. The present treatise on bark and ambrosia beetles provides a case in point — for a paperback volume of 175 pages, $8.50 will be considered expensive by most people. More unfortunately, even at the currently inflated rates, publishers expect such short-run issues to be economic failures. (The average run for California Insect Survey Bulletins is about 575.) Economic matters aside, the Bark and Ambrosia Beetles of California is a valu- able compilation of biological and distributional information. An introductory review of taxonomic characters is helpful in working the keys. Summaries of the distribution, bionomics and the ecological and economic importance of bark beetles provide a general overview of the detailed information in the species treatments. A useful adjunct to the text and index is a list of host plants and the scolytids attacking them. Keys to the genera and species are well constructed, and non-specialists should be able to use them proficiently. In a minority of couplets overlapping size ranges are used as primary characters, but are almost always combined with more easily distinguished secondary characters. I had little difficulty in correctly keying five species from different genera. The bulk of the text consists of summaries of the geographic distribution and host ranges for each species, with discussions of the biology when that information is available. For the more important economic species these biological treatments may be rather extensive, marking an important digression from most earlier Survey Bulletins, which consisted largely of keys and distributional records. If the present format is continued, the Bulletin Series should enjoy more general interest than in the past. One rather disconcerting feature is the large number of distribution maps (93), which occupy nearly 25 percent of the text space devoted to species treatments. Since the majority of species feed on conifers whose ranges in California are sim- ilar, there is much repitition of distributional patterns. On several of the maps the symbols are very difficult to discern, and on a few others as few as four to seven collection records are indicated. Illustrations include excellent photographs of typical egg galleries of at least one species of most of the genera treated. Probably of less utility is the page of line drawings of antennal clubs and funicles which is referred to in the key. My impression was that the antennal characteristics were frequently clear without visual aids, while some of the other key characters were less well defined. On the whole these criticisms are minor compared to the general completeness of the work, which will certainly be valuable to those dealing with forest insects in California. At $8.50 it may even be a bargain if the relative price of scientific monographs continues to inflate at an exponential rate. — John T. Doyen, University of California, Berkeley. 84 The Pan-Pacific Entomologist SCIENTIFIC NOTES Martarega hondurensis and Buenoa antigone as predators of mosquito larvae in Costa Rica (Hemiptera: Notonectidae). — Buenoa antigone Kirkaldy inhabits freshwater pools in southern United States, Mexico, Central and South America (Truxal, 1953 Univ. Kansas Sci. Bull. 35: 1351-1523). Martarega hondurensis Bare has been recorded from Mexico, Central America and Peru (Truxal, 1949, J. Kansas Entomol. Soc. 22: 1-24) and is found aggregated into schools in deep sheltered eddies of rivers, as is M. mexicana Truxal (Menke and Truxal, 1966 Los Angeles Co. Mus. Contr. Sci. 106: 1-6). Under certain circum- stances both species prey on mosquito larvae, and one shows potential as a biological control agent. In July, 1972, I surveyed the fauna of two dozen small water-filled depressions (2m 2 or less) in rock outcroppings along the banks of the Corobici and Tenorio Rivers, 5 km from Canas, Guanacaste, Costa Rica. Other than notonectids, these pools seemed to lack potential mosquito predators. Six pools which contained Buenoa (predominately B. antigone ) lacked mosquito larvae, while the others containing no Buenoa had large aggregations of mosquito larvae. At each pool I recorded a number of habitat parameters (size of the pool, depth and character of the bottom sediments, water turbidity, temperature, pH, vegeta- tion, amount of shade, water depth and presence of other aquatic organisms) . The distribution of backswimmers and mosquito larvae was similar with respect to these parameters, the only difference between habitats being the presence of the backswimmers and mosquito larvae themselves. Whether the distribution of Buenoa in pools is a fortuitous event or is dependent upon some undetermined factor remains unclear, but it seems probable that the absence of mosquito larvae in these pools is related to the presence of Buenoa. In Santa Rosa National Park, Guanacaste, Costa Rica, Buenoa were especially abundant and were never found together with mosquito larvae in several large ponds that were surveyed. Other potential mosquito predators in these ponds may have reduced numbers of larvae, but the high density of Buenoa suggests that the backswimmers may have had a significant part in mosquito elimination. Laboratory experiments tended to confirm the predator-prey relationship sug- gested by field observations. B. antigone were collected and kept in jars (110 ml) for 24 hours at air temperature. Each jar contained two B. antigone as well as five mosquito late instars. In twelve repetitions (six late instars, six adults) all mosquito larvae offered were eaten. Martarega hondurensis from river sites ad- jacent to the rock outcroppings where B. antigone occurred were offered mosquito larvae in the same format as used for Buenoa. After 24 hours adult Martarega had consumed an average of 2.6 of 5 larvae offered in each of 33 repetitions, while late instars ate an average of 1.4 of 5 in each of 39 repetitions. Clearly, under these conditions, M. hondurensis ate mosquitoes at a slower rate than did B. antigone. Martarega seemed more inclined to cannibalism than to mosquito predation. In 3.5 liter glass aquaria containing from 15 to 20 M. hondurensis, 15 to 20 mosquito larvae were offered but not eaten, but earlier instar M. hondurensis were often cannibalized. Under similar conditions B. antigone rapidly ate mosquito larvae, with no cannibalism. Prey preference tests showed that B. antigone ate mosquito larvae and water boatmen before ostracods, blood worms, mosquito pupae and ants (P ^ .05) Vol. 50, No. 1, January 1974 85 (Gittelman, in preparation). Probably Baenoa are capable of subsisting on a variety of prey, allowing them to be effective predators on mosquito larvae while not totally depending on them for their distribution. When offered a similar selection of prey types, M. hondurensis ate ants struggling at the water surface before all other prey species including mosquito larvae (P^.01). B. antigone is readily maintained in the laboratory but breeding under laboratory conditions has not been attempted. It is likely that these insects lay their eggs on or in plant material. In either case, eggs could be collected by harvesting plants or some artificial oviposition substrate (Toth and Chew, 1972b). Since B. antigone does not cannibalize even in crowded laboratory enclosures, little difficulty would be expected in mass rearing. In contrast, M. hondurensis would necessitate separate containers, for each individual. The suggestion that backswimmers be used for mosquito control (Toth and Chew, 1972, Ann. Entomol. Soc. Amer. 65: 1270-79; 1972. Env. Entomol. 1: 534-5; Ellis and Borden, 1970, Ann. Entomol. Soc. Amer. 63: 963-73) seems well applied to B. antigone but not to M. hondurensis. Although both species will eat mosquito larvae, M. hondurensis do so at a slower rate than do B. antigone. In addition it would appear impractical to distribute M. hondurensis as a biological control agent because it tends to inhabit moving water where mosquito larvae are not prevalent. On the other hand, B. antigone seem to prefer mosquito larvae as food, and their microhabitats generally overlap.- — -Steven H. Gittelman, Division of Biological Sciences, U-42, University of Connecticut, Storrs 06268. Biological observations on darkling ground beetles from western North America (Coleoptera: Tentyriidae). — Coelosattus fortineri Blaisdell (1927) was originally described from the Algodones Sand Dunes of California; this genus was placed in synonymy under Eusattus (Doyen, 1972: Quaest. Ent., 8: 357-376). Coelomorpha maritima Casey (1890) was originally described from “lower Cali- fornia.” This genus was later placed in synonymy under Coelus (Doyen, 1972). Subsequent to their original descriptions, these beetles have remained largely un- known in collections, although both are locally abundant. Seventy-six individuals of Eusattus fortineri were collected from the Algodones Dunes, 2 mi. N. Glamis, Imperial County, California, during April, 1972. Four of these were encountered on the sand surface between 1900-2300 hrs. (PST). The remainder were excavated from the sand around the bases of creosote bush ( Larrea divaricata Cavanilles) , at depths of 6-20 cm. The distribution of the beetles was highly aggregated, with 28 individuals buried beneath a single plant. A typical assemblage consisted of six to eight beetles, and only three groups of less than three individuals were discovered. Beetles were found only beneath plants with canopies close to the sand. Suitability of plants as shelters depends on the degree of shading of the sand about their bases, since exposed substrates in the Algodones Dunes reach midday temperatures well above 40°C during warm months. E. fortineri appears to be abundant within its restricted geographic range, which coincides with the distribution of exceptionally fine sand composing the Algodones Dunes and adjacent dunes in southwestern Arizona and northwestern Mexico. Its allopatric counterpart in other areas of western North America is the morphologically similar E. muricatus LeConte, which occurs on a variety of 86 The Pan-Pacific Entomologist sandy substrates, including dunes, over a broad geographic area from eastern Washington to Texas and northern Mexico. It has been recorded from numerous localities within 50 mi. of the Algodones Dunes, but never in actual sympatry with E. fortineri. Coelus maritima has been recorded from scattered coastal localities in northern and central Baja California, but fewer than 30 specimens were known prior to 1972. During late March, 1972 and 1973, and mid-June, 1973, approximately 500 adults of C. maritima were collected by screening sand at the maritime dunes at Santa Maria Beach, San Quintin Bay, Baja California Norte. In March both adults and larvae were extremely abundant to depths of 3 cm in the sand canopied by perennial plants, especially Abronia maritima Nuttal ex Watson, where they were associated with small numbers of other tenebrionids Cryptadius inflatus LeConte, Eusattus laevis LeConte, E. erosus Horn, and Micromes ovipennis (Horn). In June only adults were encountered, many teneral, suggesting that adult emergence peaks in late spring or early summer. The local abundance of C. maritima is probably conditioned by the characteristics of the sand in which it lives. The beach sand at San Quintin Bay is extremely fine and friable, allowing entry by small organisms such as Coelus. South of San Quintin, the dunes extend inland a distance of 3-4 miles, but the sand becomes progressively coarser and more consolidated inland. Collections away from the immediate coast contained no Coelus, but the larger species, E. laevis was present, and Micromes was abundant on the sand surface beneath plants. In extreme northern Baja California, C. maritima is replaced by C. globosus LeConte, which occurs in the same habitat. In extensive collections from dunes near Cantamar, 25 mi. S. Tijuana, C. globosus was very abundant, while C. maritima was absent, and the two species do not appear to occur sympatrically. John T. Doyen, Division of Entomology and Parasitology, University of California, Berkeley 94720. Differential predation of darkling ground beetles (Coleoptera: Tene- brionidae) by the Channel Islands Fox. — During April, 1972, 73 scats of the Channel Islands Fox ( Urocygon littoralis) were collected from the sand dunes near Wilson’s Cove on the northwestern end of San Clemente Island, Los Angeles County, California. The diet of the fox is known to be extremely varied, including berries, acorns, and carrion, as well as living invertebrates and small vertebrates (von Blocker, 1967, in Philbrick, Proceedings of the Symposium on the Biology of the California Islands, pp. 245-253) . All of these items were represented by the fragments recovered from the San Clemente scats, but insect remains were partic- ularly abundant. What is of interest is the relative numbers of some of the beetle prey (Table 1). By far the most abundant were corpses of Trigonoscuta sp. (Cur- culionidae), June beetles (Scarabaeidae) , and the tenebrionids Coelus remotus Fall and Eusattus robustus LeConte. All these species are relatively abundant on or about the sand dunes, but two of them ( Trigonoscuta and Coelus) are relatively small bodied. Large numbers would be required to sustain an animal as large as a fox. The large tenebrionid Eleodes laticollis LeConte is also abundant at Wil- son’s Cove, but was rare in the scats. Three other small tenebrionids, Apsena grossa LeConte, Coniontis latus LeConte, and Helops bachei LeConte were also uncommon in the scats, though relatively abundant on or about the dunes. With Vol. 50, No. 1 , January 1974 87 Table 1. Beetle contents of 73 scats of the Channel Islands fox. The left column of figures lists the numbers of scats containing each species. The right column lists total numbers found in all scats combined. Numbers of tenebrionids and of Trigo- noscuta were estimated by counting prothoraces, which usually remained intact. Numbers of Scarabaeidae were estimated by counting elytra. Scats Total number of individuals Tenebrionidae Coelus remotus Fall 23 196 Coniontis latus LeConte 1 1 Eusattus robustus LeConte 9 15 Apsena grossa LeConte 2 2 Eleodes laticollis LeConte 2 2 Helops bachei LeConte 3 3 Scarabaeidae ( Phobetus and Serica ) 17 49 Curculionidae ( Trigonoscuta ) 45 270 the exception of Coniontis, which was the least numerous of the beetles in the habitat, the tenebrionid species which were infrequently eaten possess quinonoid secretions which they release at the abdominal tip when disturbed. Eleodes lati- collis is particularly well supplied with secretions, which are stored in large res- ervoirs in the abdomen and forcibly ejected as a fine spray which may travel over 30 cm. Coelus, Coniontis, and Eusattus, as well as the weevil and scarabs, lack secretions. Although quinonoid secretions of tenebrionids are popularly referred to as defensive secretions (Eisner and Blumberg, 1959, An. Rec., 134: 558-559), little evidence of differential predation is available. The data presented here sug- gest that species possessing secretions are ignored, at least when other food is present. Quantitative estimates of the population densities of the various species are unavailable, and nothing is known of the searching strategy of the fox. How- ever, the differences in the numbers of individuals taken, especially between Eleodes and Coelus, are so marked that selective predation to avoid the beetles with de- fensive secretions seems likely. — John T. Doyen, Division of Entomology and Para- sitology, University of California, Berkeley 94720. A note on the Nesting Biology of Dianthidium pudicum. pudicum (Cresson) (Hymenoptera : Megachilidae).- — A nest of Dianthidium pudicum pudicum (Cresson) was discovered on 26 March, 1967, 9.3 miles south of Quartsite, Arizona. The nest contained 10 cells located about one meter off the ground in a forked branch of a creosote bush, Larrea tridentata Sesse & Mocino. The branch was in a horizontal position and about 6 mm in diameter at the nest location. The nest assumed the triangular shape of the branches (Fig. 1), with the sides 27-28 mm long. The dorsal surface extended 6 mm above the horizontal branches and the bottom extended 1-2 mm below the branches, averaging about 13 mm thick. A dark brown resin, with a strong scent of Larrea, cemented numerous small pebbles together to form the outer surface of the nest. Internally, the resin became 88 The Pan-Pacific Entomologist Fig. 1 . A nest of Dianthidium pudicum pudicum. softer and yellowish-green in color. The small pebbles were tightly embedded throughout the interior of the nest, with occasional pieces of masticated leaf material. In constructing the 10 vertically fused cells, the female bee appeared to have built the first cell in the base of the forked branches and then added rows of two, three, and four cells as she increased the triangular shape of the nest. The cells measured 5-5.5 mm in width and 9-10 mm in length. The cell walls averaged 0.5-0.75 mm in thickness. Cell surfaces adjacent to the branches had only a thin layer of resin, whereas the other surfaces contained numerous small pebbles which extended through the inner surface of resin into the cell cavity. Cell caps of the downward opening cells averaged 0.5-0.75 mm in thickness. The caps of 6 cells consisted of a circle of small pebbles with a resin filled center, while the other 4 cell caps did not show this construction pattern. The dark brown cocoons were thin walled and transparent and the bees were visible through the cocoons. Anteriorly, the cocoons were thicker and heavily interwoven with white, silk threads (0.03-0.05 mm thick) which gave rise to the outer median mammillary projection. The white mammillary projections measured 0.75 mm in length and 0.75-1.0 mm wide at the base. Cocoons ranged from 8-8.5 mm in length from the tip of the projection to the base and they were about 4 mm wide. An air space approximately 1 mm in length separated the tip of the projection and the inner surface of each cell cap. Individual orange fecal pellets were about 0.75 mm in length and 0.25 mm in width, without surface grooves or ridges. The cylindrical pellets were smeared against the inner surface of the cell walls, forming a continuous reddish-brown layer 0.25 mm thick around all but the anterior quarter of the cell. This fecal layer smooths the rough inner surface of the cell. When the nest was opened in early April, the 3 basal cells and one of the third row cells contained post-defecating larvae and the remaining cells held 4 Vol. 50, No. 1, January 1974 89 male and 2 female pupae. Three of the pupae in the outer row of cells had their heads pointed downward, whereas the other 7 bees were oriented with their heads up. Adults emerged 10-15 days later. There are several differences between the nests observed by Hicks (1927, Psyche 34: 193-198) and the nest we have described. All nests described by Hicks were attached to rocks found on the surface of the ground. The cells were larger, measuring 15 mm in length, and the cocoons were larger than those we observed. All of the cells examined by Hicks contained bees oriented downward and we observed only 3 of 10 bees oriented downward. Grigarick and Stange (1968, Bull. Calif. Insect Surv. 9: 1-113) briefly mentioned the discovery by F. D. Parker in Reno, Nevada of a nest of D. p. puclicum in the crotch of a small fruit tree. Dr. A. A. Grigarick kindly identified the Dianthidium for us. — S. L. Clement, University of California, Davis 95616 and R. W. Rust, University of Delaware, Newark 19711. 90 The Pan-Pacific Entomologist PACIFIC COAST ENTOMOLOGICAL SOCIETY J. A. Chemsak C. B. Philip M. R. Gardner P. H. Arnaud, Jr. President President-Elect Secretary Treasurer Proceedings Three Hundred and Fiftieth Meeting The 350th meeting was held Friday, February 23, 1973 in the Morrison Audi- torium of the California Academy of Sciences, Golden Gate Park, San Francisco, with President Chemsak presiding. Members present (34) : P. H. Arnaud, Jr., D. Bacon, R. G. Blair, T. Briggs, J. G. Brady, R. M. Brown, D. Burdick, J. A. Chemsak, J. G. Edwards, F. Ennik, W. E. Ferguson, L. Floyd, N. E. Gary, A. J. Gilbert, A. & A. Gillogly, B. W. Grushkowitz, K. S. Hagen, A. Hardy, L. S. Hawkins, T. E. Hewton, H. B. Leech, D. W. Moss, R. P. Papp, C. B. Philip, B. Potts, J. Powell, C. Radus, C. Rees, E. S. Ross, S. L. Szerlip, R. W. Thorp, T. Toren, M. S. Wasbauer. Visitors present (29) : L. Bezark, N. Blair, A. Bush, W. D. Carstens, M. R. Ebertz, S. Ferguson, K. Gary, M. Gilbert, D. Grushkowitz, W. Hardy, S. Jordan, A. I. B. Kaplan, G. J. Keutz, C. Kitayama, B. Langston, C. Langston, R. L. Morr- ison, G. M. Nishida, K. Nishida, N. E. Papp, W. Rawscha, R. G. Repke, M. Robey, E. Schmidt, J. Thorp, R. T. Wian, G. L. Willey, P. Willey, A. Yuaz. The minutes of the meeting held 15 December were summarized. The following names were proposed for membership in the Society. Student Membership: Clifford L. Radus, James P. Anduiza, Robert W. Pemberton, War- ren E. Savary, Laurence A. Lacey, Steven K. Ault, Larry G. Bezart, Gary L. Willey, Gordon M. Nishida, Regular Membership: Glenn M. Yoshimura, Regina M. Burke, Charles D. Hunter, Fred Punzo, Kilian Roever, Ernest Anderson and Dr. Carl 0. Mohr. The following note was presented: Polistiopsis mima Townsend (Diptera: Tachinidae) in Costa Rica. — Two species are presently assigned to the genus Polistiopsis Townsend — the type species mima Townsend 1915 (recorded from Mexico — the holotype male from Tehuantepec and a second male from Fortin de las Flores) and williamsi Arnaud 1966 (recorded from Paragray — the holotype male from Paso Yobay and the allotype female from Villarica) . Townsend considered mima to be a polistid counterfeit. When I re- vised the genus in 1966 (American Museum Novitates, No. 2241, pp. 1-12, figs. 1-20) only the four specimens cited above were available for study from the major North American entmological collections. This past summer while studying in Paris at the Museum National d’Histoire Naturelle (supported by Grant No. 6351, Penrose Fund, American Philosophical Society), it was a pleasure to discover a series of 15 specimens of Polistiopsis among the miscellaneous unstudied Tachinidae. I would like to thank Monsieur Loic Matile for their loan. A recent examination of these has shown that they represent 12 males and 3 females (the latter the first known of P. mima Townsend. This series of specimens is labeled with the data “Costa Rica/Paul Serre 1920”; unfortunately without more precise locality data. Two of the males have the ad- ditional datum that they were collected during the month of “Juillet”. Several Vol. 50, No. 1, January 1974 91 specimens of this series are displayed this evening. — Paul H. Arnaud, Jr., Cali- fornia Academy of Sciences, San Francisco. The principal speaker of the evening was Dr. Norman E. Gary, Professor of Entomology, University of California, Davis. His illustrated talk was entitled: “Magnetic Retrieval of Honeybee Labels.” Coffee and other refreshments were served during a social hour held in the entomology rooms following the meeting. — M. R. Gardner, Secretary. Three Hundred and Fifty-First Meeting The 351st meeting was held Friday, 30 March 1973 in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San Francisco, with President Chemsak presiding. Members present (36) : F. G. Andrews, C. Armin, P. H. Arnaud, Jr., J. Anduiza, F. L. Blanc, R. G. Blair, T. Briggs, R. M. Brown, D. Burdick, K. S. Corwin, D. Denning, J. Denk, J. G. Edwards, F. Ennik, M. R. Gardner, J. Guggole, R. Hall, L. S. Hawkins, T. E. Hewton, A. I. Kaplan, C. Kitayama, R. L. Langston, H. B. Leech, R. W. Pemberton, C. L. Radus, F. C. Roberts, E. S. Ross, W. Savary, M. V. Silveira, R. Stecker, T. J. Toren, R. L. Tassan, R. W. Thorp, P. Welles, G. L. Willis, D. W. Whitman. Visitors present (12) : S. Anduiza, T. Andrews, J. R. Batch, F. Blanc, A. Bush, R. Greek, R. Morrison, P. Neuenschwander, P. Peterson, T. Slay, J. Thorp, P. Willey. The minutes of the meeting held 23 March were summarized. The following note was presented: Insects and a mite associated with stored Cannabis sativa Linnaeus. — On January 23, 1970 I visited the Hall of Justice in San Francisco, California, at the request of Mr. Terry L. Coddington and Mr. John F. Willims to inspect an insect problem associated with confiscated stocks of Cannabis sativa Linnaeus. In offices adjoining room vaults in which these stocks were kept, the personnel were pestered by insects in their offices. One problem area was a trunk, thought to be from the midwestern United States, in which a plastic liner kept the plant materials moist. From this source a series of 145 specimens of Desmometopa sp. (Diptera: Milichiidae, det. C. W. Sabrosky), 11 Scatopse fuscipes Meigen (Diptera: Scatopsidae, det. Alan Stone), 20 Bradysia sp. (Diptera: Sciaridae, det. R. J. Gagne), and 1 Drosophila busckii Coquillett (Diptera: Drosophilidae, det. W. W. Wirth) were collected. These flies are pri- marily scavengers. On and in packages of the marijuana, 6 adults, 1 larva, and 1 pupal skin of Plodia interpunctella (Huhner) (Lepidoptera: Pyralidae, det. D. M. Weisman) , and specimens of six species of Coleoptera (det. F. G. Andrews) — Ahasverus advena (Waltl), the foreign grain beetle, Cryptolestes ferrugineus (Stephens), the rusty grain beetle, Cryptolestes pusillus (Schonherr), the flat grain beetle, and Oryzae- philus surinamensis (Linnaeus), the sawtoothed grain beetle of the family Cucuji- dae, Microgramme arga (Reitter) of the family Lathridiidae, and Typhaea ster- corea (Linnaeus), the hairy fungus beetle of the family Mycetophagidae were collected. This group is associated with many types of stored products. A mite Macrocheles muscadomesticae (Scopoli) (family Macrochelilae, det. R. L. Smiley) was also collected. I would like to acknowledge my thanks to the 92 The Pan-Pacific Entomologist specialists noted above for the identification of the species mentioned.— P aul H. Arnaud, Jr., California Academy of Sciences, San Francisco. The principal speaker of the evening was Dr. Werner Loher, Professor of En- tomology, University of California, Berkeley. His illustrated talk was entitled: “Acoustical communication in insects.” Coffee and other refreshments were served at a social hour in the entomology rooms following the meeting.- — M. R. Gardner, Secretary. Three Hundred and Fifty-Second Meeting The 352nd meeting was the annual field day and picnic. It was held on Sat- urday, 12 May, 1973 at Castle Rock Park near Mount Diablo, Contra Costa County, California. Members present (9) : P. H. Arnaud, Jr, R. Blair, H. V. Daly, Jr., H. V. Davis, W. E. Ferguson, M. R. Gardner, R. W. Pemberton, C. B. Philip, H. Reinhard. Vistors present (15+) : M. Arnaud, P.-H. F. Arnaud, N. Blair, K. Blair, R. Blair. Mrs. H. V. Daly, Jr., S. Ferguson, N. Milliet, C. Pemberton, W. J. Reinhard, En- tomology class of Dr. H. V. Daly, Jr. The day was warm and sunny. Picnicking and insect collecting were the major activities of the participants. Judging from the full cyanide vials, flying insects were found in abundance by the enthusiastic students. — M. R. Gardner, Secretary. Three Hundred and Fifty-Third Meeting The 353rd meeting was held Friday, 19 October 1973, in the Morrison Auditorium of the California Academy of Sciences, Golden Gate Park, San Francisco, with President Chemsak presiding. Members present (44) : R. P. Allen, F. G. Andrews, C. Armin, P. H. Arnaud, Jr., L. Bezark, R. Blair, T. Briggs, D. Burdick, J. A. Chemsak, C. R. Dutton, J. G. Edwards, B. Ehreth, F. Ennik, B. K. Eya, M. R. Gardner, A. J. Gilbert, A. R. Gillogly, B. Grushkowitz, K. Hagen, R. Hall, T. E. Hewton, D. Jamieson, C. Kita- yama, R. L. Langston, H. B. Leech, K. L. Lipps, J. McCarthy, R. Morrison, D. Moss, G. Nishida, R. P Papp, R. W. Pemberton, C. B. Philip, W. W. Pilke, H. V. Rein- hard, W. E. Savary, M. V. Silveira, S. L. Szerlip, R. W. Thorp, T. J. Toren, B. Villegas, D. Whitman, G. Willey, R. L. Wong. Visitors present (41) : A. Allen, J. Baier, G. M. Bass, Mrs. Blair, P. S. Crane, W. Craven, W. Diest, A. Edwards, S. Ferguson, T. Gabel, J. Gelhaus, R. Genberg, M. Gershery, M. Gilbert, D. Green, D. Gropman, K. S. Horn, H. W. Horn, J. S. Hjelle, H. Implom, S. Jordan, H. Kent, B. Kidwell, G. Krolmer, V. Mayer, D. Meyers, B. Michalski, K. Nishida, N. Papp, P. Pitcher, M. Roberts, G. Rolfec, J. Schager, T. Schager, J. T. Sorenson, R. Stockhill, J. Thorp, B. Tilden, J. Wells, P. Willy, E. Yasukawa. The minutes of the meetings held 30 March and 12 May were summarized. The following persons were elected to membership in the Society. Student mem- bership: Thomas Lugaskii, Thomas M. Maxwell, Mark G. Torassa, Philip D. Perkins, Bruce Tilden. Regular membership: Karen Lipps, Fred Barnes, Richard Rust, Paul Peterson, Philip Hubert. The principal speaker of the evening was Dr. Edward S. Ross. His illustrated talk was entitled: “Memorable insect encounters.” Vol. 50, No. 1, January 1974 93 Coffee and other refreshments were served during a social hour in the entomology rooms following the meeting. — M. R. Gardner, Secretary. Three Hundred and Fifty-Fourth Meeting The 354th meeting was held Friday, 16 November 1973, in the Morrison Audi- torium of the California Academy of Sciences, Golden Gate Park, San Francisco, with President Chemsak presiding. Members present (40) : P. H. Arnaud, Jr., J. W. Bass, F. R. Barnes, L. H. Bezark, R. G. Blair, R. M. Brown, J. A. Chemsak, T. Delgman, J. T. Doyen, J. G. Edwards, F. Ennik, B. Eya, W. E. Ferguson, J. R. Gabel, M. R. Gardner, J. Gug- golz, B. W. Grushkowitz, T. E. Hewton, C. Kitayama, R. L. Langston, R. Main, R. Morrison, D. Moss, G. M. Nishida, R. W. Pemberton, C. B. Philip, W. W. Pitcher, R. W. L. Potts, J. A. Powell, F. Radus, M. J. Robey, E. S. Ross, R. Schick, M. V. Silveira, E. L. Smith, C. Tauber, R. W. Thorp, B. A. Tilden, G. Willey, M. Wasbauer. Visitors present (12) : N. Blair, A. Bush, L. P. Brower, D. Grushkowitz, M. H. Lobisser, C. Moffitt, G. Roefes, D. Sherman, J. Thorp, M. Tauber, P. Willey, A. Wood. The minutes of the meeting held 19 October were summarized. Dr. Arnaud mentioned that over 100 persons were in attendance at the meeting of 19 October, although not everyone may have signed the register. President Chemsak announced the nominees for the auditing and nominating committees. For auditing committee he appointed Mr. H. Vannoy Davis Chairman, and Dr. Edward S. Ross committeeman. For nominating committee he appointed Dr. Marius S. Wasbauer chairman, and Drs. Robin W. Thorp and E. Gorton Lins- ley committeemen. The nominations were unanimously approved by the mem- bership. The following note was presented: Paradejeania rutilioides nigrescens Arnaud reared from Hemihyalea sp. (Diptera: Tachinidae; Lepidoptera: Arctiidae ). — Paradejeania rutilioides nigrescens has not had a recorded host even though it is the largest species of Cal- ifornia Tachinidae and is often collected at flowers in the late summer and fall. Essig in his book (1915. Injurious and beneficial insects of California, second edition, p. 330) stated that “Caterpillars of various species are the host,” hut no specific names are given. It has been suggested that the host would be Heteroceran (Arnaud, 1968. Pan-Pac. Entomol., 44: 85). It is now possible to record the first rearing of Paradejeania rutilioides nigres- cens from the arctiid genus Hemihyalea based on a rearing made by the late Hans Peter Allmendinger and R. Maddux. Larvae of a species of Hemihyalea were col- lected by them at Los Gatos, Santa Clara County, California on 17 June 1972 and the arctiids subsequently pupated. An adult female of nigrescens emerged from one of the Hemihyalea pupae on 1 September 1972. Mr. Allmendinger had com- mented that it was not possible to locate the exact pupa from which the nigrescens was reared since this was a mass rearing. — Paul H. Arnaud, Jr., California Acad- emy of Sciences, San Francisco. The principal speaker of the evening was Dr. Lincoln P. Brower, visiting pro- fessor of environmental studies, University of California, Davis. His illustrated talk was entitled: “East versus West Coast Monarchs — a Question of Taste.” 94 The Pan-Pacific Entomologist A social hour was held in the entomology rooms following the meeting. — M. R. Gardner, Secretary. Three Hundred and Fifty-Fifth Meeting The 355th meeting was held Friday, 14 December 1973 in the Morrison Audi- torium of the California Academy of Sciences, Golden Gate Park, San Francisco, with President Chemsak presiding. Members present (45) : J. Anduiza, F. G. Andrews, P. A. Arnaud, Jr., F. R. Barnes, J. W. Bass, L. Bezark, R. G. Blair, J. W. Brady, R. M. Brown, J. A. Chemsak, T. Delgman, D. G. Denning, J. T. Doyen, D. Durbin, J. G. Edwards, B. Ehreth, F. Ennik, W. E. Ferguson, J. R. Gabel, A. J. Gilbert, P. A. Golen, B. W. Grushkowitz, R. Hall, S. E. Heston, C. Kitayama, R. L. Langston, K. L. Lipps, J. McCarthy, A. E. Michelbacher, R. W. Pemberton, P. A. Peterson, C. B. Philip, W. W. Pitcher, J. A. Powell, H. G. Real, H. I. Scudder, M. V. Silveira, E. L. Smith, R. E. Somerby, R. E. Stecker, S. L. Szerlip, R. W. Thorp, B. Tilden, W. H. Tyson, G. L. "Willey. Visitors present (23) : F. Baker, M. Brady, T. Brum, M. A. Chemsak, J. Da- monte, L. Denning, L. Doyen, T. D. Eichlin, D. Eschen, S. Ferguson, T. Gabel, M. Gilbert, K. Ghandi, D. Green, D. Grushkowitz, K. S. Heston, P. Heston, M. Michelbacher, N. Mamas, M. A. Santana, P. Tyson, J. Thorp, P. Willey. The minutes of the meeting held 16 November were summarized. The following people were elected to membership in the Society. Regular mem- bership: Rev. Andre Larochelle and Dr. Howard E. Evans. The following note was presented: Collection of adult Deuterophlebia coloradensis Pennak at Boulder Falls, Colorado (Diptera: Deuterophlebiidae) . — The Deuterophlebiidae or mountain midges are a rarely collected family of flies, with seven named species, and several unnamed taxa, all assigned to the genus Deuterophlebia Edwards. They are known only from the northern hemisphere, with one species each from Kashmir ( D . mirahilis Edwards, 1922), Japan ( D . nipponica Kitagami, 1938), Korea ( D . tyo- senensis Kitagami, 1938) , and four species from the western United States ( D . coloradensis Pennak, 1945; D. inyoensis Kennedy, 1960; D. nielsoni Kennedy, 1958; and D. shasta Wirth, 1951) . Probably unnamed taxa have been reported from the Altai Mountains in Siberia; tributaries of the Alsea and Marys Rivers, Benton County, Oregon; Yellowstone National Park; and finally several localities in the Sierra Nevada of California (Kennedy, 1958, Trans. Amer. Miscro. Soc., 77: 201- 228). Adult Deuterophlebia are rare in collections, with most of the earlier descriptive work being based on the larval and pupal stages, with adults obtained from the dissection of pupae. Kitakami (1938. Annot. Zool. Jap., 17: 501) collected fe- males of D. nipponica from spider webs at Kurama and Mt. Hira and reported a swarm of females of this species in the early morning at Simasima on the island of Honshu. Pennak discovered on July 19, 1948 (1950. Entomol. News, 61: 36) at 8:00 A.M. 200 dying, dead and disintegrating adult males and two adult fe- males of D. coloradensis floating in small side eddies of North St. Vrain Creek, near the town of Lyons, Colorado. He suggested that the imagoes emerge in the early morning hours and flutter about feebly and briefly above the water, but soon falling to the surface of the water and dying. Kennedy (1958) using rearing cages Vol. 50, No. 1, January 1974 95 placed in Convict Creek, California over pupae of D. nielsoni, observed that ima- gines emerged between 7 to 9 A.M. (P.S.T.), and that in an hour’s flight copula- tion and deposition of the eggs had taken place. Between 8:00 A.M. and 9:00 A.M. the imagines began to die. On the field trip day for the International Congress of Systematic and Evolu- tionary Biology held on August 8, 1973, my first collecting stop was at Boulder Falls, on north Boulder Creek, about 11 km west of Boulder, Colorado, at an ele- vation of 2135 meters. On the western bank of the river, within 30 or 40 meters of the base of the falls, insects were noted in flight and by sweeping an insect net from water level to a meter and a half above the surface from the edge of the river, D enter ophlebia were collected. The area was shaded and the rays of the morning sun had not penetrated the area. Collecting was probably under way at 9:00 A.M. or shortly thereafter, but the time was unfortunately not recorded. A total of 72 males presumably of D. coloradensis were collected. The type locality for D. coloradensis (the holotype being a mature larva) is Boulder Creek, at an elevation of 1810 meters, so that it would be a few kilometers below this Boulder Falls col- lection. The series of 20 point mounted and 52 alcohol preserved specimens are deposited in the collection of the Department of Entomology, California Academy of Sciences.- — Paul H. Arnaud, Jr., California Academy of Sciences, San Francisco. The December meeting is traditionally the business meeting of the year and President Chemsak asked for the reports of the auditing and nominating com- mittees. The auditing committee report was read by Dr. E. S. Ross, and the state- ment was accepted as read. President Chemsak announced his appointments to the publication committee for 1974: Dr. E. Gorton Linsley and Dr. Marius S. Wasbauer. Dr. Marius Wasbauer, Chairman of the nominating committee, presented the slate of candidates for office in the Society for 1974: President, Cornelius B. Philip; President-elect, Howell V. Daly; Secretary, Franklin Ennik; Treasurer, Paul H. Arnaud, Jr. There were no nominations from the floor. The candidates were elected to office in the Society for 1974 by a unanimous vote. The principal speaker of the evening was the outgoing President of the Society, Dr. John A. Chemsak. His presidential address was entitled, “The Relationships of the Cerambycidae of Baja California and Mainland Mexico.” Coffee and other refreshments were served during a social hour held in the en- tomology rooms following the meeting. — M. R. Gardner, Secretary. Dr. Arnaud reports the Treasurer’s office is indebted to Mrs. V. Hawley, Office Manager of the Academy, for her dedicated attention to the Society’s accounts, to Mrs. Maria Paz Malkerson, Entomology secretary, for her attention to many letters and mailings of Society publications, and to Mr. H. Vannoy Davis, member of the firm J. K. Lasser and Company, not only for the audit of the Treasurer’s records, but also for the completion of our federal and state income tax forms. 96 The Pan-Pacific Entomologist PACIFIC COAST ENTOMOLOGICAL SOCIETY (A California Corporation) Statement of Income, Expenditures and Changes in Fund Balances Years Ended September 30, 1973 and 1972 1973 1972 Income: Dues and subscriptions $ 5,526 $ 5,231 Reprints and miscellaneous 2,999 5,174 Sales of memoirs 3,748 1,487 Advertising 333 437 Interest on savings accounts 601 574 Dividends, American Telephone & Telegraph Co. 224 208 Increase or (decrease) in value of capital stock of American Telephone & Telegraph Co. 410 470 13,841 13,581 Expenditures: Publication costs — Pan Pacific Entomologist 9,751 8,017 Memoirs 199 4,735 Reprints, postage and miscellaneous 2,075 1,442 12,025 14,194 Increase or (decrease) in fund balances 1,816 (613) Fund balances October 1, 1972 and 1971 20,219 20,832 Fund balances September 30, 1973 and 1972 $22,035 $20,219 Statement of Assets September 30, 1973 and 1972 1973 1972 Cash in bank: Commercial accounts $ 3,366 $ 2,960 Savings accounts: General fund 8,426 8,069 Memoir fund 2,189 1,928 Fall fund 2,173 1,972 Life membership fund 1,721 1,540 Total cash in bank 17,875 16,469 Investment in 80 share of American Telephone & Telegraph Co. common stock (Life membership and Fall funds) , at market value 4,160 3,750 Total $22,035 $20,219 Vol. 50, No. 1, January 1974 97 ENTOMOLOGICAL EXCHANGES AND SALES Space is available to members and non-members who wish to advertise entomo- logical exchanges, sales, announcements, etc. We can offer an ad in four issues for $1.50 per line. Ads will be arranged in order of their receipt. Contact the Advertising Manager, J. A. Chemsalc, Division of Entomology and Parasitology, University of California, Berkeley, California 94720. FOR SALE. Extensive collection of mayfly reprints. Write R. Brusca, Department of Biology, University of Arizona, Tucson, 85721. FOR SALE. Over 900 species of butterflies from the Malaysian and Indonesian Jungles, very rare moth (14" wing span), Scorpion, giant sized Indonesian flying lizard, centipedes, stick insects, all types of rhinoceros beetles and rare beetles in the families Scarabaeidae, Lucanidae, Mordellidae, Cerambycidae, etc. Write Sim Yam Seng, 21, Joon Hiang Road, Singapore, 19, Singapore. EXCHANGE. Eastern U.S. and exotic Scarabaeidae, Cerambycidae, or Cicindeli- dae for western U.S. Cicindelidae. John D. Glaser, 6660 Loch Hill Rd., Baltimore, Md. 21239. FOR SALE. Entomological literature (new and second hand) and equipment. Catalogues on request. State your interests (Lepidoptera, Coleoptera, new/second hand) . We also buy second hand books. Write to Sciences Nat, 86 Rue de la Mare, 75020 Paris, France. DICTIONARIES WEBSTER Library size 1973 edition, brand new, still in box. Cost New $45.00 Will Sell for $15 Deduct 10% on orders of 6 or more Make Checks Payable to DICTIONARY LIQUIDATION and mail to Attention: Dept. D-J46 ONTARIO TEXT EDITIONS Toronto Dominion Center Suite 1400, Fourteenth Floor Toronto, Ontario, Canada M5K 1B7 C.O.D. orders enclose 1.00 good will deposit. Pay balance plus C.O.D. shipping on delivery. Be satisfied on inspection or return within 10 days for full refund. No dealers, each volume specifically stamped not for resale. Please add $1.25 postage and handling. COMMERCIAL AND RESEARCH ENTOMOLOGISTS Obtain more accurate population counts with a light-weight motor-fan unit which sucks insects into nets D-VAC MOTOR-FAN VACUUM INSECT NETS BACK-PACKS AND HAND MODELS — also — BENEFICIAL INSECTS INSECTARY GROWN AND FIELD COLLECTED T richogramma sp. and Green Lacewings Write for Brochures D-VAC CO. P. O. Box 2095 RIVERSIDE, CALIFORNIA INSECT PINS $5. / 1000 (10 Pkgs.) (Plus Postage) 12c per 1000 CLAIR ARM IN 191 W. Palm Avenue Reedley, California 93654 ph. (209) 638-3729 PROMPT DELIVERY LOWEST PRICES IN AMERICA THE MOTHS OF AMERICA NORTH OF MEXICO THE MOTHS OF AMERICA, one of the most significant publications in the history of American entomology, will contain full-color photographs of all 10,000+ species of North American moths. The color photography has been done with excruciating care and attention to detail, and must be seen to be believed. THE MOTHS OF AMERICA, written by authorities in each group treated, constitutes a thorough review of the North American moths, with descriptions of new genera and species where appropriate. In addition to the color photographs of adults, illustrations of genitalia or other structures are included where necessary to assure accurate identification. The comprehensive text gives a full taxonomic treatment, including synonymy, plus information on distribution, foodplants, biology, and other aspects of each species. THE MOTHS OF AMERICA will be published in over 100 parts, over a period of about 20 years. Each part is individually priced, according to its size, number of color plates, and number of copies printed. Three to five parts are expected each year. Special pre-publication/ subscription prices, about 20% less than the regular list prices, apply to subscriptions for the entire work and to orders received prior to the publication of specific parts. (NOTE: The only way to get the parts already published at the reduced prices is to subscribe to the entire series.) Please start your subscription now, or order the parts you desire, before the cost be- comes prohibitive. ALREADY PUBLISHED: Fascicle 13, Part la, Pyralidae: Scopariinae & Nymphulinae, by Dr. Eugene G. Munroe. 1972. 134 p. $20.60 ($16.50 to subscribers). Fascicle 13, Part lb, Pyralidae: Odontiinae & Glaphyriinae, by Dr. Eugene G. Munroe. 1972. 116 p. $20.60 ($16.50 to subscribers). Fascicle 20, Part 2a, Bombycoidea: Saturniidae, in part ( Citheroniinae and Hemileucinae, in part), by Dr. Douglas C. Ferguson. 1972. 153 p., 11 color plates, 19 figs. $39.00 ($32.50 to subscribers). Fascicle 20, Part 2b, Bombycoidea: Saturniidae, in part (Hemileucinae, in part, and Satur- niinae), by Dr. Douglas C. Ferguson. 1972. xxi + 121 p., 11 color plates, 11 figs. $39.00 ($32.50 to subscribers). Fascicle 21, Sphingoidea, by Dr. Ronald W. Hodges. 1971. xii + 158 p., 16 plates (14 in color), 8 halftones, 19 figs. $24.00 ($19.60 to subscribers). IN PREPARATION: Fascicle 20, Part 1, Bombycoidea (excluding Saturniidae) (i.e., Lasiocampidae, Mimalloni- dae, Apetelodidae, and Bombycidae), by Dr. J. G. Franclemont. Fascicle 13, Part lc, Pyralidae: Evergestiinae, by Dr. Eugene G. Munroe (including 14 color plates for all 3 parts of Fascicle 13. 1). Fascicle 6, Part 2, Gelechioidea: Oecophoridae, by Dr. Ronald W. Hodges. Fascicle 22, Part 2, Lymantriidae, by Dr. Douglas C. Ferguson. Also in preparation: Notodontidae, Cosmopterygidae, Walshiidae, and Aegeriidae. THE MOTHS OF AMERICA NORTH OF MEXICO belongs on the shelf of every serious lepidopterist and in every significant library. Please make sure that your local library sub- scribes, so that this important reference work will be available to you and your colleagues in the future. A fully descriptive brochure, with sample color plate and order form, is available on request. 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Send orders to: Pacific Coast Entomological Society c/o California Academy of Sciences Golden Gate Park San Francisco, California 94118 Vol. 50 APRIL 1974 No. 2 THE Pan-Pacific Entomologist CARLSON AND RITCHER — A new genus of Ochodaeinae and a description of the larva of Pseudochodaeus estriatus (Schaeffer) (Coleoptera: Scarabaeidae) 99 SLOBODCHIKOFF — Notes on the biology of Therion circumflexum (L.), with a description of the immature stages (Hymenoptera: Ichneu- monidae) 111 STEFFAN — Nearctic species of Schwenkfeldina (Diptera: Sciaridae) 118 HARDY AND ANDREWS — Observations on Megasoma with behavioral notes on some lamellicorn beetles associated with sand dunes (Cole- optera: Scarabaeidae, Lucanidae) 124 CHEMSAK AND LINSLEY — Descriptions and records of Clytini from Mexico and Central America (Coleoptera: Cerambycidae) 129 HOVORE AND GIESBERT — Two new species of Cerambycidae from Southern California (Coleoptera) _ 139 PHILIP AND FLOYD — New North American Tabanidae XXL Another new Bolbodimyia from Mexico (Diptera) 145 POTTS — Revision of the Scarabaeidae: Anomalinae 1. The Genera occurring in the United States and Canada (Coleoptera) 148 SLOBODCHIKOFF — Behavioral and Morphological mimicry in a cranefly and an ichneumonid 155 HARDY — A new species of Cyclocephala Latreille from California sand dunes (Coleoptera: Scarabaeidae) 160 CHANDLER — A redefinition of the Tyrini with the addition of Anitra Casey (Coleoptera: Pselaphidae) 162 DOUTT — The genus Dicopus Enock (Hymenoptera: Mymaridae) 165 SIRI AND BOHART — A review of the genus Mellinus (Hymenoptera: Sphecidae) 169 MILLER — Two new Heliococcus species, a key to the North American species, and a list of world species (Homoptera: Pseudococcidae) 177 MILLER — Mealy bugs of San Clemente Island, California (Homoptera: Pseudococcidae) 193 SCIENTIFIC NOTES 203, 204, 205, 207 RECENT LITERATURE 154, 159, 164, 168 OBITUARY 209 SAN FRANCISCO, CALIFORNIA • 1974 Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES THE PAN-PACIFIC ENTOMOLOGIST EDITORIAL BOARD J. A. Chemsak, Asst. Editor E. G. Linsley H. V. Daly J. T. Doyen, Editor P. H. Arnaud, Jr., Treasurer E. S. Ross H. B. Leech Published quarterly in January, April, July, and October with Society Proceed- ings appearing in the January number. All communications regarding nonreceipt of numbers, requests for sample copies, and financial communications should be addressed to the Treasurer, Dr. Paul H. Arnaud, Jr., California Academy of Sci- ences, Golden Gate Park, San Francisco, California 94118. Application for membership ip the Society and changes of address should be addressed to the Secretary, Franklin Ennik, Vector Control Section, California Department of Health, 2151 Berkeley Way, Berkeley, 94704. The annual dues, paid in advance, are $7.50 for regular members of the Society, $5.00 for student members, or $10.00 for subscriptions only. Single copies are $2.50 each or $10.00 a volume. Make checks payable to Pan-Pacific Entomologist. The Pacific Coast Entomological Society Officers for 1974 C. B. Philip, President Paul H. Arnaud, Jr., Treasurer H. V. Daly, President-elect Franklin Ennik, Secretary Statement of Ownership Title of Publication: The Pan-Pacific Entomologist. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher, Publisher and Owner: Pacific Coast Entomological Society, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118. Editor: Dr. John T. Doyen, Division of Entomology & Parasitology, University of California, Berkeley, California 94720. Managing Editor and Known Bondholders or other Security Holders: None. This issue mailed 8 November 1974 Second Class Postage Paid at San Francisco, California and additional offices. ALLEN PRESS, INC. ''“"J * 0 LAWRENCE, KANSAS u. s. The Pan-Pacific Entomologist Yol. 50 April 1974 No. 2 A New Genus of Ochodaeinae and a Description of the Larva of Pseudochodaeus estriatus (Schaeffer) (Coleoptera: Scarabaeidae) 1 David C. Carlson and Paul 0. Ritcher Department of Entomology Oregon State University , Corvallis 97331 Schaeffer (1906) described Ochodaeus estriatus based on a single specimen from California. From his comments it is clear that he con- sidered the species different enough from other Ochodaeus to warrant placement in another genus. He did not do so solely because he had only the one specimen and did not wish to dissect it. Since 1906, additional specimens of Ochodaeus estriatus have been taken in California and Oregon. Study of their morphology indicates that Schaeffer was indeed correct in his original conclusions. We here- by erect the new genus Pseudochodaeus for Schaeffer’s species. Pseudochodaeus New Genus Type Species. — Ochodaeus estriatus Schaffer, 1906, p. 271; monobasic. Description. — Body elongate, convex; head, pronotum, and elytra densely setose; ventral surfaces and legs clothed with longer setae. Head transverse; mandibles and labrum visible from above; labrum transverse, densely setose and very shallowly emarginate anteriorly; anterior clypeal margin strongly reflexed. Antennae 10- segmented, first segment triangular, densely setose; club with eighth segment en- larged and cupped, receiving segments 9 and 10 (Fig. 5) . Eyes not divided by ocular canthus. Pronotum subquadrate, evenly convex except for shallow, median longitudinal impression; anterior margin shallowly and evenly indented; posterior margin sinuate on each side of midline; marginal bead entire. Scutellum setose and punctate, rounded posteriorly. Elytra long, apex of abdomen not visible from above; posterior sutural angle simple, not dentate; 1 punctate stria between median suture and humeral umbone, this stria close to median suture, becoming confluent with median suture at elytral declivity. Propygidium simple, not modified to accept apex of elytra. Pygidium and 6 visible abdominal sternites densely setose; sternites free, not connate, fifth stemite lacking stridulatory peg. Foretibia tridentate, teeth well developed. Outer apical spur on meso- and metatibia pectinate; meta- trochanter produced beyond posterior edge of femur into sharp projection. Male 1 The financial support of the National Science Foundation (Grants GB-3586, GB-6194X, and GB-31129) and of the Oregon Agricultural Experiment Station is gratefully acknowledged. Oregon Agricultural Experiment Station, Technical Paper No. 3786. The Pan-Pacific Entomologist 50: 99-110. April 1974 100 The Pan-Pacific Entomologist genitalia (Figs. 1-3) : proximal end of basal piece tapering to point; parameres symmetrical, lacking basal elongate process; V-shaped apophysis present; aedeagal sac well developed, bearing numerous sclerotized structures. The genus Pseudochodaeus possesses the following combination of characters which distinguishes it from Ochodaeus Serville: single elytral stria, pectinate metatibial spur, absence of a stridulatory peg, enlarged and cupped eighth antennal segment, unmodified propygidium, elongate elytra not modified at the apical sutural angle, and produced metatrochanters. In Ochodaeus the eighth antennal segment is not cupped (Fig. 6) and the parameres of the male genitalia each possess an elongate basal process (Fig. 4) . It should be noted that at least one species now placed in the genus Ochodaeus lacks a stridulatory peg (Carlson, 1975) . The genus Pseudochodaeus is of particular interest because it has characters in common with other Ochodaeinae such as Ochodaeus, Codocera Eschscholtz, and Chaetocanthus Peringuey, and with Pachy- plectrus LeConte, now placed in the Hybosorinae. Chaetocanthus is known only from South Africa and differs from Pseudochodaeus in having bidentate foretibia, 9-segmented antennae with the eighth seg- ment not enlarged, and pectinate tibial spurs on all legs. Chaetocanthus, however, has elytra with only one stria, a tranverse and non-emarginate labrum, and lacks the stridulatory peg. VanDyke (1904) and Fall (1904) mentioned that a new species of Ochodaeus, presumably 0. estriatus, was similar in some ways to Pachy- plectrus laevis LeConte. We have examined P. laevis and find several similarities to Pseudochodaeus and other genera in the Ochodaeinae. The male genital apparatus in Pachyplectrus laevis has a pointed basal piece, symmetrical parameres, and a well developed aedeagal sac, char- acters shared by Pseudochodaeus, Ochodaeus, Codocera, and Chaetocan- thus (Carlson, 1975). The eighth abdominal spiracle of Pachyplectrus laevis is located in the tergites and the seventh is adj acent to, but not sur- rounded by the tergite. In the Ochodaeinae both the seventh and eighth abdominal spiracles are in the tergites, however, in P seudochodaeus the seventh spiracle is close to the edge of the tergite. In Ochodaeus Figs. 1-8. Fig. 1. Male genitalia of Pseudochodaeus estriatus (Schaeffer), right lateral view. AS, aedeagal sac. Fig. 2. Male genitalia of P. estriatus, (Schaef- fer), left lateral view. AS, aedeagal sac; BP, basal piece; PM, parameres. Fig. 3. Male genitalia of P. estriatus (Schaeffer), dorsal view. AP, V-shaped apophysis; BP, basal piece; PM, parameres. Fig. 4. Male genitalia of Ochodaeus praesidii Bates, dorsal view. Fig. 5. Antennal club of P. estriatus (Schaeffer), dorsal view. Fig. 6. Antennal club of Ochodaeus pectoralis LeConte, dorsal view. Fig. 7. Fe- male, hind leg of P. estriatus (Schaeffer), ventral view. SP, setal patch. Fig. 8. Male, hind leg of P. estriatus (Schaeffer), ventral view. Vol. 50, No. 2, April 1974 101 102 The Pan-Pacific Entomologist the seventh spiracle is well within the tergite. Pachyplectrus laevis lacks pectinate tibial spurs and the stridulatory peg. Examination of several other genera in the Hybosorinae ( Hybosorus MacLeay, Phaeochrous Laporte, and Chaetodus Westwood) has revealed that the male genital apparatus has a truncate basal piece and asymmetri- cal parameres. Also, according to Ritcher (1969 a, b), only the eighth abdominal spiracle of these genera is located in the tergites. On the basis of the characters we have examined, it appears that Pachyplectrus is an intermediate form and the limits of the Ochodaeinae and Hybosorinae may need to be reevaluated. A final decision on this will have to await a more extensive study of the morphology of Pachy- plectrus and examination of Synochodaeus Kolbe and Enodognathus Benderitter, two other genera in the Ochodaeinae. Pseudochodaeus estriatus (Schaeffer) New Combination (Figures 1-3, 5, 7-9) Ochodcieus estriatus Schaeffer, 1906, p. 271. Type Material. — Holotype: Female. In the NMNH. Data on labels: Cal- ifornia, Fresno County, Millwood; Holotype; estriatus Schaef. ; Property of Mark Robinson; M. Robinson collection 1959. We have examined the holotype and find it is missing one hind leg and one antenna, but is otherwise intact. It is 9.0 mm in length and 4.2 mm in width through the midpoint of the elytra. Description. Males— Length 5.7 mm to 9.5 mm, width through midpoint of elytra 2.7 mm to 4.6 mm. Color light brown to dark reddish-brown. Vertex, pronotal disc, scutellum, and elytra covered with relatively short, yellow, semi-erect setae. Margins of pronotum and elytra with longer yellow setae. Mandibles distinctly angulate on outer margins, not evenly convex. Left mandible bidentate with large prostheca and basal molar areas; secondary tooth broad, not pointed. Right mandible bidentate with large prostheca and molar area; secondary tooth bluntly pointed. Labrum densely setose, nearly obscured by setae, dorsal surface simple. Mentum subquadrate, shallowly emarginate anteriorly, with shallow, median, longitudinal impression; setose on each side of median impression. Anterior clypeal margin reflexed, forming tubercle at center; clypeus setose. Frons im- pressed medially, setose and punctate, setae arising from punctures, anterior edge of punctures slightly elevated; median area of vertex sparsely punctate and lack- ing setae; posterior- lateral areas of vertex setose and punctate. Pronotum convex, with median longitudinal impression; disc evenly setose, each seta arising from puncture. Scutellum setose and punctate. Surface of each elytron densely setose; setae arising from punctures. Apical sutural angle of elytra simple, not dentiform. Foretibia with apical spur large, arcuate, and with apex enlarged. Meso tibia expanded laterally; outer spur pectinate; first tarsal segment longer than next three segments together. Metafemur with large tooth on posterior margin at apical one-third (Fig. 8) . Metatibia expanded laterally (Fig. 8) ; oblique row of spines running from base to inner margin at distal one-third; subapical oblique row of spines at outer margin; outer apical angle between subapical row of Vol. 50, No. 2, April 1974 103 Fig. 9. Distribution of Pseudochodaeus estriatus (Schaeffer) in California and Oregon, U. S. A. 104 The Pan-Pacific Entomologist spines and apex sparsely setose; outer spur pectinate; first tarsal segment longer than next three segments together. Male genitalia (Figs. 1—3) : aedeagal sac with 1 basal sclerotized structure and 5 to 7 large, sclerotized spines; distal two-thirds of sac covered with irregularly arranged patches of pigmented spiculi. V-shaped apophysis with distal margin rounded. Females. — L ength 5.8 mm to 9.3 mm, width through midpoint of elytra 2.9 mm to 4.3 mm. Other characters same as males except that the outer apical angle of the metatibia bears a dense patch of setae (Fig. 7) . Distribution (Fig. 9) and specimens examined (holotype, 41 $ , 49 $). California: El Dorado County : 2 2, July 1932, F. T. Scott (KU) ; 4 $, 4 2, Pacific House, July — 3 August 1931, H. Hinton (CAS); 1 $, Snowline Camp, 20 July 1948, P. D. Hurd (UCB) ; 1 $ Riverton, August 1930, H. Hinton (CAS) ; 1 2, Pollock Pines, 25 July 1938, A. T. McClay (ASU) ; 1 d , nr. Whitehall, 21 July 1931, H. Hinton (NMNH) ; 1 2, Garden Valley, 4 July 1965 (ARH). Fresno County. 1 $ 2 $ , vie. Wishon, 15 August 1971, D. Marqua (PHS) ; 1 2, vie. Wishon, 6 August 1971, F. T. Hovore (ORSU) ; 1 d, Cedar Grove, Kings River Cyn, 17 July 1952, Cazier, Gertsch, Schrammel (AMNH) ; 1 $, Huckleberry Meadow, 6500', August 1912 (CAS); 1 $, Camp Greeley, 2800', 25 July (CAS); 1 $, Millwood (NMNH). Madera County: 1 $, Sugar Pine, 21 July 1933, R. P. Allen (CAS); Id, Placer Ranger Station, NE Northfork, 30 July 1931 (CAS) ; 1 2 , Miami Ranger Station, 10 mi N Oakhurst and S of Yosemite Nat. Park, 24 June 1938 (CAS). Mariposa County: 1 2, Miami Ranger Station, 4000', 29 June 1946, H. P. Chandler (CAS). Modoc County: 1 2, Hacka- more, 17 July 1934, 0. H. Schwab (CAS) ; 2 d, Pitt River, May (CAS). Plumas County: 1 2, Walker Mine, 25 July 1930, E. F. Wohletz (CAS) ; 1 2, S end of Red Clover Valley, 8 mi N Beckworth, 28 August 1963, H. B. Leech (CAS). Shasta County: Hat Creek: 1 2, July 1948, A. S. Perry (UCB) ; 1 2 , 16 July 1951, G. Pronin (CAS) ; 1 d, 20 July 1951, G. Pronin (CAS) ; 2 d, 27 July 1951, G. Pronin (CAS); 1 d, 25 August 1951, G. Pronin (CAS); 1 2 , 25 August 1952, G. Pronin (CAS); 1 $ , 12 July 1955, J. W. MacSwain (UCB); 1 2, 19 July 1955, G. Pronin (CAS) ; 1 $ 1 2 , 29 July 1971, D. Carlson (DCC). Trinity County: 1 d, Carrville, 6 July 1950, D. Guiliani (CAS). Tulare County: 1 d , 2 mi E Posey, 21 August 1971, E. Giesbert (DCC). Tuolomne County: 1 d , trib. to Herring Creek, 1.5 mi N Pinecrest Lake, 30 August 1964, H. B. Leech (CAS) ; 1 2, Twain-Harte, 4000', July 1937, Blaisdell (CAS) ; 1 2, Straw- berry Resort, 5243', 8 August 1937, E. Herald (CAS); 1 d, Strawberry, 15 August 1962, C. A. Toschi (UCB). Oregon: Jackson County: Fourbit Ford Cmpgnd., 11 mi SE Butte Falls: 4 d 5 2 , 18 July 1973, P. O. Ritcher and D. C. Carlson (ORSU, DCC) ; 8 d 10 2,9 August 1973, D. C. Carlson and W. N. Mathis (ORSU, DCC, NMNH) ; 3 d 9 2, 29 August 1973, D. C. Carlson and R. L. Westcott (ORSU, DCC, RLW, HFH) : 1 2, Jet. Rd 3520 and S. Fork Fourbit Creek, 11 mi SE Butte Falls, 29 August 1973, D. C. Carlson and R. L. Westcott (ORSU). Lake County: 1 d, Warner Canyon, 31 July 1968, R. L. Westcott (RLW). Remarks. — T he color variation of the adults of P. estriatus is ap- parently a result of age. Specimens which we collected on 18 July 1973 were light brown with the setation intact and legs unabraded. Later Vol. 50, No. 2, April 1974 105 collections at the same site (9 and 29 August 1973) yielded all dark reddish-brown specimens which were quite worn. This species has also been recorded from Deschutes County, Oregon (Hatch, 1971) , but we have been unable to verify the record. Ochodaeus simplex LeConte is the only Ochodaeus we and others have collected in Deschutes County. We have visited four of the localities where P. estriatus occurs and have collected specimens at two of them (Fourbit Ford Campground, Jackson Co., Oregon and Hat Creek, Shasta Co., California). All of these are forested areas with sparse ground cover of shrubs and grass and all are close to streams. At Fourbit Ford Campground the follow- ing plant species were fairly abundant: Pseudotsuga menziesii (Mirb.), Pinus ponder osa Laws., Abies grandis (Dougl.), Quercus hello ggi Newb., Holodiscus discolor (Pursh), Ceanothus integerrimus Hook and Arm., Amelanchier florida Lindl., Symphoricarpos albus (L.), Mahonia sp., and Corylus sp. Bracken fern and sparse grass were also present. Included with the holotype which we borrowed from the NMNH was another specimen bearing a handwritten locality label identical to that with the holotype. This specimen is a male and we believe it to be the misplaced specimen which Schaeffer referred to in the original descrip- tion of 0. estriatus. We are depositing specimens of P. estriatus from Oregon, including males with everted aedeagal sacs, in the NMNH and collection of H. F. Howden, Carleton University, Ottawa, Ontario, Canada. Biology and Immature Stages Information on the biology of the Ochodaeinae is exceedingly scanty, which is surprising since the adults are often abundant. About all that has been known of the adults is that they fly at night, are attracted to light, and burrow in the soil. By dissection, we have determined that the mid- and hindgut of P. estriatus adults often contain numerous small spores of a basidiomycete similar to those of puff balls. The female reproductive system has 6 ovarioles in each ovary but only 1 egg develops at a time on each side. A typical fully developed egg is white, elongate-oval in shape, 2.78 mm long, and 1.44 mm wide. This is quite large for such a small beetle. Like other Ochodaeinae, the flight period of P. estriatus is quite long. Adults were taken at light in southern Oregon from July 18 to August 29, 1973. Dissection indicated that egg development proceeds slowly and that the few eggs are laid over a long period of time. 106 The Pan-Pacific Entomologist While digging for Pleocoma LeConte larvae in southern Oregon in 1961, David Fellin, then a graduate student, obtained a small scarabaeid larva with 4-segmented antennae, which was unlike any other with which the junior author was familiar. He suspected it to be an Ochodaeus since he was familiar with the larvae of all scarab genera occurring in in Oregon except those of Glaresis Erichson, Bolborhombus Cartwright, and Ochodaeus. Glaresis was eliminated because of its very small size and Bolborhombus because larvae of related Geotrupinae all have 3- segmented antennae. The main problem with this tentative identification was that adult Ochodaeus had never been collected in southern Oregon west of the Cascade Mountains. In recent years, a number of unsuccessful attempts were made to collect Ochodaeus adults at black-lights in southern Oregon, in the vicinity of Butte Falls. Finally, on 18 July 1973 we took 9 adults (4 #,5$) of Pseud ochodaeus estriatus (Schaeffer), 11 miles southeast of Butte Falls. This site is less than a mile from where the supposed Ochodaeus larva was dug in 1961. Medvedev (1960) described the larva of Codocera ferruginea Esch- scholtz, a common Eurasian species of Ochodaeinae, based on 2 un- associated larvae collected in the soil of a Ukrainian tree plantation. Unfortunately, characters of the epipharynx, mandibles, and maxillae were neither described nor figured and we were unable to borrow the specimens for study. At first, we assumed our Ochodaeus larva differed from Medvedev’s Codocerra larva because they belonged in different genera. However, Medvedev’s larva could well be a Trox Fabricius since it resembles that genus in the type of head setation, in having 3-segmented antennae, and in having fleshy anal lobes. Pseudochodaeus estriatus (Schaeffer), Third-Stage Larva (Figures 10-20) Description based on one third-stage larva collected from the soil 10 miles east of Butte Falls, Jackson County, Oregon, 28 January 1961, by David Fellin. Figs. 10-18. Larva of Pseudochodaeus estriatus (Schaeffer). Fig. 10. Head. A, antenna; C, clypeus, DES, dorsoepicranial seta; F, frons; L, labrum; PFS, posterior frontal setae. Fig. 11. Epipharynx. CPA, chaetoparia; GP, gymno- paria; NI-3, nesia; PE, pedium; PA, phobae; ZY, zygum. Fig. 12. Left mandible. PA, preartis; MA, molar area; SA scissorial area (upper surface). Fig. 13. Right maxilla (lower surface). CAR, cardo; G, galea; GU, uncus of galea; LA, Vol. 50, No. 2, April 1974 107 lacina; LAU, uncus of lacinia; MST, maxillary stridulatory teeth. Fig. 14, Labium with hypopharynx. GL, glossa. Fig. 15. Last 2 segments of antenna (ventral surface) . VSS, ventral sensory spot. Fig. 16. Right mandible (Ventral surface) . MA, molar area. SA, scissorial area; VP, ventral process. Fig. 17. Claw of prothoracic leg. Fig. 18. Left lateral view of entire third-stage larva. DL1, DL2, DL3, dorsal lobes. 108 The Pan-Pacific Entomologist Description. — Larva typically scararbaeiform, C-shaped (Fig. 18). Head light yellow-brown, thorax and abdomen whitish. Maximum width of head capsule (Fig. 10) 2.28 mm. Surface of head fairly smooth. Epicranial stem present, frontal sutures largely absent. With 1 dorso- epicranial seta on each side. Frontal area with pair of posterior frontal setae on each side, other frontal setae absent. Frontoclypeal suture mostly absent. Labrum trilobed. Antenna 4-segmented with distal segment reduced in size, distal segment less than half as long as third segment (Fig. 10) . Ventral surface of next to last antennal segment (third) with 1 apical, elliptical sensory spot; last antennal segment also with similar ventral, apical spot (Fig. 15). Apex of last antennal segment with about 7 sensory pegs. Mandibles (Figs. 12 and 16) subtri- angular in shape, each with 2 scissorial teeth and with typical ventral, accessory, articulating process. Maxilla (Fig. 13) with separate galea and lacinia. Maxillary stridulatory teeth consisting of a group of about 14 short, conical teeth. Maxillary palpus 4-segmented. Labium (Fig. 14) with symmetrical oncyli. Epipharynx (Fig. 11) with complete, symmetrical zygum. Pedium surrounded by phobae and with inner second group of phobae posterior to anterior phobae. Tormae fused, symmetrical, lacking either epitorma or pternotormae. Haptolachus with 3 nesia; 1 large sclerotized plate on each side and 1 small median sense cone. Chaetopariae with a few (10 to 11) strong chaetae on each side. Chaetopariae without microsensilla. Abdominal segments 1-7 each with 3 dorsal lobes (Fig. 18) . Anterior 2 dorsal lobes each with tranverse row of setae, posterior dorsal lobe bare. Spiracles crib- riform, inconspicuous, occurring on prothorax and first 8 abdominal segments. Prothoracic spiracle elliptical, with slight caudal emargination. Respiratory plates of abdominal spiracles all similar in size, crescent shaped, with caudo- ventral emarginations. Legs 4-segmented, each ending in well developed claw (Fig. 18) : Claws similar in size, each bearing 2 setae (Figs. 17 and 18). Ventral surface of last abdominal segment (Fig. 19) with poorly developed raster. Raster consisting of a few (about 18) irregularly distributed, rather short, slender setae. Raster extending less than one-half the distance to seventh abdominal segment; anteriorly, with broad, bilobed raised area. Anal opening Y-shaped surrounded by narrow, transverse, dorsal anal lobe and 2 larger, subtriangular, lower anal lobes (Fig. 20) . Remarks. — The larva of P. estriatus has a number of morphological characters indicating it belongs to the more primitive Scarabaeidae. These characters are similar to those of the less specialized scarabaeid subfamilies Troginae and Aphodiinae. It resembles larvae of Trox (sensu strictu) in lacking a well-developed fronto-clypeal suture, in having a haptolachus with 2 sclerotized plates and a median sense cone, and in the type of anal opening and surrounding lobes. It differs from Trox and Omorgus Erichson larvae, however, in having 4-segmented antennae (not 3 as in Troginae), a trilobed labrum (not bilobed), and a different type and arrangement of maxillary stridulatory teeth. The larva of P. estriatus will key to Aphodiinae in Ritcher’s 1966 monograph and in his 1967 paper. It can be distinguished readily Vol. 50, No. 2, April 1974 109 Figs. 19 and 20. Larva of Pseudochodaeus estriatus (Schaeffer). Fig. 19. Ventral view of last (tenth) abdominal segment. Fig. 20, caudal view of last abdominal segment. AO, anal opening; DAL, dorsal anal lobe; VAL, ventral anal lobe. from larvae of Aphodiinae by such characters as the incomplete frontoclypeal suture (Fig. 10), the complete zygum of the epipharynx (Fig. 11), and the absence of an epitorma (Fig. 11). It resembles larvae of Aphodiinae in that both have ventral, apical sensory areas on the last 2 antennal segments (Fig. 15). Although nothing is known of the subterranean habits of the larva of P. estriatus, some inferences can be drawn from its morphology. The worn mandibles, well developed legs and claws, and body setation are all indicative of an active larva, not of a larva whose food was pro- visioned by the adult. Acknowledgments We wish to thank the following for the generous loan of specimens: American Museum of Natural History (AMNH) ; Arizona State Uni- versity, Tempe (ASU) ; California Academy of Sciences (CAS) ; Oregon State University (ORSU) ; National Museum of Natural History (NMNH) ; University of California, Berkeley (UCB) ; A. R. Hardy (ARH) ; F. T. Hovore (FTH) ; P. H. Sullivan (PHS) ; and R. L. Westcott (RLW). The letters in parentheses are the abbreviations used in the text for the collections from which we borrowed specimens. We are indebted to W. C. Denison, Oregon State University botanist, for identifying the basidiomycete spores. We also thank R. B. Roberts for many helpful suggestions and Mrs. B. Hall for assistance with the larval illustrations. 110 The Pan-Pacific Entomologist Literature Cited Carlson, D. C. 1974. Taxonomic characters of the genus Ochodaeus Serville with descriptions of two new species in the 0. pectoralis LeConte species complex (Coleoptera: Scarabaeidae) . Bull. S. Calif. Acad. Sci., (in press) . Fall, H. C. 1904. In: Proc. Pacific Coast Entomol. Soc., 1:22. Hatch, M. H. 1971. The beetles of the Pacific Northwest. Part V. Univ. Washington Press, Seattle, 662 pp. Medvedev, S. I. 1952. Larvae of scarabaeoid beetles of the fauna of the USSR. Opredeliteli Faune USSR, Moscow, 47:1-343. Medvedev, S. I. 1960. Descriptions of the larva of eight species of lamelicorn beetles from the Ukraine and Central Asia. Zool. Zhurn., 39:381-393. Ritcher, P. 0. 1966. White grubs and their allies, a study of North Ameri- can scarabaeoid larvae. Oreg. State Univ. Press, Stud. Entomol., 4:1-219. Ritcher, P. 0. 1967. Keys for identifying larvae of Scarabaeoidea to the family and subfamily (Coleoptera). Calif. Dept. Agr. Occ. Papers, 10:1-8. Ritcher, P. 0. 1969a. Spiracles of adult Scarabaeoidea (Coleoptera) and their phylogenetic significance. I. The abdominal spiracles. Ann. Entomol. Soc. America, 62 (2) -.869-880. Ritcher, P. O. 1969b. Spiracles of adult Scarabaeoidea (Coleoptera) and their phylogenetic significance. II. Thoracic spiracles and adjacent sclerites. Ann. Entomol. Soc. America, 62 (6) : 1388-1398. Vandyke, E. C. 1904. In: Proc. Pacific Coast Entomol. Soc., 1:22. Vol. 50, No. 2, April 1974 111 Notes on the Biology of Therion circumflexum (L.), with a Description of the Immature Stages (Hymenoptera: Ichneumonidae) C. N. Slobodchikoff Department of Biological Sciences Northern Arizona University, Flagstaff 86001 Wasps of the genus Therion are internal parasitoids of Lepidoptera larvae. A female wasp injects one or more eggs into the body cavity of a caterpillar, and the parasitoid larva, upon hatching from the egg, begins to feed on the non-vital tissues of its host. The parasitoid larva under- goes three larval molts. When its host pupates, the larval parasitoid kills the host and pupates within the host’s pupal shell. The biology of larvae and adults of T. morio (Fabr.) has been described by Tothill (1922). The purpose of this present paper is to describe the egg, larval stages, and pupa of T. circumflexum (L.). Behavior of T. morio and T. circumflexum adults has been described elsewhere (Slobodchikoff, 1973). Materials and Methods Six T. circumflexum females were captured on 22 April 1969 at Alpine Lake, Marin County, California. Each female was immediately in- troduced into a round cardboard container 10 cm in diameter and 7 cm deep, which was covered with a plastic top. Three containers each had 5 third instar larvae of Spodoptera exigua (Hbn.), the other three each had 5 third instar larvae of Pseudaletia unipuncta (Haw.). Host larvae were obtained from the insect pathology Lepidoptera cul- tures maintained by the Division of Entomology and Parasitology, University of California, Berkeley. Female wasps were kept in the con- tainers for two days, then removed and kept in larger plastic cages (see Slobodchikoff, 1973, for a description of the cages). All con- tainers were kept together at an average temperature of 22 °C and an average relative humidity of 45 percent. A total of 9 Spodoptera and 11 Pseudaletia larvae or pupae were dissected. One Spodoptera larva was dissected immediately after the adult wasps were transferred to plastic cages. Six Therion eggs were found. On 28 April, 1969, five days after initial contact with a parasitoid wasp, 3 larvae of both Spodoptera and Pseudaletia were dissected. Two Spodoptera larvae lacked Therion larvae, while the third contained 4 first instar Therion larvae. Two Pseudaletia larvae were not parasitized, while one con- The Pan-Pacific Entomologist 50 : 111 - 117 . April 1974 112 The Pan-Pacific Entomologist 3 4 Figs. 1-4. Therion circumflexum. 1. Egg, lateral view. 2. First instar larva, enclosed in sac. Dorsal view, head pointing down. 3. First instar larva head capsule, anterior view; see explanation of fig. 4 for names of labeled structures. 4. Third instar head capsule, anterior view. Labeled structures are the following: a, antenna; c, clypeus; eps, fused epistoma, pleurostoma, and hypostoma; Ip, labial palp; Is, labial sclerite; m, mandible; mp, maxillary palp; sp, silk press; ss, stipital sclerite. Vol. 50, No. 2, April 1974 113 tained a single first instar Therion larva. Twelve days after initial con- tact (5 May, 1969), two Spodoptera and six Pseudaletia were dissected. Each Spodoptera contained a single second instar Therion larva. Three Pseudaletia larvae were unparasitized, one contained two live and seven dead larvae, one contained two live and six dead larvae, and one con- tained three live and 10 dead larvae. Nineteen days after initial con- tact (12 May, 1969), two Spodoptera larvae and one Pseudaletia larva were dissected. All three larvae each contained a single third instar Therion larva. Twenty six days after initial contact (19 May, 1969), one Spodoptera and one Pseudaletia pupa were dissected. Each con- tained a Therion pupa. One Spodoptera pupa produced an adult male Therion on 25 May, 1969. The five remaining Spodoptera larvae died before pupating. One Pseudaletia pupa produced an adult moth on 27 May, 1969. The remaining Pseudaletia died before pupating. Measure- ments were made with an ocular micrometer and a Wild M7 binocular dissecting microscope. Drawings were made with a Wild drawing tube attachment to the M7 microscope. Descriptions and discussion Egg (Fig. 1). Total length, 0.371-0.266 mm (x zz 0.306 mm, N = 6). Total width, 0.214-0.186 mm (x zz 0.199 mm) ; Color milky- white. Anterior end elongate — globose; Caudal end prolonged into a tapering petiole; Length of petiole, 0.221-0.184 mm (x zz 0.199 mm) ; Width of petiole at widest point, 0.020-0.011 mm (x = 0.014 mm); Width of petiole at caudal tip, 0.0020-0.0016 mm (x zz 0.0018 mm) . All eggs found were attached to the body wall of the host. Two eggs were found in the thoracic region of the host caterpillar, and 4 eggs were removed from the abdominal region. Based on this small sample, it appears that the female wasp does not have a preferential oviposition site, at least under confined laboratory conditions. Tothill (1922) re- ported that T. morio females lay an egg into the body wall opposite to that initially pierced by the ovipositor. This egg-laying behavior is also present in T. circumflexum (Slobodchikoff, 1971). First instar (Figs. 2-3). Body enclosed in translucent white sac; length of larva, 3.1-2.8 mm (x zz: 3.0 mm, N = 11) ; width of larva at midgut, 1.2-1.0 mm (x zz 1.0 mm) ; body slender; color white; head capsule large, quadrate, strongly sclerotized; antenna papilliform; mouthparts prognathous; mandibles large, falcate, strongly sclerotized; maxillary palp present as disc; labial palp papilliform; labral sclerite present as large, sclerotized hemispherical plate, incomplete laterally; stipital sclerite large, hemiellipsoid, open laterally; hypostoma, pleurostoma, and epistoma strongly sclerotized, hypostomal arch complete dorsally, silk press strongly sclerotized. 114 The Pan-Pacific Entomologist Figs. 5-6. Therion circumjlexum. 5. Third instar larva, lateral view. 6. pupa of T. circumflexum, lateral view. Tothill (1922) reports that the sac which encloses the first instar larva of T. morio is made up of 2 non-cellular layers. The mandibles of the larva are enclosed in this sac and are not used for lacerating the tissues of the host. Tothill postulates that the food of T. morio first instar larvae is the strained liquid of the hemolymph. The first instar larva of T. circumflexum has its mandibles protruding from the sac, and the mandibles may be used for lacerating tissue. One first instar larva was observed lacerating and ingesting its host’s fat body tissue. Another apparent difference between T. morio and T. circumflexum is Vol. 50, No. 2, April 1974 115 the location of the first instar larvae in its host. Tothill reports that T. morio larvae are always found in the caudal half of their host larva, Hypantria cunea (Drury), and in the extreme caudal portion of a host pupa. In the present study no pupae were found to contain first instar larvae, and in both Spodoptera and Pseudaletia first instar Therion larvae were found in both thoracic and abdominal regions. Results of the dissections suggest that a T. circumflexum egg develops into a first instar larva within five days in both Pseudaletia and Spodoptera. First instar larvae may persist for as long as 12 days in Pseudaletia. The dead first instar larvae of Therion found in Pseu- daletia 12 days after oviposition were represented by head capsules and portions of the enclosing sacs, suggesting that they had been killed and partially eaten by other first instar larvae. No encapsulated larvae were found. Second instar. Body free of any enclosing sac; length 6.5-5. 0 mm (x = 5.75 mm, N = 2) ; width at midlength of body 1. 3-1.0 mm (x 1.15 mm) ; color white; similar in attributes to first instar. The second instar differs from the first primarily in the lack of an enclosing sac and a larger body size. Tothill (1922) reports that the mandibles of the second instar of T. morio are smaller than those of the first instar, and are probably used for feeding on the host’s tissues. The mandibles of the second instar of T. circumflexum were not observed to be any smaller than those of the first instar. Although feeding was not observed for the second instar, it seems probable that this instar, like the first, feeds directly on the tissues of the host. In Spodoptera , the time of development from oviposition to second instar is approxi- mately 12 days. The available data suggest that the time of develop- ment from oviposition to second instar may be slightly longer in Pseudaletia. Third instar (Figs. 4-5). Length, 14.0-9.0 mm (x = 11.2 mm, N = 3) ; width at midlength, 7.2-4.0 mm (x 5.6 mm) ; color yellow-white; no apparent setae on head or body; antenna disc-shaped; antennal discs dorsomedially contiguous, separated by thin extension of dorsal head capsule sclerotization ; maxillary and labial palps disc-shaped; mandible falcate; strongly scleortized distally, broadly trapezoidal in shape proximally; clypeus pentagonal, lightly sclerotized; labrum small, ellipsoid, heavily sclerotized; labial sclerite thin, heavily sclerotized laterally, expanded into lightly sclerotized hemisphere ventromedially ; sclerotic spur absent; epistoma, pleurostoma, and hypostoma fused, expanded laterally; hypostoma strongly sclerotized; stipital sclerite ellipsoid, medially recurved toward base of mandible, thickening laterally; silk press prominent, strongly sclerotized dorsally, weakly sclerotized ventrally ; abdominal segments 2-10 each with one pair of lateral lobes. 116 The Pan-Pacific Entomologist Tothill (1922) reports that the third instar larva of T. morio is similar to the second instar, and the thoracic segments have buds indicat- ing the position of the legs and the wings. The third instar larva of T. circumflexum is very different from the second instar. The head capsule and mouthparts of the third instar are greatly reduced, the body is wider, and the abdominal segments have lateral lobes. No leg or wing buds are present on the thorax. In both Spodoptera and Pseu- daletia, third instar larvae were found after the hosts had pupated. In each case, the larva was found occupying the abdominal part of the pupa. One Spodoptera pupa and one Pseudaletia pupa had no tissue left inside the pupal integument. The other Spodoptera pupa had no tissue remaining in the abdominal region, but the thoracic re- gion was still filled with tissue. All three instar larvae removed from Spodoptera were oriented with their heads pointing toward the thorax and the head of the host. Development time from oviposition to third instar is approximately 23 days in both Spodoptera and Pseudaletia. Pupa (Fig. 6). Length, 14.0-12.0 mm (x = 13.0 mm, N = 2) ; width at widest point 3.5-3.0 mm (x 3.25 mm) ; color white-pink, eyes light purple; antennal sockets not prominent; labrum and clypeus elevated, projecting outward from head; clypeus pentagonal, height 0.75 its width; labrum reduced; mandibles bidentate, dorsal tooth 1.3 length of ventral tooth, maxilla well differentiated; postmentum conspicuous, quadrate; fore and hind wings lobiform, white, forewing equal in length to hind femur, hind wing 0.75 length of hind femur; front wing lobe covers hind wing lobe; second abdominal segment of abdomen appressed to propodeum, mesonotum, and scutellum, shortening total body length; first tarsal segment of hind leg equal in length to segments 2-3 combined; fifth tarsal segment of hind leg equal in length to segments 3-4 combined; antennae, legs appressed to body. One Therion pupa was found in a Spodoptera pupa, the other was found in a Pseudaletia pupa. In each species of host, the Therion pupa was oriented with its head in the same direction as that of the host. The pupa of each host was an empty shell, with no tissue remain- ing inside. No evidence of a cocoon spun by the Therion was found. Development time from oviposition to pupa is approximately 26 days. Adults. One male T. circumflexum emerged from a Spodoptera pupa 33 days after oviposition. As an adult, this individual lived in the laboratory on a diet of honey and water for 28 days. This indicates that, at least under the laboratory conditions to which the hosts and parasitoids were exposed, development time from egg to adult is ap- proximately one month, and the entire life cycle takes approximately two months. Field studies with marked adults of T. circumflexum (Slobodchikoff, unpublished) show that individuals may be recaptured up to 30 days after they are marked. This suggests that the life-span of Vol. 50, No. 2, April 1974 117 the male reared under laboratory conditions is comparable to, or even less than, the life span of adults in the field. Literature Cited Slobodchikoff, C. N. 1971. Patterns of differentiation and evolution in the genus Therion. Ph.D. Dissertation, University of California, Berkeley. Slobodchikoff, C. N. 1973. Behavorial studies of three morphotypes of Therion circumflexum (L.) . (Hymenoptera: Ichneumonidae) . Pan- Pacific Entomol., 49: 197-206. Tothill, J. D. 1922. The natural control of the fall webworm ( Hyphantria cnnea Drury) in Canada together with an account of its several para- sites. Canada Dept. Agric. Tech. Bull. No. 3, 107 pp. 118 The Pan-Pacific Entomologist Nearctic Species of Schwenkfeldina (Diptera: Sciaridae) Wallace A. Steffan Bishop Museum, Honolulu, Hawaii 96818 Schwenkfeldina was known only from the Palaearctic and Oriental Regions except for the extension of the Holarctic species, S. tridentata (Riibsaamen) into Canada and Greenland (Tuomikoski, 1967). Some time ago Tuomikoski suggested that the North American species Brady - sia dux (Johannsen) and B. imitans (Johannsen) belong in Schwenk- feldina (pers. commun.). While revising the key to genera of North American Sciaridae, I examined specimens of B. imitans and com- pared the drawings of B. dux with those of B. imitans and agree that they are Schivenkfeldina species. Initially, Schwenkfeldina was proposed as a subgenus of Neosciara (Frey, 1942) and contained the Palaearctic species, N. (S.) carbonaria (Meigen) and the Oriental species, TV. (S.) ponderosa (Walker). In his revision of northern European Sciaridae, Frey (1948) included both Schwenkfeldina and Neosciara as subgenera of Bradysia. Tuomiko- ski (1960) elevated Schwenkfeldina to generic rank. Steffan (1973) listed S. breviantenna Steffan, S. carbonaria (Meigen), and S. Pon- derosa (Walker) from the Oriental Region. Schwenkfeldina species are characterized by the relatively short and stout antennae, wide eyebridge, setae on the posterior pronotum, the long Ri which extends beyond the base of the medial fork and frequently has ventral macrotrichia near the apex. The biology of the Nearctic species is unknown. There is con- siderable literature on the Palaearctic species and type, S. carbonaria (Meigen). Winnertz (1867) reported adults on willow blossoms dur- ing spring and summer. Beling (1886) noted that larvae live on the forest floor, preferably in spruce stands ( Abies excelsa D. C.), under the leaves or litter from dead needles. The use of the late Dr. Jean L. Laffoon’s sciarid species file is acknowledged and greatly appreciated. I also wish to thank Ms. Suzanne Keenan and Mrs. Mary Keeler for preparing the illustrations. Schwenkfeldina Frey Neosciara ( Schwenkfeldina ) Frey, 1942, p. 32. Type species Sciara carbonaria Meigen (original designation) . Bradysia {Schwenkfeldina) , Frey, 1948, p. 51. Schwenkfeldina, Tuomikoski, 1960, p. 29; Steffan, 1973, p. 469. The Pan-Pacific Entomologist 50: 118-123. April 1974 Vol. 50, No. 2, April 1974 119 Schwenkfeldina DUX (Johannsen), new combination Sciara dux Johannsen, 1912, p. 127-128 [Type: Cornell #2084] ; Leonard, 1928, p. 747; Johnson, 1930, p. 126; Fisher, 1937, p. 413. Neosciara dux, Pettey, 1918, p. 322; Johnson, 1925, p. 89. Lycoria dux, Shaw and Fisher, 1952, p. 212. Brady sia ( Brady sia ) dux. Stone and Laffoon, 1965, p. 232. Bradysia dux, Steffan, 1966, p. 35. Type locality: Wisconsin. Distribution: Michigan (Stone & Laffoon, 1965). New York (Johannsen, 1912 and Leonard, 1928). Massachusetts (Johnson, 1930). Rhode Island (Shaw and Fisher, 1952). Wisconsin (Johannsen, 1912). S. dux is very similar to S. imitans and may be conspecific. Schwenkfeldina imitans (Johannsen) new combination Sciara imitans Johannsen, 1912, p. 128 [Type: Cornell #2085]; Garret, 1925, p. 16; Fisher, 1937, p. 413. Neosciara imitans, Pettey, 1918, p. 322. Bradysia ( Bradysia ) imitans, Stone and Laffoon, 1965, p. 233. Bradysia imitans, Steffan, 1966, p. 36. Head. Interfacetal hairs abundant, extending well beyond outer curvature of facets; eyebridge 4-5 facets wide. Anterior vertex with strong setae. Antenna: flagellomeres with short necks, about % length of flagellomeres, hairs short and dense about % width; hyaline sensilla sparse; flagellomere 4 (fig. lb) about 1.5 times longer than wide. Prefrons with 7 very long and stout median setae. Clypeus bare. Palpus 3 segmented, segment 1 with 8 dorsolateral setae and broad patch of numerous dorsal hyaline sensilla, sensory pit absent. Thorax: acrostichals and dorsocentrals strong. Posterior pronotum with setae; anterior pronotum with 7 strong setae. Proepisternum with 8 strong setae. Posterior mesepimerite rectangular in shape. Legs. Forelegs: length of coxa, 0.51 mm; femur, 0.71 mm; tibia, 0.86 mm; basitarsomere 0.45 mm; fore tibia with preapical triangular patch of dense setae (fig. lc) . Tibial spurs, 1: 2: 2; apex of hind tibia with 11 large lateral setae; hind tibial spurs distinctly longer than width of tibial apex. Pretarsal claws simple. Wing: length 3.71 mm, width 1.53 mm. Vena- tion as in fig. la. R-M index 0.8; C-M index 0.8 Ri with ventral macrotrichia near apex. Abdomen: tergal and sternal setae strong and dark. Terminalia as in fig. Id; distimere with large median protuberance, dense patch of strong spines slightly distad of protuberance, and 3-4' short preapical spines. Basimere with dense patch of mesoapical setae on distinct lobe. 2 . Essentially as male with following differences: wing length, 4.29 mm; width, 1.63 mm. Vaginal furca as in fig. le. Type locality: Friday Harbor, Washington. Distribution: S. imitans was previously known only from the type locality, Friday Harbor, Washington. I have examined specimens from Washington and Cal- ifornia as listed below. California: San Bernadino Co., Mt. Home Canyon, 8-VI-1924, F. R. Cole (C.I.S.), 2$ $, 2 2 2- Washington: Mason Co., Lake Cushman, 27-VI-1917, 120 The Pan-Pacific Entomologist Fig. 1. Schwenkfeldina imitcins. A, wing, male; B, flagellomere 4, male; C, apex of foretibia, male; D, genitalia, ventral view, male; E, vaginal furca, dorsal view, female. H. G. Dyar (USNM), 1 $; San Juan Co., Friday Harbor, 29-V-1906 (Cornell, Paratype #2085.2), 1$. S. imitans is very similar to S. dux and may be conspecific; how- ever, additional specimens of S. dux need to be examined before any decision can be made. Schwenkfeldina tridentata (Riibsaamen) Sciara variabilis Boheman (not Sciara variabilis Zetterstedt, 1838) , 1865, p. 575. Sciara atrata Holmgren (not Sciara atrata Say, 1824), 1869, p. 366. Sciara tridentata Riibsaamen, 1898, p. 107 ; Coquillett, 1900, p. 392 ; , 1904, p. 6; Lundbeck, 1901, p. 312; Nielsen, 1907, p. 371; de Meijere, 1910, p. 23; Johannsen, 1912, p. 127; Sherman, 1920, p. 15; Edwards, 1922, p. 194; Vol. 50, No. 2, April 1974 121 , 1923, p. 235; , 1935, p. 532; , 1937, p. 360; Fisher, 1937, p. 413; Bertram and Lack, 1938, p. 50; McClure, 1943, p. 16; Stone and Laffoon, 1965, p. 236; Steffan, 1966, p. 51. Sciara validicornis Lundbeck, 1898, p. 243; , 1901, p. 312. Sciara holmgreni Jacobson (not Sciara holmgreni Riibsaamen, 1894), 1898, p. 34. Lycoria ( Neosciara ) meigeni, Lengersdorf (not Sciara meigeni Riibsaamen, 1894), 1930, p. 40; Edwards, 1935, p. 532. Rhynchosciara laguncularis Lengersdorf, 1930, p. 52; Tuomikoski, 1967, p. 46. Neosciara tridentata, Pettey, 1918, p. 321; Cole, 1921, p. 170; Malloch, 1923, p. 180. Lycoriella (Hemineurina) atratci, Frey, 1942, p. 36. Brady sia ( Neosciara ) incisiforceps Frey, 1948, p. 53. Schwenkfeldina tridentata, Tuomikoski, 1966, p. 137 ; , 1967, p. 45. Type locality: Greenland. Distribution: Alaska, Pribilof Islands (Cole, 1921, “specimen .... near tri- dentata Riibs.”). Arctic: Bear Island (Edwards, 1935); Jan Mayen Island (Edwards, 1923); Spitzbergen (Edwards, 1922; Tuomikoski, 1967). British Columbia (Coquillett, 1900) ; Greenland (Riibsaamen, 1898; Lundbeck, 1898, as Sciara validicornus) . Manitoba (McClure, 1943). S. tridentata is apparently a Holarctic species primarily found in the Nearctic region with extension of its range to arctic islands east and north of Greenland. Literature Cited Beling, T. 1886. Beitrag zur Metamorphose der Zweifliigler in Gattung Sciara Meig. Wiener Entomol. Z., 5: 93-96. Bertram, G. C. L. and D. Lack. 1938. Notes on the animal ecology of Bear Island. J. Anim. Ecol., 7: 27-52. Boheman, C. H. 1895. Spetsbergens Insekt-fauna. Ofver. K. Svenska. Vetensk. Akad. Forhandl., 22: 563-577. Cole, F. R. 1921. Diptera from the Pribilof Islands, Alaska. Proc. Calif. Acad. Sci., 11: 169-177. Coquillett, D. W. 1900. Papers from the Harriman Alaska Expedition. IX. Entomological Results (3) : Diptera. Proc. Wash. Acad. Sci., 2: 389- 464. (Also published separately as Harriman Alaska Expedition, Vol. 9, P. 2, 78 pp., in New York, 1904.) Edwards, F. W. 1922. Results of the Oxford University Expedition to Spits- bergen, 1921. No. 14. Diptera Nematocera. Ann. Mag. Natur. Hist., 10: 193-215. Edwards, F. W. 1923. On the nematocerous Diptera of Jan Mayen Island. Ann. Mag. Natur. Hist., 11: 235-240. Edwards, F. W. 1935. Diptera from Bear Island. Ann. Mag. Natur. Hist., 15: 531-543. Edwards, F. W. 1937. Diptera, Oxford Expedition to North-East Land. Ann. Mag. Natur. Hist., 20: 360-362. 122 The Pan-Pacific Entomologist Fisher, E. G. 1937. A comparative study of the male terminalia of the Mycetophilidae of Nearctic America. Unpublished Ph.D. Thesis in Cor- nell University Library. 430 pp. Frey, R. 1942. Entwurf einer neuen Klassification der Miickenfamilie Sciaridae (Lycoriidae). Notul. Entomol., 22: 5^14. Frey, R. 1948. Entwurf einer neuen Klassification der Miickenfamilie Sciaridae (Lycoriidae) . II. Die nordeuroparschen Arten. Notul. Entomol., 27: 33-92. Garrett, C. B. D. 1925. Seventy new Diptera. Cranbrook Courier Print. 16 pp. Holmgren, A. E. 1869. Bidrag til kannedomen om Beeren Eilands och Spits- bergen Insekt-fauna. K. Svenska. Vetensk. Akad. HandL, (n.s.) 8: 3-55. Jacorson, G. 1898. Compte rendu de l’expedition, envoyee par l’Academie Im- periale des Sciences a Novaia Zemliaen ete 1896. IV. [Zoological ex- plorations on Novaya Zemlya. Insects of Novaya Zemlya]. Imp. Akad. Nauk St. Petersburg, Zap., Fiz.-Mat. Otd., 8: 171-244. Johannsen, 0. A. 1912. The fungus gnats of North America. The Myceto- philidae of North America. Part IV. Bull. Maine Agr. Exper. Sta., 196: 249-328. Johnson, C. W. 1925. Fauna of New England. 15. List of the Diptera or two-winged flies. Occas. Pap. Boston Soc. Natur. Hist., 7: 1-326. Johnson, C. W. 1930. A list of the insect fauna of Nantucket, Massachusetts. Publ. Nantucket Maria Mitchell Ass., 3: 174. Lengersdorf, F. 1930. Lycoriidae (Sciaridae). In E. Lindner Die Fliegen der palaearktischen Region. 43: 33-71. Leonard, M. D. 1928. A list of the insects of New York with a list of the spiders and certain allied groups. Mem. N. Y. Agr. Exper. Sta., 101: 1 - 1121 . Lundbeck, W. 1898. Diptera groenlandica. Vidensk. Medd. Naturhist. Foren. Kj0benhavn 1898: 236-314. Lundbeck, W. 1901. Diptera groenlandica. II. Vidensk. Medd. Naturhist. Foren. Kj0benhavn 1900: 281-316. McClure, H. E. 1943. Aspection in the biotic communities of the Churchill area, Manitoba. Ecol. Monogr., 13: 1-35. Malloch, J. R. 1923. Insects, arachnids, and chilopods of the Pribilof Islands, Alaska. Diptera (except Tipulidae, Rhyphidae and Calliphoridae) . N. Amer. Fauna, 46: 170-227. Meijere, J. C. H. de 1910. Die Dipteren der arktischen Inseln. Fauna Arctica, 5: 13-72. Nielsen, J. C. 1907. The insects of East-Greenland. Medd. Gr0nland, 29: 363- 414. Pettey, F. W. 1918. A revision of the genus Sciara of the family Myceto- philidae (Diptera). Ann. Entomol. Soc. Amer., 11: 319-343. Rubsaamen, E. H. 1898. Zoologische Ergebnisse der von der Gesellschaft fur Erdkunde zu Berlin unter Leitung Dr. von Drygalski’s ansgesandten Gronlands-expedition. VIII. Gronlandische Mycetophiliden, Sciariden, Cecidomyiden, Psylliden, Aphiden und Galium. In R. Leuckart & C. Chun Eds. Bibl. Zook, 20: 101-119. Vol. 50, No. 2, April 1974 123 Shaw, F. R. and E. G. Fisher. 1952. Guide to the insects of Connecticut. Part VI. The Diptera or true flies of Connecticut. Fifth Fascicle: Midges and gnats [part]. Family Fungivoridae (= Mycetophilidae) . Bull. Conn. State Geol. Natur. Hist. Surv., 80: 177-250. Sherman, R. S. 1920. Notes on the Mycetophilidae of British Columbia. Proc. Entomol. Soc. Brit. Columbia, 14: 12-15. Steffan, W. A. 1966. A generic revision of the family Sciaridae (Diptera) of America North of Mexico. Univ. Calif. Publ. Entomol., 44: 1-77. Steffan, W. A. 1973. Family Scaridae. In M. Delfinado and D. E. Hardy A Catalog of the Diptera of the Oriental Region. Vol. 1: 464-476. Univ. Hawaii Press. Honolulu. Stone, A. and J. J. Laffoon. 1965. Family Sciaridae (Lycoriidae) . In Stone, et al. A Catalog of the Diptera of America North of Mexico. Agric. Hand- book No. 271. pp. 229-236. Tuomikoski, R. 1960. Zur Kenntis der Sciariden (Dipt.) Finnlands. Ann. Zool. Soc. ‘Vanamo,’ 21: 1-164. Tuomikoski, R. 1966. Die Stellung von Sciara tridentata Rubs. (Dipt., Sciari- dae). Ann. Entomol. Fenn., 32: 137. Tuomikoski, R. 1967. 6. Mycetophilidae and Sciaridae from Spitsbergen, col- lected by Dr. J. Kaisila in 1965. Ann. Entomol. Fenn., 33: 43-51. Winnertz, J. 1867. Beitrag zu einer monographic der Sciarinen. Zool. Bot. Gessell. Wien, 1867: 1-187. 124 The Pan-Pacific Entomologist Observations On Megasoma with Behavioral Notes on Some Lamellicorn Beetles Associated With Sand Dunes (Coleoptera: Scarabaeidae, Lucanidae) Alan R. Hardy and Fred G. Andrews Insect Taxonomy Laboratory California Department of Food and Agriculture Sacramento, California 95814 The description of Megasoma sleeperi Hardy (1972) was based upon four female specimens collected in Joshua Tree National Monument, California. Since the original description, the senior author has examined 24 additional specimens of both sexes of this species, as well as the fragments of numerous additional individuals. The male of Megasoma sleeperi is described below, additional distribution records and a syn- onymy in Megasoma are presented, as well as observations of unusual behavior of M. sleeperi and Pseudolucanus mazama (LeC). Megasoma punctulatus Cartwright The following specimens have been examined: ARIZONA: Pima Co.; Madera Canyon VIII-19-1949 (1 2, Los Angeles Co. Museum). Santa Cruz Co., Pena Blanca, VII-29-1972 (2 2 2 ) ; Madera Canyon, VIII- 22-1971 (1 $,1 2) (Kane and Langston, Colins.) Megasoma elephas occidentalis Bolivar y Pieltain et al. Megasoma elephas occidentalis Bolivar y Pieltain et al. 1963:187 Megasoma mexicanus Fischer 1968:139, new synonymy From an examination of the illustration accompanying the original description of M. mexicanus , and from the geographical distributional data (type of M. mexicanus from Colima, paratypes of M. elephas oc- cidentalis from Colima), the above synonymy is apparent. Megasoma sleeperi Hardy (Fig. 6.) Male: Length 24.8 to 30.5 mm; width 12.5 to 15 mm. HEAD: Clypeal plane with close, medium punctures, becoming larger pits posteriorly. Vertex polished, smooth with few scattered medium punctures. Cephalic horn short, bifurcate, curved posteriorly apically. Prothorax coarsely punctured, punctures becoming laterally confluent, appearing rugose ; prothorax occasionally lack- ing posterior marginal bead medially, marginal bead present laterally and an- teriorly; lateral margin occasionally poorly defined anteriorly, prothorax laterally smoothly rounded from dorsal to ventral surface, without lateral carina; anterior The Pan-Pacific Entomologist 50: 124-128. April 1974 Vol. 50, No. 2, April 1974 125 Figs. 1-5. Fig. 1; Area 4.8 km NW Glarnis, Imperial Co., California, showing advancing dunes (looking S., March). Fig. 2; Coral Pink Sand Dunes area, Kane Co., Utah, showing advancing dunes (looking S., July). Fig. 3; Tracks of adult Pseudolucanus mazama on sand. Figs. 4-5; Adult P. mazama burrowing into sand at base of dead Juniper. prothoracic angles acute, not spinaform, not produced anteriorly; near eyes. Elytra with scattered fine punctures interspersed with close, very fine punctures, polished, shiny; lateral marginal bead, fading apically. Pygidium evenly convex, closely punctured with uniform, fine punctures, which are seldom confluent; punctures separated by less than their diameter; with recumbent, pale hair; ventral marginal bead present. Six visible abdominal sternites; membranous area visible between ultimate and penultimate segments; ultimate segment emarginate apically. Body ventrally clothed with long, white hairs; appendages with long, reddish hairs. The male of M. sleeperi most closely resembles the male of M. cedrosa Hardy, from which it can be easily distinguished by the absence of long dorsal pubescence. M. sleeperi is glabrous dorsally. The following specimens of Megasoma sleeperi have been examined: CAL- IFORNIA: Imperial Co.; 4.8 km (3 mi.) NW Glarnis, IX-16-1972, A. R. Hardy, M. S. Wasbauer, Collrs. (10 2 2, 3 $ $ ) ; same locale, various dates, numerous dead examples and fragments. Imperial Valley, XI-29-1937 (1 $, U.C. Berkeley Colin.). Palm Desert-Deep Canyon, VII-30-1970, E. Giesbert Collr. (1 2, Giesbert Colin.). The authors have recently made several investigations of insects in the sand dune habitats in the Southwest. On a number of occasions, insects not usually associated with sand dunes have been observed to utilize vegetation killed by the drifting sand. 126 The Pan-Pacific Entomologist Figs. 6-11. Fig. 6; Adult male Megasoma sleeperi as collected on sand. Fig. 7; Dead M. sleeperi being uncovered by wind action in trough of dune (see text). Figs. 8-11; Pseudolucanus mazama mating activities (see text). In March and April 1972, a number of dead, mummified adult Mega- soma sleeperi (Fig. 7), were collected in the troughs of the dunes, near the leading edge. Many of the specimens were fragmentary and badly broken, but a few were intact. Some individuals, which were being uncovered as the sand moved, were in a vertical position, as if they had died while digging toward the surface. The area where the Megasoma were observed (about 4.8 km NW Vol. 50, No. 2, April 1974 127 Glamis, Imperial Co., California) is an interface area, where the sand meets an area of hardpan. The hardpan is laced with washes that drain the Chocolate Mountains to the Northeast. During flash flood- ing, water moves southwest towards the shores of fossil Lake Cahuilla, (now the Salton Sea), where it encounters the sand hills. The runoff is retained, creating local pockets of dense desert vegetation (Fig. 1), which support large populations of otherwise scarce organisms. These vegetated areas are slowly buried and the plants killed as the sand moves eastward at from three to five or more feet per year. Apparently, Megasoma oviposts in the dead or dying trees, probably utilizing Palo Verde ( Cercidium floridum Benth.), Mesquite, ( Prosopis sp.), or both, and develops and pupates in the buried logs. Evidently, after emergence, the adults attempt to burrow to the surface. Many are unsuccessful in reaching the surface (which may be through as much as 30 feet of sand) , die, and are mummified. On July 6, 1972, the junior author observed similar events involving Pseudolucanus mazama (LeC.) (Coleoptera, Lucanidae) at the Coral Pink Sand dunes near Kanab, Kane County, Utah. The area is a Pinyon- Juniper association in a valley bordered by red sandstone mountains. First indications of beetle activity were noted at the north end of the dunes, about 30 minutes before dark. In this area, Juniper Trees ( Juniperus sp.) were being covered by drifting dunes (Fig. 2), which had killed the smaller trees. Near the bases of the trees and stumps, which projected from the sand, were found parts of adult beetles which were still exhibiting signs of life after a recent attack by an unidentified predator, probably vertebrate. Several beetles were seen emerging from the fine unconsolidated sand in the vicinity of projecting stumps and the bases of live trees. By nightfall, more than 100 beetles were observed on the sand in an area about 30 meters by 65 meters. First observations were of apparent random wandering, followed by copulation when males and females met. Mated pairs were observed struggling and rolling on the sand (Fig. 8), or with the male mounted while the female continued to move across the dune (Fig. 9). Males outnumbered females and often con- gregations of six to eight beetles were found, with three to four males clustered about a single female (Figs. 10, 11). It appeared that only the first male mated with the female. A preponderance of males also occurs in Pseudolucanus placidus Say (Mathieu, 1969:1059). After approximately one hour, the beetles began to burrow into the sand, and within another 15 minutes all except a few wandering males were gone. The beetles usually burrowed into the dunes within 25 cm of 128 The Pan-Pacific Entomologist a stump or tree (Figs. 4, 5), but several entered the sand at distances of from four to five meters from the nearest exposed juniper. No at- tempt was made to determine whether there was dead vegetation be- neath the sand at these remote points of entry. At the peak of evening activity, living Junipers within 100 meters of the breeding congregations, but away from the dunes, were inspected for beetles with negative results. This may indicate that either dead or dying junipers or sand or both are important to the life history of P. mazama. Mathieu (1969:1059) indicates that females of P. placidus use “a typical sandy-soil habitat” in breeding. The following morning no evidence of beetles was observed except the tracks of entering and leaving adults (Fig. 3). Several excavations at the bases of stumps where Fucanids were seen to burrow the previous night resulted in the location of a few adults, all at least 45 cm. below the surface. Fiterature Cited Bolivar y Pieltain, C., L. Jiminez-Asua and A. Martinez. 1963. Notas sobre Dynastinae neotropicales con especial referenda a especies Mexicanas (Col., Scarab.). Ciencia 22(6) : 181-190, 9 figs, 1 pi. Fischer, Heinz. 1968. Zwei neue Megasoma-arten aus Amerika. Natur-forsch. Ges. Augsburg, 22:137-142. Hardy, Alan R. 1972. A brief revision of the North and Central American species of Megasoma (Coleoptera: Scarabaeidae) . Can. Entomol. 104: 765-777. Mathieu, Juan M. 1969. Mating behavior of five species of Lucanidae ( Coleop- tera: Insecta) . Can. Entomol. 101:1054-1062. Vol. 50, No. 2, April 1974 129 Descriptions and Records of Clytini from Mexico and Central America (Coleoptera: Cerambycidae) John A. Chemsak and E. G. Linsley University of California, Berkeley There have been relatively few Clytini described from Mexico and Central America since the publications of Bates in 1885 and 1892. Casey (1912), Linsley (1935), and Chemsak and Linsley (1965) added a few new species and genera, and Fuchs (1963) described a species of Neoclytus from Colombia subsequently found in Panama. Although there are additional undescribed species at hand, only a few are de- scribed at this time in order to make the names available for other projects. The remainder will be made known when the generic limits of the Neotropical Clytini are better defined. Since many species have previously been known only from single or a few localities, we are presenting additional distribution records for a number of these. We wish to thank the National Science Foundation for support through Grant No. GB-4944X and the following persons and institu- tions for the loan of material: The American Museum of Natural His- tory; Canadian National Collection; Cornell University; H. F. Howden; Kansas State University; Kansas University; and Ohio State University. Special thanks are due Celeste Green for preparing the illustrations. Placoclytus, new genus Head with front short, not carinate, median line deep; palpi with apical seg- ments dilated; antennae stout, not spinose, scape robust, cylindrical, slightly shorter than third segment, third longer than fourth, fifth longer than third. Pronotum rounded, as broad as or broader than long; disk with a well defined flattened area which is distinctly delimited; prosternum deeply to shallowly impressed, intercoxal process narrow, not expanded at apex, coxal cavities wide open behind; mesosternum with intercoxal process gradually declivous in front, not protuberant; episternum of metathorax very gradually narrowed behind. Elytra slightly tapering posteriorly; pubescence depressed, forming bands of condensed yellowish hairs; apices rounded to truncate, unarmed. Legs slender, rather elongate; hind tibiae rather short, not compressed; hind tarsi with first segment much longer than two following together. Type species — Neoclytus championi Bates. This genus differs from others presently known in the Clytini by the flattened, dish-like disk of the pronotum. In addition to the type of the The Pan-Pacific Entomologist 50 : 129 - 138 . April 1974 130 The Pan-Pacific Entomologist genus, championi, two other species may be assigned to the genus: distortus Chevrolat, and virgulatus, new species. Placoclytus championi (Bates), new combination (Fig. 2) Neoclytus ? championi Bates, 1885, Biologia Centrali- Americana, Coleoptera, 5:302. Color black, appendages reddish. Pronotum densely clothed with short, yellow- ish, appressed pubescence; discal flat area moderately coarsely, separately punctate. Elytra with undulating fasciae yellowish; apices rounded. Body beneath with patches of dense appressed, yellowish pubescence at sides. Length, 13 mm. Type locality: Totonicapam, Guatemala, 10,000 feet. In addition to the type, we have seen one other specimen of this species from 6 miles E. San Cristobal de las Casas, Chiapas, Mexico, 2 June, 1969, on Pine (H. F. Howden). Placoclytus distortus (Chevrolat), new combination (Fig. 3) Clytus ( Plagithmisus ?) distortus, Chevrolat, 1860, Ann. Soc. Entomol. France, (3) 8:500. Neoclytus? distortus, Bates, 1880, Biologia Centrali-Americana, Coleoptera, 5:58; ibid., 1885:302. This is a large (25 mm.) robust species having the flattened pronotal disk, stout antennae, and non-costate, unarmed elytra. The ground color is black, legs reddish, and the fine transverse fasciae of the elytra are yellow. Bates (1885) noted that this species should not remain in the genus Neoclytus and characterized it with championi. Placoclytus virgulatus, new species (Fig. 1) Male: Form moderate sized, tapering slightly; integument black, parts of legs and apices of antennae reddish; pubescence rather sparse, yellowish, black on elytra except for yellow fasciae. Head with front finely punctate except for glabrous median triangle; vertex irregularly punctate; pubescence sparse, ap- pressed; antennae extending about 2 segments beyond elytra, segments from fifth densely clothed with dark appressed pubescence. Pronotum as long as broad; disk with flattened area converging toward apex, punctures dense, irregular; pubescence moderately dense, golden, appressed; prosternum rather deeply impressed, sparsely -> Figs. 1-6. Typical elytral patterns of: 1, Placoclytus virgulatus Chemsak and Linsley; 2, P. championi (Bates) ; 3, P. distortus (Chevrolat) ; Vol. 50, No. 2, April 1974 131 4, Neoclytus personatus Chemsak and Linsley; 5, N. columbianus Fuchs; 6, Xylotrechus durangoensis Chemsak and Linsley. 132 The Pan-Pacific Entomologist pubescent; meso- and metasternum minutely punctate, moderately densely pubescent, metepisternum with a patch of appressed pubescence at posterior end. Elytra more than twice as long as broad; subbasal fasciae transverse, post basal w-shaped, median arcuate, preapical rounded at suture, angulate away from suture, sutural line pubescent for most of its length; pubescence on dark areas black, short, appressed; punctures vague, minute; apices subtruncate. Legs slender; posterior femora not surpassing elytral apices; pubescence pale, sparse, appressed. Abdomen minutely punctate, rather sparsely pubescent; apex of last sternite shallowly emarginate. Length, 13 mm. Holotype male (California Academy of Sciences) from 34 miles E. Morelia, Michoacan, Mexico, 2 July, 1963 (W. A. Foster). The pattern of the fasciae of the elytra, shape of the pronotum, and longer antennae will separate this species from the two others in the genus. Neoclytus personatus, new species (Fig. 4) Male: Form moderate sized, cylindrical; integument dark brown, antennae and tibiae paler; pubescence thick, appressed. Head minutely punctate on front, a little more coarsely on vertex; pubescence fine, yellowish, appressed; antennae extending to about basal % of elytra, segments from fifth slightly thickened, scape slender, longer than third segment, third longer than fourth. Pronotum as long as broad, sides broadly rounded; median line prominently elevated, rather finely asperate, each side with slightly elevated, sparsely asperate lines; punctures vague on middle, dense and deep at sides; pubescence yellowish, appressed, dense, sparser on disk ; prosternum shallowly impressed, moderately clothed with yellowish recumbent pubescence and long white erect hairs; meso- and metasternum finely, shallowly punctate, moderately pubescent, episterna with bands of condensed yellowish pubescence at posterior halves. Elytra more than 2% times as long as broad, very slightly tapering posteriorly; base with two large obtuse gibbosities behind scutellum; punctures very fine, dense; pubescence very short, appressed on dark areas, yellowish pubescent fasciae consisting of two oblique postbasal bands extending away from near suture and vaguely curving up toward lateral margin, a sutural premedian pair expanding posteriorly away from suture, apex very broadly clothed with very thick appressed pubescence which is darker at posterior end, each side directed obliquely back to lateral margin ; apices subtruncate, outer angles dentate. Legs elongate ; middle and hind femora bidentate at apices, front femora with a row of small acute teeth along inside margins, hind pair exceeding elytral apices by almost % of their length; hind tibiae elongate, compressed, strongly concave along inner side. Abdomen minutely, densely punctate, densely clothed with appressed pubescence which becomes sparser toward apex; fifth sternite rounded at apex, lightly emarginate at middle. Length 11 mm. Female: Form more robust; antennae with outer segments more thickened; pronotum with disk inflated, middle with a broad row of transverse asperites, small asperites dispersed over sides. Abdomen with last sternite narrowly rounded at apex. Length, 14 mm. Vol. 50, No. 2, April 1974 133 Holotype male (Kansas State University) from Monteverde, Costa Rica, 10°29' N., 84°50' W., 1400 M., 23 February 1963, in malaise trap (C. W. and M. E. Rettenmeyer) . One female paratype, same data, 16 February 1963; 3 male paratypes, same data, 17-21 February 1963 (R. D. Akre) . The elytral pattern of this species somewhat resembles that of N. justini Chevrolat from Colombia. The overall aspect of the pattern of persojiatus is that of a primitive mask especially enhanced by the darker pubescence at the extreme apex. The strong basal gibbosities of the elytra, the elytral pattern, and toothed front femora make this species distinctive. Neoclytus columbianus Fuchs (Fig. 5) Neoclytus columbianus Fuchs, 1963, Senckenberg Biol., 44: 451, pi. 1. This species was described from Colombia with no other indication of locality. We have a male and female from Santa Fe, Darien Province, Panama, 6 June, 1967 (D. M. DeLong and C. A. Triplehorn) and one male from Barro Colorado Island, Gatun Lake, Canal Zone, 2 April, 1924 (J. C. Bradley). In addition to the distinctive elytral pattern, N. columbianus is characterized by the rather prominent spines at the apices of the elytra. Neoclytus irroratus (LeConte) Clytus irroratus LeConte, 1858, Jour. Acad. Nat. Sci., Philadelphia, (2) 4:26. Neoclytus irroratus, Leng, 1887, Ent. Amer., 3:6; Hopping, 1932, Ann. Entomol. Soc. Amer., 25: 550, pi. 4, fig. 12; Linsley, 1964, Univ. Calif. Pubis. Entomol., 22: 139, fig. 42. Rhopalopachys irroratus, Horn, 1894, Proc. Calif. Acad. Sci., (2) 4: 339. Clytus ( Rhopalopachys ) morosus Chevrolat, 1860, Ann. Soc. Entomol. France, (3)8: 501. Rhopalopachys morosus, Bates, 1880, Biologia Centrali-Americana, Coleoptera, 5: 58. The following are additions to the previously listed Mexican records for this species: 1 female, Samalayuca, Chihuahua, 24 June, 1947 (D. Rockefeller Exp., Michener) ; 1 male, Sta. Barbara, Chihuahua, 6 May, 1947 (C. M. Bradt) ; 1 female, Monterrey, Nuevo Leon, 6 May, 1950 (Hernandez) ; 1 male, V. de Santiago, Nuevo Leon (A. Salazar) ; 3 females, Jalapa, Veracruz (W. Schauss) ; 1 male, 3 miles NW Cuernavaca, Morelos, 24 May, 1959 (H. E. Evans). Neoclytus smithi Bates Neoclytus smithi Bates, 1892, Trans. Entomol. Soc., London, 1892: 164, pi. 6, fig. 9. 134 The Pan-Pacific Entomologist Known previously from a unique specimen from Rincon, Guerrero, Mexico. We have 1 female from Jesus Maria, Nayarit, 27 July, 1955 (B. Malkin). Neoclytus justini Chevrolat Neoclytus justini Chevrolat, 1861, Ann. Soc. Entomol. France, (4) 1: 382. Originally described from Colombia, 1 female is at hand from Santa Fe, Darien Province, Panama, 29 May, 1967 (D. M. DeLong and C. A. Triplehorn). Ocliraethes nigrescens, new species Female: Form moderately robust; ground color black, elytra vaguely tinted with brown; pubescence fine, pale, moderately long, appressed, with scattered long erect pale setae on the head, pronotum, base of elytra and ventral surface. Head coarsely confluently punctured above, more finely on front, lower face and mouthparts shining, punctation fine; antennae not attaining middle of elytra, second segment longer than broad, fourth segment shorter than third or fifth, fifth segment longest. Pronotum broader than long; sides rounded, dorsal sur- face coarsely, contiguously and somewhat confluently punctate, sides finely densely punctate; pubescence thin, not obscuring surface; pro-, meso-, and metasterna finely punctate; scutellum very finely punctate, thinly pubescent. Elytra more than twice as long as basal width ; contiguously and somewhat confluently punctate basally, very finely punctate apically; apices obliquely truncate. Legs slender, finely punctate, thinly pubescent, posterior tarsi more than half as long as tibiae. Abdomen finely densely punctate, thinly pubescent except for a band along pos- terior margins of first four sternites. Length 13 mm. Holotype female (American Museum of Natural History) from Palos Colorados, Durango, Mexico, 8000 ft., 5 August, 1947, David Rockefeller Expedition (Gertsch) and 15 paratypes, all female, same locality and date, four collected by Michener, four by Cazier, three each by Schrammel and Gertsch and one by Spieth. Two additional paratypes, also female, are from near Otinapa, Durango, Mexico, 13 August, 1953, collected by C. and P. Vaurie. This species, judging from the females in the type series, is extremely variable. The pubescence varies from grayish white to bright golden yellow and the ground color of the elytra from black tinted lightly with brown to uniformly reddish-brown to reddish-brown broadly clouded with dark brown basally in the past-scutellar area along the suture, trans- versely behind the middle, and more narrowly and obliquely subapically and apically. The antennae may extend over the basal one-fourth of the elytra or reach nearly to the middle. However, the body length ranges only from 11 to 13 mm. in the 18 examples at hand. The species does not appear to be closely related to other described Ochraethes, and its general appearance, although variable, is quite distinctive. Vol. 50, No. 2, April 1974 135 Trichoxys longipes, new species (Fig. 7) Male: Form moderately robust, tapering; ground color black, antennal flagella, tibiae, tarsi and elytra centrally and posteriorly rufotestaceous. Head moderately finely, densely somewhat confluently punctate on vertex, frons and mandibles, pubescence white and depressed on lower face, mandibles and underside with long erect pale cilia intermixed, eyes encircled with dense yellow pubescence which extends between the upper lobes and between the lower lobes as a yellow band, pubescence of vertex between eyes fine, appressed, brown with an inter- mixture of long erect setae; antennae slender, extending a little beyond middle of elytra, segments finely punctate, thinly pubescent, ciliate at apices, fourth segment barely shorter than third, distinctly shorter than fifth, eleventh segment a little longer than tenth. Pronotum wider than long, sides rounded, base some- what tubular, surface finely, densely, contiguously punctate, clothed with fine appressed brown pubescence, with scattered long erect pale setae, anterior and basal margins lined with dense pale yellow pubescence; scutellum margined with yellow pubescence; prosternum clothed with white pubescence, thinner at sides; mesosternum clothed with white pubescence, a patch of dense yellow pubescence on each side, intercoxal process prominent, polished, elevated above the coxae, vertical in front ; metasternum thinly clothed with white pubescence, a patch of dense yellow pubescence on each side on anterior margin behind mesocoxae and a patch along posterior margin on each side in front of metacoxae; metepisternum densely yellow pubescent ; metacoxae with a dense patch of white pubescence laterally. Elytra two and one half times as long as basal width, narrowing posteriorly, suture depressed, apices shallowly emarginate; surface very finely punctate, ground color black over basal one-third except suture, rufotestaceous basally along suture and most of apical two-thirds except for an elongate-oval oblique black area near middle and a brownish-black cloud over apices; pubescence brown except for a narrow U-shaped, yellow-pubescent band running from suture near base to lateral margin, a yellow patch in humeral cavity, a dense yellow band surrounding oblique median dark areas, a thin yellow band along suture from basal one-fourth to apical one-third and a dense oblique white patch of pubescence extending from the suture toward the lateral margins narrowing laterally and posteriorly. Legs slender; posterior femora extending well beyond apices of elytra; posterior tarsi slightly more than half as long as tibiae. Abdomen thinly clothed with white pubescence, with a dense yellow triangular patch of yellow pubescence at sides of first two stemites; fifth sternite emarginate. Length 12 mm. Female: Antennae extending over basal one-fourth of elytra, outer segments abbreviated, thickened; fifth abdominal sternite entire. Length 13.5 mm. Holotype male, allotype (California Academy of Sciences) and two male para- types from Guelatao, Oaxaca, Mexico, elev. 1800 m., 15 November, 1964. This species differs from all others presently assigned to Trichoxys in the long hind legs with the femora surpassing the elytral apices and the unique pattern of elytral coloration and pubescence (Fig. 7). It is not closely related to any other described form. 136 The Pan-Pacific Entomologist Fig. 7. Trichoxys longipes Chemsak and Linsley, Vol. 50, No. 2, April 1974 137 Xylotrechus durangoensis, new species (Fig. 6) Female: Form moderate sized, cylindrical; integument dark reddish brown; pubescence rather sparse, white, condensed into patches on pronotum and under- side, elytra with sparsely pubescent white fasciae. Head with indistinct narrow, flattened frontal carinae on each side of median line; vertex confluently, rugosely punctate, concave behind antennal tubercles; palpi with apical segments not dilated; pubescence rather sparse, short and appressed and long and erect, long hairs more numerous below eyes and under mouth ; antennae short, attaining about basal % of elytra, basal segments with a few white appressed hairs toward base. Pronotum wider than long, sides subangulate; disk impressed on each side of middle, elevated, covered with short, irregular, transverse carinae, remainder of surface asperate; white appressed pubescence forming indistinct lines along apex, base and obliquely up from base around disk to middle, apex with 2 indistinct short patches on each side of middle and middle broadly suffused; prosternum shining, impressed at apex, transversely rugulose, sparsely pubescent; meso- and metasternum subopaque at sides, sparsely punctate at middle, metepi- sternum with a condensed patch of white pubescence posteriorly, appressed pubescence suffused over middle. Elytra less than 2% times as long as broad; pale areas connecting and encircling round and oblique areas near suture; punctures very fine, dense, subopaque; apices obliquely truncate, outer angles obtusely dentate; scutellum densely white pubscent. Legs rather short; femora finely rugulose, lacking white appressed pubescence on clavate portions, posterior pair not attaining elytral apices. Abdomen minutely, sparsely punctate, moderately densely clothed with white appressed pubescence; last sternite rounded at apex. Length, 17 mm. Holotype female (Canadian National Collection) from 24 miles W. la Ciudad, Durango, Mexico, 2 July 1964 (H. F. Howden). The pubescent pattern of this species is unlike that of any other North American Xylotrechus. The indistinct narrow frontal carinae of the head also make it very distinctive. Xylotrechus quercus Schaeffer Xylotrechus quercus Schaeffer, 1905, Bull. Brooklyn Inst. Arts Sci., 1 (7): 163; Hopping, 1932, Ann. Entomol. Soc. Amer., 25: 544, pi. 4, fig. 1; Linsley, Knull, and Statham, 1961, Amer. Mus, Nov., 2050: 19; Linsley, 1964, Univ. Calif. Pubis. Entomol., 22: 108. One female, Arroyo del Nido, 35 miles SW Gallego, Chihuahua, Mexico, 23 June, 1957 (W. C. Russell) ; 1 male, 1 female, 8 miles W. El Palmito, Sinaloa, Mexico, 29 July, 1964, on Quercus (J. A. Powell, H. F. Howden) ; 1 female, 8 miles W. El Palmito, July 1972, reared ex. Quercus (J. A. Chemsak) . Xylotrechus sagittatus subsp. Three specimens (1 male, 2 females) are at hand from 6 miles NE El Salto, Durango, Mexico, 10 August, 1947 (D. Rockefeller Exp., Spieth, Cazier, Gertsch) . 138 The Pan-Pacific Entomologist More material is necessary before this population can be linked to X. sagittatus chiricahuae Chemsak from southeastern Arizona. Xylotrechus SARTORII Chevrolat Clytus ( Xylotrechus ) Sartorii Chevrolat, 1860, Ann. Soc. Entomol. France, (3)8:492, pi. 9, fig. 7; Bates, 1880, Biologia Centrali-Americana, Coleoptera, 5:58. This species was originally described from “Mirador,” Veracruz and subsequently recorded from Paso del Macho, Veracruz by Bates. We have a series of 7 specimens (3 males, 4 females) from Chipinque Mesa, 5400 ft., nr. Monterrey, Nuevo Leon, Mexico, 22 and 30 July, 1963, on Quercus (H. and A. Howden) . Mecometopus hogei Bates Mecometopus hogei Bates, 1880, Biologia Centrali-Americana, Coleoptera, 5:56. Originally described from Paso del Macho, Veracruz. New records from Mexico: 2 males, 2 females, Tamazunchale, San Luis Potosi, 13 July, 1956 (D. H. Janzen) ; 1 male, 20 miles E. Xicotepec, Puebla, 23 June, 1964 (C. D. Johnson). Clytopsis dimidiaticornis (Chevrolat) Clytus ( Clytus ) dimidiaticornis Chevrolat, 1860, Ann. Soc. Entomol. France, (3)8:487. Clytus dimidiaticornis, Bates, 1880, Biologia Centrali-Americana, Coleoptera, 5:54, ibid., 1885:299. Clytopsis dimidiaticornis, Casey, 1912, Memoirs on the Coleoptera, 3:73. 2 females, Pedregal, Distrito Federal, 30 September, 1947 (J. Hendricks) ; 1 male, 4 miles E. Toluca, Mexico, 17 August, 1954 (Univ. Kansas Mex. Expedition) : 1 male, Tlalpam, Mexico, 4 October, 1942 (W. F. Foshag) ; 1 male, 1 female, 11 miles SE San Juan del Rio, Queretero, 21 August, 1965 (C. Slobodchikoff) ; 1 male, 4 miles SW Cuidad Mendoza, Veracruz, 16 August, 1962 (H. E. Milliron) ; 1 male, nr. Espiritu Santo, Rio Verde, Chihuahua, 8 August, 1952 (J. D. Lattin) ; 1 male, Tzintzuntzan, Michoacan, 8 August, 1954 (Ryckman, Spencer, Christian- son) ; 1 female, 10 miles NW Tamazulapan, Oaxaca, 22 August, 1959 (Menke, Stange) ; 1 male, 2 females, Oaxaca, August, 1920. Literature Cited Bates, H. W. 1879-1886. Biologia Centrali-Americana, Coleoptera, Longicornia, 5: 1-436. Bates, H. W. 1892. Additions to the Longicornia of Mexico and Central America, with remarks on some of the previously recorded species. Trans. Entomol. Soc. London, 1892: 143-183. Casey, T. L. 1912. Memoirs on the Coleoptera, 3: 1-286. Chemsak, J. A. and E. G. Linsley. 1965. New genera and species of North American Cerambycidae. Pan-Pacific Entomol., 41: 141-153. Fuchs, E. 1963. Neue neotropische Clytini aus der Sammlung des Sencken- berg-Museums. Senck. Biol., 44: 451-456. Linsley, E. G. 1935. Studies in the Longicornia of Mexico. Trans. Amer. Entomol. Soc., 61: 67-102. Vol. 50, No. 2, April 1974 139 Two New Species of Cerambycidae from Southern California (Coleoptera) Frank T. Hovore Placenta Canyon Nature Study Center 19152 W . Placenta Canyon Road, Newhall, CA 91321 Edmund Giesbert 9780 Drake Lane Beverly Hills, CA 90210 The Cerambycid genera Clytus Laicharting and Atimia Haldeman are each represented in California by four species or subspecies. In each case, three occur predominantly in the northern and central por- tions of the state. Two species, Clytus planifrons LeConte and Atimia confusa (subspecies dorsalis LeConte) have been recorded from throughout the Pacific coast region, and are the only species in either genus which are known to occur in California south of the Tehachapi Mountains. Intensive collecting in recent years in the San Gabriel and San Bernardino mountain ranges has produced new species in both genera, one of which, Atimia gannoni, described below, also oc- curs in the Sierra Nevada as far north as Tuolumne County. Atimia gannoni, new species (Figs. 1 & 3) Male. — Form moderately elongate, robust, only slightly tapering posteriorly; color black; vestiture coarse, appressed, luteus, with scattered long, erect, pale hairs on head, pronotum, elytra and undersurface of body. Head densely clothed with appressed hairs; vertex and frons densely, coarsely punctate, with glabrous, im- punctate median line; antennae attaining apical fourth of elytra, scape moderately robust, subconical, more than twice as long as broad, third segment twice as long as second segment, fourth segment distinctly longer than third, fifth segment longest, outer segments subequal in length to third segment, cylindrical, not flattened nor expanded. Pronotum. slightly wider than long, sides very feebly, obtusely rounded, widest before apex, subapical angles obtusely rounded; surface coarsely, regularly punctate medially, lateral punctures smaller, confluent to rugose, disk densely pubescent, pubescence obscuring surface, without impunctate or glabrous vittae; scutellum longer than broad, rounded posteriorly, densely pubescent. Elytra finely, irregularly punctate, with scattered large punctures, each with a single long erect hair, surface densely pubescent except for irregular denuded areas; denuded areas small, impunctate, pattern variable, usually consisting of an irregular, inter- rupted subsutural vitta, a postmedian spot and a vague, subapical oblique line; apices broadly, separately rounded or feebly subtruncate. Legs black, finely punctate, pubescent; posterior tarsi moderately broad, first segment shorter than follow- The Pan-Pacific Entomologist 50: 139-144. April 1974 140 The Pan-Pacific Entomologist Fig. 1 . Atimia gannoni Hovore and Giesbert, male. Vol. 50, No. 2, April 1974 141 ing two together, second segment about one and one-half times as long as broad. Abdomen with sternites finely, closely punctate, densely pubescent, fifth tergite narrowly rounded at apex, fifth sternite feebly truncate or emarginate at apex (Fig. 3). Length: 7-9.5 mm. Female.- — Form larger and more robust than male; antennae slightly shorter, attaining apical one-third of elytra; pronotum with sides slightly more obtuse than in male; fifth abdominal tergite and sternite broadly rounded or subtruncate at apex. Length: 8-11 mm. Holotype male, allotype (California Academy of Sciences) and 54 paratypes (37 males, 17 females), from Little Mt. Gleason, approximately 5 miles W Mill Creek Summit, Los Angeles County, California, ex. pupal cells in CaU.ocedrus decurrens (Torr.) , 21 February 1971 to 27 March 1971 (F. T. Hovore, M. T. Gannon, E. Giesbert). Additional paratypes: 13 males, 4 females, same locality as holotype, 27 February 1971 to 6 March 1971, 25 February 1972 to 6 March 1972 (D. G. Marqua, A. E. Lewis) ; 1 male, 2 females, Fallsvale, 6000 ft, San Bernardino County, California, 1 February 1970 (F. T. Hovore). Also represent- ing this species but not designated as paratypes: 1 male, Stevenson Creek, Sierra N. F., Fresno Co., Calif., 27 May 1915, “ Libocedrus decurrens Lot 173” (R. Hopping Collection, California Academy of Sciences) ; 1 male, Long Barn, Tuolumne Co., Calif., 5 June 1931 (Van Dyke Collection, California Academy of Sciences) . Paratypes are deposited in the collections of the following institu- tions and individuals: California Academy of Sciences; California Insect Survey Collection, Berkeley; Los Angeles County Museum of Natural History; California State Department of Agriculture, Sacramento; F. T. Hovore; E. Giesbert; D. G. Marqua; A. E. Lewis; J. N. Knull; W. T. Tyson; J. Cope. Discussion. — The short third antennal segment, separately rounded or feebly truncate elytral apices and lack of pronotal polished areas readily separate Atirnia gannoni from the described species of North American Atimia , with the exception of Atimia hoppingi Linsley. From A. hoppingi , to which it seems most closely related, A. gannoni may be distinguished by the coarser, denser, luteus, appressed pubes- cence (grayish-white in A. hoppingi ), glabrous, impunctate median line on the vertex and frons, and the very feebly truncate or emarginate apex of the fifth abdominal sternite in the male (broadly, deeply emargin- ate in male A. hoppingi) . (Fig. 2). Atimia chinensis Linsley is known to us only from the original de- scription of the female holotype (Linsley, 1939: p. 76, pi. 14, fig. 3), which seems to bear a strong superficial resemblance to A. gannoni. In addition to the considerable geographic separation (the type locality of A. chinensis is in Chekiang Province, China), females of A. gannoni differ from the description of A. chinensis in the black instead of dark brown integument, long erect hairs on the discal surfaces of the pro- notum and elytra, denser appressed pubescence, and more elongate scutellum. 142 The Pan-Pacific Entomologist Figs. 2-3: Fig. 2. Atimia hoppingi Linsley. Ventral view, fifth abdominal sternite and tergite (pubescence and punctation omitted) ; Fig. 3. Atimia gannoni Hovore and Giesbert. Ventral view, fifth abdominal sternite and tergite (pubescence and punctation omitted). Biology . — Atimia gannoni infests the thin outer bark of living or very recently dead Incense Cedar, most commonly in young trees, rang- ing from two to ten inches in basal diameter. Infested trees show no external signs of damage due to the presence of the Atimia. The larvae construct flat, meandering galleries, filled with fine reddish frass, in the recently dead layers of bark adjacent to the living inner bark tissues. The shallow, oval pupal cell is placed vertically, with the exit hole cut by the larva to the loose outer bark and plugged with a thin wad of stringy frass. Pupation and transformation probably occur in the late summer or fall since mature adults have been collected from their pupal chambers in February and March. The life cycle may require two or more years, as adults and several sizes of larvae have been found together in a single infestation. Atimia helenae Linsley has also been observed overwintering in both larval and adult stages (Frankie and Jensen, 1971), and from this evidence it has been suggested that A. helenae also has a two-year life cycle. It is also possible that A. gan- noni , like Atimia confusa dorsalis (Linsley, 1936, 1939), is double- brooded in the southern portions of its range. This species is named for the late Michael T. Gannon, with whose valuable assistance much of the above material and observations were gathered. Clytus chemsaki, new species (Fig. 4) Male. — Form elongate, slender; integument piceous to black, legs and antennae dark brown ; entire surface of body sparsely clothed with erect pale hairs ; pronotum margined with a band of coarse, appressed, pale yellowish- white hair; pale pubes- cent pattern of elytra consisting of small, tranverse subhumeral bars, narrow, deeply arcuate, lunate antemedian fasciae, extending forward to sutural and Vol. 50, No. 2, April 1974 143 Figs. 4-5: Fig. 4. Clytus chemsaki Hovore and Giesbert, male elytral pattern. Fig. 5. Clytus planifrons LeConte, male elytral pattern. lateral margins, transverse postmedian fasciae at apical third and oblique apical fasciae which extend anteriorly along suture to postmedian fasciae. Head coarsely, densely punctate, eyes and vertex margined with coarse yellowish-white hair; antennae extending over basal third of elytra. Pronotum slightly longer than broad, sides somewhat narrowly rounded, surface closely, coarsely punctate; mesepisterna and metepisterna clothed posteriorly with a dense patch of appressed pale hairs; pro-, meso-, and metesterna shining, sparsely clothed with pale ap- pressed hairs; scutellum densely clothed with pale appressed hairs. Elytra slightly more than 2 1 /j times as long as broad, parallel-sided, surface shining, closely, finely punctate, clothed with fine, brownish, appressed pubescence except on fasciae; apices feebly truncate. Legs slender; femora feebly clavate. Abdomen with sternites shining, sparsely punctate, clothed with pale appressed pubescence; fifth sternite subequal in length to fourth, apex narrowly, evenly rounded. Length: 8-10 mm. Female. — Form more robust than male, antennae slightly shorter; legs and basal antennal segments testaceus; pale appressed pubescence yellow; pubescent bands on pronotum wider than in male; undersurface of body more densely pubescent than in male; pronotum more broadly rounded at sides; abdomen with fifth sternite slightly longer than fourth, apex broadly, evenly rounded. Length: 7-11.5 mm. Holotype male, allotype (California Academy of Sciences) and 12 paratypes 144 The Pan-Pacific Entomologist (2 males, 10 females), from Mt. Wilson Road, 5000 ft, Los Angeles County, California, 16 to 18 June 1971 (F. T. Hovore, E. Giesbert) . Paratypes are de- posited in the collections of the following institutions and individuals: California Insect Survey, Berkeley; Los Angeles County Museum of Natural History; F. T. Hovore; E. Giesbert. Discussion. — Clytus chemsaki will key to C. planifrons LeConte (Fig. 5) in Linsley ’s (1964) key to North American Clytus. From C. planifrons it may be immediately distinguished by the narrower elytral fasciae, the apical elytral fasciae which are directed anteriorly along the suture, typically forming a “T” with the postmedian fasciae, the darker appendages, and the smaller, less robust form. From Clytus pacificus (Van Dyke), to which it seems morphologically closest, and the Californian species, C. blaisdelli Van Dyke and C. clitellarius (Van Dyke), C. chemsaki may be separated by the narrow, strongly arcuate antemedian elytral fasciae, which are directed anteriorly at the sutural and lateral margins, the configuration of the apical mark- ings, the dense bands of appressed pubescence on the fourth and fifth abdominal sternites in both sexes, and the pale yellowish-white appressed pubescence in the male. Biology. — The type series of Clytus chemsaki was collected in the late afternoon from the blossoms of Ceanothus sp. on the steep north slope of Mt. Wilson in the San Gabriel Mountains. The larval host for this species is not yet known, but it probably infests dead branches of coniferous trees. Acknowledgments We wish to extend our gratitude to Dr. John A. Chemsak and Dr. E. Gorton Linsley, University of California, Berkeley, for reviewing the manuscript; to Dr. Paul Arnaud, California Academy of Sciences, for the loan of comparative material; and to D. G. Marqua and A. E. Lewis for the use of specimens from their personal collections. Literature Cited Frankie, G. W & G. L. Jensen. 1971. Studies on the distribution and biology of Atimia helenae Linsley on two California Cupressus species. Pan- Pac. Entomol., 47 : 287-293. Linsley, E. G. 1936. Observations on the habits of some western longhorn beetles. Pan-Pac. Entomol., 12: 199. Linsley, E. G. 1939. The longicorn tribe Atimiini. Bull. S. Calif. Acad. Sci., 38: 63-80. Linsley, E. G. 1964. The Cerambycidae of North America, Pt. V ; Taxonomy and classification of the subfamily Cerambycinae, tribes Callichromini through Ancylocerini. Univ. Calif. Publ. Entomol., 22: 1-197. Vol. 50, No. 2, April 1974 145 New North American Tabanidae XXI. Another New Bolbodimyia from Mexico (Dipt era) Cornelius B. Philip California Academy of Sciences, San Francisco 94118 AND Linda Floyd Californa State University, San Francisco 94132 Bolbodimyia comprises a New World genus of strikingly marked species mostly distributed in the northern Neotropical Region. There are rare Nearctic records of B. atrata (Hine) in Arizona as far north as the type locality in famed Oak Creek Canyon. This is the only species that has been reared (Burger in Goodwin and Murdock, 1974). Philip (1954) described the male of B. atrata from Jalisco, Mexico. Species of the genus, revised by Stone (1954), are never taken in large numbers as adults, which suggests peculiar habits. Since the unique type male of the brightly marked new species described below was taken not far below the arbitrarily established northern neotropical boundary in Chihuahua, it is possible that a more northern occurrence of this species will eventually be discovered. The present specimen was among miscellaneous Mexican tabanids kindly submitted for study by Dr. R. 0. Schuster of the University of California, Davis, where the type will be on deposit. Bolbodimyia lampros, new species (Fig. 1) A robust, contrastingly orange and black species with orange pollinose cheeks and face; pleura orange with concolorous pile; legs blackish brown, tibiae swollen; wings evenly fumose with reduced apical hyaline crescents. Holotype male. — length 15.0 mm; wing length 11 mm. Head subhemispherical, eyes bare, facets in upper two-thirds enlarged, dull tan, small facets black, extended up occipital margin to vertex. Tubercle in occipital notch small and moderately depressed. Frontal triangle bare, dull grayish pollinose in apex, mostly shining, dark brownish black and swollen around antennal fossae (Fig. la) . Cheeks slightly swollen and face only moderately depressed, entirely orange pollinose with con- colorous hairs below. Scapes greatly enlarged, a little more so below, shiny and dark mahogany to black with sparse black hairs; flagella reddish, the styles about one-half lengths of plates; latter rather slender with obtuse dorso-basal angles. Palpi porrect, the basal segments and hairs orange; apical segments thicker, ovoid, brown with orange and black hairs (Fig. lb). The Pan-Pacific Entomologist 50: 145-147. April 1974 146 The Pan-Pacific Entomologist Fig. 1. Bolbodimyia lampros, male. a. Profile of head and antenna show- ing protuberant frontal triangle around antennal fossae and swollen scape, b. Palpus, c. Outer wing with reduced apical hyaline crescent. Unlined notum and scutellum subshiny black with sparse concolorous hairs; chest and vestiture similar. Pleura and prealar tubercles bright orange with concolorous pile. Legs dark mahogany to black, mostly black-haired, some yellow hairs basally on coxae. Wings evenly fumose with hyaline crescent restricted to apex between veins R+ and R 5 (Fig. lc) . Abdomen almost entirely bright orange, with short coarse black hairs dorsally; venter similarly colored but with orange hairs. Edges of dorsum with a narrow black line and tufts of black hairs. Type Locality. — Mexico: Chihuahua, Cuiteco, 30 August 1969. T. A. Sears, R. C. Gardner, and E. C. Glasser. This new species is one of the more robust representatives of the genus. It is likely that the unknown female will show sexual dichroma- tism, familiar to that found in some other species in the group. Though B. lampros is structurally somewhat similar to B. atrata and B. celeroides Stone, it differs from both in having the scape less produced below and tibiae a little less inflated. Both sexes of the former are entirely black, but only the females of celeroides from Peru are available for com- parison; the wide geographic separation as well as the tinctorial dif- ferences in the abdomens and flagella reinforce the doubt that B. lampros could be the dichromatic male, unless unexpected intervening intergrada- tion is eventually discovered. Presumably, the still unknown male of B. dampfi Philip from Mexico will be distinguished at least by shining black face and cheeks. It may be noted that Fairchild (1964) reported the male of B. philipi Stone as taken “at light” in Panama. The name lampros (Gr., lamp or lantern) is assigned for the bright, almost luminous, contrasting orange abdomen. Vol. 50, No. 2, April 1974 147 References Fairchild, G. B. 1964. Notes on Neotropical Tabanidae (Diptera) IV. Further new species and new records for Panama. J. Med. Entomol., 1: 169- 185. Goodwin, J. T. and W. P. Murdoch. 1974. A study of some immature Neo- tropical Tabanidae (Diptera). Ann. Entomol. Soc. Amer., 67: 85-133. Philip, C. B. 1954. New North American Tabanidae (Diptera) Part V. A striking new Bolbodimyia from Mexico. Wasmann J. Biol., 12: 29-33. Stone, A. 1954. The genus Bolbodimyia Bigot (Tabanidae, Diptera) . Ann. Entomol. Soc. Amer., 47 : 248-254. 148 The Pan-Pacific Entomologist Revision of the Scarabaeidae : Anomalinae 1. The Genera Occurring in the United States and Canada ( Coleoptera ) Robert W. L. Potts California Academy of Sciences, San Francisco, California It appears necessary in a consideration of the generic arrangement of this subfamily that we remember that taxonomic divisions above the species are necessarily artificial, with no objective existence, and are formed largely as a matter of convenience. Nonetheless, an organized taxonomy demands a certain stability based on the work of previous authors, and while their arrangements cannot be ruled sacrosanct merely on a basis of age or previous acceptance, such factors should not be lightly dismissed. In the Anomalinae we find an exceptionally large and remarkably interrelated group of species. Except for a relatively few genera, with some that may be intermediate to adjacent subfamilies, the Melo- lonthinae and Rutelinae, it is often difficult or impossible to separate genera in a manner consistent with the usual practice of assembling species only when they share several characters reasonably disparate to the species assembled in other genera. Despite a number of variable and distinctive characters, intermediates are the rule, not the exception. A veritable wonderland of generic and subgeneric names have been proposed in the subfamily. A strict application of what most systematists would consider necessary to a valid generic grouping would synonymize a majority of these into the single genus Anomala, with thousands of species, worldwide, and dozens, if not even hundreds of secondary homonyms. This, most certainly, would not be more convenient, and it would scarcely contribute to the ease of identification. Compromise, with a minimum of violence to a few questionable, but nonetheless long-estab- lished and widely-used genera probably becomes necessary. However, it is also obvious that a number of the proposed groupings were without true value from their inception, merely following in the mould of an early-day taxonomy that regarded any large genus as unmanageable until infinitely divided, no matter how precariously. We believe the following best represents the status of the Anomalinae of our area: The Pan-Pacific Entomologist 50 : 148 - 154 . April 1974 Vol. 50, No. 2, April 1974 149 Tribe Anomalini AnOMALA Samouelle 1819:191. Type species: Scarabaeus aeneus DeGeer, 1774, Arrow designation, 1917:126. syn. Anomalepta Casey 1915:8. Type species: Anomala semilivida LeConte, 1878, Machatschke designation, 1957:30. syn. Anomalopus Casey 1915:40. Type species: Anomala rhizotrogoides Blanchard, 1850, Casey designation, 1915:40. syn. Blitopertha Reitter 1903:85. Type species: Melolontha lineata Fabricius, 1798, Arrow designation, 1917:127. syn. Callistethus Blanchard 1850:198. Type species: Mimela auronitens Hope, 1835, Arrow designation, 1917:127. syn. Hemispilota Casey 1915:45. Type species: Melolontha lucicola Fabri- cius, 1798, Machatschke designation, 1957:30. syn. Oliganomala Casey 1915:38. Type species: Strigoderma exigua Schwarz, 1878, monobasic. syn. Pachystethus Blanchard 1850:201. Type species: Popillia vidua Newman, 1838, monobasic. syn. Paranomala Casey 1915:12. Type species: Melolontha binotata Gyllen- hal, 1817, Machatschke designation, 1957:31. syn. Phyllopertha Stephens 1830:223. Type species: Scarabaeus horticola Lin- naeus, 1758, Arrow designation, 1917:126. syn. Rhombonalia Casey 1915:5. Type species: Anomala cavifrons LeConte, 1867, Machatschke designation, 1957:31. syn. Rhombonyx auctorum (not Rhombonyx Hope 1837:106). Type species: Melolontha holosericea Fabricius, 1787, Arrow designation, 1917:126. syn. Spilota Burmeister 1844:266. Type species: Melolontha marginata Fab- ricius, 1792, Arrow designation, 1917:126. Only the synonyms with application in our area are included; many others are necessary in other sections of the world. All have been synonymized previously, however several are still accepted and in present use by some systematists. Blitopertha Reitter and the much older Phyllopertha Stephens must be included because of our introduced species, Anomala orientalis Waterhouse, which is variously placed in one or the other. Other species included in the genera are Asiatic and European, though species from Mexico and Central America have been included. Both genera were established on a combination of characters that have been subject to a confusing emendation and change by almost every author that has used them. No single character is common to all species usually included, and all are found duplicated in species not included, where they are sometimes carried to an even greater extreme. To com- pound this confusion, most authorities have failed to agree on their lists of species properly belonging in the two genera. Our introduced species, orientalis , does not differ in any material 150 The Pan-Pacific Entomologist way from our native species, and can be separated on characters of no more than species-level significance. Spilota Burmeister, Callistethus Blanchard and Pachystethus Blanchard are all largely based on the development of a prominent intermesocoxal process. This might have a certain validity for the species with the most extreme development, except that there is no place where an arbi- trary limit might be placed logically in a perfectly intergraded series. Moreover, the character also appears in some Strigoderma and in species placed in Epectinaspis. The three genera have also been subject to varying application and have never had universal acceptance. The type species of Callistethus is of Himalayan origin, and Ameri- can species probably should never have been assigned to the genus, however Machatschke, 1957:93, somehow, and quite mistakenly gave Melolontha marginata Fabricius as the type, crediting the designa- tion to Arrow 1917:127, whereas Arrow actually designated Mimela auronitens Hope as the type, and designated Melolontha marginata Fabricius as the type of Spilota , which immediately preceded Callistethus in his list. But Machatschke, 1957 :92, also gives marginata as the type of Spilota , thus incorrectly indicating the same species as the type of both genera. Among our own species sometimes variously placed in one or another of these three genera, such as marginata Fabricius, the intermesocoxal character is quite variable, from a prominent boss to a large porrect process, and selected specimens might be fitted into each of the three genera. Unfortunately, this forces the change of the name of a distinctive Florida species described by Mark Robinson, his name becoming a secondary homonym of floridana Casey 1915:33. We propose: Anomala robinsoni, new name, for Pachystethus floridana Robinson 1941:133. A majority of the Casey names were proposed as subgenera, almost entirely on characters of merely species-level importance, and admit- tedly in an attempt to fraction a large genus. But whether as genera or subgenera, merely adding species from no greater distance than Mexico and Central America immediately demonstrates the lack of validity of his groupings. Inexplicably Oliganomala Casey was misplaced as a synonym of Strigoderma by Machatschke 1957:139 and 1972:216, possibly on the basis of the fact that the unique species, exigua, equally inexplicably was described as a Strigoderma by Schwarz, although it met none of the key Vol. 50, No. 2, April 1974 151 criteria. Casey, 1915:38, made it clear it definitely was not a Strigo- derma, but the characters cited are not of sufficient importance to justify its separation from Anomala. The genus Rhombonalia Casey has had a certain acceptance, however, and for some time we struggled to maintain the concept, since it is based on a notable character, the simple protarsal claw, and does include an obviously closely-related group of species. But, it would necessarily need enlarging to include other species with the same character and eventually breaks down in a way similar to the case of genera pre- viously discussed. The simple protarsal claw is highly variable and there is no point in an intergrading series where it may be established. More- over, it was found that even cavifrons LeConte, the type, may have distinctly cleft claws not too uncommonly, and that the simple claw occurs occasionally in many of our species. It is moderately com- mon in one species that rather obviously should not be assigned to the genus. Finally, a species that is very closely related to cavifrons in its other major characters almost always has cleft claws. Other char- acters that have been cited for the genus proved valueless, and it clearly becomes necessary to accept the judgment of those who had already reduced the name to the synonymy. STRIGODERMA Burmeister 1844:310. Type species: Strigoderma sulcipennis Burmeister, 1844, Casey designation, 1915:48. syn. Alamona Casey 1915:61. Type species: Alamona parviceps Casey, 1915, Casey designation, 1915:61. syn. Strigodermella Casey 15:59. Type species: Melolontha pygmaea Fab- ricius, 1798, monobasic. Considering the entire genus and its separation from worldwide Anomalinae, the definition is far from satisfactory, with no single character that is not duplicated in other genera, both in the Americas and in the Old World. Placement in the genus depends on general habitus almost as much as on any distinct character, while the assign- ment of a number of intermediate species is entirely arbitrary. Also, several Asiatic species would have to be included in the genus by any definition except distribution. Nonetheless, in our area the genus is easily distinguished on the characters noted in the key. It has had a very long-standing acceptance in the literature, with many species de- scribed in the genus, and it would only create greater confusion to return it to Anomala , regardless of its weakness. However, the Casey attempt to split the genus with two additional genera is indefensible at any level. The small justification the genera 152 The Pan-Pacific Entomologist had when applied to our few species disappears entirely with the addi- tion of species from south of our borders. Alamona has been misplaced by Blackwelder, 1944:242, as a sub- genus and synonym of Anomala , but does not belong with that series at all, being a typical Strigodermid. Tribe Anisopliini AnOMALACRA Casey 1915:10. Type species Anomalacra cuneata Casey, 1915, monobasic. While this genus has been synonymized with Anomala by Machatschke, 1957:30 and 1972:83, and there has been confusion as to the species belonging in the group, in the present state of our knowledge it is a very distinct and valid genus. It is closely related to the tropical Callirhinus, with the clypeus remarkably thinned and the labrum reduced and withdrawn posteriorly. These characters are also common to the Old World Anisopliini, to which Tribe it would necessarily be assigned. Charles Schaeffer described our first North American species as Anomala clypealis , and despite its remarkable differentiation from other Anomala the species is still found assigned to that genus in all published lists, but it must be reassigned to Anomalacra. Tribe Popilliini POPILLIA Serville 1825:367. Type species: Trichilis bipunctata Fabricius, 1787, Arrow designation, 1917:58. Represented in our area by the single introduced species, Popillia japonica Newman, although the genus is large and diverse. It is primarily an Asiatic and African development, and has elements of be- ing parallel to the Strigoderma development in the Americas. It is generally given a separate Tribal status, however, this may be question- able as strong intergradation to the Anomalini certainly exists. Tribe Lepothopliini, new Tribe LEPTOHOPLIA Saylor 1935:132. Type species: Leptohoplia testaceipennis Saylor, 1935, monobasic. This genus is apparently intermediate to the Melolonthinae where it was originally placed as belonging with the Hopliini. Disagreement as to its relationship still exists, however we believe it belongs in the Anomalinae, where it was reassigned by Howden and Hardy, 1971:337. Vol. 50, No. 2, April 1974 153 The very distinctive tumid and penicillate ligula; the exceptionally thick anterior face of the clypeus, some two or more times as thick as normal in the Anomalinae; the obsolescent epipleurae; and the reduction of the minor protarsal claw to half or less the length of the the major claw indicate a disparity sufficient for the erection of a separate Tribe. Withal the major differences, the genitalia are of the generalized anomaline type, closely similar to Anomala binotata and flavipennis. Key to the Genera of Anomalinae of the United States and Canada 1. Ligula and undermouth parts essentially glabrous, with no more than a a few distant hairs, or, rarely, with some close-set flexuous hairs pos- teriorly 2 Ligula penicillate, with thick erect bristles; California (Imperial Val- ley) Leptohoplia 2. Clypeus anteriorly thickened, with apical face more or less vertical, round- ing under or sloping forward to distinct labrum beneath 3 Clypeus, as viewed from side, apically reflexed and thinned, without thick- ened anterior face; labrum withdrawn and not visible; Arizona Anomalacra 3. Basal pronotal margin posteriorly rounded or sinuate, always projecting backward over scutellum 4 Scutellum projecting forward into an arcuate emargination of basal pronotal margin ; Eastern States, 1 introduced species Popillia 4. Mesosternal epimera antero-dorsally raised and swollen, narrowly projecting before elytral humeri ; pronotum subquadrate, distinctly narrower than elytra; elytra more or less tapering posteriorly from near humeri .... Strigoderma Epimera reduced, obliquely rounding under covering humeri; pronotum obviously broader than long, as broad as elytra or nearly; elytra widest near middle or toward apex Anomala Literature Cited Arrow, G. J. 1917. Fauna of British India, Coleoptera Lamellicornia, Vol. 2. Blackwelder, Richard E. 1944. Checklist of the coleopterous insects of Mexico, Central America, The West Indies and South America, Part 2, U. S. Nat. Mus. Bulletin 185, pp. 189-341. Blanchard, Emile. 1850. Catalogue de la collection entomologique Coleopteres, Musee d’histoire naturelle de Paris. Burmeister, Hermann C. C. 1844. Handbuch der Entomologie, Berlin, Vol. 4, Pt. 1. Casey, Thos. L. 1915. Memoirs on the Coleoptera, Vol. 6, A Review of the Ameri- can species of Rutelinae, Dynastinae and Cetoniinae. Hope, Frederick W. 1837. The Coleopterist’s Manual, Vol. 1. Howden, H. F. and A. R. Hardy. 1971. Generic placement and adult behavior of the genus Leptohoplia Saylor. Proc. Entomol. Soc. Wash., 73: 337- 341. 154 The Pan-Pacific Entomologist Machatschke, Johann W. 1957. In Wytsman, Genera Insectorum, Fasc. 199, Coleoptera Lamellicornia, Family Scarabaeidae, Subfamily Rutelinae. Machatschke, Johann W. 1972. Coleopterorum Catalogus Supplementa, Part 66, Fasc. 1, Scarabaeoidea: Melolonthidae, Rutelinae. Reitter, Edmond. 1903. Yerh. Nat. Ver. Brunn, 41: 85. Robinson, Mark. 1941. Studies in the Scarabaeidae of North America, Trans. Amer. Entomol. Soc., 67: 127-136. Samouelle, George. 1819. The entomologists useful compendium, etc., London. Stephens, James F. 1830. Illustrations of British Entomology, Mandibulata, Vol. 3. RECENT LITERATURE Frederick Valentine Melsheimer, Parent of American Entomology. Robert Snetsinger, 86pp. (Melsheimer Entomological Series, No. 12.), 1973. Available from the Entomological Society of Pennsylvania, 106 Patterson Building, Uni- versity Park, Penna. 16802. $3.75 plus $.25 for mailing. Although most Americans prefer to consider Thomas Say as the male parent of entomology in the United States, F. V. Melsheimer certainly played a prominent role in the early history of the science. This volume contains a brief biography, followed by a facsimile reproduction of Melsheimer’s catalogue of the insects of Pennsylvania.- — Editor. Vol. 50, No. 2, April 1974 155 Behavioral and Morphological Mimicry in a Cranefly and an Xchneumonid C. N. Slobodchikoff Department of Biological Sciences Northern Arizona University, Flagstaff 86001 Field observations in Costa Rica (Slobodchikoff, unpublished), Mexico (P. Arnaud, personal communication), and Arizona (J. Wick, personal communication) suggest that mimetic relationships between craneflies and hymenopterans are fairly common. However, apparently such relationships have not been described (C. P. Alexander, personal communication) nor have such relationships been subjected to experi- mentation to determine whether Batesian or Mullerian mimicry is in- volved. The purpose of this paper is to describe an apparent mimetic situation between a crane fly, Ctenophora vittata angustipennis Loew, and an ichneumonid wasp, Therion circumflexum (L) . The observations reported here were made at Alpine Lake, Marin County, California, during May of 1971-1973. The vegetation sur- rounding Alpine Lake may be described as mixed coniferous forest interspersed with small patches of oak-grassland. Occasional streams are present, draining into Alpine Lake. The oaks in the oak-grassland are primarily live oaks, Quercus agrifolia. Both the cranefly and the ichneumonid are found mainly around the oak trees. Mimicry in these two species apparently involves two factors : similarity in flight behavior and similarity in coloration. Flight behavior in Therion circumflexum has been described by Slobodchikoff (1973). Female T. circumflexum fly comparatively slowly, at a speed of approximately 10 cm/sec, about one meter above the top of grass stems in shady areas underneath oak trees. Males, on the other hand, fly twice as fast as females, approximately 20 cm/sec, around the perimeter of the upper three-quarters of the oak canopy. When the male and female ichneumonids fly, their third pair of legs are stretched out 20 degrees below the horizontal plane of the body, their abdomen is held up 20 degrees above the plane of the body, the femora and tibiae of the first pair of legs are pressed close to the thorax, and the femora of the middle legs are pressed to the thorax while the tibiae of the middle legs are held out away from the body. Both the male and female fly with their antennae extended forward. Male and female craneflies exhibit the same flight patterns as the ichneumonid males. The craneflies fly around the perimeter of oak The Pan-Pacific Entomologist 50 : 155 - 159 . April 1974 156 The Pan-Pacific Entomologist canopies, at a speed equivalent to that of male ichneumonids. Males and females both fly with their hind legs stretched out downward and back, and with the abdomen elevated 15 to 20 degrees above the horizon- tal plane of the body. The middle femora are pressed to the thorax, while the middle tibiae are held away from the body. The front femora are kept pressed to the thorax, while the front tibiae are extended for- ward, beyond the head, apparently simulating the ichneumonid’s anten- nae. In flight, the craneflies are very difficult to separate from the ichneumonids. Apparently, neither male nor female craneflies mimic the flight behavior of female ichneumonids. No Ctenophora have been observed flying at the speed or the level of the female Therion. The similarity in coloration of the ichneumonid and cranefly is very marked. Both the Therion and the Ctenophora have a yellow, orange, and black overall color pattern. The ichneumonid (fig. 1) has a black thorax, with yellow scutellum dorsally and irregular orange splotches laterally. The head is black, with long orange antennae. The abdomen is orange, with an incomplete black stripe dorsally, and the three termi- nal abdominal segments are entirely or mostly black. The hind and mid- dle coxae are black, the front coxae are yellow. The hind femora and tibiae are yellow in their proximal 0.7, black in their distal 0.3. The middle and front femora and tibiae are yellow, as are all tarsal segments. The cranefly (fig. 2) has a black thorax, with faint yellow in the scutellar region and irregular yellow splotches laterally. The head is black, the short antennae are orange. The abdomen is orange, with a longitudinal black stripe dorsally and a reddish-black longitudinal stripe laterally. The basal third of the first abdominal segment is black, creating the illusion of a constriction or petiole when the abdomen is seen in side view against a dark background. The terminal three visible segments of the abdomen are reddish black. All coxae are black, and all trochanters are yellow. The distal ends of the trochanters of the first pair of legs are pointed dorsally, assuming the same position in flight as the yellow coxae of the first pair of legs of Therion. All femora and tibiae are yellow-orange in their proximal 0.8, and black in their distal 0.2. When seen against a dark background, such as the dark green of oak leaves, the black on the legs creates the illusion that the legs are shorter than they are in actuality. Even though all the cranefly ’s legs are longer than the ichneumonid’s legs, the cranefly ’s legs in flight appear to be no longer than those of the ichneumonid. The first tarsal segment of each leg is orange, the remaining tarsal segments are blackish-orange. This also contributes to the illusion of Vol. 50, No. 2, April 1974 157 Fig. 1. Male T. circumflexum, lateral view, shown in position of flight. Fig. 2. Female C. vittata angustipennis, lateral view, shown in position adopted during flight. shorter legs in flight. Since the orange front tibiae are held out in front of the cranefly in flight, an illusion of orange ichneumonid an- tennae is created. The geographical distribution of the cranefly coincides with the distribution of the ichneumonid. Ctenophora vittata angustipennis has been recorded from British Columbia, Washington, Oregon, and Cal- ifornia (Stone et al., 1965). Although T. circumflexum occurs through- 158 The Pan-Pacific Entomologist out Canada and the United States in transition life zones, 19 percent (31 localities) of the total localities (163) from which T. circumflexum has been recorded occur in the Pacific Northwest (Slobodchikoff, 1971). Neither the Therion males nor the Ctenophora are present in very large numbers. During May 1971 and May 1972 males of Therion and both sexes of the Ctenophora were captured, marked with enamel paint, and released. Marking and subsequent attempts at recapture were carried out for 30 days in May 1971, and 21 days in May 1972. A total of 32 male Therion were captured, marked, and released in 1971, and 27 were captured, marked, and released in 1972. Seven Ctenophora were captured, marked, and released in 1971, and 15 were captured, marked, and released in 1972. In this two year sampling period, Therion represented 73 percent of the total number of ichneumonids and craneflies caught, while Ctenophora represented 27 percent of the total captures. Each year, after the first 5 days of sampling, more than 50 percent of the Therion caught on a given day were recaptures of previously marked individuals. No marked Ctenophora have been recaptured. The mimicry may be either Batesian or Mullerian. The lower per- centage of craneflies present suggests that mimicry may be of the Batesian type, with the ichneumonid serving as the model. Protection from predation is probably provided by some distasteful quality. Sup- porting this conclusion are: 1) male ichneumonids lack an ovipositor that could be used for defensive purposes; 2) both the ichneumonid and cranefly have orange, yellow, and black coloration, colors that frequently have an aposematic function in insects (Rettenmeyer, 1970). Acknowledgments I would like to thank Drs. R. Baida and C. Holm, Northern Arizona University, for their helpful criticisms of this manuscript. Literature Cited Rettenmeyer, C. W. 1970. Insect mimicry. Ann. Rev. Entomol., 15: 43-74. Slobodchikoff, C. N. 1971. Patterns of differentiation and evolution in the genus Therion (Hymenoptera: Iclineumonidae) . Ph.D. dissertation, University of California, Berkeley. Slobodchikoff, C. N. 1973. Behavioral studies of three morphotypes of Therion circumflexum (L). (Hymenoptera: Ichneumonidae) . Pan-Pa- cific Entomol., 49: 197-206. Vol. 50, No. 2, April 1974 159 Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson. 1965. A catalog of the Diptera of North America. U.S.D.A. Agriculture Handbook No. 276, 1696 pp. RECENT LITERATURE The Anisodactylines (Insecta: Coleoptera: Carabidae: Harpalini) : Classification, Evolution, and Zoogeography. Gerald R. Noonan. Questiones entomologicae, 9: 266-480. 1973. This revision contains keys to genera and subgenera, with complete taxonomic treatment of Anisotarus, Notobia, and Gynandrotarsus. Nearly 50 pages are de- voted to discussion of the zoogeography and phylogeny of the Anisodactylina. The application of Hennig’s methods of analysis to a group of insects with no fossil record and few characters which can intrinsically be considered plesiomorphic or apomorphic should be of interest to many systematists. — Editor. 160 The Pan-Pacific Entomologist A New Species of Cyclocephala Latreille from California Sand Dunes (Coleoptera: Scarabaeidae) Alan R. Hardy Insect Taxonomy Laboratory Department of Food and Agriculture Sacramento, CA 95814 In the course of investigations of the sand dune habitat in the western United States, the following undescribed species of Cyclocephala was collected from black light in series with Leptohoplia testaceipennis Saylor, a strikingly similar species. Cyclocephala wandae, new species Holotype Male: length 7.3 mm; width (elytral humeri) 3.0 mm. Head rufo- testaceous to testaceous, rest of body pale flavotestaceous; clypeus with angles acute, upturned, sides divergent to base, disc glabrous, with scattered medium punctures, posterior clypeal margin raised above bisinuate suture; front and vertex with scattered coarse punctures, those of vertex with short erect setae; eye large, anterior edge visible in front of canthus from dorsal aspect; antennae 9 segmented, club three segmented, approximately twice as long as basal segments. Pronotum widest just behind anterior angles; sides straight behind and in front of angle; an- terior and posterior angles broadly rounded ; marginal line complete except medially posteriorly; margin broad anteriorly, narrowing laterally and posteriorly; disc with scattered fine to medium punctures. Scutellum with large shallow scattered punctures; punctures anteriorly and laterally each with a single posteriorly in- clined seta. Elytra without easily discernable striae or intervals; surface with very short setae, setae barely visible (best seen in side view). Pygidium closely, shallowly rugosely punctate; provided with long, fine, scattered hairs, these hairs longer and denser basally. Mentum with a dense, anteriorly directed brush made up of many closely set erect bristles. Anterior tibiae with three, acute, widely separated teeth on outer margin, basal tooth median; anterior tarsal claws with larger claw not split at tip, smaller claw reduced to a short stub. Hind tibia with median carina poorly defined, evident as a dark pigmented, setose, poorly elevated line, apical marginal carina missing; posterior tarsal claws fine, straight, acute. Male genitalia: figure 1. Variation in paratypical series: length: 6.6 mm to 7.5 mm; width 2.7 mm to 3.0 mm. Generally agreeing with type. Holotype male (CAS # 11941) and 2 male paratypes, California, Imperial Co., 3 mi. NW Glamis, IX-15/16-1972, M. Wasbauer and A. Hardy (USNM, HFH) ; 1 male paratype, California, Imperial Co., Regina, approx. 20 mi. SE Niland, VII-4-1971, Sand Dunes, Hardy et al. Collrs. (ARH). Paratypes are de- posited in the following collections: USNM, Howden and that of the author. I take pleasure in naming this species for my wife, Wanda, who participated in the first collection of this species. The Pan-Pacific Entomologist 50: 160-161. April 1974 Vol. 50, No. 2, April 1974 161 This species is similar to Cyclocephala arenosa Howden and Endrodi in the form of the apex of clypeus and general body shape and color, although smaller. C. wandae keys to C. puberula LeConte or C. crepuscu- laris Martinez in Endrodi’s key (1966:63) (however, not fitting either couplet), but may be readily distinguished by the male genitalia (fig. 1 ) . The male genitalia closely resembles that of C. seditiosa LeConte, however seditiosa has a 10-segmented antennae vs. 9-segmented for wandae , and is larger (10-12 mm vs. 6. 6-7. 5 mm). This species exhibits a number of characteristics which appear to be adaptations to the Sand Dune habitat. The well developed brush of hairs on the mentum is present in Leptohoplia (Howden and Hardy, 1971 ) , and an undescribed species in Anomalini. The light color is present in Pseudocotalpa andrewsi Hardy; an undescribed species of Anomala ; and Cyclocephala arenose H. & E., in addition to the above. Literature Cited Endrodi, Sebo. 1966. Monographic der Dynastinae. Teil I. [Cyclocephalini] . Entomol. Abhandl., 33: 1-460. Howden, H. F. and Endrodi, S. 1966. Five new species of Cyclocephala Latreille from North and Central America (Coleoptera: Scarabaeidae) . Can. Entomol., 98(1) : 295-302. Howden, H. F. and Hardy, A. R. 1971. Generic placement and adult behavior of the genus Leptohoplia Saylor. Proc. Entomol. Soc. Washington, 73(3): 337-341. 162 The Pan-Pacific Entomologist A Redefinition of the Tyrini with the Addition of Anitra Casey (Coleoptera: Pselaphidae) Donald S. Chandler Ohio State University, Columbus, 43210 A small series of a pselaphid was recently collected at Molino Basin in the Santa Catalina Mountains of Arizona by Karl Stephan on March 7, 1970. The beetles were found under a large, flat rock and were as- sociated with a colony of ants warming brood. The ants were determined as Paratrechina melanderi (Wheeler). The pselaphids were identified as Anitra glaberula Casey upon com- parison with the male holotype (USNM #38748). Anitra was initially placed in the Ctenistini by Casey (1893) and has been retained there by succeeding workers (Raffray 1908, Park 1953). Detailed examination of the specimens convinced me that the genus was misplaced and should be removed to the Tyrini. Casey placed the genus in the Ctenistini when the differences between that tribe and the Tyrini were not very obvious to him (p. 499). His detailed description gives no indication as to why the genus was placed in the Ctenistini rather than the Tyrini. Raffray (p. 331) questioned Casey’s placement and suggested that the genus might best be placed in the Hybocephalini. However, Park (p. 316) still retained Anitra in the Ctenistini long after Raffray ’s suggestion. The Ctenistini are placed in the same subfamily, the Pselaphinae, and the same division, the Macroscelia, as the Tyrini, but the two tribes can be separated easily by two characters. The Ctenistini are characterized by the presence of squamiform setae, particularly in the foveae and sulci of head, thorax and the articular surfaces of the abdominal segments. Also, the epistome is large, expanded and usually prominent laterally. The Tyrini are characterized by moniliform setae and the epistome may he large, but is neither expanded nor prominent. The Hybocephalini, which was referred to by Raffray, also possess squamiform setae. Anitra possesses no squamiform setae. On the basal prontal projec- tion there are some short, thick setae which were clearly not squamiform when a specimen mounted on a slide was examined. A similar grouping of short, thick setae occurs also in Juxtahamotopsis Park of the Hamotini (Park 1945), a tribe very closely related to the Tyrini. The epistome is smoothly fused to the frons and is neither expanded nor prominent. On The Pan-Pacific Entomologist 50 : 162 - 164 . April 1974 Vol. 50, No. 2, April 1974 163 Fig. 1. Right maxillary palpus. Fig. 2. Right mandible. the basis of these characters, it is apparent that Anitra does not be- long in the Ctenistini or the Hybocephalini and should be placed in the Tyrini. Anitra does not fit within the Tyrini as it is currently defined, but the differences can be explained as reduction of characters due to the myrmecophilous habits of the genus. The Tyrini as currently defined have four-segmented maxillary palpi and one to three basal pronotal foveae. Anitra has two-segmented maxillary palpi (Fig. 1) (Casey said three segments) and has no basal pronotal foveae. The genus is certainly myrmecophilous because of the circumstances of its collection and probably functions as a synoekete. Reduction of the number of palpal segments is common to myrmecophilous genera, for example the two-segmented maxillary palpi of Biotus Casey (Ctenistini) and the one- or two-segmented palpi of the Clavagerinae. The reduced teeth on the mandibles (Fig. 2) are a characteristic common to myrmecophilous beetles. The lack of basal pronotal foveae is not too surprising in a tribe where the number varies from one to three Because of the absence of squamiform setae and the lack of a promi- nent epistome, I conclude that Anitra should be placed in the Tyrini rather than the Ctenistini or Hybocephalini. I am also expanding the limits of the Tyrini so that the number of the segments of the maxillary palpus vary from two to four and the number of basal pronotal foveae vary from zero to three. I would like to thank Dr. Charles Gaspar of the Institut Agronomique de Tetat Gembloux, Belgium, for the identification of the ants, Mr. T. J. Spilman of the United States National Museum for permission to examine the Casey collection, and Dr. C. A. Triplehorn of the Ohio State University for checking the manuscript. 164 The Pan-Pacific Entomologist Literature Cited Casey, T. L. 1893. Coleopterological notices, V. Ann. New York Acad. Sci., 7: 281-606. Park, 0. 1942. A study in neotropical Pselaphidae. Northwestern Univ. Studies Bio. Sci. Med., no. 1, 403 pp. Park, 0. 1953. Discrimination of genera of pselaphid beetles of the United States. Bull. Chicago Acad. Sci., 9: 299-331. Raffray, A. 1908. Pselaphidae. Genera insectorum, 64th fascicule, P. Wytsman, ed. Bruxelles. 487 pp. RECENT LITERATURE The Insects of Virginia: No. 6. Horseflies and deer flies of Virginia (Diptera: Tabanidae). L. L. Pechuman. 92pp. (Research Division Bulletin 81, Virginia Polytechnic Institute and State University), 1973. Available on an exchange basis from the Research Division Mailing Room, Hutcheson Hall, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061. This faunal treatment ostensibly covers only a small geographic area, but will probably be useful as a guide to the horseflies of much of the central Atlantic coast. Included are short sections on the biology and ecology of tabanids, their taxonomic characters, distribution, and economic significance, as well as lengthier taxonomic treatment of the 95 species known to occur in Virginia. Keys are pro- vided for separating the genera and the species within each genus (separate keys for males and females). For each species a concise summary is given of the distribution in Virginia, and the seasonal occurrence, along with biological notes. — Editor. Vol. 50, No. 2, April 1974 165 The Genus Dicopus Enock (Hymenoptera: Mymaridae) Richard L. Doutt University of California, Berkeley A century has passed since Enock in England became interested in the Mymaridae. He quaintly described his initial fascination with these parasitic wasps as follows (1909) : “ in 1872 I commenced my own observations from a simple exhibit by Mr. Frederick Fitch at the Quekett Microscopical Club. Under the microscope was shown ‘A Fairy Fly in a Spider’s Web.’ It was illuminated on a black background, which gave it the most wonderful appearance, every limb, each tiny hair and the long cilia resembling brilliant silver. It was a most fasci- nating object to any one, and speaking for myself, I had never before seen anything to compare with its delicate fairy-like structure. I then and there determined to know more about its relatives. Next day I searched the spiders’ webs in my garden at Holloway and found quite a number of several species, which I mounted in Canada balsam.” “On removal in 1882 to Woking I very soon found I was in the land of plenty for Mymaridae, which simply swarmed on the windows of the house and in a very small conservatory — where I caught seven of the new genera which I am introducing for the first time tonight.” One of the new genera in the paper which Enock read to the Entomological Society of London was Dicopus. The type species is Dicopus minutis- sirnus Enock which he described from a female, the male being unknown. In 1911 A. A. Girault received a single male mymarid from Quebec “so minute as almost to be lost in the medium in which it was mounted.” At first glance he thought it was an Alaptus but quickly recognized that it belonged to another genus and he made a very astute analysis (1911) : “There is an English genus, however, recently described by Enock for Dicopus minutissima Enock, which closely resembles Alaptus Haliday in form, but which differs in bearing two more antennal segments; the male of this genus is unknown. Now, this Canadian species precluded from being an Alaptus must belong to Dicopus, at least until we know to the contrary. It is one of the smallest North American Mymaridae and because of this and also because of its characteristic appearance, I believe it is incumbent on me to describe it rather than risk its being lost.” This single male individual, the type of Dicopus halitus Girault, was the sole record for the genus on the North American continent until the specimens described herein were collected in California. The Pan-Pacific Entomologist 50 : 165 - 168 . April 1974 166 The Pan-Pacific Entomologist In September 1911, while en route to Australia, Girault stopped for a few hours at Suva, Fiji and he “managed to collect a few parasitic Hymenoptera.” One of these was captured on a window pane in a woodworker’s shop. Girault recognized it as a male Dicopus and named it D. psyche. “This Fijian species, like the American form, is very minute, also extremely delicate, and was extraordinarily difficult to capture; it was moving slowly over the pane of a window, but I was not able to keep sight of it for more than the fraction of a second at a time. This fact, taken in conjunction with its fragility, made it necessary to spend three-quarters of an hour in effecting its capture.” (Girault, 1912). In 1912 Ricardo Mercet described Dicopus citri from Spain, and Girault later added 3 Australian species by describing D. victoria and D. maximus in 1920, and D. hidentiscapus in 1931. Also in 1931 Claude Morley in England described D. cervus from a male hymenop- teran which “ flew on to the book I happened to be reading at 5 p.m. on July 17th, 1921, while sitting upon the lawn beside the moat which surrounds Monks’ Soham House in Suffolk.” This extraordinary detailed collection datum for a mymarid was rendered ineffectual in 1955 when Alejandro Ogloblin indicated that D. cervus Morley is not a mymarid. In the same paper Ogloblin pro- posed four new genera from Argentina all closely related to Dicopus. These are Chromo dicopus, Callodicopus, Dicopulus and Dicopomorpha. Ogloblin suggested that some of the species referred to Dicopus by Mercet and Girault might possibly be transferred to other genera. From my examination of Girault ’s types at the Queensland Museum I believe that D. victoria Girault and D. maximus Girault are incorrectly placed in Dicopus and instead probably belong to one of Ogloblin’s genera or to a closely related but undescribed genus. The genus Dicopus is therefore presently constituted of D. minutis- simus Enock, D. halitus Girault, D. psyche Girault, D. citri Mercet, D. hidentiscapus Girault and the California species described as follows: Dicopus pygmaeus, new species Female. — Body length 0.22 mm; forewing length 0.44 mm; wing blade 0.30 mm, apical marginal cilia 0.14 mm; head, thorax, abdomen brown, appendages pale testaceous, eyes and ocelli deep maroon. Toruli high on face near frontal carina of vertex; mandibles elongate. Vertex finely granulate, lateral carinae well de- veloped, thick, interrupted, in 3 sections. Face with several prominent bristles. Antennae 1.5 X body length. Funicle segments 6, 7 distinctly wider than preced- ing funicle segments, club elongate with 2 terminal projections (Fig. 2) . Abdomen sessile, mesophragma penetrates % its length, ovipositor short, not exserted. Legs Vol. 50, No. 2, April 1974 167 Figs. 1-3. Antennae. 1. Dicopus pygmaeus male. 2. D. pygmaeus female. 3. D. enocki female. longer than body, fore femora and tibiae slightly swollen, fore coxae largest. Foretarsus with basal segment barely exceeding segment 2, apical segment (5) longest. Male. — Similar to female in size and color. Antenna (Fig. 1) 12 segmented, funicle segments 6, 7, 8, 9 largest, widest ; club with terminal spine. Hind trochanters elongate. Holotype female, Mecca, Imperial County, California, on sugar beet, 19 June 1964, H. R. Moffitt. Allotype, same data as holotype except collected 21 May 1964. Types at Division of Biological Control, University of California, Albany. The male antenna of D. pygmaeus is similar to D. halitus , but the terminal spine of the club and the elongate hind trochanters are dis- tinctive. There are also color differences as D. halitus is sooty black. The female of D. halitus is unknown. Dicopus enocki, new species Female. — Body length 0.35 mm; forewing length 0.73 mm; wing blade 0.46 mm, apical marginal cilia 0.27 mm. Head, thorax, abdomen brown; legs and an- tennae light brown; eyes red. Scutum, postscutellum lightly reticulate; scutellum, propodeum smooth. Head large, as wide as thorax. Vertex with gross carinae. Mandibles elongate, single toothed. Toruli high on face, near frontal carina of vertex. Scape long with two distinct tooth-like projections bearing spines on ventral aspect (Fig. 3). Funicle segments 1-6 slender, elongate; segment 7 shortest, widest. Club elongate with 2 terminal projections. Legs slender; fore coxae largest; foretarsus with basal segment distinctly longer than segment 2. Mesophragma penetrates Vs abdomen; ovipositor short, not exserted. Male. — Unknown. Holotype female, Kingsburg, Fresno County, California, suction trap in vineyard, 23-24 April 1965, J. Nakata; 2 paratype females same data as holotype except collected 15-16 May 1965 and 25-26 April 1965; 1 2 paratype; Caswell State Park, Stanislaus County, California, on Rubus, 29 April 1963, R. L. Doutt. Type series at Division of Biological Control, University of California, Albany. 168 The Pan-Pacific Entomologist D. psyche Girault is similar to D. enocki but differs by the lack of tooth-like projections on the scape and a wider funicle segment 6. D. pygjnaeus differs in its antennal structure (compare Figs. 2 and 3), in its shorter basal segment of the foretarsus, and in the larger develop- ment of the mesophragma. The name D. enocki is in centennial recogni- tion of the contributions of Enock to the study of Mymaridae. Literature Cited Enock, F. 1909. XI New genera of British Mymaridae. Trans. Entomol. Soc. London, 1909, pt IV, 449-459. Girault, A. A. 1911. The probable occurrence of the Mymarid genus Dicopus Enock in North America (Hymen.). Entomol. News, 22: 347-349. Girault, A. A. 1912. On the probable occurrence of the Mymarid genus Dicopus Enock in Fiji. Proc. Entomol. Soc. Wash., 14: 22-23. Girault, A. A. 1920. New genera and species of Australian Mymaridae. Insec. Inscit. Menstr., 8: 96-100. Girault, A. A. 1931. A new habit in an old insect. Homo pudicus and new Eurytomidae. Privately published, Brisbane, 4 pp. Mercet, Ricardo. 1912. Mimaridos nuevos de Espaha. Boll. R. Soc. Esp. Hist. Nat., Madrid, 12: 331-337. Morley, Claude. 1931. New Oxyura from Britain. The Entomologist, 64(812) : 14-16. Ogloblin, Alejandro. 1955. Los nuevos representantes de la famn. Mymaridae (Hym.) de la Republica Argentina. Boll. Lab. Zool. Gen. Agr. Filippo Silvestri, Portici, 33: 377-397. RECENT LITERATURE Cerambycidae of North America. Part VI, No. 1. Taxonomy and classifi- cation of the subfamily Lepturinae. E. G. Linsley and J. A. Chemsak, Uni- versity of California Publications in Entomology, 69: 1-138, 1973. The excellence of the first five volumes is continued in this work which treats the tribes Desmocerini and Necydalini, along with a portion of the Lepturini. As in the preceding parts, the text is well illustrated with habitus drawings and distributional maps. — Editor. Vol. 50, No. 2, April 1974 169 A review of the genus Mellinus (Hymenoptera: Sphecidae) M. L. Siri and R. M. Bohart Department of Entomology, University of California, Davis Mellinus Fabricius is a small genus occurring in the Palearctic Re- gion and in North America as far south as Guatemala. These wasps form an apparently relict group best placed in the Nyssoninae. Gen- eralized features are the short tongue, evident notauli, simple pro- podeum, no omaulus, two midtibial spurs, recurrent veins widely sepa- rated at their forward ends, and a long triangular marginal cell. The principal specialization is the petiolate gaster which results in a body form somewhat like that of the philanthine genus Trachypus. Morphological details useful at the specific level are few except for size and color. Helpful are clypeal shape, male antennal form, petiole shape, tibial setation, and male genitalia. Studies of male genitalia have revealed dramatic differences among several species (figs. 13-16) and strong similarities among others. In abdominalis, a peculiarity is a dorsal arm of the digitus which opposes a fingerlike extension from the enlarged gonostyle (fig. 15). The three species of the rufinodus group, rufinodus , bimaculatus and imperialis , have nearly identical genitalia. In these, the two branches of the aedeagus are loosely connected by membrane which seems capable of great expansion (fig. 16). Also unusual in the rufinodus group are the overlapping distal lobes of the gonostyle. In palearctic species, ground nests are stocked with flies, especially Muscoidea. Since arvensis is rather common, it has been observed frequently in the act of pouncing like a cat on flies attracted to fresh cow droppings. Huber (1961) has presented a detailed exposition of known biology. We are treating Mellinus as including 10 species of which three are palearctic. One of the seven New World forms is described as new. Several of the species are rare in collections. We have seen no ma- terial of obscurus , and females only of alpestris and satanicus. Since markings are diagnostic for known species, males of the last two should be readily recognized when found. Material has been borrowed from several institutions, but we are especially grateful to curators at the U.S. National Museum, Cornell University, American Museum of Natural History, California Academy of Sciences, and Canadian National Collection. The Pan-Pacific Entomologist 50: 169-176. April 1974 170 The Pan-Pacific Entomologist Key to species of Mellinus 1. Propodeum generally rugose posteriorly, enclosure medially ridged and sculptured but with a smooth outer area (fig. 4), palearctic 2 Propodeum weakly sculptured posteriorly except toward abdominal in- sertion, enclosure evenly shagreened (fig. 5), New World 4 2. Tergal markings yellow arvensis (L.) Tergal markings whitish, female legs reddish 3 3. Tergum V banded, scutellum with a whitish spot crabronea Thunb. Tergum V all dark, scutellum all dark obscurus Handl. 4. Abdomen all dark _ 5 Abdomen with pale markings 6 5. Female clypeus without a median discal projection, scutellum lightly sha- greened and somewhat polished between punctures, pronotal ridge yellow, fe- male flagellomere I about 2.5 times as long as broad (fig. 8) _ pygmaeus Handl. Female clypeus with median discal projection (figs. 3, 6), scutellum closely shagreened and dull between punctures, pronotum all dark, female flagel- lomere I about 2.8 times as long as broad (fig. 7) satanicus Siri and Bohart 6. Tergum III with a complete pale band or with closely approximated pale spots 7 Tergum III at most with widely separated pale spots 8 7. Petiole and legs largely red, both scutellum and metanotum with pale spots, terminal flagellomeres about half yellow and half black, tergum IV usually with pale spots rufinodus Cr. Petiole and legs largely brown, both scutellum and metanotum dark, ter- minal flagellomeres dark, tergum IV banded imperialis Bohart 8. One or more abdominal segments beyond petiole red, male flagellomeres XXI distinctly enlarged and black (fig. 9), median lobe of male clypeus protruding and truncate (fig. 1) abdominalis Cr. No abdominal segments beyond petiole red, no asymmetry or enlarge- ment of male flagellomeres, male clypeus flat and with a tridentate apex or at least one distinct median tooth 9 9. Flagellomeres dark, femora and tibiae black, tarsi brown with some yel- low, petiole black with yellow markings alpestris Cam. Flagellomeres dark dorsally but yellow ventrally, femora and tibiae light red or yellow, petiole dark red to light red bimaculatus Pack. Mellinus abdominalis Cresson (Figs. 1, 9, 15) Mellinus abdominalis Cresson, 1882. Trans. Amer. Entomol. Soc., Proc. 9:xxxix. Lectotype $, Montana (Acad. Nat. Sci. Philadelphia). Mellinus abdominalis personatus W. Fox, 1894. Entomol. News 5:202. Holotype 2 , Montana (Acad. Nat. Sci. Philadelphia) . This is the most distinctive species of Mellinus. It is the only one in which the abdomen is mostly or all red with pale markings reduced Vol. 50, No. 2, April 1974 171 Figs. 1-12. Fig. 1. Mellinus abdominalis, male face. Fig. 2. M. rufinodus female face, showing color pattern. Fig. 3. M. satanicus , female face. Figs. 4-5. Propodeum and enlarged detail. Fig. 4, arvensis. Fig. 5, satanicus. Fig. 6. M. satanicus head, lateral. Figs. 7 -8. Pedicel and flagellomeres I-II. Fig. 7, satanicus female. Fig. 8, pygmaeus female. Figs. 9-11, crabronea. Fig. 12, M. satanicus, petiole and base of abdominal segment II. 172 The Pan-Pacific Entomologist or absent. The male is especially odd with respect to the slender and gradually expanded abdomen, sculptured clypeal form (fig. 1), special- ized antenna (fig. 7), prominent hair tuft on last two visible sterna, and complex genitalia (fig. 15). We have seen 19 specimens all from localities west of the 100th meridian as follows: Montana: Platte Co.; Wyoming: Carbon Co.; Nebraska: Valentine, Thedford, Alliance, Halsey: Colorado: Mesa Verde, Great Sand Dunes. Mellinus alpestris Cameron Mellinus alpestris Cameron, 1890. Biol. Centr.-Amer. Hymen. 2:85. Holotype 2, Xucumanatlan, Guerrero, Mexico (Brit. Mus. Nat. Hist. London). We have seen females only, 11 specimens in all, including the holo- type. Morphologically, alpestris and bimaculatus are quite similar and must be separated by color characters as given in the key. All of our specimens have a pair of yellow spots on the propodeal enclosure, contrary to the case in bimaculatus. Also, our alpestris have the clypeus all yellow but this character is known to be variable in other species. Collections are from Mexican States as follows: Michoacan: El Pueblito; Guerrero: Xucumanatlan; Mexico: Texcoco; Morelos: Cuernavaca, Amecameca. Mellinus arvensis (Linnaeus) (Figs. 4, 13) V espa arvensis Linnaeus, 1758. Syst. Nat. Ed. 10:573. Holotype 2, “Europa,” Linnaean Coll., London. The complicated synonymy is not given here; see Bohart and Menke (in press). We have studied 52 specimens of arvensis. In determining species limits, reliance has been placed on the discussions given by Handlirsch (1888) and Beaumont (1964). This is the largest species of Mellinus , surpassing rufinodus on the average. Some females attain a length of 15 mm. The propodeal sculpture is characteristic of European Mellinus as shown in fig. 4. Genitalia are depicted in fig. 15. The species is widespread in Europe and is sometimes abundant in moun- tainous areas. Specimens from the Alps tend to have the legs some- what reddish, especially in females. Mellinus bimaculatus Packard Mellinus bimaculatus Packard, 1867. Proc. Entomol. Soc. Philad. 6:419. Syntype 2, Dublin, New Hampshire (Mus. Comp. Zool. Harvard); syntype 2, Bruns- wick, Maine (location of type unknown). Mellinus ivolcotti H. S. Smith, 1908. Entomol. News 19:299. Holotype 2, Beaver Island, Michigan (Univ. Nebraska) . Vol. 50, No. 2, April 1974 173 Figs. 13-16. Male genitalia, ventral, and enlargement from aedeagal area. Fig. 13, arvensis. Fig 14, pygmaeus. Fig. 15, abdominalis. Fig. 16, imperialis. We have seen 18 specimens. The face may be nearly all black as in Packard’s syntypes, variously mottled, or mostly yellow as in Smith’s wolcotti. The species is similar to alpestris but differs in markings as indicated in the key. The petiole is ordinarily black but occasionally 174 The Pan-Pacific Entomologist may be dull red and rarely is yellow blotched. M. bimaculatus average a little smaller than rufinodus, 8-9 mm long rather than 9-12 mm. The male genitalia are similar to those of rufinodus but the distal lobes of the gonostyle are somewhat more slender and tapering. This is the only species known to occur in eastern United States. However, it ranges into Mexico. Localities are: Maine: Brunswick; New Hampshire: Dublin, Franconia; Massachusetts; New York: Huntington, New Jersey; Ohio: Stark Co.; Michigan: Clare Co., Midland Co., Beaver Island, Douglas Lake. In Mexico it has been collected in Jalisco: La Primavera; and Oaxaca: Macuilxo- chitl. Mellinus crabronea (Thunberg) (Fig. 11) Crabo sabulosus Fabricius, 1787, nec Sphex sabulosus Linnaeus, 1758, now in Ammophila. Lectotype 2, Copenhagen, Denmark (Univ. Copenhagen Zool. Mus.) . Sphex crabronea Thunberg, 1791, new name for Crabo sabulosus Fabricius (Art. 59c, ICZN) . We have seen only 10 specimens of crabronea and key characters have been based on discussions (under sabulosus ) by Handlirsch (1888) and Beaumont (1964). The subserrate form of the male an- tenna is shown in fig. 11. The serrations seem to be a little more pro- nounced than in arvensis but it must be said that the two species are structurally almost identical. Separation on the basis of the white markings in crabronea is not entirely satisfactory, particularly in males. Recorded localities are in Europe and central Asia. Mellinus imperalis R. Bohart (Fig. 16) Mellinus imperialis Bohart, 1968. Pan-Pac. Entomol. 44:235. Holotype £, Bard, California (Univ. California, Davis). The brown and extensively deep yellow color pattern distinguishes this species from rufinodus. However, in the absence of structural dif- ferences there remains the possibility that it is an early season (Feb- ruary to April) Sonoran desert subspecies of rufinodus (June to Sep- tember). We have seen a total of 11 specimens: California: Bard; Sonora (Mexico) : Guaymas, Cocorit, Alamos. Mellinus obscurus Handlirsch Mellinus obscurus Handlirsch, 1888. Sitz. Akad. Wiss. Wien 96:288. Holotype $, Korea (Radoszkowsky Coll., Warsaw). Mellinus tristis Perez, 1905. Bull. Mus. Paris 1905:156. Holotype 2, Japan, “Japan central” (Nat. Hist. Mus. Paris). Vol. 50, No. 2, April 1974 175 Since we have seen no specimens of obscurus , reliance has been placed on the original descriptions referred to above. The possibility exists that obscurus is a dark variety or subspecies of crabronea from eastern Asia. Mellinus rufinodus Cresson (Fig. 2) Mellinus rufinodus Cresson, 1865. Proc. Entomol. Soc. Philad. 4:475. Leetotype $, Colorado (Acad. Nat. Sci., Philadelphia). The mostly red legs and petiole are a fairly constant color character but the femora may be all dark. As in other Mellinus , facial markings are variable. Some rufinodus have the face mostly yellow (fig. 2), others mostly black. We have seen two specimens with a yellow band on tergum IV as in imperialism but the more extensively pale underside of the flagellum in the former species is distinctive. We have seen 83 specimens. All United States material has been from west of the 100th meridian. Nebraska: Thedford; Texas: Valentine; Colorado: Den- ver, Dumont, Poudre Canyon; New Mexico: Pecos, Organ Mts., Raton; “Mon- tana”; Utah: Box Elder Co., Farmington, Provo; Arizona: Chiricahua Mts., Huachuca Mts., Santa Rita Mts. Mexican localities by States are: Chihuahua: Arroyo Mestino, 7600 ft.; Durango: 10 mi. w. Durango; Guerrero: 40 mi. n. Queretaro; Mexico: Teotihuacan; Aguascalientes: Penvelos. Mellinus pygmaeus Cameron (Figs. 8, 10, 14) Mellinus pygmaeus Handlirsch, 1888. Sitz. Akad. Wiss. Wien 96:289. Syntypes, 1 $, 2 2, Orizaba, Mexico (Nat. Hist. Mus. Vienna). This small dark species is similar to satanicus but smaller, about 7 mm long instead of 12.5, and more polished, especially on terga I-II. The female antenna is perceptibly more clubbed in pygmaeus and a pair of yellow spots are nearly always present on the pronotal ridge. The clypeus is flat and may be black or with considerable yellow. Genitalia (fig. 14) are distinctive among known forms. The gonostyle is relatively simple and the aedeagus bears stout spines. We have seen 20 specimens, one from Panajachel, Guatemala and the others from Mexico: Michoacan: Puerta Garnica; Veracruz: Coscomatepec, Orizaba (holotype) ; Mexico: Texcoco; Puebla; Cuetzalan; Morelos: Amecameca. Cuernavaca; Chiapas: San Cristobal. Handlirsch (1888) reported a pair of pygmaeus in the Dresden Museum (now destroyed) from “Brasilien.” The oc- currence of the species in South America needs confirmation. 176 The Pan-Pacific Entomologist Mellinus satanicus, new species (Figs. 3, 5, 6, 7, 12) Female: Length 12.5 mm. Black with yellow in narrow spots along inner orbit, scape in front, faint lateral dots on clypeus, and most of foretibia anteriorly; wings lightly stained. Pubescence pale fulvous, moderate, short but erect and thick beneath thorax as well as under trochanters and femora. Body closely microsculptured in a reticulate pattern, scutum posteriorly with short and fine longitudinal ridging, mesopleuron and scutellum with minute punctures scattered over microsculpture. Face (figs. 3, 6) with a sharp, compressed, hornlike, median, projection at lower one third of disc; antenna slender, flagellomere I about 2.8 times as long as broad (fig. 7) ; propodeum finely sculptured (fig. 5) ; petiole 4.0 times longer than broad, one third as broad as tergum II (fig. 12). Outer margin of hindtibia with numerous relatively long spines set on low protuberances. Holotype female, Zontehuitz near San Cristobal, Chiapas, Mexico, 9600 ft., June 25, 1969 (W. R. M. Mason, Canadian National Collection, Ottawa). The peculiar horned clypeus (figs. 3, 6) separates the species im- mediately. In addition, the extreme reduction of yellow is remarkable. This should lead to recognition of the male when it is discovered. References Cited Beaumont, J. de. 1964. Insecta Helvetica fauna 3. Hymenoptera: Sphecidae. 169 pp. Soc. Entomol. Suisse, Lausanne. Bohart, R. M. and A. S. Menke (in press) . Sphecid wasps, a generic revision, U. C. Press, Berkeley. Dalla Torre, C. G. de. 1897. Catalogus Hymenopterorum, etc., vol. 8. Fossores. viii + 749 pp. G. Engelmann, Lipsiae. Fox, W. J. 1894. Studies among the fossorial Hymenoptera. III. Synopsis of the N. Am. species of the genus Mellinus Fab. Entomol. News, 5: 201- 203. Handlirsch, A. 1888. Monographie der mit Nysson und Bembex verwandten Grabwespen. Sitz. Akad. Wiss. Wien. Math. -Nat. Classe, 96: 219-311. Huber, A. 1961. Zur biologie von Mellinus arvensis. Zool. Jahrb. Abt. Syst. Oekol. Geol. Tiere, 89: 43-118. Vol. 50, No. 2, April 1974 177 Two New Heliococcus Species, a Key to the North American Species, and a List of World Species. (Homoptera: Coccoidea: Pseudococcidae) Douglass R. Miller Systematic Entomology Laboratory, IIBIII, USDA, ARS, Beltsville, Maryland 20705 In this paper two new species of Heliococcus are described for in- clusion in a study on the mealybugs of San Clemente Island. Because no comprehensive work has been published on Heliococcus, a diagnosis of generic characters and a list of known species are given. Heliococcus Sulc, 1912. Type species.— Heliococcus bohemicus Sulc, 1912, by monotypy. Diagnosis. — Adult female: anal lobes large, strongly protruding, normally sclerotized; ventral-lobe sclerotization normally elongate and conspicuous; 2-18, normally 17 or 18, pairs of cerarii; 2 or 3 conical setae in each cerarius; multi- locular pores, when present, ventral only (except on H. phaseoli) , normally re- stricted to areas near vulva, sometimes absent; quinquelocular pores restricted to venter (except on H. dorsiporosus) , confined to medial and mediolateral areas; trilocular pores present in all areas except medial and mediolateral areas of venter; small discoidal pores sparce on both surfaces; crateriform tubular ducts present; oral-collar tubular ducts normally present, restricted to venter; anal ring with 3 pairs of setae and 3-5 rows of pores; dorsal and ventrolateral setae normally conical; circulus normally present; hind legs with or without translucent pores; tarsal digitules not apically capitate; claws each with large denticle; claw digitules capitate; antennae normally 9-segmented; with 2 pairs of ostioles. Notes. — Heliococcus is most closely related to Saliococcus Kanda and T akahashicoccus Kanda in that all three genera possess the characteristic crateriform tubular ducts. Saliococcus and T akahashicoccus differ from Heliococcus in possessing cerarii which each contain more than 3 conical setae. In Heliococcus all but the ocular cerarii contain less than 4 conical setae. List of World Species of Heliococcus Except for one questionably placed Ethiopian species, H. phaseoli, Heliococcus is exclusively Holarctic. The following list has been com- piled from the literature and includes 39 species and subspecies, eight from the New World and 31 from the Old World. The Pan-Pacific Entomologist 50: 177-192. April 1974 178 The Pan-Pacific Entomologist 1. adenostomae McKenzie, 1960: 707 (California) 2. artemisiae Ter-Gigorian, 1967: 134 (Armenian SSR) 3. atraphaxidis Bazarov, 1963: 38 (Tadzhik SSR) 4. atriplicis McKenzie, 1964: 235 (California) 5. bambusae (Takahashi) (Taiwan) Phenacoccus bambusae Takahashi, 1930: 6 Heliococcus bambusae (Takahashi), Goux, 1934: 171 6. bohemicus Sulc, 1912: 39 (“Bohemia”; France; Germany; USSR) 7. caucasicus Borchsenius, 1949: 279 (Armenian SSR) 8. cinereus Goux, 1934: 164 (France) 9. clemente Miller, new species (California) 10. cydoniae Borchsenius, 1949: 272 (USSR) 11. deserticola Miller, new species (Mexico; Arizona, California, Nevada, New Mexico, Texas) 12. destructor Borchsenius, 1941: 6 (Kirgiz SSR, Kazakh SSR, Tadzhik SSR, Turkmen SSR, Uzbek SSR) 13. dorsiporosus Danzig, 1971: 382 (Eastern USSR) (It is questionable that this species belongs in Heliococcus because it has dorsal quinquelocular pores.) 14. glycinicola Borchsenius, 1956: 678 (Korea) 15. halocnemi Borchsenius, 1949: 277 (Uzbek SSR) 16. herbaceus Borchsenius, 1956: 678 (Korea) 17. insignis (Lobdell) (Kansas, Louisiana, Mississippi) Phenacoccus insignis Lobdell, 1930: 210 Heliococcus insignis (Lobdell), Ferris, 1950: 97 18. kehejanae Ter-Grigorian, 1967: 136 (Armenian SSR) 19. kurilensis Danzig, 1971: 386 (Eastern USSR) 20. marginalis Goux, 1953: 104 (France) 21. maritimus Danzig, 1971: 388 (Eastern USSR) 22. minutus (Green) (Channel Island of Guernsey, England) Phenacoccus minutus Green, 1925: 519 Heliococcus minutus (Green), Williams, 1962: 29 23. montanus Borchsenius, 1949: 274 (Tadzhik SSR, Uzbek SSR) 24. nivearum Balachowsky, 1953: 238 (France; USSR) 25. nivearum austriacus Balachowsky, 1953 : 240 (Austria) 26. oligadenatus Danzig, 1972: 333 (Mongolian Republic) 27. osborni (Sanders) (Canada; Colorado (?), Indiana, Iowa, Louisiana, Missouri, New York, Ohio, Virginia; Egypt (?) ) Phenacoccus ( Paroudablis ) osborni Sanders, 1902 : 284 Phenacoccus pettiti Hollinger, 1917 : 281 Heliococcus osborni (Sanders), Ferris, 1950: 99 28. pavlovskii Borchsenius and Tereznikova, 1959: 491 (Maritime Territory, USSR) 29. phaseoli (Laing) (Sierra Leone) Phenacoccus phaseoli Laing, 1929: 475 Heliococcus phaseoli (Laing), Goux, 1934: 171 (Based on the large number of dorsal multilocular pores, it is possible that phaseoli does not belong in Heliococcus.) 30. radicicola Goux, 1931: 113 (France; Germany; Poland; Sweden; Cwmea) Vol. 50, No. 2, April 1974 179 31. salviae Borchsenius, 1949: 282 (Tadzhik SSR) 32. saxatilis Borchsenius, 1949: 276 (Armenian SSR) 33. Slavonicus Borchsenius and Tereznikova, 1959: 492 (Ukrainian SSR) 34. stachyos (Ehrhorn) (California) Phenacoccus stachyos Ehrhorn, 1900: 313 Heliococcus stachyos (Ehrhorn), Goux, 1934: 171 35. sulcii Goux, 1934: 167 (France; Germany; Ukrainian SSR) 36. szetshuanensis Borchsenius, 1962: 232 (China) 37. tes quorum Borchsenius, 1949: 284 (Kazakh SSR) 38. wheeleri (King) (Texas) Dactylopius wheeleri King, 1902: 285 Pseudococcus wheeleri (King), MacGillivray, 1921: 133 Heliococcus wheeleri (King), Ferris, 1953: 363 (Described from immatures. The identity of this species is uncertain. Ferris (1953) believed that it might be a senior synonym of insignis.) 39. zizyphi Borchsenius 1958: 161 (China) List of Species Erroneously Placed in Heliococcus 1. hystrix (Baerensprung) = Phenacoccus hystrix (Baerensprung) Coccus hystrix Baerensprung, 1849: 174 Phenacoccus hystrix (Baerensprung), Lindinger, 1912: 293 Heliococcus hystrix (Baerensprung), Thiem, 1930: 140 2. malvastrus McDaniel = Ferrisia virgata (Cockerell) Heliococcus malvastrus McDaniel, 1962: 323; McKenzie, 1967: 181 3. multipori Takahashi Heliococcus multipori Takahashi, 1951: 18 (This species does not belong in Heliococcus because it lacks crateriform tubular ducts. It probably should be placed in Phenacoccus, but without specimens I cannot be positive.) 4. sakai Takahashi Heliococcus sakai Takahashi, 1951:16 (This species does not belong in Heliococcus because it lacks crateri- form tubular ducts. It probably should be placed in Phenacoccus, but without specimens I cannot be positive.) 5. takae (Kuwana) = Saliococcus takae (Kuwana) Dactylopius takae Kuwana, 1907 : 184 Phenacoccus takae (Kuwana) , Kuwana, 1917 : 6 Saliococcus takae (Kuwana), Kanda, 1934: 309 Heliococcus takae (Kuwana), Kanda, 1935: 70 (In 1934 Kanda chose takae to be the type species of Saliococcus Kanda. In 1935 he stated that Saliococcus was a junior, subjective synonym of Heliococcus and placed takae in that genus. The description of Salio- coccus tokyoensis (1959a) indicates that he changed his mind about the status of Saliococcus. The presence of cerarii with more than 3 conical setae easily distinguishes Saliococcus from Hel.iococcus.) 6. takahashii Kanda — Takahashicoccus takahashii (Kanda) Heliococcus takahashii Kanda, 1935: 73 Takahashicoccus takahashii (Kanda), Kanda 1959b: 239 180 The Pan-Pacific Entomologist 7. ivilmattae (Cockerell) — Phenacoccus wilmattae Cockerell Phenacoccus wilmattae Cockerell, 1901: 57 Heliococcus wilmattae (Cockerell), Goux, 1934:171 (This species was placed in Heliococcus because of a misidentification of wilmattae by Bueker (1930).) Key to Adult Females of North American Species of Heliococcus 1 1. With 15 or more pairs of cerarii 3 With less than 10 pairs of cerarii 2 2(1). Large-sized crateriform tubular ducts surrounded by cluster of small- sized ducts; multilocular disk pores absent clemente Miller, n. sp. Large-sized crateriform tubular ducts without cluster of small-sized ducts; multilocular disk pores present near vulva atriplicis McKenzie 3(1). Crateriform tubular ducts of at least 2 sizes 5 Crateriform tubular ducts generally of 1 size, occasionally with 1 or 2 smaller ducts in dorsomedial areas of abdomen 4 4(3). Venter with multilocular disk pores and oral-collar tubular ducts; with at least 10 trilocular pores associated with each cerarius insignis (Lobdell) Venter without multilocular disk pores and oral-collar tubular ducts; with 5 or fewer trilocular pores associated with each cerarius adenostomae McKenzie 5(3). Large-sized crateriform tubular ducts without associated smaller ducts; multilocular disk pores normally present 6 Large-sized crateriform tubular ducts along body margin with as- sociated clusters of smaller ducts; mutilocular disk pores ab- sent deserticola Miller, n. sp. 6(5). Small-sized crateriform tubular ducts much more abundant than large- sized ducts; normally with a cluster of oral-collar tubular ducts near anterolateral margins of abdominal segment IX; oral-collar tubular ducts normally present on venter of abdominal segments VIII-V or IV osborni (Sanders) Small-sized crateriform tubular ducts less abundant or equal in number to large-sized ducts; without a cluster of oral-collar tubular ducts on abdominal segment IX; oral-collar tubular ducts normally restricted to abdominal segments IX or VIII, rarely on VII stachyos (Ehrhorn) Heliococcus clemente, new species (Fig. 1) Suggested common name. — San Clemente mealybug. Type material. — Holotype adult female, Pyramid Head, San Clemente Island, Los Angeles County, California, 12 May 1973, collected under rock, D. R. Miller and A. S. Menlte (USNM). Adult female. — Holotype, mounted, 2.9 mm long, 1.8 mm wide. Body oval; anal lobes strongly protruding and sclerotized, with 3 large-sized crateriform ducts on each lobe. 1 H. ivheeleri is not included because adult females are not available. Vol. 50, No. 2, April 1974 181 Fig. 1. Heliococcus clemente, adult female 182 The Pan-Pacific Entomologist Dorsum with 3 pairs of cerarii on abdomen, 1 pair on head. Anal-lobe cerarii each with 2 elongate, conical setae, 2 or 3 trilocular pores, and large area of basal sclerotization. Remaining abdominal cerarii with progressively smaller conical setae, with or without trilocular pores, without basal sclerotization. Ocular cerarii each with 1 large and 1 small conical seta, 1 or 2 associated trilocular pores, no basal sclerotization. Multilocular disk pores and quinquelocular pores absent. Trilocular pores scattered over surface. Discoidal pores present, few. Crateri- form tubular ducts of 2 sizes: larger size scattered over surface, with 1-4 setae at base of each sclerotized crater, surrounded by 13-35 (av. 25) small-sized ducts; smaller size present in clusters around larger ducts and singly in medial, medio- lateral, and rarely lateral areas; small-sized crateriform ducts without associated setae. Body setae conical. Anal ring bent over apex of abdomen, with 4 or 5 rows of pores; each of 6 anal-ring setae noticeably longer than greatest diameter of ring. Venter with anal-lobe sclerotization elongate. Multilocular disk pores absent. Quinquelocular pores abundant in medial and mediolateral areas, absent else- where. Trilocular pores present in mediolateral and lateral areas. Discoidal pores present in small numbers. Crateriform tubular ducts of same sizes as on dorsum, clusters composed of 1 large- and many small-sized ducts, clusters restricted to near body margins; small-sized ducts lightly scattered over surface, least abundant on medial areas of abdomen. Oral-collar tubular ducts with conspicuous, sclero- tized dermal opening, restricted to mediolateral areas of abdominal segments VIII and VII. Body setae of 2 kinds: bristle-shaped setae present medially and laterally; conical setae present mediolaterally and laterally. Circulus present. Legs with small translucent pores on dorsal surface of hind tibiae; hind tibia/tarsus ratio 1:3.1 and 1:3.2; hind tibia -f- tarsus length 604 and 616 /*; claws with large denticle. Antennae 9-segmented, 823 and 854 g long. Notes. — This is the most unusual known species of Heliococcus in North America because of the large clusters of crateriform tubular ducts. It is most closely related to H. deserticola, herein described as new. H. clemente is also similar to H. atriplicis McKenzie in that both species have less than 15 pairs of cerarii. H. atriplicis differs in lacking crateriform tubular-duct clusters and in having a few multilocular disk pores around the vulva. H. clemente has many crateriform tubular- duct clusters and lacks multilocular disk pores. H. clemente shows a remarkable resemblance to the Russian species H. cydoniae. Both species possess large clusters of crateriform tubular ducts and have small-sized crateriform ducts that lack associated setae. They can be readily separated as follows. H. cydoniae has multilocular disk pores surrounding the vulva, translucent pores on the hind coxae, no pores on the hind tibiae, and 18 pairs of cerarii. H. clemente has no multilocular disk pores, no translucent pores on the hind coxae, pores present on the hind tibiae, and no more than 4 pairs of cerarii. Vol. 50, No. 2, April 1974 183 Heliococcus cleserticola, new species (Fig. 2-5) Suggested common name. — Desert crateriform mealybug. Type material. — Holotype adult female, 10 mi. N. Alamo, Lincoln County, Nevada, 5 July 1970, on Ambrosia dumosa (Compositae) , D. R. Miller, (USNM). Paratypes are as follows: Arizona: Welton, Yuma County, 27 March 1940, on Encelia farinosa (Compositae), R. C. Dickson. California: Niland, Imperial County, 19 March 1971, on Ambrosia dumosa, R. D. Goeden and D. W. Ricker; Picacho Peak, Imperial County, 14 December 1970 and 13 April 1971, on Ambrosia ilicifolia, R. D. Goeden and D. W. Ricker; 3 mi. N. Shoshone, Inyo County, 12 June 1963, on Encelia frutescens and Eriogonum foliolosum (Polygonaceae) , D. R. Miller; Palm Desert, Riverside County, 28 April 1971, beaten from Am- brosia dumosa, R. D. Goeden and D. W. Ricker; Palm Springs, Riverside County, 29 April 1971, beaten from Ambrosia dumosa, R. D. Goeden and D. W. Ricker; Morongo Valley, San Bernardino County, 9 June 1963, on Baccharis sp. (Com- positae), D. R. Miller; Twenty -nine Palms, San Bernardino County, 25 March 1970, 20 May 1970, 20 May 1971, Ambrosia dumosa, R. D. Goeden and D. W. Ricker; Yucca Valley, San Bernardino County, 20 May 1970, Ambrosia dumosa, R. D. Goeden and D. W. Ricker. Nevada: same data as for holotype. Texas: Presidio, Presidio County, 17 May 1951, on Viguiera stenoloba (Compositae), J. H. Russell. Mexico: 10 mi. W. Zacapu, Michoacan, 6 March 1972, under rock, D. R. Miller and F. D. Parker. The 33 paratypes will be placed in the following collections: British Museum (Natural History), London; California Department of Agriculture, Sacramento; Florida State Collection of Arthropods, Gainesville; Museum National D’His- toire Naturelle, Paris, France; Mexican National Collection, Mexico City; Uni- versity of California, Davis; University of California, Riverside; University of Hawaii, Honolulu; U. S. National Museum (Natural History), Beltsville, Mary- land; Virginia Polytechnic Institute and State University, Blacksburg; Zoological Institute, Academy of Sciences, Leningrad, USSR. Adult female (Fig. 2). — Holotype, mounted, 2.6 mm long (paratypes 2. 1-3.8), 1.8 mm wide (paratypes 1. 2-2.7) . Body oval, anal lobes strongly protruding and heavily sclerotized, with 3 large-sized crateriform tubular ducts on each lobe (paratypes with 3-5, av. 3.5). Dorsum with 17 pairs of cerarii. Anal-lobe cerarii each with 2 elongate, conical setae, 4 or 5 trilocular pores, and large area of sclerotization. Remaining cerarii each with 2 somewhat smaller conical setae, 2-4 trilocular pores, and little or no basal sclerotization; ocular cerarii each with 3 conical setae. Multilocular disk pores and quinquelocular pores absent. Trilocular pores abundant. Discoidal pores sparce. Crateriform tubular ducts of 3 sizes: large-sized ducts scattered over surface, most abundant near body margin, with 2-5 associated setae, marginal ducts sometimes surrounded by cluster of 4-11 (av. 7) small-sized crateriform tubular ducts; medium-sized ducts present in medial and mediolateral areas, with 1 or 2 associated setae; small-sized ducts in clusters associated with larger ducts only, normally without associated setae. Body setae conical. Anal-ring apical, bent over apex of abdomen, with 3 or 4 rows of pores; each of 6 anal-ring setae noticeably longer than greatest diameter of ring. 184 The Pan-Pacific Entomologist Fig 2 Heliococcus deserticola, adult female Vol. 50, No. 2, April 1974 185 Venter with conspicuous, elongate anal-lobe sclerotization. Multilocular disk pores absent. Quinquelocular pores present on medial and mediolateral areas. Trilocular pores present in mediolateral and lateral areas, and in medial areas of abdominal segments X-VI. Discoidal pores sparce. Crateriform tubular ducts of 2 sizes, same as larger and smaller sizes on dorsum: larger size always with associated cluster of small-sized ducts, restricted to lateral areas, with 2-6 associated setae; smaller size present in mediolateral and lateral areas, sometimes in clusters around larger ducts, normally without associated setae. Oral-collar tubular ducts with small dermal rim, restricted to mediolateral areas of abdominal segments VIII and VII. Body setae of 2 kinds: bristle-shaped setae present medially and laterally; conical setae present mediolaterally and laterally. Circulus present. Legs with very small translucent pores restricted to dorsal surface of hind tibiae; hind tibia/tarsus ratio 1:3.2 and 1:3.3 (para types 1:3.3- 1:3.6, av. 1:3.5) ; hind tibia tarsus length 573 and 567 /x (paratypes 573-763 /x, av. 671) ; claws each with denticle. Antennae 9-segmented, 713 and 732 /x long (paratypes 683-946 /x, av. 836). Variation . — Paratypes differ from holotype as follows: with 15 to 18 pairs of cerarii, normally with 18, with 2-4' setae, normally 2, in each ocular cerarius, some- times with crateriform tubular-duct clusters more numerous along margin of dor- sum, sometimes with small-sized crateriform ducts sparcely scattered over dorsal surface, occasionally with 1 associated seta, with 2-16 (av. 7) smaller ducts in clusters around larger ducts, with variable numbers of crateriform tubular-duct clusters along body margin, sometimes with ventromedial trilocular pores restricted to posterior 3 abdominal segments, without oral-collar tubular ducts on 1 specimen. Notes . — The above description is based on 16 specimens from 8 localities. H. deserticola is most closely related to H. clemente but differs in pos- sessing 15-18 pairs of cerarii, crateriform tubular-duct clusters which are restricted to the body margin, crateriform clusters which are com- posed of 2-16 (av. 7) small-sized ducts, setae associated with some small-sized crateriform ducts, 3 sizes of crateriform ducts. H. clemente has 3 pairs of cerarii, crateriform tubular-duct clusters which are scattered over the dorsum, crateriform clusters which are composed of 13-35 (av. 25) small-sized ducts, no setae associated with the small- sized ducts, and 2 sizes of crateriform ducts. H. deserticola has been confused with H. stachyos. These species can be separated as follows: H. stachyos normally has multilocular disk pores near the vulva and lacks clusters of crateriform tubular ducts ; H. deserticola lacks multilocular pores and has clusters of crateri- form tubular ducts. H. deserticola shows a remarkable resemblance to the Russian species H. kurilensis in that both have clusters of small-sized crateriform tubular ducts surrounding a single larger duct. Unlike H. clemente and H. 186 The Pan-Pacific Entomologist Fig. 3. Heliococcus deserticola, third instar female. Vol. 50, No. 2, April 1974 187 Fig. 4. Heliococcus deserticola, second instar male 188 The Pan-Pacific Entomologist cydoniae, H. deserticola and H. kurilensis have only small clusters of these ducts. H. kurilensis differs from H. deserticola in having the crateriform clusters scattered over the entire dorsum and in having multilocular disk pores near the vulva; H. deserticola has the crateri- form clusters restricted to the marginal areas and has no multilocular disk pores. Third instar female (Fig. 3). — Same as for adult female except as follows: mounted, 1. 4-2.1 mm long, 0.9-1. 3 mm wide. Anal lobes each with 2 large-sized crateriform tubular ducts. Dorsum with 18 pairs of cerarii. Anal-lobe cerarii each with 2-5 trilocular pores. Discoidal pores absent or sparce. Crateriform tubular ducts of 3 sizes: larger size with marginal ducts sometimes with 2-10 (av. 5.1) associated smaller- sized crateriform tubular ducts; medium-sized ducts present in medial areas of some posterior abdominal segments; small-sized ducts associated with larger ducts and scattered in small numbers in mediolateral and lateral areas. Venter with crateriform tubular ducts of 2 sizes: larger size present on margin of abdominal segment VII, surrounded by cluster of small-sized ducts. Oral- collar tubular ducts absent. Legs without translucent pores; hind tibia/tarsus ratio 1: 1.7-1: 1.8 (av. 1:1.8) ; hind tibia + tarsus length 311-415 (av. 366) /a. Antennae 7- or 8-segmented, 421- 519 (av. 458) ^ long. Notes . — The above description is based on 2 specimens from 2 localities. Second instar male (Fig. 4). — Same as for adult female except as follows: mounted 1.1 mm long, 0.7 mm wide. Anal lobes each with 1 large-sized crateri- form tubular duct. Dorsum with 17 pairs of cerarii. Anal-lobe cerarii each with 2 or 3 trilocular pores. Remaining cerarii sometimes indefinite, with widely separated cerarian setae. Crateriform tubular ducts of 2 sizes: large-sized ducts in small numbers over surface, with 1-4 associated setae, marginal ducts occasionally with 1-3 as- sociated small-sized ducts; small-sized ducts scattered over surface. Oral-collar tubular ducts of 2 sizes: shorter ducts restricted to lateral areas of anterior ab- dominal segments, thorax, and head, of same size and shape as those on dorsum; longer ducts present medially near legs. Legs without translucent pores; hind tibia/tarsus ratio 1:1.4; hind tibia -f- tarsus length 226 (jl . Antennae 7-segmented, 317 /j. long. Notes . — The above description is based on 1 specimen. First instar (Fig. 5; Sexes not distinguished). — Same as for adult female ex- cept as follows: mounted, 0.5 mm long, 0.2-0.3 mm wide. Anal lobes lightly sclerotized, without crateriform tubular ducts. Dorsum with 16—18 indefinite cerarii. Anal-lobe cerarii each with 1 trilocular pore. Remaining cerarii with 1 associated trilocular pore of same size and shape as other triloculars; ocular cerarii normally absent, when present, with 2 conical setae. Trilocular pores scattered over surface, arranged in 4 or 5 pairs of longi- tudinal lines on abdomen. Discoidal pores and crateriform tubular ducts absent. Body setae arranged in 4 pairs of longitudinal lines on abdomen and head, median lines sometimes with additional row on thorax. Anal ring with 2 rows of pores. Vol. 50, No. 2, April 1974 189 190 The Pan-Pacific Entomologist Venter with quinquelocular pores normally distributed as in figure 5, sometimes with only 1 pore near mouthparts, occasionally with 3 pores near anterior spiracle and 2 near posterior spiracle. Trilocular pores nearly always present near spiracles, rarely with 1 or 2 present near body margin. Discoidal pores restricted to medio- lateral areas. Crateriform and oral-collar tubular ducts absent. Body setae bristle shaped. Legs without translucent pores; hind tibia/tarsus ratio 1: 1.0-1: 1.1 (av. 1:1.1) ; hind tibia -f- tarsus length 168-188 (av. 177) /x. Antennae 6-segmented, 225- 243 (av. 234) /x long. Notes. — The above description is based on 8 specimens from 2 localities. First instar nymphs of North American Heliococcus species have not been described previously. The first instars of Heliococcus deserticola , Phenacoccus dearnessi King (see Miller and Appleby, 1971), and Heterococcus spp. (see Miller, in press) are similar. The first instar of H. deserticola differs from that of P. dearnessi and Heterococcus spp. in having a circulus, conspicuous ostioles, and one size of trilocular pore. Acknowledgments I thank Robert 0. Schuster, University of California, Davis, Richard D. Goeden and Donald W. Ricker, University of California, Riverside, for the loan of specimens. I thank my wife, Judy, for her assistance in typing the drafts of this manuscript and for her comments and criticisms. To Louise M. Russell, Ronald W. Hodges, Robert W. Carlson, and Richard E. White, System- atic Entomology Laboratory, ARS, USD A, I give thanks for their criticisms of this manuscript. Literature Cited Baerensprung, F. von 1849. Beobachtung fiber einige einheimische Arten aus der Familie der Coccinen. Ztg. Zool. Zoot. Palaeozool., 1: 165-170, 173-176. Balachowsky, A. S. 1953. Sur un Heliococcus Sulc nouveau, vivant a 3600 metres d’altitude dans les Alpes du Dauphine et du Tyrol. Mitt. Schweiz. Entomol. Ges., 26: 236-240. Bazarov, B. 1963. Two new species of coccids from Tadzhikistan. (In Russian.) Akad. Nauk Tadzhikskoi SSR Dokl., 6: 38-42. Borchsenius, N. S. 1941. New pest of mulberry — the central Asiatic cochineal, Heliococcus destructor Borchs. from the Republic of Middle Asia. (In Russian.) Guidebook for Plant Quarantine, 1: 6-7. Borchsenius, N. S. 1949. Fauna of USSR. Homoptera, Pseudococcidae. (In Russian.) Akad. Nauk Zool. Inst. (n.s. 38), 7: 1-383. Vol. 50, No. 2, April 1974 191 Borchsenius, N. S. 1956. Notes on the Coccoidea of Korea. Entomol. Obozr., 35: 671-679. Borchsenius, N. S. 1958. Notes on the Coccoidea of China. II. Descriptions of some new species of Pseudococcidae, Aclerdidae and Diaspididae. (In Russian.) Entomol. Obozr., 37: 156-173. Borchsenius, N. S. 1962. Notes on the Coccidea of China. XI. New genera and species of Pseudococcidae. (In Russian.) (Akad. Nauk SSSR Zook Inst. Trudy, 30: 221-244. Borchsenius, N. S. and E. M. Tereznikova. 1959. Two new species of mealy bugs of the genus Heliococcus Sulc. (In Russian.) Zool. Zhur., 38: 491-494. Bueker, E. D. 1930. Phenacoccus wilmattae Ckll. Can. Entomol., 62: 93-94. Cockerell, T. D. A. 1901. The New Mexico Coccidae of the genus Ripersia. Ann. Mag. Nat. Hist. (ser. 7), 8: 51-57. Danzig, E. M. 1971. New and little-known species of mealy-bugs from the far east of USSR. (In Russian.) Entomol. Obozr., 50: 366-390. Danzig, E. M. 1972. Contribution to the fauna of the white flies and scale in- sects of Mongolia. (In Russian.) Nasekomye Mongolii, Mongol Orny Shav’zh pp. 325-348. Ehrhorn, E. M. 1900. New Coccidae from California. Can. Entomol., 32: 311-318. Ferris, G. F. 1950. Atlas of the scale insects of North America. Series V. Pseudococcidae (Part I). Stanford Univ. Press, California, 278 pp. Ferris, G. F. 1953. Atlas of the scale insects of North America. Volume VI. The Pseudococcidae (Part II). Stanford Univ. Press, California, pp. 279-506. Goux, L. 1931. Notes sur les coccides de la France. (3 e note). Description d’une espece nouvelle. Bull. Soc. Entomol. Fr., 1931: 113-118. Goux, L. 1934. Notes sur les coccides de la France. (9 e note) . Contribution a l’etude du genre Heliococcus avec description de deux especes nou- velles. Bull. Soc. Entomol. Fr., 39: 164^171. Goux, L. 1953. Notes sur les coccides de la France (42 me note). Etude d’un Heliococcus nouveau. Bull. Mens. Soc. Linn. Lyon, 22: 104^-109. Green, E. E. 1925. Notes on the Coccidae of Guernsey (Channel Islands), with descriptions of some new species. Ann. Mag. Nat. Hist., 16: 516- 527. Hollinger, A. H. 1917. A new species of Phenacoccus. Can. Entomol., 49: 281-284. Kanda, S. 1934. Two new genera of Japanese Coccidae. Insect World, 38: 308-313. Kanda, S. 1935. On the genus Heliococcus. Annot. Zool. Jap., 15: 70-77. Kanda, S. 1959a. Two new species of the Pseudoccidae. Kontyu, 27: 177-180. Kanda, S. 1959b. Two new genera and two new species of the Pseudococcidae. Kontyu, 27: 239-243. King, G. B. 1902. Two new ants’-nest coccids from Texas. Can. Entomol., 34: 285-286. Kuwana, S. I. 1907. Coccidae of Japan, I. A synoptic list of Coccidae of Japan with descriptions of thirteen new species. Imp. Centr. Agric. Exp. Stn. Bull., 1: 177-207. 192 The Pan-Pacific Entomologist Kuwana, S. I. 1917. A check list of the Japanese Coccidae. Yokohama, Japan. 19 pp ‘ Laing, F. 1929. Descriptions of new and some notes on old species of Coccidae. Ann. Mag. Nat. Hist., 4: 465-501. Lindinger, L. 1912. Die Schildlause Europas, Nordafrikas und Vorderasiens, einschlieblich der Azoren, der Kanaren und Madeiras. Ulmer, Stuttgart. 388 pp. Lobdell, G. H. 1930. Twelve new mealybugs from Mississippi. Ann. Entomol. Soc. Amer., 23: 209-236. MacGillivray, A. D. 1921. The Coccidae. Scarab Co., Illinois. 502 pp. McDaniel, B. 1962. A new species of scale insect from the Rio Grande Valley, Texas. Tex. J. Sci., 14: 323-327. McKenzie, H. L. 1960. Taxonomic study of California mealybugs with descrip- tions of new species. Hilgardia, 29: 681-770. McKenzie, H. L. 1964. Fourth taxonomic study of California mealybugs, with additional species from North America, South America, and Japan. Hilgardia, 35: 211-272. McKenzie, H. L. 1967. Mealybugs of California with taxonomy, biology, and control of North American species. Univ. California Press, California. 525 pp. Miller, D. R. 1974. A systematic revision of the genus Heterococcus , with a diagnosis of the genus Brevennia. (in press) Miller, D. R. and J. E. Appleby. 1971. A redescription of Phenacoccus dear- nessi. Ann. Entomol. Soc. Amer., 64: 1342-1357. Sanders, J. G. 1902. A new Phenacoccus on Platanus occidentalis. Ohio Nat., 2: 284-286. Sulc, K. 1912. Heliococcus bohemicus n. gen. n. sp. Cesk. Spolec. Entomol. Casopis., 9: 39-48. Takahashi, R. 1930. Observations on the Coccidae of Formosa. Part II. Dept. Agric. Gov. Res. Inst. Formosa, Rpt. no. 48, 45 pp. Takahashi, R. 1951. Some mealy bugs from the Malay Peninsula. Entomol. Soc. India (1950), 12: 1-22. Ter-Grigorian, M. A. 1967. Two new species of the genus Heliococcus Sulc from Armenia. (In Russian.) Dokl. Akad. Nauk Arm. SSR, 44: 134- 138. Thiem, H. 1930. Die wichtigsten einheimischen Schildlause. Dtsch. Landwirtsch. Presse., 57 : 140. Williams, D. J. 1962. The British Pseudococcidae. Bull. Br. Mus. (Nat. Hist.) Entomol., 12: 1-79. Vol. 50, No. 2, April 1974 193 Mealybugs of San Clemente Island, California (Homoptera: Coccoidea: Pseudococcidae) Douglass R. Miller Systematic Entomology Laboratory, USDA, ARS, Beltsville, Maryland 20705 In earlier papers (Miller, 1971, 1973) I described the mealybug faunas of Santa Cruz and San Miguel Islands and presented brief dis- cussions relating to the Southern California Island biota. In April 1973 Dr. Arnold Menke and I collected on San Clemente Island for eight days. Although this was an inadequate amount of time to thoroughly sample the island pseudococcids, the material ob- tained should allow some tentative comparisons of the mealybug fauna of this island with those of Santa Cruz and San Miguel Islands. San Clemente Island is located approximately 49 statute miles from the nearest point on the mainland and is 21 miles from the closest island, Santa Catalina, which intervenes between the nearest mainland point and San Clemente. The island is approximately 21 miles long, about 4 miles across at its widest point, and approximately 56 square miles in land area (Philbrick, 1967). The long axis of the island is oriented approximately northwest, nearly parallel to the near mainland coast. Of the several geological studies of the island, two of the more detailed ones were published by Olmstead (1958) and Smith (1898). Geologically, San Clemente is the emergent portion of a tilted, slightly arched block of the earth’s crust. A major fault is present on the sea floor at the northeastern edge of the San Clemente block, and, although movement of this fault is thought to be primarily horizontal, nearly the entire northeastern side of San Clemente Island is a single, large fault escarpment. The northeastern side of the island is very steep in contrast to the southwestern side which generally slopes gently. Both sides of the island are deeply cut by geologically young, often precip- itous canyons. The highest point is Thirst (elevation 1,967 feet), located near the northeastern escarpment. The southwestern side of the island, up to 1,500 feet, shows some of the best examples of marine terracing on the Southern California Islands, with as many as 20 separ- ate terraces visible on some slopes. The island is principally com- posed of volcanic rocks of probable Miocene age. Sedimentary marine deposits (Miocene), sand deposits (Pleistocene? and Recent), and alluvial deposits (Recent) overlying the volcanic rocks are scattered locally on the island. Normally the soil covering is deposited in rela- tively thin layers and has a high clay content. San Clemente is apparently The Pan-Pacific Entomologist 50 : 193 - 202 . April 1974 194 The Pan-Pacific Entomologist no older than the Miocene and may be as recent as the Early to Mid- Pliocene. Most evidence suggests that this island has never been con- nected with the mainland or the other islands, although at times of lower sea levels, particularly during the Pleistocene, San Clemente undoubtedly was closer to other land masses than at present. As is evident from the lack of marine deposits and marine terraces above 1,500 feet elevation, the areas of the island above this level have probably been continually emergent since Miocene or Pliocene times. Data on the weather of the island are scant, and have been sum- marized by Raven (1963) as follows: “ . . . the east (northwestern) side of the island is apparently moister than the west (northeastern), and it is likely that the main tracks of the storms from north to south are often deflected down along the east side of the island. Generally the wind is from the sea, over the island and onto the mainland; but during the Santa Ana cycles of winter, violent and sustained winds blow off the mainland and from Santa Catalina Island toward the east side of San Clemente. These Santa Ana winds may obviously be of importance in transporting seeds (or first instar mealybugs) to the is- land.” Most of the rainfall occurs during the winter months, but mois- ture producing fogs are common during rain-free times of the year. No permanent streams are present although intermittent flows are common during the rainy season. Olmstead (1958) stated that the deep plunge pools formed by intermittent waterfalls contain water throughout the year. Additional small impoundments have been con- structed, but basically the island is without water during the dry months. The predominant habitat types on the island are flat grassland areas, hilly cactus areas, deep precipitous canyons, steep cliffs, volcanic re- gions often with many small caves, recent sand dunes, fossil sand dunes, sand beaches, and shingle beaches. The island reptile fauna includes only two lizard species, one belongs to an endemic genus known from San Clemente, San Nicolas, and Santa Barbara Islands ; there are no amphibians ( Savage, 1967 ) . The indigenous terrestrial mammal fauna of the island is composed of four bat species, three mice species including one endemic sub-species, and one fox species which is an insular endemic with an endemic race on San Clemente (von Bloeker, 1967). The mammals introduced by man are pigs, goats, cats, and deer. The aquatic mammal fauna consists of three species of pinnipeds (Bartholomew, 1967) and the sea otter (von Bloeker, 1967). Diamond (1969) reported 24 species of land birds on San Clemente in 1968. To my knowledge no detailed study of the Vol. 50, No. 2, April 1974 195 insect fauna of San Clemente has been published, although several short papers have appeared in the series “Contributions from the Los Angeles Museum — Channel Islands Biological Survey.” Most of the native vegetation of the island has been ravaged by the feeding of a large population of feral goats. Because of this, the is- land is sparsely vegetated, and introduced herbs, mostly grasses, pre- dominate. Plant species once apparently widespread are now either extinct or are restricted to small populations in areas inacessible to goats. Some of the best refuges on the island are located in the pre- cipitous canyons, particularly on the northeastern side. The upper reaches of these canyons are of particular interest because they have not been inundated by the sea since Miocene or Pliocene times. A flora of the island published by Raven (1963) is the source of much valuable information. Additions to the flora have been published by Raven (1965, 1967) and Thorne (1969). Of the Channel Islands, San Clemente has the most interesting flora, despite the larger land area and seemingly greater habitat diversity of some of the other is- lands. It supports more insular endemics (43, 13 of which occur ex- clusively on San Clemente) than the other Channel Islands, evidently because its isolation has allowed little floral interchange with other land areas. San Clemente Island Mealybugs Knowledge of the unusual San Clemente flora and of the similarities between the dispersal mechanisms of scale insects and plants (Miller, 1971), encouraged me to look to San Clemente as the most promising source of unusual mealybugs in the Southern Channel Islands. Un- fortunately, collecting mealybugs on this island is difficult because most of the more interesting plants occur in nearly inaccessible areas and the heavy clay soil generally present is unsuitable for most pseu- dococcid species. The following list includes 9 genera and 11 species. No mealybugs have been reported from this island previously. Plant names are as listed by Raven (1963). Chorizococcus cibroniae McKenzie 1. Shifting Sands, 14 May 1973, Abronia. umbellata (roots and decumbent stems) Discococcus simplex Ferris 1. 0.25 mi. N. Boulder, 10 May 1973, Stipa sp. and Avena sp. (on crown) 2. canyon N. Boulder, 11 May 1973, Stipa sp. (in sheaths near crown) 196 The Pan-Pacific Entomologist 3. China Canyon, 13 May 1973, Stipa sp. (in sheaths) 4. Eel Point, 8 May 1973, Stipa sp. (in sheaths near crown) 5. Pyramid Head, 12 May 1973, Stipa sp. (on crown) 6. Seal Cove, 9 May 1973, Stipa sp. (in sheaths) 7. near mouth of Wall Rock Canyon, 9 May 1973, Stipa sp. (on crown) 8. near White Rock, 10 May 1973, Stipa sp. and Avena sp. (in sheaths near crown) 9. Wilson Cove, 8 May 1973, Avena sp. (on crown) Distichlicoccus salinus ( Cockerell ) 1. China Point, 13 May 1973, Distichlis spicata (on leaf blade) 2. Shifting Sands, 14 May 1973, Distichlis spicata (on leaf blade) Heliococcus clemente Miller 1. Pyramid Head, 12 May 1973, under rock Paludicoccus distichlium (Kuwana) 1. China Point, 13 May 1973, Distichlis spicata (in leaf blade sheath) Phenacoccus eschscholtziae McKenzie 1. China Point, 13 May 1973, Lotus argophyllus subsp. ornithopus (?) (on main roots) solani Ferris 1. S. West Cove, 7 May 1973, Franseria chamissonis (decumbent branches) 2. 2 mi. SE West Cove, 11 May 1973, Astragalus miguelensis and Atriplex semibaccata (roots) Puto yuccae (Coquillett) 1. Eel Point, 8 May 1973, Suaeda calif ornica (on roots) Spilococcus corticosus McKenzie 1. canyon N. Boulder, 11 May 1973, Artemisia calif ornica (under bark on stems) keiferi McKenzie 1. China Point, 13 May 1973, Lotus argophyllus subsp. ornithopus (?) and Suaeda californica (on roots) 2. Eel Point, 9 May 1973, Astragalus miguelensis, Atriplex semibaccata, and Suaeda californica (on roots) 3. Pyramid Head, 12 May 1973, Phacelia floribunda (?) and, Suaeda cal- ifornica (on roots) , and under rock 4. near mouth of Wall Rock Canyon, 9 May 1973, Suaeda californica (on roots) 5. 2 mi. SE West Cove, 11 May 1973, Suaeda californica (on roots) Trionymus smithii (Essig) 1. Wilson Cove, 7 May 1973, Avena sp. (leaf sheath) Vol. 50, No. 2, April 1974 197 Discussion Perhaps my most exciting find on San Clemente was the endemic mealybug, Heliococcus clemente Miller. I believe this is the first of several endemics which may be discovered when the canyons on the northeastern side of the island are intensively collected. H. clemente is most similar to Heliococcus deserticola Miller, a species that occurs in the deserts of the southwestern U. S. and is relatively close to another desert species, Heliococcus atriplicis McKenzie. H. clemente appears to be more similar to xeric southern desert species than to the more common and widespread mesic species (such as H. adenostomae Mc- kenzie, H. osborni (Sanders), or H. stachyos (Ehrhorn). An unidentified species, apparently a second endemic, has been en- countered on both San Clemente and Santa Cruz Islands. On Santa Cruz it was collected at several localities ranging from the higher eleva- tions of the north ridge to the coastal areas near Coches Prietos. It has been collected in spring on a varied range of hosts, on rootlets in heavy, clay soil. On San Clemente, the apparent same species was found near Boulder, in heavy, clay soil on small roots of Malacothrix foliosa (Compositae) , Lupinus bicolor (Leguminosae) , and other small dicoty- ledonous plants. I found only first instar nymphs at both locations, and suspect that adults occur much later in the year. Until adult fe- males can be found, this pseudococcid should remain unnamed, but it is so different from the known first instar nymphs of any other North American mealybug, that I am fairly certain it is a Channel Island insular endemic. It is quite probable that this mealybug represents an endemic genus. Using the work of McKenzie (1967) as a source for California pseudococcid distribution patterns, the 10 nonendemic species occurring on San Clemente are distributed on the mainland as follows: one is present throughout most of continental California, three occur exclu- sively in coastal mountain ranges, one occurs in the coastal ranges and the southern California deserts, two occur in the coastal ranges and the ocean beach areas, two occur in saline areas, and one occurs exclusively on ocean beach areas. These data suggest that most mealybug species known to occur on San Clemente Island are most closely allied to the aggregation of species found in coastal areas on the California main- land; in fact, all of the known San Clemente species except H. clemente are known to inhabit coastal areas in part. One species shows a disjunct distribution pattern. Discococcus sim- 198 The Pan-Pacific Entomologist plex is not known on the portion of the mainland closest to San Cle- mente Island but occurs approximately 190 miles to the north. With the limited amount of material available, it has not been possible to make general statements concerning the number of species on San Clemente compared to a similarly sized area on the mainland. How- ever, the absolute number of individual specimens on the island is small compared to the mainland because an average of only three mealy- bug collections per day was made on San Clemente, whereas I find an average of about ten mealybug collections per day on the California mainland. Discococcus simplex is unusually abundant on San Clemente. Pre- viously, this species was known by less than 10 specimens from two mainland localities. On San Clemente, with some diligence, D. simplex can be collected in most grassland areas on the island; it is normally quite common on Stipa spp. The same species was collected once on another island (Santa Cruz), where apparently it is rare. Spilococcus keiferi is also unusually abundant. It is a fairly com- mon coastal mainland pseudococcid which seems to prefer the roots and decumbent branches of Franseria chamissonis in sandy soils. It has occasionally been found on a few other hosts and has been collected inland in two instances. Under normal circumstances only one or two specimens of S. keiferi are collected at mainland localities, and they usually are mixed with specimens of Chorizococcus abroniae. On San Clemente this species seems to behave differently. Next to Discococcus simplex, it was the most commonly encountered mealybug on the is- land. It was collected on five plant species which are all new host records. Although populations of Franseria chamissonis were examined exten- sively, S. keiferi was never found on this host. Infestations of S. keiferi normally were very large, and they never were mixed with Chorizococcus abroniae. On San Miguel and Santa Cruz Islands S. keiferi did not show characteristics of the San Clemente populations. Similar insular differences in relative abundance and host preference are known to occur in island populations of Puto yuccae. On the main- land P. yuccae occurs on a variety of hosts and is common in many habitats including the Sierra Nevada up to 6,000 feet, in chaparral areas of the coastal mountain ranges, and in sand areas near the ocean. On San Clemente the species was collected only once, even though pre- ferred hosts were examined extensively. On San Miguel P. yuccae was much more abundant. Although it apparently is restricted to sandy Vol. 50, No. 2, April 1974 199 beach situations, it occurs on a variety of hosts and is quite common. On Santa Cruz it is more abundant than on the other two islands. It occurs in several types of habitats and has been recorded on seven dif- ferent host plant species. The mechanisms affecting differences in habitat preference, host diversity, and relative abundance of mealybug species on islands appear to be complex. A study of these mechanisms would undoubtedly give insight into aspects of pseudoccid biology and ecology not presently understood. In previous papers on the Southern California Islands, I have not mentioned morphological variation of insular pseudococcid species, although some island populations are highly variable. At times this variation has caused problems in placing the island species, but this problem was anticipated. If the Southern California Islands possess endemic forms at generic and specific levels, then they should contain examples of endemism at intraspecific levels. In fact, this endemism should be more common than at higher levels. Variation observed in three mealybug species bears mention. Material identified as Puto yuccae has shown considerable variation from island to island and from island to mainland. Some insular specimens possess features of both Puto echinatus McKenzie, a coastal mainland species, and P. yuccae. The greatest amount of variation was noted on specimens from San Miguel Island where typical representatives of both P. ecliinatus and P. yuccae were found, in addition to many intermediate individuals. On Santa Cruz variation was less marked, but still considerable. On San Clemente P. yuccae possessed key characters of both species. With- out more material, particularly from the mainland coastal areas, the identity of this species must remain tentative. There are a number of possible explanations for this type of in- sular variation. For example, P. echinatus may be a recent coastal de- rivative of the inland species P. yuccae , or vice versa, with intermediate forms persisting on the islands; P. echinatus and P. yuccae may be the same species; annectant forms may be hybrids; and there are other possibilities. Variation seen in insular populations of Spilococcus keiferi and Trionymus smithii show the same general pattern as mentioned above. S. keiferi varies between typical S. keiferi and individuals approaching S. atriplicis (Cockerell) or S. nototrichus Miller and Mc- Kenzie. T. smithii varies between specimens typical of California forms of T. smithii and material approaching T. winnemucae McKenzie or T. nanus Cockerell. Although annectant specimens cause concern to 200 The Pan-Pacific Entomologist the taxonomist, they point out possible relationships which otherwise might go undetected. Comparison of the Mealybug Faunas of San Clemente, San Miguel, and Santa Cruz Islands Notable similarities exist in the three pseudococcid faunas. (1) All are depauperate aggregations of species similar to the pseudococcid fauna on the adjacent southern California mainland. (2) All are pri- marily composed of coastal or coastal mountain species. (3) All contain representatives of the following species: Chorizococcus abroniae, Dis- tichlicoccus salinus, Paludicoccus distichlium, Puto yuccae, and Spilo- coccus keiferi. The mealybug fauna of San Clemente most closely resembles the fauna found on Santa Cruz. Of the 12 species (including the unde- scribed form) known to occur on San Clemente, nine occur on Santa Cruz while only six are known on San Miguel. The pseudococcids of both San Clemente and Santa Cruz, with the exception of Discococcus simplex on San Clemente, occur in localized areas; whereas on San Miguel, several species are widespread and common. Both San Clemente and Santa Cruz have representative species which do not occur on the adjacent mainland but are found farther north in California, and both islands have specimens of the apparent insular endemic known only from immatures. A feature found within the mealybug faunas of San Clemente and San Miguel, but not within the Santa Cruz fauna, is the presence of species which show more diversity in habitat and host preference than they do on the mainland. The San Clemente pseudococcid fauna is unique in the presence of an endemic species. Its resemblance to species occurring in the desert areas of the southwestern U. S. suggests the presence of southern xeric elements within the San Clemente fauna. Summary Nine genera and 11 species of mealybugs are known on San Clemente Island. Heliococcus clemente is endemic, and another probable insular endemic known only from immatures, is reported from San Clemente and Santa Cruz Islands. The pseudococcid fauna of San Clemente is most like the aggregation of mealybug species found on the adjacent mainland, but contains elements of both northern and southern faunas. The flora and mealybug fauna of San Clemente have many similar attributes. Both show affinities to their respective counterparts on the Vol. 50, No. 2, April 1974 201 adjacent mainland, both contain endemics, both possess species with disjunct distribution patterns, and both contain small components of northern and southern biotas. Acknowledgments I am grateful to Lt. Commander W. G. Kay, Officer in Charge, San Clemente Island Facility, for his assistance and cooperation during my stay on San Clemente. Thanks are also due to Chief West who made some of the arrangements for transportation on the island, pointed out areas which might be of special interest, and allowed us to collect in normally inaccessible regions. Special appreciation is expressed to Dr. Arnold S. Menke for his assistance in collecting scale insects while on the island. I also thank my wife Judy Miller for her help in typing the drafts of this article and for her comments on its content. I appreciate the help of Louise M. Russell, Raymond J. Gagne, Douglas C. Ferguson, and Ronald W. Hodges, Systematic Entomology Laboratory, ARS, USDA, for their careful criticisms of this manuscript. Literature Cited Bartholomew, G. A., 1967. Seal and sea lion populations of the California Islands. Pp. 229-244 in R. N. Philbrick ( ed. ) , (For complete citation see Philbrick) . Diamond, J. M. 1969. Avifauna equilibria and species turnover rates on the Channel Islands of California. Proc. Nat. Acad. Sci., 64(1) : 57-63. McKenzie, H. L. 1967. Mealybugs of California with taxonomy, biology, and control of North American species. Univ. Calif. Press, Berkeley, 525 pp. Miller, D. R. 1971. Mealybugs of Santa Cruz Island, California. Pan-Pac. Entomol., 47 (4) : 293-303. 1973. Mealybugs of San Miguel Island, California. Pan-Pac. Entomol., 49(3) : 264-269. Olmstead, F. H. 1958. Geologic reconnaissance of San Clemente Island, Cal- ifornia. Geol. Surv. Bull., 1071-B: 55-68. Philbrick, R. N. Introduction. Pp. 3-8 in R. N. Philbrick (ed.), Proceedings of the Symposium on the Biology of the Californa Islands. Santa Barbara Botanic Garden, Calif., 341 pp. Raven, P. H. 1963. A flora of San Clemente Island, California. Aliso, 5(3) : 289-347. 1965. Notes on the flora of San Clemente Island, California. Aliso 6: 11. 1967. The floristics of the California Islands. Pp. 57-67 in R. N. Philbrick (ed.), (For complete citation see Philbrick). Savage, J. M. 1967. Evolution of the insular herpetofaunas. Pp. 219-227 in R. N. Philbrick (ed.), (For complete citation see Philbrick). 202 The Pan-Pacific Entomologist Smith, W. S. T. 1898. A geological sketch of San Clemente Island. U. S. Geol. Surv., Ann. Rpt. 18, (1896/97) pt. 2, pp. 459-496. Thorne, R. F. 1969. A supplement to the floras of Santa Catalina and San Clemente Islands, Los Angeles County, California. Aliso, 7 : 73-83. von Bloeker, J. C., JR. 1967. Land mammals of the Southern California Islands. Pp. 245-263 in R. N. Philbrick (ed.), (For complete citation see Philbrick ) . DICTIONARIES WEBSTER Library size 1973 edition, brand new, still in box. Cost New $45.00 Will Sell for $15 Deduct 10% on orders of 6 or more Make Checks Payable to DICTIONARY LIQUIDATION and mail to Attention: Dept. D-146 ONTARIO TEXT EDITIONS Toronto Dominion Center Suite 1400, Fourteenth Floor Toronto, Ontario, Canada M5K IB 7 C.O.D. orders enclose 1.00 good will deposit. Pay balance plus C.O.D. shipping on delivery. Be satisfied on inspection or return within 10 days for full refund. No dealers, each volume specifically stamped not for resale. Please add $1.25 postage and handling. Vol. 50, No. 2, April 1974 203 SCIENTIFIC NOTES The American Toad as Champion Carabid Beetle Collector. — The Ameri- can Toad ( Bufo americanus Holbrook) has been reported as a predator of Carab- idae (Coleoptera) by Kirkland (1897. Hatch Exp. Sta. Mass. Agric. Bull., 46: 1-30; 1904. U. S. Dept. Agric. Farmer’s Bull., 196: 1-16), Garman (1901. Kentucky Agric. Exp. Sta. Bull., 91: 60-68), Stoner (1937. Florida Entomol., 19: 49-53), Smith and Bragg (1949. Ecol., 30: 333-349), and also by Bush (1959. Herpetol., 15: 73-77). From May to October 1973, I collected and analyzed the stomach contents of 342 American toads from southern Quebec in order to determine the extent of predation on carabid beetles. Immediately after capture, the animals were killed in hot water, their stomachs removed and the contents emptied into a small dish of water. The total number of species of carabid beetles in the toads was 98, and the maximum of individuals in one toad was 17. In addition, remains of 19 species were found in toads’ excreta. The diet of the American toad is de- termined mostly by relative availability of prey items (Smith and Bragg 1949), and selectivity of prey is probably a factor of minor importance. The amphibians live, on the whole, in the same habitats as carabid beetles, and both are predomi- nantly active nocturnally, partly explaining the large consumption of Carabidae by toads. The number of each species of prey is listed below. Numbers in parentheses indicate additional individuals found in excreta. Agonurn sp., 6 (28) , A. creni- striatum Leconte, 47, A. cupripenne Say, 12 (1), A. decentis Say, 1, A. harrisi Le- conte, 1, A. melanarium Dejean, 8, A. placidum Say, 41 (6), A. propinqiium Gem- minger and Harold, 1, A. puncticeps Casey, 1, A. retractum Leconte, 6, Amara sp., 40 (8), A. aenea DeGeer, 41 (8), A. angustata Say, 2, A. apricciria Paykull, 2, A. avida Say, 3, A. convexa Leconte, 2, A. discors Kirby, 2, A. familiaris Duftschmid, 1, A. impuncticollis Say, 3, A. latior Kirby, 4 (1), A. littoralis Mannerheim, 1, A. musculis Say, 5, A. obesa Say, 14 (1). A. quenseli Schonherr, 33 (8), A. rubrica Haldeman, 2, Anisodactylus discouleus Dejean, 1 , A. harrisi Leconte, 5, A. kirbyi Lindroth, 1, A. rnerula Germar, 21, A. nigerrimus Dejean, 2, A. nigrita Dejean, 1, A. rusticus Say, 3, A. sanctaecrucis Fabricius, 6, Anisotarsus nitidipennis Leconte, 2, A. terminatus Say, 2, Badister notatus Haldeman, 1 , Bembidion frontale Leconte, 2, B. nitidum Kirby, 3, B. patruele Dejean, 2, B. punctato striatum Say, 1, B. quadri- maculatum oppositum Say, 8, B. stephensi Crotch, 1, B. tetracolum Say, 2, Bradycel- lus sp., 1, B. nigriceps Leconte, 1, B. rupestris Say, 1, B. semipubescens Lindroth, 1, Calathus sp., 1, C. gregarius Say, 2, C. ingratus Say, 1 (2), Calleida punctata Le- conte, 3, Calosoma calidum Fabricius, 1 (1), Carabus nemoralis Muller, 4, C. ser- ratus Say, (1), Chlaenius lithophilus Say, 2, C. pensylvanicus Say, 1, C. tomentosus Say, 4, C. tricolor Dejean, 1, Clivina sp., 1, C. fossor Linne, 13, Cymindis americana Dejean, 1, C. borealis Leconte, 1, C. cribricollis Dejean, 3 (4), C. pilosa Say, 1, Dicaelus politus Dejean, 1, Diplocheila obtusa Leconte, 2, Dromius piceus Dejean, 1, Dyschirius globulosus Say, 4, Geopinus incrassatus Dejean, 22, Harpalus sp., 6 (3), H. affinis Schrank, 72 (11), H. bicolor Fabricius, 9, H. caliginosus Fabricius, 1, H. egregius Casey, (1), H. erraticus Say, 12, H. fallax Leconte, 1, H. fuliginosus Duft- schmid, (1), H. funerarius Csiki, 1, H. herbivagus Say, 13, H. indigens Casey, 7, H. lewisi Leconte, 3, H. opacipennis Haldeman, 5, H. pensylvanicus DeGeer, 13, H. pleuriticus Kirby, 2 (2), H. viduus Leconte, 1, Loricera pilicornis Fabricius, 8, 204 The Pan-Pacific Entomologist Metabletus americanus Dejean, 3, Notiophilus aquaticus Linne, 1, Olisthopus par- matus Say, 1, Omophron americanum Dejean, 3, Patrobus longicornis Say, 1, Pseudamara arenaria Leconte, 1 , Pterostichus sp., (6), P. adstrictus Eschscholtz, 1, P. corcicinus Newman, 1 (1), P. corvinus Dejean, 9 (1), P. lachrymosus New- man, 1, P. leconteianus Lutshnik, (2), P. luctuosus Dejean, 4, P. lucublandus Say, 27 (30), P. melanarius Illiger, 11, P. mutus Say, 10, P. pensylvanicus Leconte, 7 (2), P. tristis Dejean, 1, Selenophorus gagatinus Dejean, 3, Sphcieroderus lecontei Dejean, 1 , Stenolophus comma Fabricius, 3, S. conjunctus Say, 4, Synuchus im- punctatus Say, 11, Tachys incurvus Say, 1, undetermined species, 31 (17). These data suggest that the American toad may not be entirely beneficial as a predator of noxious insects, since many Carabidae, especially larger species, are important predators of lepidopterous larvae and other destructive insects (Blatch- ley, 1910. The Coleoptera or Beetles known to occur in Indiana; Balduf, 1935. The Bionomics of Entomophagous Insects). I am grateful to the following persons for their technical assistance: Pierre Cloutier, Stephane Couture, Jean Dube, Paul Duval, Claude Gelinas, Jean-Pierre Grenier, Raymond Hutchinson, Frangois Marcotte, Herve Perrault, Jean-Pierre Pilotte, Paul Phaneuf and Jean-Pierre Tchang. — Andre Larochelle, College Bourget, C. P. 1000, Rigaud, Quebec. Notes on the general biology of the flatheaded fir borer Melanophila drummondi Kirby reared from ponderosa pine (Coleoptera: Buprestidae). — Melanophila drummondi Kirby is a common metallic wood borer of dying, slow growing, freshly cut, or wind-thrown Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, and western hemlock, Tsuga heterophylla (Raf.) Sarg., in Washington state. Chamberlin (1926, The Buprestidae of North America exclusive of Mexico) has cited many other host species. Barr (1971, in Hatch, The beetles of the Pacific Northwest, Part V. University of Washington Press, Seattle, Washington) adds white spruce, Picea glauca (Moench) Voss, to Chamberlin’s list of hosts for M. drummondi. Anderson (1966, Forest and shade tree entomology) mentions pine as being a host for M. drummondi, but does not indicate which species, nor could I find any other reference identifying pine as a host for this buprestid. I recently found M. drummondi to occur on ponderosa pine, Pinus ponderosa Laws. Infestation of host material by M. drummondi occurs shortly after spring cut- ting or windfall and extends through early fall at low elevations. At higher eleva- tions, infestation may be delayed by a month or more due to slower maturation of larvae and later emergence of adults. Maturation of larvae generally occurs within a year, but may take several years when live trees are attacked (Anderson, 1966). On March 15, 1973, I placed several 1 m long bolts of ponderosa pine cut from a tree felled in the spring of 1972, 2 miles northeast of Liberty, Kittitas County, Washington, into a plywood rearing chamber maintained at 20.6 °C and 78% relative humidity. The first M. drummondi emerged on April 22, and emergence continued until July 4, 1973, yielding a total of 12 males and 10 females. Melanophila gentilis LeConte, a common buprestid inhabitant of ponderosa pine, was also reared from the ponderosa pine bolts (30 $ $ , 22 $ $ ) . The first insects noted to emerge were two individuals of the parasitoid Vol. 50, No. 2, April 1974 205 Hymenoptera, Coeloicles brunneri Viereck on April 12. The emergence of bark- or wood-inhabiting insects continued through July 16, 1973. Although C. brunneri is primarily an ectoparasite on bark beetle larvae, it is sometimes associated with other larvae as well. Four adults of the hymenopteran Atanycolus longifemoralis Shenefelt emerged between May 7 and May 27. A. longifemoralis is a primary ectoparasite on both M. drummondi and M. gentilis (Shenefelt, 1943. Res. Stud. State Coll. Wash., 11: 51-163.). Freshly emerged Melanophila adults were placed in cylindrical plastic con- tainers for observation and were fed fresh ponderosa pine twigs and needles. Pine bark and wood chips were provided for the beetles to walk on. A small specimen vial cap was filled with tap water and placed in each container to supply moisture and drinking water. Nearly all individuals of both M. drummondi and M. gentilis fed on fresh ponderosa pine needles or twig phloem when needles became scarce. In a feeding-choice experiment I found that adults of M. drummondi, reared from Douglas-fir, fed on needles of Douglas-fir, western hemlock, or ponderosa pine. However, closer examination of the needles of the three species revealed feeding to be greater on Douglas-fir than on the others. Although this evidence is by no means conclusive, it is conceivable that feeding preference is directly re- lated to the host tree species in which the brood develops. — Donald W. Scott, College of Forest Resources, University of Washington, Seattle, 98195. Tolerance of Neotoma and Rattus to the Feeding of Triatoma . — Evidence is accumulating that wild rodents can tolerate feedings of large numbers of blood sucking triatomid bugs without lethal effects. These hematophagous insects normally inhabit the houses of wood rats, Neotoma, with associated species of white- footed mice, Peromyscus. Wood (1943, Am. J. Trop. Med. 23:315-320) reported survival of an adult 2 Neotoma a. albigula after 14 feedings by a total of 1016 Triatoma during 168 days of laboratory confinement. The maximum number of bugs fed in one day was 148. Experiments reported here using a larger subspecies of Triatoma protracta indicate larger volumes of blood consumed in toto and for all stadia. A wild 2 Neotoma fuscipes captured 1 August 1970 in a horse barn in Juniper Hills, northern Los Angeles County, California, was transferred to the Los Angeles City College laboratory. It was fed water and rat chow with carrots, lettuce and Bermuda grass supplied at irregular intervals. Except for removal to hardware cloth cylinders for bug feedings, the wood rat was confined in a cage 28 X 21 X 22 cm resting on SAN-I-CEL bedding in an open cardboard box until released alive 13 August 1971 near the original site of capture. During this time 1789 Triatoma protracta navajoensis (513 $, 644 2 and 612 nymphs) fed on this wood rat at 39 different times at approximately weekly intervals. The largest number of bugs fed at one time was 201 on 9 September 1970, the smallest 13 on 12 January 1971, the average was 49.5 per feeding. Adult bugs averaged 30.1 per feeding, nymphs 15.6 at any one time. The 232.5 g weight of the wood rat before first feeding on 3 August 1970 did not indicate an abnormal size but on 17 August it weighed 248.5 g and on 24 August 269 g. One young born on 30 August was found dead. Weights before and after the 36 bug feedings after birth of the young averaged 218.5 g (range 209.5 to 230) before and 214 (207- 206 The Pan-Pacific Entomologist 221.5) after feeding revealing an average weight loss of 4.5 g per feeding. Greatest weight loss from bug feedings was 12.5 g 11 January 1971, the smallest was 0.5 g 16 July 1971. Although most of the weight loss must have been from the obvious loss of blood, some would be due to natural metabolic processes. I have used an engorgement formula of 5 quantities as follows: fully engorged, %, %, % and %. The formulae for the 1789 bugs fed were as follows: 1177 adults (790: full, 133:%, 128:%., 61:% and 65:%) ; 349-5th instars (54-60-72- 102-61) ; 151-4th instars (28-19-37-41-26) ; 74-3rd instars (6-7-11-33-17) ; 27-2nd in- stars (7-3-5-8-4) and 11-lst instars (8-0-3-0-0). Triatoma classified as full or % were successful in molting so as noted above 114-5th, 47-4th, 13-3rd, 10-2nd and 8-lst instars molted in the laboratory. The 923 engorged adults produced numerous eggs after feeding. Xenodiagnosis for Chagas’ trypanosome was negative in 65 Triatoma (3 $, 5 2, 54-5th and 3-4th instars) examined in August 1970. Laboratory temperatures at the time of 27 feedings averaged 23 °C, the range being 19° to 29 °C. Exposure times for Neotoma for 36 feedings varied from 60 to 390 min, the average being 168. Degree of irritability of the rat in the hard- ware cloth feeding cylinders may be indicated by the number of fecal pellets released during feeding. Extremely nervous rats may defecate more frequently. This 2 Neotoma was remarkable in releasing only 6 pellets during the 39 day- time feeding exposures. One 85.5 g $ Neotoma fuscipes was killed 13 June 1968 by exposure to the feeding of 14 $ and 40 2 T. p. navajoensis. Weight loss was 6.5 g, exposure time was 240 min and there were no fecal pellets in the feeding chamber. Linsdale and Tevis (1951, The Dusky-footed Wood Rat, U. C. Press, Berkeley) noted that wood rats being handled released fecal pellets as a sign of nervousness. They reported that pellets were released at night when the wood rats were active and one female studied for 24 days averaged 124 pellets per 24 hour period (range 7 to 208) . One 94 g $ Neotoma fuscipes was killed 13 June 1968 by the feeding of 33 $ and 19 2 T. p. navajoensis and 10 <5,2 2 and l-5th instar T. rubida uhleri. Weight loss was 8.5 g, exposure time 150 min and there were 27 fecal pellets released. Wood (1947, Bull. So. Calif. Acad. Sci. 46:144-155) reported survival of albino Rattus norvegicus to feedings of 130 Triatoma over 7 hours, 168 over 3.4 days and 317 over 59 days. Recently, one 305 g $ CFN (Carworth, Inc.) albino Rattus norvegicus survived the feedings of 4 $ , 97-5th, 84-4th, 12-3rd and 1-lst instar nymphs of T. p. navajoensis on 13 August 1969 and 38 $ , 62 2 and 19-5th instar nymphs of T. p. navajoensis and 2 $ and 5 2 T. rubida uhleri on 18 August 1969. Weight losses were 38 and 27 g, exposure times 510 and 360 min and pellet counts were 12 and 34, respectively. Another 358 g S CFN rat sur- vived the feeding of 64 $ and 41 2 T. p. navajoensis on 11 September 1969. Weight loss was 15.5 g, exposure time 180 min, and pellet count 15. The capability of wood rats in providing blood meals for numerous adult and 5th instar Triatoma is important in replenishing stocks of bugs in nature. Recent experiments with Peromyscus also verify a supporting role of these mice as hosts for early instars of Triatoma (unpublished data). I thank Jack C. von Bloeker, Jr., for specific identification of the adult wood rat. — Sherwin F. Wood, 614 W. Shenandoah St., Thousand Oaks, California 91360. Vol. 50, No. 2, April 1974 207 Ovipositional Behavior of Chloealtis aspasma (Orthoptera: Acrididae). — At the time of its description the ovipositional behavior of Chloealtis aspasma Rhen and Hebard was unknown. The structure of the ovipositor suggests that this species might oviposit in wood, a phenomenon common to some other species in the genus (Jago, 1969, Proc. Acad. Nat. Sci. Philadelphia, 121: 229-335). The ovipositional behavior of C. aspasma was studied 16 miles northeast of Ash- land, Jackson County, Oregon. The study site is an old logged-over clear-cut area surrounded by mixed conifer forest. In recent years selective cutting has continued immediately adjacent to the clearing. Consequently the area is littered with slash, logs and stumps, all in varying stages of decay. Vegetation in the clearing consists of grasses, forbs and one perennial shrub, elderberry, Samhucus caerulea Raf. On July 19, 1971 a number of elderberry stakes were driven into the ground to serve as nesting sites for solitary bees. When the stakes were examined in April, 1972 the presence of grasshopper eggs in the center pith was noted. On July 6, 1972 a large number of elderberry stakes were placed in the study area and monitored periodically in an effort to identify and study the grasshopper. The first female C. aspasma visited the stakes and oviposited on July 18. Ovi- position continued throughout the study period but declined in frequency before the study was terminated on August 11. Each female spent 22-46 minutes in- serting her ovipositor and depositing a single pod of eight eggs. No female was observed to deposit successive egg-pods. Periodic surveys were conducted to determine the distribution of C. aspasma within the area and to identify oviposition sites. Females and nymphs were found throughout the clear-cut area. They were less frequently encountered in the brushy interface between the clearing and forest. Males were apparently restricted to the brushy interface area. They were not found in either the cleared or forested area. Potential ovipositional sites in the area were examined. These included all available types of dead wood, the stems of dead herbaceous plants and elderberry stems. Females were attracted only to the exposed pith of the numerous dead and broken stems around the base of each elderberry plant. Examination of old stems revealed ovipositional scars from previous years. No such marks were evident on the other materials examined. Undisturbed habitats in the immediate vicinity were also surveyed. Elderberry was the only ovipositional substrate utilized in un- disturbed areas. Earlier literature (Gurney, Strohecker and Heifer, 1964, Trans. Amer. Entomol. Soc., 84: 119-137) suggests that large series of C. aspasma are not common. How- ever, the population in the study area was large. The numerous females and nymphs were conspicuous and easily collected. It is plausible that clearcutting and the subsequent large-scale invasion of elderberry has enhanced the population’s de- velopment. In adjacent, undisturbed habitats elderberry plants are widely scat- tered and restricted to open, south-facing slopes. Grasshopper populations in un- disturbed areas are much smaller than in the study area. Stakes used in the study were allowed to overwinter in the field. They were re- turned to the laboratory in late April, 1973. Examination of and rearings from this material revealed that two insects commonly prey upon the eggs of C. aspasma. The larva of a chloropid fly, Goniopsita oophaga Sabrosky, destroyed an average of 67.1 percent of the eggs examined while the larva of a clerid beetle, Trichodes oregonensis Barr, destroyed 5.3 percent. I wish to thank Drs. Ashland B. Gurney and Curtis W. Sabrosky, both of the 208 The Pan-Pacific Entomologist National Museum of Natural History, Washington, D. C., for their respective iden- tifications of C. aspasma and G. oophaga, and Dr. W. F. Barr, University of Idaho, who helped with certain aspects of the field work. — David E. Foster, Entomology Section, College of Agriculture, Texas Tech University, Lubbock, 79409. ENTOMOLOGICAL EXCHANGES AND SALES Space is available to members and non-members who wish to advertise entomo- logical exchanges, sales, announcements, etc. We can offer an ad in four issues for $1.50 per line. Ads will be arranged in order of their receipt. Contact the Advertising Manager, J. A. Chemsak, Division of Entomology and Parasitology, University of California, Berkeley, California 94720. FOR SALE. Extensive collection of mayfly reprints. Write R. Brusca, Department of Biology, University of Arizona, Tucson, 85721. EXCHANGE. Eastern U.S. and exotic Scarabaeidae, Cerambycidae, or Cicindeli- dae for western U.S. Cicindelidae. John D. Glaser, 6660 Loch Hill Rd., Baltimore, Md. 21239. FOR SALE. Entomological literature (new and second hand) and equipment. Catalogues on request. State your interests (Lepidoptera, Coleoptera, new/ second hand) . We also buy second hand books. Write to Sciences Nat, 86 Rue de la Mare, 75020 Paris, France. Beginning January, 1975 (Vol. 51, No. 1), The Pan-Pacific Entomologist will no longer carry advertisements. No new advertisements will be accepted. Refunds will be made for any advertisement which would normally run beyond the October, 1974 issue. INSECT PINS $5. / 1000 (10 Pkgs.) (Plus Postage) 12c per 1000 PROMPT DELIVERY LOWEST PRICES IN AMERICA CLAIR ARM IN 1 91 W. Palm Avenue Reedley, California 93654 ph. (209) 638-3729 Vol. 50, No. 2, April 1974 209 Ralph Ward Dawson. Dr. Dawson was born in Lincoln, Nebraska, May 1, 1887, and died from a heart attack at Folsom, California, March 1, 1974. He at- tended public schools at Lincoln and obtained a B. S. degree at the University of Nebraska in 1912. He was instructor and assistant professor respectively in ento- mology at Nebraska until 1922, when he attended graduate school at the University of Minnesota, obtaining his doctorate in 1929. Within a year he was asked to join the Department of Zoology at Minnesota, where he remained until his re- tirement. He then lived in Colorado and Arizona and in 1959 moved to Pullman, Washington, where he accepted an appointment as Collaborator in Entomology at Washington State University. In June of 1973 he joined a retirement com- munity at Folsom. He never married and is survived by a niece in Cleveland. Dawson’s tenure of almost 14 years as Collaborator at WSU was of tremendous benefit to our Department. In his honorary appointment without pay, he worked regularly running Malaise traps, sorting and mounting material and making de- terminations of groups within his own specialty. His acquisitions to our insect collection were substantial. In addition, he prepared considerable class material, occasionally assisted in the laboratory and gave a few lectures. Unfortunately, during his last few years he experienced declining vision which reduced much of this activity, but his mental acuity remained excellent until the end. Professor Dawson was one of the vanishing breed of old time naturalists. He had a most profound knowledge of natural history but especially insects and birds. He was also an accomplished nature photographer, a gifted pianist and a weaver of Navaho rugs. At the University of Minnesota where he had charge of huge beginning classes in Zoology, he had the reputation of being the best lecturer on the entire Minnesota campus. Ralph was reserved and on the quiet side but a most interesting and stimu- lating conversationalist when discussing his principal interests — birds and insects. He maintained a wide correspondence with his former students and materially aided those in financial need. He made lasting friendships and was well liked. His taxonomic interests were quite broad. He published a number of papers 210 The Pan-Pacific Entomologist on the genus Serica (Scarabaeidae) . His extensive personal collection of these beetles was donated to the California Academy of Sciences a number of years ago. He also maintained an active interest in the Syrphidae, bumblebees, butterflies, meloids and other groups. H. S. Telford, Washington State University, Pullman, WA 99163. COMMERCIAL AND RESEARCH ENTOMOLOGISTS Obtain more accurate population counts with a light-weight motor-fan unit which sucks insects into nets D-VAC MOTOR-FAN VACUUM INSECT NETS BACK-PACKS AND HAND MODELS -also- BENEFICIAL INSECTS INSECTARY GROWN AND FIELD COLLECTED Trichogramma sp. and Green Lacewings Write for Brochures D-VAC CO. P. O. 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Send orders to: Pacific Coast Entomological Society c/o California Academy of Sciences Golden Gate Park San Francisco, California 94118 Vo!. 50 JULY 1974 No. 3 THE Pan-Pacific Entomologist AYALA AND DOBZHANSKY — A new subspecies of Drosophila pseudo- obscura (Diptera: Drosophilidae) 211 POWELL — Occurrence of prolonged diapause in ethmiid moths (Lepidoptera: Gelechioidea) 220 TORCHIO — Mechanisms involved in the pollination of Penstemon visited by the masarid wasp, Pseudomasaris vespoides (Cresson) (Hymenoptera: Vespoidea) 226 MOCKFORD AND GARCIA — Two new synonymies and a new name in North American Lachesilla (Psocoptera: Lachesillidae) 235 DOUTT — Chaetogramma, a new genus of Trichogrammatidae (Hymenoptera: Chalcidoidea) 238 HARDY — Two new species of Pseudocotalpa Hardy (Coleoptera: Scar- abaeidae) 243 PENROSE— A new subspecies of Crossidius humeralis LeConte from Texas with a redescription of the species (Coleoptera: Cerambycidae) 248 ANDERSON, OLKOWSKI AND HOY — The response of tabanid species to C0 2 -baited insect flight traps in northern California (Diptera: Tabanidae) 255 ALINIAZEE — Contribution to the bionomics of the grape leaffolder, Desmia funeralis (Hubner) : a laboratory study with field observations (Lepi- doptera: Pyralidae) 269 ALEXANDER — A new species of Idiognophomyia from California (Diptera: Tipulidae) 279 BYERS — Larva and pupa of Idiognophomyia ennihi Alexander (Diptera: Tipulidae) 282 MIDDLEKAUFF — Larva of the wood-boring sawfly Syntexis libocedrii Rohwer (Hymenoptera: Syntexidae) 288 FISHER AND ORTH— A new species of Sepedon Latreille from Oregon (Diptera: Sciomyzidae) 291 BURNETT- — A new cynipid wasp from California (Hymenoptera) 298 SCIENTIFIC NOTES 303, 304, 305, 307 BOOK NOTICES 234, 237, 254 NEW JOURNAL 247 ZOOLOGICAL NOMENCLATURE 278 SAN FRANCISCO, CALIFORNIA • 1974 Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES THE PAN-PACIFIC ENTOMOLOGIST EDITORIAL BOARD J. A. Chemsak, Asst. Editor E. G. Linsley H. V. Daly J. T. Doyen, Editor P. H. Arnaud, Jr., Treasurer E. S. Ross H. B. Leech Published quarterly in January, April, July, and October with Society Proceed- ings appearing in the January number. All communications regarding nonreceipt of numbers, requests for sample copies, and financial communications should be addressed to the Treasurer, Dr. Paul H. Arnaud, Jr., California Academy of Sci- ences, Golden Gate Park, San Francisco, California 94118. Application for membership in the Society and changes of address should be addressed to the Secretary, Franklin Ennik, Vector Control Section, California Department of Health, 2151 Berkeley Way, Berkeley, 94704. The annual dues, paid in advance, are $7.50 for regular members of the Society, $5.00 for student members, or $10.00 for subscriptions only. Single copies are $2.50 each or $10.00 a volume. Make checks payable to Pan-Pacific Entomologist. The Pacific Coast Entomological Society Officers for 1974 C. B. Philip, President Paul H. Arnaud, Jr., Treasurer H. V. Daly, President-elect Franklin Ennik, Secretary Statement of Ownership Title of Publication: The Pan-Pacific Entomologist. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher, Publisher and Owner: Pacific Coast Entomological Society, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118. Editor: Dr. John T. Doyen, Division of Entomology & Parasitology, University of California, Berkeley, California 94720. Managing Editor and Known Bondholders or other Security Holders: None. This issue mailed 24 January 1975 Second Class Postage Paid at San Francisco, California and additional offices. IN ALLEN PRESS, INC. LAWRENCE, KANSAS The Pan-Pacific Entomologist Vol. 50 July 1974 No. 3 A New Subspecies of Drosophila pseudoobscura (Diptera: Drosophilidae ) 1 Francisco J. Ayala and Theodosius Dobzhansky Department of Genetics , University of California, Davis, California 95616 Drosophila pseudoobscura Frolowa, which was earlier confused with a European species, Drosophila obscura Fallen, was distinguished by its chromosomal complement and some morphological traits by Frolowa and Astaurov (1929). It proved to be one of the most common and ecologically versatile species in western United States, western Canada, Mexico, and Guatemala (Dobzhansky and Epling 1944). Throughout its enormous geographical distribution area, it is morphologically uniform (except for variation in body size, which is due mostly to abundance or scarcity of food and to temperature during the larval development) . Most populations are chromosomally polymorphic, and populations from remote localities can sometimes be distinguished by the gene arrangements in their chromosomes (Dobzhansky and Epling 1944, Powell, Levene and Dobzhansky 1972). The interpopulational crosses produce fully fertile hybrids, and there is no evidence of prefer- ence for mating with partners of the same geographic origin (Anderson and Ehrman 1969). In the Hooker Lecture read before the Linnean Society of London in 1962, D. pseudoobscura was given as an example of a unified, genetically undifferentiated species (Dobzhansky 1963). In 1960-1962, Prof. Alice S. Hunter found D. pseudoobscura far from its previously known distribution area, namely in the highlands above Bogota, Colombia, at elevations of 2200-3280m. The Bogota colony seems to be separated from the nearest point of the main distribu- tion area of the species, in Guatemala, by a gap of some 2400Km. A study of strains from Bogota showed that in chromosomal polymorphism they resembled most the populations from Guatemala, but lacked some of the chromosomal variants that are not uncommon in Guatemala. The Bogota population is also remarkable by having the lightest genetic load of any population of the species (Dobzhansky et al., 1963). Prakash (1972) discovered that male hybrids in the Fi of the cross 1 Supported by AEC Contract AT (04-3) -34. The Pan-Pacific Entomologist 50 : 211 - 219 . July 1974 212 The Pan-Pacific Entomologist Bogota $ X USA $ are completely sterile, while the reciprocal cross gives full fertility; hybrid females are all fertile. He also found no ethological isolation between Bogota and USA flies, and concluded that an incipient reproductive isolation, namely unilateral sterility of hybrid males, has arisen despite “the absence of apparent genic differentiation.” Dobzhansky (1974) confirmed the findings of Prakash as far as the F x male sterility is concerned, but found a more complex situation in the backcrosses where different chromosomes were marked by suitable mutant genes. One or more genes determining sterility vs. fertility are present in both limbs of the X-chromosome, in the second, and in the third chromosomes. Moreover, while in the Fi hybrids sterility or fertility are all-or-none phenomena (the sons of Bogota mothers and USA fathers are always sterile, those of USA mothers and Bogota fathers always fertile), in the backcross males the sterility is a threshold character. Some males with identical chromosomal constitution are fertile and other sterile. It remains to be discovered whether this is caused by environmental variations, or by gene modifiers present in the strains used, or by a combination of both. Prakash and Dobzhansky both found that Bogota and USA strains mate at random, at least when tested in experiments in laboratory environments. It is unprofitable to discuss whether the genetic differentiation between the Bogota and USA races of D. pseudoobscura is large or small, unless it can be at least approximately measured. Fortunately, the techniques of gel electrophoresis provide a tool for such analysis. Allelic variation can readily be identified in genes coding for enzymes and other soluble proteins. In recent years, these techniques have become widely used in the fields of population and evolutionary genetics. Gel electrophoresis can also be a powerful tool in systematics. Ayala and Powell (1971) pointed out that allozymes (i.e., enzyme or protein variants coded for by different alleles of the same gene) are good diagnostic characters of sibling species of Drosophila, which can hardly or not at all be morphologically distinguished. Ayala (1973) discovered that allozymes are also good diagnostic characters of Drosophila subspecies. In the genus Drosophila, sub- species rarely are morphologically distinguishable, although they may exhibit incipient reproductive isolation, generally in the form of partial hybrid sterility. Allozymes were in fact the diagnostic characters used in the formal description of two new subspecies, Drosophila willistoni quechua and D. equinoxialis caribbensis (Ayala, 1973). In the present note, we use allozymes — as well as certain karyotypic differences — to describe a new subspecies, Drosophila pseudoobscura bogotana. Vol. 50, No. 3, July 1974 213 0 .10 .20 .30 .40 .50 .60 .70 .80 .90 1.0 Genetic Similarity Fig. 1. Frequency distribution of 44 loci relative to genetic similarity (/, after Nei, 1972) for comparisons between the subspecies Drosophila pseudoobscura pseu- doobscura and D. p. bogotana. Enzyme Differentiation Using standard techniques for gel electrophoresis and enzyme assay (Hubby and Lewontin, 1966; Ayala et al., 1972) 44 gene loci coding for enzymes and soluble proteins have been studied in natural popula- tions of Drosophila pseudoobscura. Twenty five loci were studied in our laboratory; data for the other 19 loci are taken from Prakasb et al. (1969). Five natural populations of D. p. pseudoobscura from the United States, and the Bogota population of D. p. bogotana were sampled. Genetic similarity, /, and genetic differentiation, D , between the subspecies have been calculated using Nei’s (1972) method. The mean values are I = 0.824, and D = 0.194; that is, on the average about 19 electrophoretically detectable allelic substitutions have occurred for every 100 loci in the separate evolutions of the two subspecies. In contrast, little genetic differentiation exists between local populations of D. p. pseudoobscura. I = 0.993, D = 0.007. Figure 1 shows the distribution of loci relative to genetic similarity between the two sub- species. At 29 loci (66% of the total) , the two subspecies have essentially identical genetic constitutions, I between 0.95 and 1.00. At two loci 214 The Pan-Pacific Entomologist (5%), the differentiation between the subspecies is essentially com- plete, I < 0.05. The similarity between the subspecies at the other 13 loci (30%) ranges from 0.10 to 0.95. The degree of genetic differentiation between the two subspecies of D. pseudoobscura is similar to, although somewhat smaller than, that found between other subspecies of Drosophila. Between D. w. willistoni and D. w. quechua, I = 0.808, D = 0.214; between D. e. equinoxialis and D. e. caribbensis, I = 0.782, D = 0.246. (Ayala and Tracey, 1974; Ayala et al., 1974) . Laboratory tests give no evidence of sexual isolation between these subspecies. Crosses between D. w. willistoni females and D. w. quechua males yield fertile females and males; crosses between D. w. quechua females and D. w. willistoni males produce fertile females but sterile males. Laboratory crosses between D. e. equinoxialis and D. e. caribbensis yield fertile females but sterile males, independently of the subspecies of the male parent. Table 1 gives the allelic frequencies of the six loci which are most different in the two subspecies of D. pseudoobscura. Using the method of Ayala and Powell (1972) it is possible to calculate the probability of correct diagnosis of the subspecies of a single individual of known genotype. This probability ranges from 0.999 ( Acph-1 locus) to 0.978 (a -Amy-1) for the six diagnostic loci listed in Table 1. Using jointly the six loci, the probability of incorrect diagnosis of the subspecies of a single individual is negligible, 5 X ICC 15 . Karyotypic Differentiation As pointed out above, the chromosome pool of the Bogota population is a depauperate variant of the Guatemala populations. Only two gene arrangements in the third chromosomes have been found in Bogota: the TL and SC gene sequences. These are common also in Guatemala and in Mexico (Michoacan), but in Guatemala CU and OA gene ar- rangements are also found (Dobzhansky and Epling 1944) . It has been known for a long time (Dobzhansky 1937) that at least four cytologically distinguishable kinds of Y-chromosomes are found in D. pseudoobscura , and that each of them has a geographic distribution different from the others. The Y-chromosomes of Bogota strains re- semble those of the Guatemala populations. They clearly differ from the chromosomes of most of the United States populations, particularly those on the Pacific coast (Figure 2). These cytological findings make it utterly unprobable that the Bogota population arose from recent introduction by man, presumably with Vol. 50, No. 3, July 1974 215 Table 1. Allelic frequencies at six diagnostic loci in two subspecies of Drosophila pseudoobscura. Alleles are designated by the relative electro- phoretic mobility of the proteins which they code for. The symbols for the loci refer to the proteins coded, as follows: Acph-1 = acid phos- phatase; Hk-2 = hexokinase; Pt-10 and Pt-8 = larval proteins; Est-6 — esterase; a- Amy-1 = alpha amylase. Data for Pt-10, Pt-8 and a- Amy-1 after Prakash et al. (1969). Probability of correct diagnosis Subspecies Locus and alleles of the subspecies Acph-1 95 98 100 104 106 D. p. pseudoobscura .038 .000 .947 .000 .014 > .999 D. p. bogotana .000 .056 .000 .111 .833 Hk-2 92 94 96 98 100 102 D. p. pseudoobscura .000 .000 .033 .007 .957 .003 > .999 D. p. bogotana .111 .667 .222 .000 .000 .000 Pt-10 102 104 106 D. p. pseudoobscura .012 .840 .147 > .989 D. p. bogotana .000 .000 1.000 Pt-8 80 81 83 85 D. p. pseudoobscura .011 .441 .534 .013 > .981 D. p. bogotana .870 .100 .030 .000 Est-6 100 102 D. p. pseudoobscura .800 .200 > .980 D. p. bogotana .000 1.000 a-Amy-1 74 84 100 D. p. pseudoobscura D. p. bogotana .010 .000 .208 1.000 .782 .000 > .978 fruits or vegetables from the United States. Far more likely, it came by passive transport, winds or hurricanes, perhaps many thousands or even millions of years ago, from the geographically nearest source, which is Guatemala (Dohzhansky, 1973). 216 The Pan-Pacific Entomologist Fig. 2. Spermatogonial metaphase chromosomes of Drosophila pseudoobscura bogotana (left) and Drosophila pseudoobscura pseudoobscura (right). Drosophila pseudoobscura Frolowa This species becomes the nominate subspecies, Drosophila pseudo- obscura pseudoobscura Frolowa. Drosophila pseudoobscura bogotana, new subspecies. Morphologically indistinguishable from Drosophila pseudoobscura pseudoobscura as described by Frolowa (1929), but differing from it by enzyme and protein patterns in electrophoretic assays for following. (1) Acid Phosphatase-1: in buffer system of pH 8.65, adult flies exhibit band migrating anodally more than most common bands exhibited by D. pseudoobscura. (2) Hexokinase-2 : in buffer system of pH 7.0, adult flies exhibit bands migrating anodally less than most common band exhibited by D. p. pseudoobscura. (3) Protein-10 : in buffer system of pH 8.9, larvae exhibit band migrating anodally more than most common band exhibited by D. p. pseudoobscura. (4) Protein-8: in buffer system of pH 8.9, larvae exhibit band migrating anodally less than most common band exhibited by D. p. pseudoobscura. (5) Esterase-6: in buffer system of pH 8.65, adult flies exhibit band migrating anodally more than most common band exhibited by D. p. pseudoobscura. (6) a-Amylase-1 : in buffer system of pH 8.9, adult flies exhibit band migrating anodally more than most common band exhibited by D. p. pseudoobscura. The Y-chromosome is large, submetacentric, apparently like Y’s of Guatemala populations, and unlike smaller, submetacentric Y’s of most United States populations. Holotype male, laboratory reared, original stock from Colombia: collected near Bogota by net sweeping over fruit baits in 1968 by Alice S. Hunter, reared at Department of Genetics, University of California, Davis by F. J. Ayala and T. Dobzhansky, killed July 11, 1974. Allotype and 821 paratypes (397 males, 424 females), same data as holotype, with paratypes killed July 8-11, 1974, either point mounted or preserved in alcohol. Holotype and allotype, point mounted, Vol. 50, No. 3, July 1974 217 deposited in the California Academy of Sciences, Department of Entomology, Type Number 12072. Paratypes deposited in the collections of the California Academy of Sciences, National Museum of Natural History, Washington, D.C., and University of California collections Berkeley, Davis, and Riverside. The original stock consisted of at least six inseminated females whose progenies were maintained in separate cultures. Crosses made among six cultures. The holotype and paratypes are progenies from these crosses. Discussion The analogies between Drosophila willistoni willistojii and its sub- species quechua , and between Drosophila pseudoohscura pseudoobscura and its subspecies bogotana are striking and worthy of note. The major nominate subspecies are in both cases very common, widely distributed, and only weakly differentiated populations. The minor subspecies quechua and bogotana are confined to relatively very small areas, and are isolated from the main bodies of their species by geographic barriers. In the case of quechua the barriers are the high Andes and the extremely arid coastal zone; D. willistoni does not live at high elevations or in parched deserts. The isolation of bogotana is due presumably to the intervening tropical lowlands; while D. pseudoobscura is abundant on the Pacific Coast of the United States, it withdraws to higher elevations in Mexico and Guatemala. The hybrid sterility is in both species restricted to only one sex, the male, and to only one of the reciprocal crosses, that in which the minor, narrowly distributed and geographically isolated subspecies, is the female parent. In neither case is there any trace of ethological isolation detectable — the subspecies in the laboratory interbreed freely, despite the production of some sterile hybrids. This is in accord with many previous findings: the premating and the postmating isolating mecha- nisms are genetically independent. Furthermore, the postmating mechanism (sterility of one sex of hybrids) appears first. So long as the subspecies giving sterile hybrids are securely isolated geographically, there is no need for premating isolation. Females and males of the major and minor subspecies do not meet in nature. Then sterile hybrids are formed solely in laboratory experiments. Only when and if they would expand their areas and come in contact, would there arise a stimulus for natural selection to limit or to interdict their hybridization. Can the sub- species bogotana and quechua be regarded as incipient species? Only in the sense in which any geographically confined population may conceiv- ably differentiate and diverge from the ancestral stock to become a new species. In this sense any human tribe, especially in preliterate times, 213 The Pan-Pacific Entomologist was an incipient species. And yet Homo sapiens has preserved its spe- cific unity. Among incipient species there is an enormous “childhood mortality.” Summary A new subspecies, Drosophila pseudoohscura bogotana, is described. This subspecies lives in the highlands above Bogota, Colombia, and is separated by a gap of some 2400Km from the nearest point of distribu- tion of the widely distributed subspecies, D. p. pseudoohscura. Male Fi hybrids between the subspecies are sterile when their mothers are D. p. bogotana. The subspecies are genetically differentiated; on the average about 19 allelic substitutions for every 100 loci have occurred in the separate evolution of the subspecies. D. p. bogotana also differs from most populations of D. p. pseudoohscura in the configuration of the Y- chromosome in spermatogonial metaphase preparations. The subspecies of a single individual of known genotype can be unambiguously iden- tified using the electrophoretic patterns for six gene loci coding for soluble proteins. Literature Cited Anderson, W. W., and L. Eiirman. 1969. Mating choice in crosses between geographic populations of Drosophila pseudoohscura. Arner. Midi. Natur., 81: 47-53. Ayala, F. J. 1973. Two new subspecies of the Drosophila willistoni group (Diptera: Drosophilidae) . Pan-Pacific Entomol., 49: 273-279. Ayala, F. J. and J. R. Powell. 1972. Allozymes as diagnostic characters o.f sibling species of Drosophila. Proc. Nat. Acad. Sci., USA, 69: 1094- 1096. Ayala, F. J., J. R. Powell, M. L. Tracey, C. A. Mourao, and S. Perez-Salas. 1972. Enzyme variability in the Drosophila willistoni group. IV. Genic variation in natural populations of Drosophila willistoni. Genetics, 70: 113-139. Ayala, F. J. and M. L. Tracey. 1974. Enzyme variability in the Drosophila willistoni group. VIII. Genetic differentiation and reproductive isolation between two subspecies. J. Heredity, 64: 120-124. Ayala, F. J., M. L. Tracey, L. G. Barr, and J. Ehrenfeld. 1974. Genetic and reproductive differentiation of the subspecies, Drosophila equi- noxialis carihhensis. Evolution, 28: 24^41. Dobzhansky, Th. 1937. Further data on the variation of the Y-chromosome in Drosophila pseudoohscura. Genetics, 22: 340-346. Dobzhansky, Th. 1963. Species of Drosophila. Proc. Linnean Soc. London, 174: 1-12. Dobzhansky, Th. 1973. Active dispersal and passive transport in Drosophila. Evolution, 27 : 565-575. Vol. 50, No. 3, July 1974 219 Dobzhansky, Th. 1974. Genetic analysis of hybrid sterility within the species Drosophila pseudoobscura. Hereditas, 77: 81-88. Dobzhansky, Th., and C. Epling. 1944. Contributions to the Genetics, taxonomy, and ecology of Drosophila pseudoobscura and its relatives. Carnegie Inst. Washington. Publ., 554. Dobzhansky, Th., A. S. Hunter, 0. Pavlovsky, B. Spassky, and B. Wallace. 1963. Genetics of natural populations. XXXI. Genetics, 48: 91-103. Frolowa, S. L., and B. L. Astaurov. 1929. Die Chromosomengarnitur als systematisches Merkmal. Zeit. Zellf. Mikr. Anatomie, 10: 201-213. Powell, J. R., H. Levene, and Tii. Dobzhansky. 1972. Chromosomal poly- morphism in Drosophila pseudoobscura used for diagnosis of geographic origin. Evolution, 26: 553-559. Prakash, S. 1972. Origin of reproductive isolation in the absence of apparent genic differentiation in a geographic isolate of Drosophila pseudoobscura. Genetics, 72: 143-155. Occurrence of Prolonged Diapause in Ethmiid Moths (Lepidoptera: Gelechioidea) Jerry A. Powell University of California, Berkeley Diapause in its broadest sense, an arrested state of growth and development, has been studied from many standpoints. Attempts have been made to explain causes of its initiation and termination and to more precisely define, predict and classify the various kinds. One aspect that seems to have been slighted is prolonged diapause, the maintenance of a dormant state for more than one year. Yet, this must be an impor- tant evolutionary strategy in insects, where so many kinds are dependent for food and shelter upon the inconstant occurrence of specific organ- isms. Not only must insects cope with extremely variable factors in the physical environment, such as seasonal changes in Temperate Zones, but many species must display a precise timing with their hosts as well, whether they be plants or other insects. Thus it is not surprising that in a diversity of insects the arrest in development has been shown to sometimes last more than a year. Examples are known in eggs of Orthoptera (Kreasky, 1960; Rentz, 1973) ; prepupal larvae of gall midges (Barnes, 1952), sawflies (Prebble, 1941; Sullivan & Wallace, 1967), and anthophorid bees (MacSwain, 1958) ; in pupae of various moths and butterflies; and in adults of chrys- omelid beetles (Ushatinskaya, 1967). Prolonged diapause usually is expressed by a small fraction of a one- season diapausing population (e.g., Dohanion, 1942; Ushatinskaya, 1967). It seems to be an expression of genetic heterogeneity, and the few late individuals emerge synchronously with their normal season but more than one year later. Most researchers have not been sufficiently fascinated with the phenomenon to invest years for the chance of meager results. Thus experimental data are lacking to show whether some kind of fixed polyphenic expression of the genotype is involved as a kind of natural buffer against extraordinary extremes or whether dormant individuals are responding to suboptimal conditions, or whether an interaction between genetic heterogeneity and variable microenviron- mental factors is the cause. Some insects, by contrast, in exploiting marginal and extreme con- ditions seem to have evolved prolonged diapause to cope with typically irregular seasonal and/or biotic conditions. In such examples whole The Pan-Pacific Entomologist 50 : 220 - 225 . July 1974 Vol. 50, No. 3, July 1974 221 populations may undergo dormancy for more than one year as in certain high elevation grasshoppers (Kreasky, 1960) and yucca moths (Powell, 1973a, unpublished data). A distinction needs to be made between the two classes of prolonged diapause, but experimental studies with yucca moths indicate that the distinction may lie in the degree of fitness to the optimum in environmental conditions rather than in clear cut classes of insect species. Most of the literature on this phenomenon for Lepidoptera is limited to chance rearings in situations remote from natural populations. Usually successful emergence has not been recorded beyond the third year, although there are a few reports of longer duration, up to 6-7 years in Lasiocampidae and Geometridae (Standfuss, 1896) and 8 years in Rothschildia (Saturniidae) (Rowley, 1923). By contrast, more detailed studies in other insects indicate prolonged diapause is not limited to chance situations occasioned by highly unnatural conditions. For example, Barnes (1943, 1952) showed that larvae of gall midges pass up to 10-12 winters in diapause and that the modality of longevity varies from one year’s collection to another; while Prebble (1941) found that diprionids remain in dormancy 1-6 years with the number of years variable in different parts of the geographic distribution. In Microlepidoptera prolonged diapause is known in Tortricidae and in prodoxine Incurvariidae, the yucca moths. Dohanion (1942) found that 5% of prepupal larvae of Melissopus (Laspeyresiini) overwintered 3 years, and the Douglas-fir cone moth, Barbara (Eucosmini), is re- ported by Keen (1958) to undergo dormancy lasting 3 years. Hedlin (1960a, b) indicates that a large proportion of Barbara pupae remain in diapause during some years. In yucca and agave moths of the genera Prodoxus and Agavenema, prepupal larvae wait in the dry inflorescence stalks commonly up to 4 years, and successful emergence has been obtained after 8 years (Powell, 1973a, unpublished data). Thus the ability to delay for protracted periods occurs among scattered moth taxa, and although it has not been demonstrated for many Gelechioidea, I have previously speculated that facultative delay of 2 or more seasons may be a widespread means by which Ethmiidae cope with seasonal drought (Powell, 1973b: 40). This idea was based on an early report that emergence after 1 and 2 winters had been obtained in a European species of Ethmia (dejoannis, 1926), and on the fact that up to the time of my writing (December, 1970) I had observed living pupae of 3 species of this genus that had been in diapause for 16 months or more. During the subsequent 3 years a few adults of 2 of these species have continued to emerge (Table 1) , tending to confirm my speculation. 222 The Pan-Pacific Entomologist Table 1. Number of moths emerging following storage of pupae through winter in various situations: R = Russell Reserve shed; U = Unheated room; L = Lab (mobile trailer) with fluctuating temperature above 18°C. Winter: (prior to) 1969-70 1970-71 1971-72 1972-73 Housing: L R U R R/L scylla - 0 2 1 2/- semilugens 3 0 0 2 0/1 The data are given here, as enumeration of such observations is a neces- sary step to more detailed analysis of prolonged diapause. The general biologies of the 2 species and details on the particular collections (lot JAP 69D60 for Ethmia scylla and 69E78-79 for E. semilugens) have been recorded elsewhere (Powell, 1971). Ethmia scylla Powell This univoltine, diurnal species is known only from the inner Coast Range in central California. Larvae collected in April, 1969, were pro- vided with small blocks of dry yucca stalk pith, owing to the widespread habit of members of this genus to bore into soft woody substrates for pupation. The yucca blocks were stored in a translucent, plastic box in an open-sided shed at the U. C. Russell Biological Reserve in inland Contra Costa County, California, from August, 1969, to the end of Febru- ary, 1970. This place has been used for several years to expose overwin- tering Microlepidoptera from various parts of California to greater winter extremes than available at coastal Berkeley. 1 The Russell Reserve is located about 85 airline km northwest of the scylla collection site and has comparable winter conditions. However, no scylla adults emerged in 1970, which is difficult to explain, particularly because at the same time a collection of a closely related species, E. plagiobothrae Powell (lot 69D58, see Powell, 1971) taken on the same day in April, 1969, at a site 7 km west of the scylla colony, was housed in the same conditions and all individuals successfully emerged in February, 1970. The following year some of the scylla pupae were taken for study and description (Powell, 1971), and while I was away on sabbatic leave the remainder of the collection was stored in an unheated room at Walnut Creek, Contra Costa County. Temperature and humidity conditions were 1 Temperature and humidity data at the shed will be summarized in a study of prolonged diapause in yucca moths, in preparation. Vol. 50, No. 3, July 1974 223 not monitored but certainly were much more uniform than at the Russell Reserve shed. Nonetheless, two individuals (1 <$ , 1 2) emerged between September, 1970, and August, 1971. The box was again housed at the Russell shed during the 1971-72 and 1972-73 winters, from October to April. The lot was watered at the end of March and was transferred to a mobile trailer lab at Berkeley with natural lighting conditions during spring and summer months. One male eclosed in March, 1972, and in late February, 1973, the final 2 scylla ( $ S ) emerged, after 45 months in diapause. At least these later emergences occurred within the natural flight period, late February to early April (Powell, 1973b). Subsequent examination of the pupation substrate revealed that no pupae remained unhatched. Unfortunately, in addition to 3 pupae har- vested in 1970 and the 5 moths that successfully emerged, 8 more indi- viduals pupated and developed but on eclosion were trapped in the emergence tackways (Powell, 1971: 17), possibly by fungal sealing of the outer silken caps. Thus additional completion of diapause occurred during one or more years, possibly including 1970, after one year dormancy. All development took place in a translucent container in shallow galleries a few mm into the porous yucca pith, so photoperiod as well as other factors could have been involved in diapause break. Ethmia semilugens (Zeller) This nocturnal species is widespread in the southwestern Nearctic in desert-margin habitats and other arid places. Based on collection rec- ords of adults and my rearing data, I concluded that the species is facul- tatively doubled brooded, presumably in response to rainfall because the known hosts are 2 species of annual Phacelia (Powell, 1971, 1973b). Overwintering is accomplished by pupae, and most flight records are for March and April. Resulting progeny then either complete develop- ment and emerge in July, August or September or remain in diapause. Larvae collected in Inyo County, California, in May, 1969, were allowed to pupate in folds of soft paper toweling. Three moths completed development during 1969 under laboratory conditions, the first emer- gence occurring in July, after about 6 weeks pupation. Subsequent housing for the remainder of the collection was similar to that described for E. scylla. Cocoons of semilugens in several translucent containers (plastic vials and polyethylene bags) were placed at Russell Reserve during the 1969-70 winter, and no moths matured in the spring. Part of the lot was harvested for study of the pupae in late 1970 (Powell, 1971), and the remainder were housed at Walnut Creek during the 1970-71 winter, which also produced no emergence. Overwintering in 224 The Pan-Pacific Entomologist 1971-72 was the same as for scylla, and the cocoons were watered on March 30, then transferred to Berkeley on April 10; 30 days later 2 adults (1 $ , 1 $ ) emerged. For the 1972-73 winter the collection was divided, part going to the Russell shed and part remaining at the trailer lab in Berkeley, under natural lighting, fluctuating temperature above 18 °C, and essentially dry conditions. No more individuals completed development in the Russell shed segregate, but in Berkeley 1 female emerged in September, 1973, after 52 months in diapause. Examination of the pupation substrate in October, 1973, showed that in addition to 2 pupae that died in immature stages and 3 moths pro- ceeding through development in 1969, 3 pupae harvested for study in 1970, and 3 which matured to adults in 1972-73, 2 other pupae remained in unopened cocoons, possibly still living. The emergence of 2 moths in 1972, 40 days following artificial watering, after 33 months dormancy, suggests that sporadic rainfall in desert situations may be a critical factor in breaking diapause. Literature Cited Barnes, H. F. 1943. Studies of fluctuations of insect populations X. Prolonged larval life and delayed subsequent emergence of the adult gall midge. J. Anim. Ecol., 12: 137-138. Barnes, H. F. 1952. Studies of fluctuations of insect populations XII. Further evidence of prolonged larval life in the wheat blossom midges. Ann. Appl. Biol., 39: 370-373. Dohanion, S. M. 1942. Variability of diapause in Melissopus latiferreanus. J. Econ. Entomol., 35: 406-408. Hedlin, A. F. 1960a. Diapause in the Douglas-fir cone moth, Barabara colfaxiana (Kft.) . Canad. Dept. Agric., Div. Forest Biol., Bi-Mon. Progr. Rept., 16(6) : 4. Hedlin, A. F. 1960b. On the life history of the Douglas-fir cone moth, Barbara colfaxiana (Kft.) (Lepidoptera: Olethreutidae) , and one of its parasites Glypta evetriae Cush. (Hymenoptera: Ichneumonidae) . Canad. Entomol., 92: 826-834. de Joannis, J. 1926. Longevite chez Ethmia ( Psecadia ) decemguttella Hb. Bull. Soc. Entomol. France, 1926: 151. Keen, F. P. 1958. Cone and seed insects of western forest trees. U.S. Dept. Agric., Tech. Bull., 1169; vi -f- 168 pp. Kreasky, J. B. 1960. Extended diapause in eggs of high-altitude species of grasshoppers, and a note on food-preferences of Melanoplus bruneri. Ann. Entomol. Soc. Amer., 53: 436-438. MacSwain, J. W. 1958. Longevity of some anthophorid bee larvae (Hyme- noptera: Apoidea). Pan-Pacific Entomol., 34 : 40. Powell, J. A. 1971. Biological studies on moths of the genus Ethmia in California (Gelechioidea) . J. Lepid. Soc., 25, suppl. 3, 68 pp. Powell, J. A. 1973a. Facultative longevity and polyphenism in diapause of Vol. 50, No. 3, July 1974 225 yucca moths. Presented at 24th annual meeting of The Lepidopterists’ Society, Sarasota, Fla., June 22, 1973. Powell, J. A. 1973b. A systematic monograph of New World ethmiid moths (Lepidoptera: Gelechioidea) . Smithson. Contr. Zool., 120; 302 pp. Prebble, M. L. 1941. The diapause and related phenomena in Gilpinia polytoma (Hartig). V. Diapause in relation to epidemiology. Canad. J. Res., D, 19: 437-454. Rentz, D. C. 1973. The shield-backed katydids of the genus Idiostatus. Mem. Amer. Entomol. Soc., 29; 211 pp. Rowley, R. R. 1923. Extended pupal duration. Canad. Entomol., 55: 198. Standfuss, M. 1896. Handbuch der Palaarktischen Gross-Schmetterlinge fur Forscher und Sammler. Second ed. Fischer; Jena, Germany [not seen]. Sullivan, C. R. and D. R. Wallace. 1967. Interaction of temperature and photoperiod in the induction of prolonged diapause in N eodiprion sertifer. Canad. Entomol., 99: 834^850. Ushatinskaya, R. S. 1967. The extended diapause of the Colorado beetle ( Leptinotarsa decemlineata Say) and the conditions for its occurrence. Rev. Appl. Entomol., 55: 112 [abstract]. Mechanisms Involved in the Pollination of Penstemon Visited hy the Masarid Wasp, Pseudomctsciris vespoides (Cresson) (Hymenoptera: Vespoidea) Philip F. Torchio Bee Biology and Systematics Laboratory, Agr. Res. Serv., USD A Logan, Utah 84322 Introduction Pseudomasaris is a North American representative of the vespoid wasp subfamily Masarinae. This subfamily, which is easily distin- guished by clavate antennae, is unique among foraging wasps in that it provisions cells with pollen and nectar rather than with other arthro- pods. Bequaert (1940) and Cooper & Bequaert (1951) reviewed the flower records of masarids and listed the host plants visited by these wasps, but it was Cooper (1952) who first discussed the oligolectic habits expressed within Pseudomasaris. He concluded that these wasps collect pollen and nectar primarily from three hosts [ Phacelia , Eriodictyon (Hydrophyllaceae) and Penstemon (Scrophulariaceae) ] and suggested that wasp size was associated with the flower size found on host plant species. Therefore, the smallest species [Pseudomasaris edwardsii (Cresson) ] visit Phacelia flowers, the largest species [P. vespoides (Cresson)] are restricted to Penstemon , and one inter- mediate-sized species [P. wheeleri Bequaert] visits both Eriodictyon and Penstemon. Cooper’s tenets have since been confirmed by Straw (1956a, b), Richards (1962), Crosswhite & Crosswhite (1966), and Torchio (1970). Pennell (1935) pointed to the fact that Penstemon is probably not surpassed by any other genus of flowering plants in the expression of different flower forms between species. Straw (1956b) described and figured the flowers of three species of Penstemon [section Peltanthera (Keck, 1937)] and discussed how the differences in form and color between them were associated with their pollinators. Thus, P. cen- tranthifolius Bentham is pollinated by hummingbirds, P. palmeri Gray is pollinated by bumblebees ( Bombus spp.) and carpenter bees ( Xylocopa spp.), and P. spectabilis Thurber ex Gray is pollinated by P seudomasaris vespoides. Straw’s discussion of wasp-pollinated Penstemon species was the The Pan-Pacific Entomologist 50: 226-234. July 1974 Vol. 50, No. 3, July 1974 227 first attempt to describe the adaptive features involved in the pollinat- ing mechanism of these flowers. He recognized the importance of pol- linators fitting the form and size of flowers closely and learned that the lateral position of the nectaries in conjunction with the particular angulation of the flower serves to protect the ovary from damage by the probing proboscises of pollinators. He did not, however, discuss other important pollinating mechanisms of these flowers, and his in- terpretation of function for particular flower structures was incomplete. The present paper describes the pollinating mechanisms found in Penstemon species visited by Pseudomasaris wasps. Methods Numerous observations of pollen and nectar collections from Penstemon by Pseudomasaris vespoides were made during biological studies in a greenhouse at Logan, Utah. Wasps were captured in the field and released in a greenhouse supplied with bouquets of Penstemon sepalulus A. Nels. and P. cyananthus Hook, and with potted plants of the latter. Fresh bouquets were supplied daily, and the wasps nested successfully. Dissections of these flowers in conjunction with addi- tional field observations demonstrated that flower and wasp structures are synchronized to effect successful pollination. Adaptive Floral Mechanisms Staminode: Penstemon is distinctive from other Scrophulariaceae in its retention of the fifth stamen as a prominent, sterile filament. This filament, or staminode, arises on the inner-dorsal surface of the sympetalous corolla at various points (species dependent) along the basal third of the tube and bisects the throat of the flower at various angles until it turns and parallels the inner-ventral surface of the corolla tube where it ends at or near the mouth of the flower. The apical area of the staminode in some Penstemon species (P. cyananthus ) is orna- mented with shaggy hairs, whereas the staminode is devoid of hairs in other species ( P . sepalulus ). The outer-dorsal surface of the corolla tube directly above the insertion of the staminode is demarked by a linear depression in those Penstemon species studied (Fig. 1). This surface also possesses a narrow, keel-shaped ridge that arises immediately anteriad of the depression and terminates near the apical tip of the tube (Fig. 1). The inner-dorsal surface of the corolla directly be- neath the ridge is inverted into an elongate depression. The style is positioned dorsally so that the sub-basal section is adjacent to the 228 The Pan-Pacific Entomologist Fig. 1 . Outer-dorsal view of Penstemon cyananthus flower showing: [a] ridge above style insertion; and, [b] corolla depression at staminode insertion. staminode insertion, and its apical section is normally fitted into the linear depression of the inner-dorsal surface of the corolla tube. Delpino (1875) presumed that the staminode serves to aid pollinators in locating nectaries; Kerner (1876) and Errera (1878) concluded that the dorsal to ventral position of this filament traversing the throat of flowers imposes a barrier that prevents or strongly reduces the in- cidence of nectar robbing by small insects with short tongues. Straw (1956b) discounted Delpino’s theory, but he accepted the Kerner and Errera proposal. The staminode in Penstemon flowers visited by Pseudomasaris wasps may actually function as a nectar guide (see discussion) and, at the same time, may serve as a mechanical barrier that reduces nectar robbing by smaller arthropods. Its main function, however, is to trigger pollination. This mechanism is initiated when the wasp enters the flower and pushes against the nearly vertical basal section of the staminode. When the wasp extends its proboscis to probe the flower for nectar, it habitually jerks its body back and forth as it rams the filament repeatedly with its head. If sufficient force is applied to the filament during any particular thrust by the wasp, the staminode is pushed inward (basad) a short distance, thus placing stress on the area of staminode insertion. This area functions as a fulcrum for the translocated pressure (due primarily to the angulation of the filament and the weaker corolla structure surrounding the depressed insertion area). Thus, outward pressure is applied to the basal section of the staminode insertion, and an equal but opposite pull-force develops Vol. 50, No. 3, July 1974 229 across the anterior section. Invariably, the pull-pressure is sufficiently great to bend a short transverse section of the corolla tube anteriad of the insertion area downward as the staminode is pushed slightly back- wards. Since the sub-basal section of the style is always found ap- pressed against the inner-dorsal surface of the corolla tube adjacent to the staminode insertion, it too is pushed downward when the small section of corolla tube in front of the staminode insertion is forcibly indented. If the stigma is receptive (anterior section of style bent downward) , the downward movement of the style causes the stigma to touch the pollen-laden scutum of the wasp. The style springs back to its original position when pressure is removed from the staminode, but it will move downward each time the wasp rams the staminode. This adaptive mechanism not only assures pollination, but it also facilitates a higher probability for cross pollination. Stamens : Protandry as a mechanism to better faciliate outcrossing is well known in Penstemon. Clements and Long (1923, plate 12) il- lustrated the typical maturation process of these flowers, and Straw (1956b) discussed the sequential development of flower structures during anthesis. In those species studied, in Utah, the anterior pair of anthers mature shortly after the flower opens, but dehiscence by the shorter pair (upper) is delayed 12-15 hours. The stigma begins to turn downward to its receptive position 6-12 hours after the shorter pair of anthers begins to dehisce, which decreases the probability of selfing. Most of the pollen grains in Penstemon flowers visited by Pseudo- masaris wasps are retained in the anthers after dehiscence, and only a few are released each time the dehisced anthers are vibrated. Pollen dispersion, therefore, occurs after the wasp enters the flower and positions its scutum and scutellum to face the elongate openings of the dehisced anthers. Each opening is guarded by a row of short, stiff, strong spines that rub across the thoracic surface of the wasp when the insect begins to move backward and forward during its nectar collect- ing activities. The scraping of thecal spines across the rough surface of the insect’s thorax (sometimes heard as a clicking sound in flowers of P. cyananthus ) initiates vibration of the anthers, which results in the release of pollen grains directly onto the wasp’s scutellum. Therefore, this mechanism has varied functions: (1) it releases pollen onto ap- propriate pollinators; (2) it deposits pollen directly on that surface of the insect which is subsequently touched by the stigma; (3) it per- mits pollen from each theca to be deposited on a number of wasps over an extended period to better facilitate outcrossing (the delayed 230 The Pan-Pacific Entomologist dehiscence of the shorter anther pair also increases the period of pollen release) ; and, (4) it serves to prevent or impair pollen robbing by other Penstemon-v isiting insects because of the position of the anthers, the long narrow dehiscence line of each theca, and the guard spines. Obviously the pollinators of Penstemon also possess adaptive features that insure successful pollination. Straw (1956b) discussed such ap- parent points as body size and form. However, two additional features should be mentioned in the case of Pseudomasaris. One is the elongate, nearly flat-surfaced scutum that is covered laterally with sparse, erect hairs. This structure should be considered adaptive because its ex- treme length accommodates at least the anterior anthers, irrespective of variation in the size of individual flowers or wasps; and the erect hairs function to accept and hold pollen grains in place, which pre- vents pollen dispersion over a large surface area. The scutellum is also elongated and could be considered a posterior extension of the thoracic surface that receives Penstemon pollen. The transfer of pollen from the wasp’s mesonotum to the proboscis with specific preening move- ments of the front legs should also be considered adaptive. This combing activity with specialized hairs transfers most of the pollen, but it also mixes a residue of the pollen grains collected from different flowers across the surface of the thorax, thus increasing the probability of outcrossing. Style-. Clements and Long (1923, plate 12) illustrated how the style of Penstemon glaber Pursh elongates as the open flower matures, but this structure is fully extended as flowers open in P. cyananthus and P. sepalulus. The length of the style in these two species is also variable. Thus, the stigma on the shortest style is located slightly distad of the upper (shorter) stamens, and the longest style has the stigma positioned between or slightly distad of the longer stamens. Further, variation in style length occurs on the same or different racemes of the same plant in either species. These variations, coupled with variation in in the size of flowers and wasps, increase the effective surface area of the wasp’s thorax that is utilized for pollen transfer to the stigma; and at the same time, they increase the probability of cross pollination. The primary mechanism that forces the style to move downward (infolding of corolla tube distad of staminode insertion) has already been described, but additional flower structures should be included as supportive mechanisms of this movement. The groove on the inner- dorsal surface of the corolla tube in which the style is inserted is one such structure. Its presence prevents the style from swinging back and forth laterally whenever the flower is moved, thus reducing the chance Vol. 50, No. 3, July 1974 231 of its entanglement with the anthers or staminal filaments. If entangle- ment occurs, the style can no longer move vertically; and its normal function is lost. Lateral migration of stamens also occurs before the style fully ma- tures. When the flower first opens, the stamens are positioned with their anthers nearly touching each other. By the time dehiscence is initiated, the anthers have migrated a short distance away from each other and the resulting space is sufficiently wide to permit the style to pass between them. Straw (1956b) discussed how the position of the ovary below the midline of the corolla and the position of the nectaries at the outer bases of the upper stamens served to protect the ovule against me- chanical damage when pollinators probe flowers for nectar. The direc- tion and position of both pairs of stamens may serve the same function. The shorter pair arises laterad of the ovary and converges in front of it. These stamens then diverge as they appress against the upper wall of the corolla tube. The longer stamens arise below the ovary and nearly parallel the shorter stamens as they converge in front of the ovary. Together, the four stamens almost completely surround the ovary, thus protecting it against mechanical damage by pollinators and other arthropods. Discussion The foregoing descriptions suggest that those species of Penstemon visited by Pseudomasaris wasps have evolved particular floral mech- anisms that are well adapted to these oligolectic pollinators. In addi- tion, Straw (1956b) discussed various floral mechanisms found in many Penstemon species that are particularly well adapted to other pollinators including hummingbirds and bees. Yet, the position and form of most flower structures in this taxon are sufficiently similar to indicate that particular functions are retained at the generic level irrespective of most adaptive mechanisms associated with oligolectic or monolectic pollinators. The staminode, for example, has the primary function of triggering pollination in those Penstemon species visited by Pseudomasaris wasps. Conversely, the staminode does not possess this function in Penstemon flowers pollinated by hummingbirds, bombyliid flies, and some bees; though the filament is retained as a prominent floral structure in the genus. Therefore, the staminode is a multifunctional structure that has evolved in some species as an adaptive mechanism to trigger or facilitate pollination, and at the same time, performs other functions that are expressed in all Penstemon flowers. To date, no direct evidence has been presented to explain what these 232 The Pan-Pacific Entomologist functions are, but Delpino (1875) suggested that the sterile filament serves as a nectar guide, and Kerner (1876) and Errera (1878) pro- posed that it acts as a barrier to small, nectar-robbing arthropods. Observations during the present study tend to support Delpino’s contention that the staminode does indeed act as a nectar guide. Also, the filaments of the upper stamens, in conjunction with the staminode, serve as nectar guides once the wasp enters the flower and touches its head against the staminode. The midline of the wasp’s head ap- presses against the filament, which is automatically positioned directly in front of the pollinator’s mouthparts. When the wasp begins to ex- tend its proboscis into the throat of the flower, the tongue hits the staminode, which in turn causes it to be deflected to one side or the other in the general direction of the laterally positioned nectaries. When the tongue is further extended, it normally touches one of the two filaments of the upper stamens each of which is angulated in such a way as to further guide the tongue toward a nectary. Additionally, observations indicated that the staminode does act as a barrier to some bee visitors (two species of Osmia and one species of Andrena) , but not to others (one species of Hylaeus, one species of Osmia , and one species of Halictus ). Apparently, the workability of the barrier is dependent upon two factors: (1) the size of each opening in the corolla tube that is present on either side of the staminode, and (2) the angulation of that staminode in relation to the particular form of each Penstemon flower. If the opening of the corolla tube on either side of the staminode is larger than the diameter of the arthropod visitor, that insect can crawl directly to the nectaries and rob them. Conversely, if either opening is smaller than the diameter of the visitor’s body, the staminode bars its entrance. Angulation of the staminode is sometimes important because it pre- vents “robbing” by non-adaptive pollinators whose tongues are suf- ficiently long to reach the nectaries should their heads touch the stami- node insertion (some Osmia, Hoplitis, etc.). However, the heads of these long-tongued bees are usually more elongate and/or broader than those of Pseudomasaris and require a larger corolla tube opening in front of the staminode insertion. The size of this space is limited dorsally by the corolla tube and ventrally by the staminode filament. Therefore, the particular angulation of the filament determines the vertical diameter of this space (the greater the angulation of the staminode, the greater the. vertical diameter of the space). Invariably this opening proves too narrow to accommodate full entrance by long- tongued, non-adaptive bee visitors, and robbing is thus thwarted. Vol. 50, No. 3, July 1974 233 Information presented here is derived from the study of only two wasp-pollinated species of Penstemon in a limited geographical area. The genus, however, is a relatively large, North American taxon that is associated with many specific pollinators. Consequently, additional pollination studies must be completed before all the adaptive floral mechanisms expressed in the taxon can be understood. With a de- finitive understanding of these pollination mechanisms in Penstemon (a popular introduced plant in other parts of the world), we should be better able to determine which species groups can be successfully established in areas outside their normal ranges. Acknowledgments R. Shaw, Utah State University, Logan, and G. Baker, Lewis-Clark State College, Lewiston, Idaho, reviewed the manuscript, and Jane Barnes inked Figure 1. Literature Cited Bequaert, J. 1940. Notes on the distribution of Pseudomasaris and on the food plants of the Masaridinae and Gayellinae (Hymenoptera, Vespidae). Bull. Brooklyn Entomol. Soc., 35: 37-45. Clements, F. E., and F. L. Long. 1923. Experimental pollination. Carneg. Inst. Wash., Publ., 336. 274 p. Cooper, K. W. 1952. Records and flower preferences of masarid wasps. II. Polytrophy or oligotrophy in Pseudomasaris ? (Hymenoptera: Vespi- dae). Amer. Midland Nat., 48: 103-110. Cooper, K. W., and J. Bequaert. 1951. Flower records and flower preference of masarid wasps. Psyche, 57 : 137-142. Crosswhite, F. S., and C. D. Crosswhite. 1966. Insect pollinators of Penstemon Series Graciles (Scrophulariaceae) with notes on Osmia and other Megachilidae. Amer. Midland Nat., 76: 450-467. Delpino, F. 1875. Ulteriori osservazioni e considerazioni sulla dicogamia nel regno vegetale. Atti. Soc. Ital. Sci. Nat., 12: 21-141, 179-223; 16: 151-349; 17: 266-407. Errera, L. 1878. Penstemon gentianoides et Penstemon hartvegii. Bull. Soc. Roy. Belg., 17: 182-243. Keck, D. D. 1937. Studies in Penstemon V. The section Peltanthera. Amer. Midland Nat., 18: 790-829. Kerner, A. 1876. Die schutzmittel der Bluten gegen unberufene Gaste. Festschr. Zool.-Bot. Ges. Wien, 189-261. Richards, O. W. 1962. A revisional study of the masarid wasps (Hymenoptera, Vespoidea). British Museum (Nat. Hist.), London. 294 p. Pennell, F. W. 1935. Scrophulariaceae of eastern temperate North America. Acad. Nat. Sci. Monogr., No. 1. 650 p. Straw, R. M. 1956a. Floral isolation in Penstemon. Amer. Midland Nat., 90: 47-53. 234 The Pan-Pacific Entomologist 1956b. Adaptive morphology of the Penstemon flower. Phytomorphology, 6: 112-119. Torchio, P. F. 1970. The ethology of the wasp, Pseuclomasaris edwardsii (Cresson) and a description of its immature forms (Hymenoptera; Vespoidea, Masaridae). Los Angeles Co. Mus. Nat. Hist. Contrib. Sci., No. 202. 32 p. BOOK NOTICE Biological Control by Natural Enemies. Paul Debach. Cambridge Uni- versity Press, London and New Tork. 323 pp. 1974. Cloth, $14.95; paper, $5.95. This semi-technical book should attract a wide audience among non-entomolo- gists and entomologists alike. Written in a clear, informative style, and appended by a functional but not overwhelming bibliography, it provides a useful overview of the field, while allowing enjoyable reading at the same time. The first and last chapters discuss pesticide usage and problems. Relatively short sections describe the biological characteristics of natural enemies of pests (insects and weeds), applied ecology, the role of science and the public, and a survey of cultural, genetic, and other non-chemical methods of control. About one third of the book relates case histories of the more important biological control projects. Especially fascinating are the accounts of early foreign exploration. The dedication and endurance of many of the pioneering workers, who usually labored under ex- tremely primitive conditions, recall the earlier biological expeditions of the 18th and 19th centuries. Perhaps the fervor of these hardy individuals set a historical precedent which partly explains the zeal of many contemporary prac- titioners of biological control. — Editor. Two New Synonymies and a New Name in North American Lachesilla (Psocoptera: Lachesillidae) Edward L. Mockford Illinois State University, Normal, Illinois And Alfonso Neri Garcia Aldrete Universidad Nacional Autonoma de Mexico, Mexico City The new synonymies reported herein result in removal of one name from the list of North American Caecilius. Correct association of sexes of Lachesilla pacifica Chapman produces a synonymy of L. sil- vicola Chapman with L. pacifica and necessitates one new name. Caecilius nigrotuberculatus Curran (Curran, 1925:292).- — The senior author examined the type of C. nigrotuberculates (type No. 1543 in Canadian National Collection, Ottawa, Canada) and found it to be Lachesilla pedicularia (Linnaeus). Therefore, by priority, the name C. nigrotuberculatus falls into the extensive synonymy of L. pedic- ularia. For complete synonymy of L. pedicularia to date see Smithers, 1967:62. Lachesilla silvicola Chapman. — This is a synonym of L. pacifica Chapman. The latter species was described by Chapman (1930) in page 353 and is based on female material only. L. silvicola is described in the same paper on page 361. The holotype is a male. We find that the male of L. silvicola is the male of L. pacifica and therefore, by page priority, L. silvicola falls into the synonymy of L. pacifica. Our evidence for associating the sexes as indicated above is as follows: (1) Association of sexes in field collections. — Of 66 lots of L. pa- cifica and L. silvicola that have come to our attention, the male and female associated by Chapman under the name L. silvicola have been found together in only one lot — the type material of L. silvicola. In this lot there were also females of L. pacifica. In 14 lots (including the one just referred to) there were males of L. 4 silvicola ’ with females of L. pacifica. In 22 lots of L. silvicola $ that have come to our at- tention there were males associated with only one lot (the type ma- terial) and females alone in the other 21. (2) Comparative morphology. — Garcia Aldrete (1974) has studied the systematics above the species level in Lachesilla and has estab- lished a series of species groups for this genus. In his classification, The Pan-Pacific Entomologist 50 : 235 - 237 . July 1974 236 The Pan-Pacific Entomologist L. pacifica is placed in the pedicularia group on basis of both male and female characters. The female assigned by Chapman to L. silvicola (called L. sp. R-2 by Garcia Aldrete) goes to the rufa group, which is rather far from the pedicularia group in characters of both sexes. The female of L. silvicola Chapman is now un-named, and we pro- pose the name L. yakima new species for this species. Chapman’s allotype is here designated holotype (type locality Seattle, King County, Washington; type material in P. J. Chapman collection, Ge- neva, New York). Records include the female records of L. silvicola cited by Chapman, female records of L. silvicola cited by Sommer- man (1946) plus the following (collector is E. L. Mockford except where indicated otherwise) : Canada: British Columbia: Vancouver, 16 August 1963, beating Douglas fir ( Pseudotsuga taxifolia Britton) 1 $ , R. C. Rounds. United States: California: El Dorado County: 1 mile south of Camp Richard- son, State Highway 88, 28 August 1953, 5 $ ; Humboldt County: 9 miles east of Blue Lake, Highway 299, 30 July 1969, altitude 900 feet, beating hemlock ( Tsuga heterophylla Sargent) and western red cedar ( Thuja plicata Donn) , 1 2,2 nymphs; King County: 8 miles east of Issagoah on Highway 10, altitude 825 feet, 4 August 1969, beating western red cedar, vine maple ( Acer circinatum Pursch) and blackberry ( Rubus sp.), 1 2 ; Mariposa County: Yosemite National Park, Glacier Point Road, altitude 7174 feet, 6 September 1953, beating pines ( Pinus spp.) and red fir ( Abies magnified Murray) 1 2 ; Monterey County: Redwood Gulch, 7 August 1962, 4 2 , E. Schlinger; Tulare County: Generals Highway to Sequoia National Park, altitude 7500 feet, 15 miles north of park boundary, 7 September 1953, beating red fir, 2 2. Utah: Utah County: Alpine Loop Road off State Highway 189 near Heber City, altitude 5800 feet, 24 August 1953, beating firs (Abies spp.), especially dead branches, 1 2. Washington: Clallam County: Lake Sutherland, 10 August 1927, 1 2,1 nymph, C. R. Crosby; Jefferson County: Hoh River Road, 4 miles east of Highway 101, 14 August 1966, beating Sitka spruce ( Picea sitchensis Carriere) 1 2 , sifting litter of maple leaves, 1 2- Wyoming: Teton County: 17 miles southwest of Jackson, Highway 26, 5 August 1966, beating spruces (Picea sp.) , Snake River Valley, 1 2,3 nymphs; Highway 89, 5 miles south of Yellowstone National Park, altitude 6900 feet, 18 August 1969, beating Douglas fir, 1 2 • Note. — Since 54 adult females of this species have been collected to date and the male remains unknown, we suspect that the species is partheno genetic. Acknowledgements. — The senior author wishes to thank the author- ities of the Division of Entomology, Department of Agriculture of Canada, and especially Mr. J. E. H. Martin for the loan of the type of Caecilius nigrotuberculatus. Some of the material cited in this study was borrowed from the Department of Entomology, Univeristy of Cal- ifornia, Riverside. We wish to thank Dr. Saul Frommer of that insti- Vol. 50, No. 3, July 1974 237 tution for arranging the loan. We also wish to thank Dr. P. J. Chap- man, Geneva, New York, for loan of material. Literature Cited Chapman, P. J. 1930. Corrodentia of the United States I. Suborder Isotec- nomera. Jour. N. Y. Entomol. Soc., 38: 219-290, 319-403. Curran, C. N. 1925. Descriptions of two insects found in imported foodstuffs. Canad. Entomol., 57 : 292-293. Garcia Aldrete, A. N. 1974. A classification above species level of the genus Lachesilla Westwood (Psocoptera:Lachesillidae) . Folia Entomol. Mex., 27: 1-88. Smithers, C. N. 1967. A catalog of the Psocoptera of the world. Austr. Zool., 14: 1-145. Sommerman, K. M. 1946. A revision of the genus Lachesilla north of Mexico. Ann. Entomol. Soc. Amer., 39: 627-661. BOOK NOTICE Insects in Flight. Werner Nachtigall. Translated by Harold Oldroyd, Roger Abbott and Marguerite Biederman-Thorson. McGraw-Hill, New York. 153 pp. 1974. $12.95. The informal style and extensive use of photographs and line drawings give this account of the morphology and physiology of insect flight mechanisms the flavor of a Life Nature Guide Volume. The author has entwined strands of personal and scientific philosophy with factual material, and emphasizes analogies between organisms and machines to clarify explanations of physiological processes. The 44 chapters cover subjects ranging from the fine structure of insect flight musculature to orientation in foraging honey bees and discussions of the instru- mentation required for studies of aerodynamics. — Editor. Chaetogramma, a new genus of Trichogrammatidae (Hymenoptera: Chalcidoidea) Richard L. Doutt 1 University of California, Berkeley Minute Hymenoptera form a common component of the aerial in- sect fauna, consequently the suction traps designed to sample this aerial plankton frequently capture large series of trichogrammatids and my- marids. Although this collecting technique does not reveal host rela- tionships, it does yield information on the seasonal flight activities and dispersal patterns of these parasitic wasps. Very importantly it adds interesting new taxa to these groups, for some species have not otherwise been taken. For example, two of the species described herein are known only from suction trap collections in South Africa, and most of the California specimens representing the third species were also taken in a suction trap. Chaetogramma, new genus This genus keys out near Xipho gramma in the classification of Trichogram- matidae proposed by Doutt and Viggiani (1968), hut it differs in lacking the large sabre-shaped valvulae and greatly exserted ovipositor so characteristic of Xipho- gramma (Fig. 9) . While the forewings of both genera have abundant discal setae the vein track patterns are different (compare Figs. 2 and 8). In Xipho- gratnma there are two distinct funicle segments while in Chaetogramma the funicle is either a single segment (Fig. 3) or one that is partially divided (Fig. 1). Chaetogramma has a single annellus while Xiphogramma has a minute second an- nellus fused to the base of the first funicle segment (Fig. 7). Both Xiphogramma and Chaetogramma have toruli low on face near lower ocular line ; vertex wrinkled, with prominent setae (Figs. 5 and 6) ; maxillary palpi one segmented, elongate; hypogynium well developed. Doutt and Viggiani (1968) in discussing Xiphogramma mention that “A species from California exhibits all the characteristics of Xipho- gramma except the long ovipositor and sabre-shaped valvulae. This species probably represents a new subgenus, but until more material is available the genus Xiphogramma is not being divided.” I now believe this species represents a new genus, Chaetogramma , and it is the type species described as follows: Chaetogramma occidentalis, new species Female.- — Body color dark brown, propodeum bright yellow, apical abdominal tergite yellow, antennae light brown, eyes red, face and vertex golden, legs dark 1 Present address: 9240 S. Riverbend Ave., Parlier, CA 93648. The Pan-Pacific Entomologist 50: 238-242. July 1974 Vol. 50, No. 3, July 1974 239 brown except apices of femora, tibiae pallid. Venation brown except for light band across marginal vein; wings hyaline except brown under venation. Antennae (Fig. 1), scape and club subequal length, pedicel slightly longer than funicle, latter partially divided, basal portion with oblique sensorium. Head (Fig. 6) rounded, convex frontally but face impressed to receive antennal scapes; toruli low near lower ocular line; compound eyes with prominent ommatidia, relatively fewer than in Xiphogramma (compare Figs. 5 and 6) ; vertex with coarse, wrinkled sculpture; mandibles with two sharp teeth, third blunt; maxillary palpi one segmented. Scutum and scutellum with large prominent bristles; forewings broadly rounded apically, vein tracks as shown in Fig. 2. Ovipositor originating at base of abdomen, between hind coxal bases, strongly developed but only slightly exserted ; hypogynium prominent. Male. — Similar to female, except abdomen entirely dark brown. 240 The Pan-Pacific Entomologist 6 0.2 mm Fig. 5. Xipho gramma anneckei , head. Fig. 6. Chaetogramma occidentalis, head. This species is distinguished by its partially divided funicle segment, dark brown body color, and broadly rounded forewings. Holotype female, Parlier, Fresno County, California, suction trap in citrus, 1 September 1973, R. L. Doutt. Allotype, two female paratypes, same data as holotype except collected 19 August 1973. Two female paratypes, same data, collected 17 August 1973. Additional paratypes include two females, by suction trap, U. C. Botanical Garden, Berkeley, Alameda County, California, 15 June 1966, and one female 16 June 1966, F. E. Skinner. Two females, on Distichlis spicata, Altamont Pass, Alameda County, California, 16 April 1964, R. L. Doutt. One female by sweeping, Graton, Sonoma County, California, 11 April 1947, R. L. Doutt. One female on hopleaf, Wheatland, Yuba County, California, 9 July 1954, K. S. Hagen. Two males, one female, on low vegetation, Edison, Kern County, California, 25 May 1953, C. E. Kennett. Holotype, allotype to be deposited California Academy of Sciences, paratypes at U. S. N. M., and Division of Biological Control, University of California. Chaetogramma pretoriensis, new species Female. — Basic body color light yellow with 4 brown bands on abdomen, brown markings on thoracic pleurae, propodeum, base of ovipositor. Head yellow with brown markings in malar space, eyes and ocelli bright red. Club and funicle golden; brown band across scape and base of pedicel. Tarsi brown, tibiae light brown, trochanters and apices of coxae and femora pale. light band across mar- ginal vein; apex of marginal and stigmal vein brown; forewing from base to end of venation light brown, remainder hyaline. Antenna (Fig. 3) with single funicle segment bearing two obliquely oriented Vol. 50, No. 3, July 1974 0.2 mm 241 7 Figs. 7-9. Xiphogramma anneckei. Fig. 7. Antenna. Fig. 8. Forewing. Fig. 9. Body, lateral view. sensoria. Scape longer than club. Forewing (Fig. 4) with 3 distinct vein tracks, abundant discal setae. Ovipositor half length of abdomen, not exserted. Male. — Unknown. This species is distinguished from C. occidentalis by its yellow color, abdominal cross bands, undivided funicle, shorter ovipositor, forewings not broadly rounded, and scape longer than club. Holotype female. Pretoria, South Africa, in suction trap, April 1957, D. P. Annecke. One paratype female same data as holotype. One paratype female same data but collected March 1958. Holotype at Plant Protection Research Institute, Pretoria. Paratypes at Divi- sion of Biological Control, University of California. Xiphogramma anneckei, new species Female. — Body basically yellow with brown markings. Anterior half scutum, entire scutellum light brown; thoracic pleurae edged with dark brown. Abdomen with brown markings on basal tergite, two distinct dark bands across mid-abdomen, two large brown blotches on each side of abdomen near apex. Valvulae dark 242 The Pan-Pacific Entomologist brown, in strong contrast with yellow venter of abdomen. Head yellow gold, eyes and ocelli crimson; malar area brown, mandibles brown. Ventral aspect of scape pale, rest of antenna dark brown except pale, obliquely oriented sensoria on funicle segments. Legs testaceous. Wings hyaline with smoky area beneath brown venation. Head (Fig. 5) with wrinkled vertex, prominent bristles, toruli low on face but slightly above line of lower eye margin, maxillary palpi of single segment, mandibles with 3 teeth. Antennae (Fig. 7) ; scape slender, shorter than club, pedicel with prominent dorsal spines; two annelli, 2nd closely appressed to funicle 1 ; two funicles, first slightly longer than second, each with prominent, obliquely oriented sensorium. Forewings (Fig. 8), wing blade densely setose. Abdomen (Fig. 9) much longer than thorax; ovipositor very long extending forward between hind coxal bases, strongly exserted and curved upward. Two large, exserted valvulae characteristic of genus. Hypogynium well developed extending to apex of abdomen. Fore coxae with bristles. Male. — Color similar to female, except scutellum golden, thoracic pleurae darker brown, apex of abdomen brown. Antennae similar to female. This genus has been known only from four female specimens of X. holorhoptra Nowicki taken on window panes of an electric railway, Warsaw, Poland (Nowicki, 1940). X. anneckei differs from X. holor- hoptra by its yellow color and pattern of brown markings, head as long as wide, scape slender and not swollen at base, toruli slightly above lower eye margin, and marginal vein not widened apically. This large, handsome African species is dedicated to the distinguished South African entomologist, Dr. David P. Annecke. Holotype female. Pretoria, South Africa, in suction trap, March 1958, D. P. Annecke. Allotype same data as holotype except collected April 1957. Female paratypes same data except collected as follows: nine in April 1957, two in February 1957, five in April 1958, and two in March 1958. Holotype and allotype at Plant Protection Research Institute, Pretoria. Para- types at California Academy of Sciences, U.S.N.M., and Division of Biological Control, University of California. Literature Cited Doutt, R. L. and G. Viggiani. 1968. The classification of the Trichogrammat- idae (Hymenoptera: Chalcidoidea) . Proc. Calif. Acad. Sci., 35(20): 477-586. Nowicki, S. 1940. Descriptions of new genera and species of the family Tri- chogrammidae (Hym. Chalcidoidea) from the Palearctic region, with notes — supplement. Zeit. angew. Entomol., 26(4) : 624-663. Two New Species of Pseudocotalpa Hardy (Coleoptera: Scarabaeidae) Alan R. Hardy Insect Taxonomy Laboratory Department of Food and Agriculture Sacramento, California 95814 Since the publication of the description of Pseudocotalpa Hardy (1971:238), two additional undescribed species of this group have been collected. This necessitates a slight redefinition of the genus, as follows. The illustrations were prepared by Ellen Parker, to whom I extend my deep appreciation. Genus Pseudocotalpa Hardy Body form robust; eye large, globular, antennae 10-segmented, with three- segmented club, clypeus concave, mandible with outer edges rounded, without external teeth ; maxillary palp four-segmented with fourth segment enlarged, longer than other three segments, apical segment with deeply impressed groove on outer edge, running length of segment. Postcoxal prosternal spine poorly developed, not present as well developed knob. Flight wings well developed ; six free abdominal segments visible from below. Without well developed process between meso- coxae. Anterior tibia tridentate, with or without apical spur. Posterior tibia en- larged at apex, with one or two spurs. All tarsal claws simple, not cleft, all chelate. The genus Pseudocotalpa may be distinguished from closely related genera in the Areodina by the distinctive shape of the clypeus, Pseudo- cotalpa being the only genus in which the clypeus is deeply concave; and by the poorly developed prothoracic post-coxal spine or knob, which is well developed and evident in other genera. The enlarged, deeply grooved maxillary palp is also characteristic. Pseudocotalpa giulianii , new species Holotype male: Greatest length 23 mm, width at elytral humeri 11.5 mm. Dorsal surface light colored, prothorax and head with a deeper yellow colora- tion than elytra, apex of clypeus, margins of prothorax and posterior margin of scutellum dark reddish brown. Legs same color as prothorax; abdomen, apex of tibiae and all tarsi dark reddish brown. Upper surface glabrous except for margins of prothorax, which are provided with fine, long, erect pale hairs and a dense net of shorter recumbent hairs on anterior and posterior margins. Ventral surface of thorax densely clothed with erect, long, fine white hairs. Clypeus con- cave, margins elevated, closely, shallowly punctured; suture arcuate (Fig. 2). Front with scattered fine punctures. Prothorax with uniformly spaced medium punctures; punctures separated by own diameter or slightly less. Lateral pro- The Pan-Pacific Entomologist 50: 243-247. July 1974 244 The Pan-Pacific Entomologist Figs. 1-6. Pseudocotalpa spp. P. andrewsi ; Fig. 1, head; Fig. 4, prothorax. P. giulianii; Fig. 2, head; Fig. 5, prothorax; P. sonorica ; Fig. 3, head; Fig. 6, pro thorax. thoracic margin with slight emargination anterior to posterior angles (Fig. 5) . Elytra with fine marginal bead laterally, indistinct or absent apically and basally. Elytral apex subacute at sutural margin. Elytral striae faintly indicated; disc with scattered medium and fine punctures. Pygidium with complete apical margin; rugulose with scattered fine punctures. Meso- and meta-tibiae with two apical spurs. Allotype female: Greatest length 20.5 mm, width at elytral humeri 10 mm. Differs from male in these respects: Antennal club and maxillary palp very slightly shorter than that of male. Anterior tibiae with teeth thinner, spine-like; tarsal claws thinner. Variation in type series: Males; length 17 mm to 25 mm, width 7.5 mm to 11 mm. Females; 14 mm to 22 mm, width 7 mm to 10 mm. Unlike P, andrewsi. Vol. 50, No. 3, July 1974 245 Figs. 7-12, Pseudocotalpa spp. P. andrewsi; Fig. 7, male genitalia; Fig. 10, anterior tibia. P. giulianii ; Fig. 8, male genitalia ; Fig. 11, anterior tibia. P. sonorica ; Fig. 9, male genitalia; Fig. 12, anterior tibia. (All tibiae are of males). all specimens examined have two posterior apical tibial spurs. Most noticeable variation is in coloration, which is darker in many specimens, apparently the result of different amounts of brown pigment. Male genitalia Fig. 8. Holotype (Calif. Acad. Sci. No. 11942) and 42 paratypes (34 males and 8 females) ; Nevada, Nye Co., Amargosa Desert, Sand Dunes, 26-IV-1972, Derham Giuliani Collr. (CAS) ; allotype and 6 paratypes, same data except 17-V-1972 (CAS); 23 paratypes (18 males, 5 females), same data except 28-IV-1973, A. J. and M. E. Gilbert Collrs. (AJG) ; 85 paratypes (75 males, 10 females), same data except 29-IV-1973, R. McPeak Collr.; 31 paratypes (23 males, 8 females), same data except 30-IY-1973, R. McPeak Collr.; 113 paratypes (86 males, 27 females), same data except l-V-1973, R. McPeak Collr. (RMcP ; AMNH; CNC; USNM; LACM; MCZ; FMNH; ARH; HFH) . The type locality is located in South Central Nye Co., near the border with California and Death Valley National Monument, and is known as Big Dune. The first series of this species was collected by Derham Giuliani (for whom this species is named) by utilizing the information on behavior which accompanied the original (1971) description of Pseudocotalpa andrewsi Hardy. Giuliani reports that the behavior of the two species is similar. 246 The Pan-Pacific Entomologist Pseudocotalpa sonorica, new species Holotype male: Greatest length 21 mm, width at elytral humeri 10 mm. Dorsal surface light colored, prothorax and head with a slightly deeper yellow coloration than elytra. Antennae, palps, posterior margin of scutellum and all tarsi dark reddish brown. Disc of prothorax, elytra, and pygidium with many scattered, very fine, long, erect, white hairs; lateral prothoracic margins with denser pubes- cence; anterior and posterior prothoracic margins with shorter, very dense semi- recumbent white hair. Ventral surfaces of head and thorax densely clothed with erect, long, fine white hairs. Clypeus concave, margins elevated; closely rugosely punctured; suture abruptly arcuate medially (Fig. 3). Front with dense medium punctures, punctures often with long erect hairs ; prothorax with complete marginal bead; all angles broadly rounded (Fig. 6). Elytral margin with poorly developed bead; sutural apex spiniform. Elytral striae faintly indicated; disc with scattered fine punctures. Pygidium with complete apical marginal bead; disc rugulose with scattered punctures. Meso- and meta-tibia with two apical spurs. Male genitalia Fig. 9. Holotype male (CAS No. 11943) ; Mexico, Sonora, 50 mi. S. W. Sonoyta. III-12-1973, F. G. Andrews, A. R. Hardy Collrs. The type locality is approximately 10 mi. N.E. of Puerto Penasco on Highway 8, and is the long high dune mass lying one or two miles to the north of the highway. Two additional individuals have been examined, both dead and badly broken, which are excluded from the type material. So far as can be determined, both agree and are conspecific with the type. One specimen (female?) from the type locality; the second (female?) was collected 4 mi. N.E. Sotello, a railroad stop on the Ferrocarril del Pacifico, between El Doctor and Puerto Penasco, and is approximately 10-20 mi. N. or N.W. of Puerto Penasco. The type specimen was collected at dusk, as it flew over the surface of the sand. An additional specimen was observed at the type locale, but could not be collected. It thus appears likely that this species ex- hibits the same type of behavior as the other known species of this genus. Pseudocotalpa andrewsi Hardy Pseudocotalpa andrewsi was originally described from 120 specimens. Since that time I have collected 51 males and 31 females on IV-22-1971, at the type locality. I have seen an additional specimen from 3 mi. N.W. Glamis, Imperial Co., California. It should be noted that through an error on the author’s part, the captions to the figures in the original description were reversed. Figure 1 represents the male holotpye, not the female allotype as stated. Figure 2 represents the female allotype. The other captions are correct. Vol. 50, No. 3, July 1974 247 A Key to the Species of Pseudocotalpa 1. Hind angles of the prothorax broadly rounded (Fig. 6) ; elytra and pro- thorax with erect fine pubescence medially; size larger (20 mm-)-) an- terior tibiae with basal tooth distal of midpoint (Fig. 12) ; from Sonora, Mexico Pseudocotalpa sonorica new species 1' Hind angles of the prothorax well defined, often with a slight emargina- tion anterior to the angle (Figs. 4 and 5) ; elytra and pro thorax glabrous medially; size smaller or larger (14 mm to 25 mm) ; anterior tibiae with basal tooth either distal of midpoint (Fig. 10) or distinctly medial (Fig. 11) ; from the United States 2 2. Frontal- cly peal suture evenly arcuate (Fig. 2) ; anterior tibiae with basal tooth distinctly medial (Fig. 11) ; size larger (17 mm to 25 mm) ; from Nevada Pseudocotalpa giulianii new species 2' Frontal-clypeal suture abruptly arcuate medially (Fig. 1) ; anterior tibiae with basal tooth distal of mid-point (Fig. 10) ; size smaller (13.5 mm to 17.5 mm) ; from California Pseudocotalpa andrewsi Hardy Literature Cited Hardy, Alan R. 1971. The North American Areodina with a description of a New Genus from California. Pan-Pacific Entomol., 47(3): 235- 242, illus. NEW JOURNAL Paleobiology. T. J. M. Schopf and R. G. Johnson, editors. Department of the Geophysical Sciences, University of Chicago, Chicago, Illinois 60637. Published quarterly, approximately 400 pp. per year. Initial issue, Spring, 1975. $8.00 (members of the Paleontological Society) ; $12.00 (non-members) . This journal will focus on ecological and evolutionary processes, especially as seen in the paleontological context. The first issue will contain articles dealing with rates of evolution and extinction, growth rates in corals, and theoretical morphology of plants with spiral growth. Contributions are accepted from neon- tologists as well as paleontologists. A New Subspecies of Crossidius humeralis LeConte from Texas with a Redescription of the Species (Coleoptera: Cerambycidae) R. L. Penrose Oregon Department of Agriculture, Salem A recent systematic study of the species of Crossidius occurring north of Mexico revealed a new subspecies of C. humeralis from the Texas gulf coast. Discovery of this subspecies and study of more specimens of the nominate form from southeastern New Mexico pro- vided a basis for redescription of the species and a discussion of its variability. The taxonomic format is essentially that utilized by Linsley and Chemsak (1962) in their revisionary work on the Cerambycidae of North America. Collection abbreviations are those of Arnett and Samuelson (1969). Special thanks are extended to Mr. W. H. Tyson, Fresno, California, and Dr. J. W. Tilden, San Jose, California, for making specimens of the new subspecies available for study, and Dr. P. 0. Ritcher, Oregon State University, and Mr. R. L. Westcott, Oregon Department of Agriculture, for review of the manuscript. Crossidius humeralis LeConte Crossidius humeralis LeConte, 1858:24. MALE — Form robust; head pale, black or bicolored; antennae as long as or slightly longer than body; pronotum pale, immaculate or infuscated laterally, or with five black spots; elytra light brown or orange-yellow, immaculate or with sutural and humeral vittae, vittae distinct or confluent apically; venter pale to entirely black. Pronotum inflated, rounded at sides; surface with an anterior discal callosity on each side of the midline, usually with an elongate impunctate area on basal third at middle, punctures rather dense, pubescence erect or appressed. Elytral surface densely punctate, punctures decreasing in size posteriorly, clothed with short, pale, suberect or subrecumbent pubescence which may partially obscure punctation. Length: 8-19 mm. FEMALE — Antennae reaching to apical one-third of elytra. Pronotum paler than in male and with 2, 3 or 5 spots. Elytra with sutural and humeral vittae, the latter sometimes broken into a short basal portion and an elongate apical portion, the apical portion rarely absent. Length: 9-19 mm. C. humeralis is easily distinguishable from all other species of Crossidius by its unique pronotal sculpturing and pattern of elytral maculations. DISTRIBUTION — Southeastern New Mexico, western and gulf coast region of Texas. LARVAL HOSTS — Haplopappus. The Pan-Pacific Entomologist 50: 248-254. July 1974 Vol. 50, No. 3, July 1974 249 This species is separable into two well defined, completely allopatric sub- species as follows: Crossidius humeralis humeralis LeConte (Figs. 1, 2) Crossidius humeralis LeConte, 1858:24; LeConte, 1873:197; Horn, 1885:176; Leng, 1886:119; Linsley and Chemsak, 1961:48; Linsley, 1962:161. MALE — Moderate-sized; head dark orange-red, sometimes with frons and posterior edge just in front of pronotal margin black; antennae and legs dark orange-red; pronotum orange-red, immaculate or with lateral areas clouded with black; elytra light brown or yellow-brown, immaculate, or with sutural and humeral vittae present, sutural vitta short, narrow, its width decreasing gradually to suture near or before middle, humeral vittae extending a short distance behind humeri; venter usually pale. Head coarsely punctate, densely clothed with moderately long, mostly recumbent golden hairs. Pronotum coarsely punctate, largest punctures located laterally, becoming finer and more sparse on disk, clothed with appressed golden pubescence which usually partially obscures punctures. Elytral surface rather densely punctate and pubescent. Venter mod- erately densely pubescent. Length: 11-19 mm. FEMALE — Pronotum paler than in male, dorsal callosites usually darkened or black; elytra with sutural vittae frequently present, humeral vittae narrow, usually broken into a short basal portion and an elongate apical portion, the latter markings infrequently narrowly coalesced on basal sixth. Length: 11-19 mm. Type locality: Llano Estacado, Texas and New Mexico. DISTRIBUTION — Southeastern New Mexico and western Texas. All material examined is from New Mexico. Inclusion of western Texas as part of the range is based on a single male labeled “S.W. Texas” cited by Linsley and Chemsak (1961). LARVAL HOST — Haplopappus heterophyllus (Gray). ADULT HOSTS — Same as larval host and also Chrysothamnus. FLIGHT PERIOD — Late June to early September. Most specimens examined were collected during the first two weeks of August. The eai'liest record seen was June 19 at four miles south of Loving, the latest September 11 at Carlsbad, both localities in Eddy County, New Mexico. VARIATION — Diversity in prothoracic and elytral markings is ex- hibited by this subspecies. Prothoracic maculations of the female are usually present in the form of five black or darkened areas on the pro- notum. The two anterior markings and median marking correspond with the dorsal callosities and the basal impunctate area, respectively. A pair of vaguely dark areas, one on each side of the basal third, completes the pattern. In some specimens these two basal markings may be absent, as may be the one corresponding to the median im- punctate area. The males exhibit a more variable and in some cases a more melanic prothoracic coloration. In the most reduced state the marking consists of a narrowed anterior transverse prosternal band which extends obliquely up the sides and onto the pronotal surface 250 The Pan-Pacific Entomologist just behind the anterior margin. A progressive widening of this band results in a condition in which most of the prosternal and lateral areas become black. Increased melanization of the ventral surface is ac- companied by lateral infuscation of the pronotum. The dorsal mark- ings, one on each side of the midline, frequently exhibit a bilobed ap- pearance with the lobes partially enveloping the discal callosities. Fusion of the ventral and dorsal black areas occurs laterally on each side of the prothorax to give the appearance of a single maculation. Geographic variation in the presence of prothoracic markings is expressed in the males examined. Individuals with immaculate pronota predominate in collections made at southerly localities, particularly in the Carlsbad and Loving areas of Eddy County, New Mexico. To the north near Roswell and Artesia, New Mexico, beetles with protho- racic markings are the most frequently observed phenotype. Sexual dichromatism also is evident in the nature and extent of elytral markings. In the male the markings consist of short humeral and a narrow sutural vittae located on the basal half (Fig. 1). The humeral vittae are longer in the female and are usually broken into a short basal and an elongate apical portion (Fig. 2). These markings may be present or absent, but when present their lengths vary considerably. The humeral vittae are the most frequently expressed maculations in the males at hand. Slightly over two-thirds (68%) of the specimens examined exhibit this character. The remain- ing 32 percent are immaculate. The sutural vittae are present less frequently than the humeral markings and are exhibited by slightly more than one-half of the males which have humeral maculations. The humeral and sutural vittae occur more uniformly in the females and are present on nearly all specimens at hand. The elongate apical portions of the former vittae, which are not found in the males of this subspecies, are present in three-fourths of the specimens examined. These are narrowed and generally restricted to the apical two-thirds of the elytra. Approximately 10 percent of the specimens have the basal and apical segments coalesced. The lengths and distinctness of these vittae vary dependently. Speci- mens displaying short, pale and vaguely defined humeral markings exhibit similar sutural vittae. Conversely, individuals with longer, blacker and well defined humeral markings display a longer and more clearly defined sutural vitta. SPECIMENS EXAMINED— NEW MEXICO: CHAVES CO: ROSWELL, 15 mi. S., 13 August 1950, on Chrysothamnus, J. W. MacSwain (1 $ , 1 $, UCRC) ; 7 mi. E., 5 August 1969, on H. heterophyllus, D. E. Foster, L. S. Hawkins, Jr., Vol. 50, No. 3, July 1974 251 R. L. Penrose (11 $ , 4 2, UICM) ; 6 mi. E., 5 August 1969, on H. heterophyllus, D. E. Foster, L. S. Hawkins, Jr., R. L. Penrose (17 $,10 2, UICM). EDDY CO: LOVING, 2 August 1969, on H. heterophyllus, D. E. Foster, L. S. Hawkins, Jr., R. L. Penrose (5 $, UICM); 16 August 1950, on Chrysothamnus, J. W. MacSwain (1 $, UCRC) ; 4 mi. S., 19 June 1968, on stems and in the roots of H. heterophyllus, S. M. Hogue, R. L. Penrose (9 $, UICM); ARTESIA, 6 mi. NE., 1 August 1968, on H. heterophyllus, R. L. Penrose (26 $, 1 2, UICM) ; 5 mi. NE., 2 August 1969, on H. heterophyllus, D. E. Foster, L. S. Hawkins, Jr., R. L. Penrose (13 $, 52, UICM); LAKEWOOD, 1 mi. S., 31 July 1968, on H. heterophyllus, R. L. Penrose (6 $ , 5 2 , UICM) ; MALAGA, .5 mi. E., 2 August 1969, on H. heterophyllus, D. E. Foster, L. S. Hawkins, Jr., R. L. Penrose (3 $ , UICM) ; 1 mi. N., 2 August 1969, on H. heterophyllus, D. E. Foster, L. S. Hawkins, Jr., R. L. Penrose (3 $, UICM); HARROAN LAKE, 2 August 1968, on H. heterophyllus, R. L. Penrose (1 $ , 1 2, UICM) ; CARLS- BAD, 3 August 1969, on H. heterophyllus, D. E. Foster, L. S. Hawkins, Jr., R. L. Penrose (8 $, 2 2, UICM) ; 10, 11 September 1969, on H. heterophyllus, R. L. Penrose (8 $, UICM). Crossidius humeralis quadrivittatus, new subspecies (Figs. 3, 4) MALE — Moderate sized; head black; prothorax bicolored, prosternum with anterior two-thirds black, prosternal process and regions laterad of coxae pale, sides orange-red, each with a moderately large anterior and small posterior black spot, pronotum orange-yellow with five black spots located as follows; a large submedian pair on anterior half, two large sublateral spots and a smaller medial one on basal third; elytra orange-yellow, sutural vittae narrowed, enclosing apical portion of scutellum, gradually diminishing to suture on apical sixth, humeral vittae wide, sinuate along the basal fourth, slightly expanded apically and extending to near apex but remaining distinct from sutural vittae; appendages piceous; meso- and metathoracic sterna and abdomen prodominately black. Head densely punctate, clothed with pale, mostly erect pubescence which does not obscure the punctures. Pronotal surface moderately densely punctate, punc- tures contiguous and confluent, clothed with shortened, thin, pale, mostly erect to suberect pubescence which does not obscure the punctation. Elytral surface densely punctate, punctures nearly contiguous to confluent, clothed with thin, pale, mostly semi-erect pubescence which does not obscure surface features. Length: 12 mm. FEMALE — Prothorax with prosternum pale, sides yellow-orange, lateral spots small, vaguely defined; pronotal spots smaller than in male; elytra with sutural and humeral vittae expanded laterally, humeral vittae enveloping apex, coalesced with sutural vittae on apical sixth. Length: 14 mm. Holotype male and allotype from WELDER [WILDLIFE] REFUGE, SAN PATRICIO COUNTY, TEXAS, 25-30 October 1968 (W. H. Tyson), deposited on indefinite loan at the California Academy of Sciences. Paratypes: (12 males, 23 females), same locality data, date and collector; (48 males, 27 females), same locality, 17 October 1970 (J. W. Tilden) ; (1 male), same locality, 2 November 1963 (J. W. Tilden) , deposited in the collections of the California Academy of Sciences, University of Idaho, J. W. Tilden, W. H. Tyson and the 252 The Pan-Pacific Entomologist Figs. 1-2: Elytral patterns of Crossidius h. humeralis LeConte. 1) male female. Figs. 3-4: Elytral patterns of C. h. quadrivittatus Penrose. 3) male female. Vol. 50, No. 3, July 1974 253 author. Additional specimens not designated as paratypes include: (1 male, 1 female), Corpus Christi, Nueces County, 10 October 1905 (F. C. Pratt); (3 males), same locality, 11 November 1969, on Compositae (C. W. Griffin); (2 males, 2 females), Arroyo City, Willacy County, 6 October 1973 (J. E. Wappes). The type series was collected from the flowers of an unidentified species of Haplopappus which is presumably the larval host. H. phyllocephalus (Gray) is an inhabitant of the Texas coastal plain and is the probable larval host. C. h. quadrivittatus is separable from the nominate subspecies by its black head, sparser pronotal punctation and pubescence and by its expanded, quadrivittate elytral pattern. C. h. quadrivittatus is anatomically quite distinct and, judging from available distributional data, geographically well isolated from the nominate populations of C. humeralis. Treatment as a subspecies is based on the fact that character differences by which the two phenotypes can be separated are known to vary geographically in other species of Crossidius. Males in the type series range from 8 to 13 mm. in length. Females from 9 to 14 mm. Considerable variation in the prothoracic and elytral markings is evident, as in the nominate form. The pronotal spots of the female may be distinct, as in the allotype, or the large anterior and posterior spot on each side may be fused. Additionally, the pleural spots are often enlarged and quite distinct. The holotype exhibits the basic male pattern of black prothoracic markings. Most males have the pronotal and pleural spots and prosternal band expanded and fused which presents an appearance quite similar to that described for the most melanic individuals of the nominate form. The humeral and sutural vittae are nearly always distinct in the male (Fig. 3) and always fused apically in the female (Fig. 4). In some females the widening and posterior fusion between these two vittae is extensive and the basic elytral coloration appears black with the ground color remaining as a narrow, pale, medially located vitta which tapers from the base to slightly beyond the midpoint. Literature Cited Arnett, R. H. and G. A. Samuelson. 1969. Directory of Coleoptera Collec- tions of North America (Canada through Panama). Ann Arbor, Michigan, Cushing-Malley, Inc., vii -f- 123 pp„ Horn, G. H. 1885. Descriptions of some new Cerambycidae with notes. Trans. Amer. Entomol. Soc., 12: 173-197. LeConte, J. L. 1858. Catalogue of Coleoptera of the regions adjacent to the 254 The Pan-Pacific Entomologist boundary line between the United States and Mexico. Jour. Acad. Nat. Sci. Philadelphia, 4: 9-42, 1 pi. 1873. New species of North American Coleoptera Part II. Smithsonian Misc. Coll., (264): 169-240. Leng, C. W. 1886. Synopses of Cerambycidae (in part) . Entomol. Amer., 2: 118-120. Linsley, E. G. and J. A. Chemsak. 1961. A distributional and taxonomic study of the genus Crossidius (Coleoptera, Cerambycidae) . Misc. Publ. Entomol. Soc. America, 3(2): 25-64. Linsley, E. G. 1962. The Cerambycidae of North America. Part III. Taxonomy of the subfamily Cerambycinae, tribes Opsimini through Megaderini. Univ. Calif. Publ. Entomol., 20: 1-188. BOOK NOTICE Insects in Relation to Plant Disease. Second edition. Walter Carter. John Wiley & Sons, Inc., New York, xiv -f- 759 pp. 1973. $39.50. Although the author states that this work “does not presume to impinge on the subject matter of the highly specialized virology texts,” the 431 pages com- prising this last part of the book are devoted exclusively to viruses. Included are a general introduction to plant virology, followed by more detailed discussions of modes of virsus transmission, virus-vector relationships, the clinical aspects of plant virus diseases, and the ecology and control of plant viruses. The first part of the book contains separate chapters dealing with the other groups of organisms implicated in plant disease. The second part deals with various aspects of phytotoxemias. Despite the author’s modest disclaimers, this text must rank as a most authoritative account of a complex, multidisciplinary subject. — Editor. The Response of Tabanid Species to CCb-Baited Insect Flight Traps in Northern California (Diptera: Tabanidae) J. R. Anderson 1 , W. Olkowski 1 , & J. B. Hoy 2 Following the initial reports of the attraction and collection of many species of Tabanidae in dry ice-baited traps (DeFoliart et al. 1965, Otsuru et al. 1965 & Wilson, Tugwell & Burns 1965), a basic objective in our 1966 studies was to determine if blood-sucking species of snipe flies (Diptera: Rhagionidae, Symphoromyia ) also could be captured in this manner. As reported (Anderson & Hoy 1972), dry ice-baited insect flight traps of the Malaise-type were very successful in catching large numbers of Symphoromyia species, as well as non-bloodsucking, but host-seeking, Cephenemyia females ( Diptera :Oestridae) (Ander- son & Olkowski 1968) . In unbaited Malaise traps, Smith, Breeland & Pickard (1965) caught nearly 25 tabanids/trap (7,057 in 6 months), representing 9 species in 5 different genera. As anticipated from this and the 1965 reports of DeFoliart et al., Otsuru et al., & Wilson et al., our experimental baiting of insect flight traps with CO 2 proved very efficient in catching many species of Tabanidae as well as many other hematophagous species in other families (Anderson & Hoy 1972). Since the success of C02-baited Malaise-type traps for catching tabanids was first reported (Olkowski, Anderson & Hoy 1967), several other workers also have used C02-baited Malaise-type traps to trap Tabanidae (Blume et al. 1972, Knudsen & Rees 1968, Roberts 1970, 1971, 1972, Thornhill & Hays 1972). As only a summary of our results concerning Tabanidae were reported in our 1967 abstract, and only a few species were mentioned, we herein report the complete results for comparison with later studies in Cali- fornia and results from other areas. Methods Because the primary emphasis of our research with C02-baited traps concerned species of Symphoromyia, a detailed description of the study area, etc., has been published. For details of the study area, CO 2 release and other methodology, trap design, aerial photographs 1 Professor of Entomology and Graduate Research Assistant, respectively, Department of Ento- mological Sciences University of California, Berkeley 94720, U.S.A. 2 Research Entomologist, Agricultural Research Service, U.S. Department of Agriculture, Fresno, California 93727, U.S.A. The Pan-Pacific Entomologist 50 : 255 - 268 . July 1974 256 The Pan-Pacific Entomologist of the trap sites and deer pens, etc., readers are referred to Anderson & Hoy (1972). This research was conducted at the University of Cali- fornia field station at Hopland (Mendocino Co.). The topography here is characterized by rolling hills interspersed with ravines; eleva- tion ranges from 200m to nearly 1,000m. The climate consists of winter rains and a summer drought. All trapping was done in a woodland-grass habitat dominated by oak trees, and we used white nylon mesh insect flight traps of the Malaise-type, 2.74m high X 2.44m wide. These traps, operated from 0800 to 1700 hrs. (Pacific Standard Time), were baited with dry ice held in a polystyrene foam insulated container 3 set on the ground next to the center support pole of the trap. All traps were baited with 6.8kg of dry ice each day from 27 May through 15 June, 1966. Sub- limated CO 2 escaped through 2cm holes on each side of the box. CO 2 emission rates of 2.04kg/trap/day (about 2.0 liters/min) were de- termined from 9-hour weight losses of the insulated boxes. In the experimental design, each of 4 traps (I, II, IV & V) was placed in the corners of a grid about 124m on a side, and 1 trap (trap III) was set in the center of the square. A sixth trap (trap VI) was set up 0.77km SE from the center of the grid and 187m WNW from a group of penned, tame black-tailed deer ( Odocoilius hemionus columbianus ) that were observed throughout the study. This design allowed comparison of the catch of the center trap in the 5-trap grid with that caught in each of the 4 corner traps and the isolated trap VI near the tame deer. Aerial photographs of the 5-trap grid, Trap VI and the deer pens were used to determine the percentage canopy cover within circles hav- ing radii of 5, 15, 30 and 60m from each trap site. Temperatures and wind direction and occasionally velocity 4 , were taken at trap site I at 0800, 1100, 1400 and 1700 hours. These data later proved to be es- sentially the same as those recorded at one of the Field Station weather stations 2.4km SE of our trapping grid. For the principal study comparing flies caught in the 5-trap grid, at Trap VI and at deer, the traps were operated from 27 May through 15 June 1966. Each day the traps were serviced in numerical order with the dry ice containers being rotated from trap to trap to avoid possible bias among different containers. On 16 June traps II, III and IV were baited with dry ice and traps I, V and VI were operated 3 Freez/Safe®, Mfgd. by Polyfoam Packers Div., glo-brite Foam Plastics, Chicago, 111., U.S.A. 4 FloRite air velocity meter, model MRF, Becharach Industrial Instrument Co., Pittsburgh, Pa., U.S.A. Vol. 50, No. 3, July 1974 257 without dry ice. On 17 June traps I, V and VI were baited with dry ice and traps II, III and IV were without. Between 18 June and 26 June several traps also were operated sporadically with dry ice on a survey basis. In preliminary studies in April and early May a smaller trap (1.52m wide X 1.37m high) was operated with dry ice for 10 days and for 20 days without. At the apex of a trap flies were collected in a removable 1.4 liter styrene container through which they entered via a 15mm hole in a screen cone which formed the inner base of the collecting container. The removable collecting containers were collected and replaced with empty ones at 1100, 1400 and 1700 hrs each day, except for 4 and 12 June when they were replaced at 1.5-hr intervals (beginning after 1100 hr on 12 June). On 15 June the traps were operated for an addi- tional period from 1700-1930 hrs. Between 0800 and 1700 hr on each day that traps were operated, J. B. H. observed the fly activity at 4 tame deer. Various tabanid species were sporadically aspirated from, or squeezed and collected by hand from the deer. The deer were held in two pens about 20 X 70m each which enclosed the same type of woodland-grassland habi- tat as at the trap grid site. All trap catch data first were assessed by analysis of variance on the basis of the completely random design. For treatments showing significant and highly significant differences, the means of different species caught at different trap sites were compared by Duncan’s New Multiple Range Test (Duncan 1955). The original data from trap catches were transformed to the log (n + 1) prior to conducting analysis of variance. Results COi-Emission Rates. An analysis of variance (Anderson & Hoy 1972) showed that the major source of variation for CO 2 emission at trap sites was chance error (52%). Day-to-day variation (42.8%) was next, with site-to-site variation accounting for only 5.1% of the variation. There was no significant difference in COo emission among trap sites or among days. Effect of Temperature on Studies. Data on the feeding behavior of Symphoromyia species during 1964 and 1965 (Hoy 1966) indicated that minimum host-seeking temperatures would not be reached on days when the temperature was below 12.2°C at 0800 hr. Hence, neither host observations were made nor traps operated on 29 and 30 May and 6 June 1966. No flies were seen at hosts on 28 and 31 May nor 258 The Pan-Pacific Entomologist on 1 and 2 June; respective daily temperature maximums were 18.9, 19.4, 16.7 and 17.2°C. The 6 traps, operated for full trapping periods on 31 May and 1 and 2 June, and until noon on 28 May, caught the following tabanids: 28 May — 1 Hybomitra aasa, 1 Tabanus punctifer , 1 T. kesseli; 31 May — 1 T. similis ; 1 June — -1 T. similis ; 2 June — 1 H. aasa. Numbers of Tabanids Caught in CO^-baited Traps vs Unbaited Traps. Like other results for tabanids (Bennett & Smith 1968, Roberts 1970, Wilson, Tugwell & Burns 1966), our CCVbaited traps caught significantly greater numbers of Symphoromyia species (Anderson & Hoy 1972) and tabanids than unbaited traps. Only female tabanids were caught in the traps. On the 10 days the small trap was operated with CO 2 between 6 April and 21 May, the combined catch of Sym- phoromyia and tabanids averaged nearly 25 females/day. During 20 days of operation without dry ice in April and early May this trap caught only 1 T. similis. When the 6 larger traps were operated with and without CO 2 on 16 and 17 June (see methods), the 3 traps baited with CO 2 caught 24 tabanids on 16 June and 33 tabanids on 17 June. The 3 unbaited traps caught only 1 tabanid on 17 June and none on 16 June. Species Caught in CO^-baited Traps. A total of 18 species of Tabani- dae were caught in C02-baited traps operated between 6 April and 26 June 1966; 14 species (Table 1) were trapped during the period of the primary study shown in Table 2, one species ( H . procyon) was caught in C02-baited traps operated in April and 3 species ( H . mel- anorhinus, Chrysops proclivis proclivis , C. furcatus ) were trapped in other studies conducted between 8 and 26 June 1966. Only C. pechu- mani , C. hirsuticallus , S. gigantulus and A. incisuralis were not col- lected between 18 and 26 June. A total of 1094 females of 14 species of Tabanidae were caught in the 6 C02-baited traps during our 13 day study (Table 1) comparing catches in the 5 grid traps versus isolated Trap VI, 0.77 km away. Silvius notatus and Tabanus similis together made up 57% of the total catch, and the 4 species of which we caught more than 100 speci- mens each comprised 78% of the total catch. The Relationship Between Temperature and the Tabanid Fauna Captured. The catch data in Tables 2 and 4 shows that activity oc- curred within a well-defined temperature range. The beginning of activity in the morning was suppressed or delayed by low tempera- tures, and activity usually was diminished from mid- to late afternoon and finished earlier on cool days. Joyce and Hansens (1968) also Vol. 50, No. 3, July 1974 259 Table 1. The species and total numbers of female Tabanidae caught at all 6 trap sites during 13 trap-days. a Species 15 Number % of total flies caught No. of days caught Mean temp. ranges captured at c Silvius notatus (Bigot) d 355 32.45 12 19.4-40.6 Tabanus similis Macquart 3 266 24.31 13 16.7-40.6 Tabanus kesseli Philip 3 124 11.33 13 17.8-40.6 Apatolestes comastes Brennan 3 112 10.24 11 21.7-40.6 Chrysops surdus Osten Sacken 3 78 7.13 10 22.2-38.3 Hybomitra aasa Philip 42 3.84 12 15.6-36.1 Chrysops coquillettii Hine 42 3.84 12 19.4^40.6 Tabanus punctifer Osten Sacken 3 39 3.56 9 15.6-38.3 Chrysops coloradensis Bigot 3 13 1.19 5 23.9-30.0 Silvius gigantulus (Loew) 3 11 1.00 4 22.2-25.6 Chrysops pechumani Philip 3 7 0.64 3 19.4-24.4 Chrysops asbestos Philip 3 0.27 3 24.4-31.7 Chrysops hirsuticallus Philip 1 0.09 1 23.9 Atylotus incisuralis (Macquart) 1 0.09 1 31.7 ■ a Same dates as in Table 2. 13 The H. aasa and C. surdus were identified by Dr. C. B. Phillip who also confirmed the identifications of representative specimens of most other species. e Includes days other than the 13 primary trapping days. 11 Species seen or collected while feeding on deer. reported that temperature was one of the major factors affecting the activity (numbers of flies caught on traps) of T. nigrovittatus and T. lineola. The minimum temperature threshold of host seeking activity for most species was between 20-21. 7°C. Including the 4 trapping days (not included in the tables) when the daily maximum was below 19.4° C, only 2 T. punctifer, 3 specimens each of H. aasa, T. kesseli and 4 T. similis were captured at temperatures below 20° C. Although a few specimens of C. pechumani and S. notatus and 1 C. coquillettii were captured during one mid-day period having a mean temperature of 19.4°C (Table 4, and 3 June), we feel that the latter 2 species were caught during the hour when the temperature was at 21.1 and 21.7°C because none were caught during any other periods or on any days hav- ing temperatures below 21.1 °C. Overall, the smallest catches during the 13 favorable trapping days occurred on the 3 days having daily maximums below 26.7°C (Tables 2 and 4). Only T. kesseli and T. similis were caught on 7 June (Table 2). Even on warmer days only 9 specimens (including H. aasa , C. 260 The Pan-Pacific Entomologist Table 2. The numbers and times at which all tabanid species were caught at 6 trap sites during first 10 and last 3 trap-days. Date Wind a Duration of trapping periods (hrs) Total flies 0800-1100 1100-1400 1400-1700 27 May SW 10 (12) b 46 (56) 26 (32) 82 3 June NW 2 (10) 14 (70) 4 (20) 20 4 June W 2 (1) 128 (59) 86 (40) 216 5 June SE 11 (ID 62 (63) 25 (26) 98 7 June SW 1 (ID 7 (78) 1 (ID 9 8 June NW 28 (30) 46 (50) 19 (20) 93 9 June NW 5 (6) 53 (68) 20 (26) 78 10 June NW 1 (3) 20 (53) 17 (45) 38 11 June N 3 (3) 44 (49) 43 (48) 90 12 June N 24 (17) 72 (51) 46 (32) 142 Subtotals 86 (10) c 492 (57) 288 (33) 866 13 June W 31 (50) 16 (26) 15 (24) 62 14 June W 41 (55) 12 (16) 22 (29) 75 15 June w 45 (49) 19 (21) 27 (30) 91 Subtotals 117 (51) d 47 (21) 64 (28) 228 Totals all clays 203 (19) 539 (49) 352 (32) 1094 a Predominant direction from whicli wind was blowing. b Percent of total days catch. e Per cent of total flies caught during 1st 10 days. a Per cent of total flies caught during last 3 days. pechumani, T. kesseli, T. punctifer and T. similis) were caught during the first trapping period on the 5 days when the mean temperature for period I was 20°C or lower (Tables 2 and 4). As the mean tem- perature of the first trapping period increased, so too did the numbers and species of tabanids caught. Thus, during the first 10 days, except for 9 June, from 12 to 30% of a day’s total catch was caught during the first period when the mean temperature was greater than 21.1°C. Maximum numbers of tabanids were caught during the first period only on the 3 days when the mean temperature for this period ex- ceeded 24.4°C (Tables 2 and 4) . The temperature range at which all species were most active was 23.9-32.2 °C, but activity seemed affected by the time of day the lower figure was reached. Thus at 24.4 and 25 °C during the middle trapping period more than 50% of the day’s catch was caught, but at 24.4 °C during the first period only 17% of the total day’s catch was caught (Tables 2 and 4). Below mean temperatures of 32.2°C most flies were Vol. 50, No. 3, July 1974 261 Table 3. The numbers and times at which the 5 most abundant tabanid species were caught during the first 10 and last 3 trap-days. Date Duration of trapping periods (hrs) Total Flies Species 0800-1100 1100-1400 1400-1700 27 May S. notatus 7 (3) a 181 (72) 65 (26) 253 through T. similis 26 (11) 111 (46) 106 (44) 243 12 June T. kesseli 21 (19) 49 (44) 42 (37) 112 A. comast es 12 (15) 47 (59) 21 (26) 80 C. s urdus 3 (5) 39 (68) 15 (26) 57 Subtotals 69 (9) b 427 (57) 249 (33) 745 13 June S. notatus 36 (35) 33 (32) 33 (32) 102 through T. similis 11 (48) 4 (17) 8 (35) 23 15 June T. kesseli 2 (17) 0 10 (83) 12 A. comastes 26 (81) 4 (13) 2 (6) 32 C. surdus 14 (67) 0 7 (33) 21 Subtotals 89 (47) c 41 (22) 60 (32) 190 Total 158 (17) 468 (50) 309 (33) 935 a Percent of total days catch. b Percent of total flies caught during 1st 10 days. c Percent of total flies caught during last 3 days. caught during the middle trapping period, regardless of temperatures (Tables 2—4), but the middle period usually was the warmest (Table 4). The usual late afternoon decline in activity of most species (Tables 2 and 3, period III) seemed related to the usual cooling temperatures during the last 2 hrs of the third trapping period. Excepting the 3 hot days of 13-15 June, on all days except 27 May and 12 June, the temperature was 20°C or lower, by 1700 hrs. On 4, 10 and 11 June, when the mean temperature of the 3rd period was nearly the same as that of the 2nd period and closer to the daily maximum than on the first 10 other days in Table 4, there was little difference in the numbers of flies caught during the 2nd and 3rd trapping periods. However, when the traps were operated for an additional period from 1700- 1930 hrs on 15 June, 24 tabanids were caught during this 4th period compared to 27 caught during the 3rd period. This indicates that when temperatures remain within a favorable range activity may continue until nearly dusk. The nearly 20% decline in numbers caught during the 3rd period as opposed to the 2nd period on 12 June, seemed associated with a temperature above 32.2 °C during most of the 3rd period (Tables 2 262 The Pan-Pacific Entomologist Table 4. Temperature data for the 13 days on which CCh-baited traps were operated in the 5-trap grid. Mean temp. a during 3 hr periods from: Daily max. Daily min. Date 0800-1100 1100-1400 1400-1700 27 May 22.2 26.1 25.0 27.8 7.8 3 June 15.6 19.4 16.7 21.7 1.7 4 June 19.4 24.4 23.9 26.7 3.9 5 June 20.6 24.4 23.3 26.7 9.4 7 June 17.8 21.1 20.0 23.3 6.1 8 June 22.2 25.6 23.3 27.2 10.0 9 June 21.7 25.0 23.9 26.7 8.9 10 June 18.9 22.8 21.7 24.4 8.3 11 June 20.0 25.0 25.6 27.2 5.0 12 June 24.4 30.0 31.7 32.8 9.4 13 June 30.6 37.8 38.3 40.0 15.6 14 June 33.9 41.1 40.6 42.8 18.9 15 June 32.8 37.8 36.1 40.6 18.9 a Mean temperatures were derived from the average of temperatures at the beginning and end of each trapping period. and 4). This was the first trapping period having temperatures above 32.2 °C, and as indicated by the subtotals in Tables 2 and 3 and the data in Table 4, the host-seeking activity of most species was markedly altered on hot days (13—15 June). During the middle part of hot days activity of all species was suppressed when the temperature rose above 32.2°C, but after several hours of temperatures between 32.2 to over 37.8° C most species exhibited a slight increase in activity dur- ing the 3rd period (Tables 2—4). As the temperatures of the 3rd periods on 13-15 June were nearly identical to those of the 2nd periods, the nearly normal percentage level of activity for the 3rd period (see subtotals of Tables 2 and 3) suggests that most species gradually ac- climated to the high temperatures. This also is suggested by the gradual increase in numbers of flies caught from 13-15 June. But since collecting containers were only collected at the end of the period, a sudden burst of fly activity with cooler temperatures preceding sunset also was possible. All of the first 10 species in Table 1, except H. aasa, were collected during periods having mean maximum tem- peratures over 37.8°C. The way in which the 5 most common species reacted on the 3 hot days, as contrasted with their daily activity on the 10 most productive trap days between 27 May and 12 June, is summarized in Table 3. The Vol. 50, No. 3, July 1974 263 Table 5. Mean number of S. notatus females caught per day per trap site during 13 days of trapping. a Distance From Center Of Trap Site Trap sites IV II III VI V I 2.38 2.92 3.08 4.85 5.31 8.77 5 nr 0 b 5 0 2 51 35 15 m 0 16 0 25 70 60 30 m 14 25 4 47 55 32 60 m 26 32 12 65 45 23 a Totals underscored by the same line are not significantly different at the 5% level of con- fidence when compared by Duncan’s multiple range test. 15 Total percentage canopy cover within a circle with the indicated radius. larger catches of T. similis and kesseli (than the other species) in the first period during the first 10 days is indicative of the lower minimum temperature thresholds at which they exhibited host-seeking activity. However, the number of A. comastes caught in the first period on these days (Table 3) is misleading when summarized in this manner because all 12 specimens were caught on days having a mean maximum tem- perature of 21.7°C, or higher, for the first period; 10 were caught on 12 June. By contrast, the first period catches of T. similis and kesseli on these 10 days were spread out over 6 and 7 days, respectively, 4 of which had mean maximum temperatures of 20° C or lower for the first period. If one excludes the data for 12 June, then S. notatus , C. surdus and A. comastes all showed similar patterns of activity during the first part of the study which were different from the daily pattern of activity exhibited by T. similis and kesseli (Table 3). However, on the 3 hot days the daily patterns of activity did not fall into 2 well-defined cate- gories (Table 3). Instead, (1) the activity of A. comastes was largely confined to the first period during 13-15 June, as one might have ex- pected from its response in the first period on 12 June; (2) the princi- pal activity of T. kesseli unexpectedly occurred in the 3rd period; (3) S. notatus exhibited a uniform level of activity throughout the day; and (4) C. surdus and T. similis exhibited bimodel activity peaks with somewhat greater activity in the first period. Tabanids Caught at Various Trap Sites. After transformation of the numbers of the 5 most abundant species caught/trap/day to the log (n + 1), analyses of variance revealed: (1) no significant dif- ference among site means for T. similis ; (2) a significant difference 264 The Pan-Pacific Entomologist Table 6. Mean number of T. kesseli females caught per day per trap site during 13 days of trapping. 51 Trap sites' 3 III VI II IV V I 0.69 0.92 1.46 1.46 2.00 3.00 a Totals underscored by the same lines are not significantly different at the 5% level of con- fidence when compared by Duncan’s multiple range test. b See Table 5 for the total percentage canopy cover within circles having radii of 5, 15, 30 and 60 m from each trap site. among site means for C. surdus and A. comastes; and (3) highly significant differences among site means for S. notatus and T. kesseli. For the numbers of S. notatus captured (Table 5, trap I was signifi- cantly different from all other traps except V and VI, but there was no significant difference among traps II— VI. For T. kesseli (Table 6), the number of flies caught in trap I was significantly different from only the number caught in traps III and VI, and there was no signifi- cant difference in the number of flies caught in traps II— VI. Although the more sensitive F test revealed a significant difference among trap sites for both A. comastes and C. surdus , there was no significant dif- ference among trap site catches of these at the 5% level of confidence when compared by Duncan’s multiple range test. In contrast to the Symphoromyia species studied, whose catches were influenced by the percentage canopy cover surrounding trap sites (Anderson & Hoy 1972), the 4 corner traps of the grid did not signifi- cantly decrease the number of most tabanid species caught in the center trap (Trap III). Only the numbers of S. notatus and T. kesseli caught in the center trap were significantly less than the numbers caught in one of the 4 corner traps (Tables 5 and 6). Wind direction also had no marked effect on tabanid catches in different traps as it did for the Symphoromyia species (Anderson & Hoy 1972). Thus, the more uniform occurrence of tabanids than snipe flies in all traps probably was related to their being stronger fliers than snipe flies and to their known visual response to traps and targets of different colors (Bracken, Hanec & Thorsteinson 1962, Morris 1963, Thorsteinson, Bracken & Hanec 1965). Although white is not very attractive to most tabanids (Barrass 1960, Bracken, Hanec & Thorsteinson 1962, Hansens 1947), the contrast between the white traps and the surrounding green grass, trees and shrubs made them very conspicuous. It also is commonly known that many species of tabanids, particularly the larger ones, disperse throughout unsheltered pastures where they attack livestock. Vol. 50, No. 3, July 1974 265 Table 7. A comparison of tabanids caught in traps and observed at hosts from 27 May through 15 June 1966. Fly No. flies per Adjusted no. flies Site sampling unit a per sampling unit b Traps 183 183 Deer 9 18 a Mean number of flies/trap or host. b Adjustment for hosts is 2X the actual number observed/sampling unit. This is based arbitrarily on the fact that tabanids commonly require about 4 minutes to feed (e.g. Philip 1931, and personal observations) and instantaneous fly counts were made on a host only at 10 minute in- tervals between 0800 and 1700 hrs. Tabanids Feeding on Deer. Females of 9 of the 14 species listed in Table 1 fed on deer, at least occasionally. Of the more common species trapped only H. aasa and C. coquillettii were not collected from or specifically recognized when feeding on deer. None of the last 3 species in Table 1 were collected from or seen feeding on deer. Tabanus punctifer, readily distinguished from the other local fauna, commonly was observed feeding on deer, whereas the remaining 8 species in Table 1 all were collected while feeding on deer. A total of 37 tabanids was observed feeding on deer from 27 May through 15 June; they fed at temperatures between 21.1-37.8°C. Two specimens fed between 0800-1100 hrs, 26 between 1100-1400 hrs, and 9 between 1400- 1700 hrs. Other species caught feeding on deer were H. procyon in March and April, and C. proclivis proclivis in June. All Chrysops species fed on the face and ears of deer, whereas the 2 Silvius species most commonly attacked the rear legs. The larger, more robust T. kesseli and T. punctifer usually fed on the neck, but occasionally on the back and rarely on the side of the face. Like the Chrysops species, H. procyon fed on the faces of deer, and recently it was found to be the principal vector of the arterial worm, Elaeophora schneideri , to deer in the Hopland study area (Anderson & Weinmann, Weinmann, et al. 1973). The above feeding sites are essentially the same as those observed for related fauna in eastern Canada (Smith, Davies & Golini 1970). Trap Efficiency for Tabanids. In general, the traps caught more tabanids than would be expected from concurrent observations of the deer. On average, a trap caught about 10 times as many tabanids as were observed at a host deer (Table 7), whereas about the same num- ber of Symphoromyia species were caught in a trap as were seen at a host (Anderson & Hoy 1972). Everett & Lancaster (1968) and Wil- 266 The Pan-Pacific Entomologist son (1968) also caught far more tabanids in CCU-baited traps than were seen attacking cattle. With respect to the number of blood meals taken from deer in 1964 and 1965, Hoy (1966) found that tabanids were outnumbered by Symphoromyia species by 40 or 50 to one. On the 13 trapping days from 27 May through 15 June 1966, 1369 Symphoromyia species were seen feeding on deer (Anderson & Hoy 1972) versus only 37 tabanids. Summary A total of 18 species of tabanid females was caught in C02-baited traps between 6 April and 26 June 1966. The baited traps caught about 10 times as many female tabanids as were observed at deer, and there were fewer significant differences among trap site catches for the most abundant tabanid species than for species of Symphoromyia previously studied. Wind direction and the percentage canopy cover surrounding traps had little effect on tabanid catches at various trap sites. The ratio of tabanids caught in dry ice-baited versus unbaited traps was 57:1. Daily host-seeking activity of various tabanid species occurred within well-defined temperature ranges; for all species activity was suppressed below 23.9 and above 32.2 °C. Normal host-seeking times for most species were markedly altered on hot days (daily maximum temperature above 32.2°C). Eleven of the 18 species trapped fed on deer at temperatures between 21.1 and 37.8°C; species of Chrysops fed on the face and ears, Silvius most commonly on the rear legs, Hyhomitra on the face, and Tabanus on the neck, back, and rarely the face. The ratio of Symphoromyia: tabanid species feeding on deer was about 40:1. Acknowledgments We thank Dr. W. M. Longhurst (University of California, Davis), Resident Zoologist, Mr. A. H. Murphy, Supervisor, and Mr. G. Con- nolly, Research Technician, all of the University of California Hopland Field Station, for their aid in making this study possible. We also thank Dr. H. F. Heady and Dr. D. Carneggie, School of Forestry and Conservation, University of California, Berkeley, for their provision of and assistance with aerial photographs, and Dr. C. B. Philip, Cali- fornia Academy of Science, San Francisco, California, who identified some of the tabanid species and confirmed the identification of most others. Vol. 50, No. 3, July 1974 267 Literature Cited Anderson, J. R. and J. B. Hoy. 1972. Relationship between host attack rates and CCL-baited insect flight trap catches of certain Symphoromyia species. J. Med. Entomol., 9: 373-93. Anderson, J. R. and W. Olkowski. 1968. 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Methods used in Utah for sampling tabanid populations. Mosq. News, 28: 356-61. Morris, K. R. S. 1963. A study of African tabanids made by trapping, Acta Tropica, 20: 16-34. Olkowski, W., J. R. Anderson and J. B. Hoy. 1967. Relationship between host attack rates and C0 2 -baited Malaise trap catches of certain tabanid species. Proc. 35th Ann. Conf. Calif. Mosq. Control Assoc., 35: 77. 268 The Pan-Pacific Entomologist Otsuru, M., Y. Saito, Y. Ohmori, S. Saito, S. Mizuno and H. Abe. 1965. The breeding places and some habits of the mountainous tabanid flies (Diptera:Tabanidae) . Jap. J. Sanit. Zool., 16: 123-32. Roberts, R. H. 1970. Tabanidae collected in a Malaise trap baited with C0 2 . Mosq. News, 30: 52-53. 1971. The effect of amount of COa on the collection of Tabanidae in Malaise traps. Mosq. News, 31: 551-58. 1972. The effectiveness of several types of Malaise traps for the collec- tion of Tabanidae and Culicidae. Mosq. News, 32: 542-47. Smith, G. E., S. G. Breeland and E. Pickard. 1965. The Malaise trap — a survey tool in medical entomology. Mosq. News, 25: 398-400. Smith, S. M., D. M. Davies and V. I. Golini. 1970. A contribution to the bionomics of the Tabanidae (Diptera) of Algonquin Park, Ontario: seasonal distribution, habitat preferences, and biting records. Can. Entomol., 102: 1461-73. Thornhill, A. R. and K. L. Hays. 1972. Dispersal and flight activities of some species of Tabanus (Diptera : Tabanidae) . Environ. Entomol., 1: 602-06. Thorsteinson, A. J., G. K. Bracken and W. Hanec. 1965. The orientation behavior of horse flies and deer flies (Tabanidae :Diptera) . III. The use of traps in the study of orientation of tabanids in the field. Entomol. Exp. Appl., 8: 189-92. Weinmann, C. J., J. R. Anderson, W. M. Longhurst and G. Connolly. 1973. Filarial worms of Columbian black-tailed deer in California. 1. Observations in the vertebrate host. J. Wildlife Dis., 9: 213-20. Wilson, B. H. 1968. Reduction of tabanid populations on cattle with sticky traps baited with dry ice. J. Econ. Entomol., 61: 827-29. Wilson, B. H., N. P. Tugwell and E. C. Burns. 1965. Attractiveness of dry ice to tabanids and horn flies under field conditions in Louisiana. Bull. Entomol. Soc. Amer., 11: 151, 174. Wilson, B. H., N. P. Tugwell and E. C. Burns. 1966. Attraction of tabanids to traps baited with dry ice under field conditions in Louisiana. J. Med. Entomol., 3: 148-49. Contribution to the Bionomics of the Grape Leaffolder, Desmia funeralis (Hubner) : A Laboratory Study with Field Observations (Lepidoptera: Pyralidae) M. Taskeen AliNiazee 1 Department of Entomology University of California, Davis 95616 The grape leaffolder Desmia funeralis (Hubner), which occurs on wild grapes from southern Florida to Canada and from northeastern Mexico to the Atlantic seacoast was probably accidently introduced to California in the late 19th century (Doutt et ah, 1969). Since its introduction, it has been a secondary pest of grapes in California, and in recent years has become an important pest in certain areas, par- ticularly in the eastern San Joaquin Valley. Barnes (1944) reported that in central California, the grape leaf- folder completes three full generations per year. In favorable years, a partial fourth generation may occur. Little additional biological in- formation concerning this insect is available. The present study was initiated to obtain life history information for use in developing pest management programs. Methods and Materials The insects were obtained from laboratory culture of the grape leaf- folder reared on fresh grape leaves. The culture was maintained in one-gallon ice cream cartons at the San Joaquin Valley Agricultural Research and Extension Center, Parlier. An ample supply of fresh leaves was provided five times a week. Insects were reared under controlled laboratory conditions at 23.9° ± 1° C and about 35% — 15%RH. Eggs less than 24 hour old were obtained by placing fertilized fe- males in ice cream cartons containing green grape shoots with two to five leaves. Mated females were released about 4 p.m. every afternoon and eggs were collected the following morning and were isolated in small plastic vials. Newly hatched larvae (less than 14 hours old) were isolated in small plastic vials supplied with about 10 one inch diameter circles of grape leaf. Observations were continued throughout the larval development and changes in the feeding habits, body measure- ments, molting and morphological appearance were recorded. After 1 Presently Assistant Professor of Entomology, Oregon State University, Corvallis, Oregon. The Pan-Pacific Entomologist 50 : 269 - 278 . July 1974 270 The Pan-Pacific Entomologist larvae pupated, they were again isolated in the small plastic vials for adult emergence. Adults were fed on a 10% sugar solution and their behavior, mating, and oviposition were observed in the laboratory. Egg production data were obtained from twice daily observations of freshly emerged pairs of adults in one-gallon ice cream cartons, pro- vided with sugar solution and fresh grape leaves for feeding and ovi- position, respectively. Oviposition preference experiments were con- ducted by releasing pairs of males and females in the ice cream cartons provided with different surfaces for oviposition. Field studies utilized pheromone trap and sweep samples to estimate the number of generations and daily flight patterns of the grape leaf- folder. Since the discovery of a sex pheromone in grape leaffolder (AliNiazee and Stafford, 1973), the virgin female traps have become an important survey tool for detecting low populations of this insect. Field trapping studies were conducted for six months (April 15th through October 18th) using closed sticky traps (AliNiazee and Staf- ford, 1971). Each trap was provided with one virgin female as a pheromone source. Virgin females were fed on a 10% sugar solution and changed twice per week. Average counts from three virgin female traps are reported in this paper. The sweep samples were collected at two hour intervals on six con- secutive days in August, 1971, using an insect net with 12 inch diam- eter rim. Ten sweep samples were collected in each of 4 different 0.25 acre blocks. Averages of ten samples are given in figure 9. Laboratory Results EGG STAGE : The eggs of the grape leaffolder are deposited singly, occasionally touching each other, but mostly with some space in be- tween. In the laboratory, eggs were laid on both surfaces of the leaves, mostly along the veins and vein angles. Both surfaces of the Thompson seedless grape leaves were equally attractive for oviposition, perhaps because the surface pubescence is less prominent in this variety. In the laboratory, almost all eggs were deposited during the dark period of a 16 hr. light 8 hr. dark cycle. This indicates that in nature oviposition occurs predominantly during the evenings and nights. Eggs are flat, shiny, mostly round to oval in shape, measuring 0.6 to 0.9 mm in length (Fig. 1). They are loosely attached to the leaf surface. The developmental period of the eggs is correlated with tem- perature. Observations during the summer of 1971 indicated that under natural conditions eggs hatch in about four to seven days depending upon the weather. Under laboratory conditions the average incubation Vol. 50, No. 3, July 1974 271 50i 40 -a « 3 °- _i S 20 M U 10 123456789 10 Days after Emergence Fig. 1. Eggs of grape leaff older. Fig. 2. Frequency of egg deposition by the grape leaffolder adults. period was 5.3 days (range 3.5 to 9; n = 202) at 23.9 ±1° C. About 84% of the eggs hatched within first eight days. Numbers of eggs laid by individual females varied from 6 to 431 (average 199; n = 24). A majority of the eggs were laid between the second and fifth day after emergence (Fig. 2). The average preoviposition period was 1.5 days, the oviposition period 6.7 days, and postoviposition period 1.3 days. Female moths failed to oviposit on a variety of artificial substances, in- cluding aluminum foil, plastic sheets and Saran® wrap, glass plates and smooth plywood sheets. LARVAL STAGE: Soon after hatching, the young larvae move to protected places and feed by webbing two or more leaves, initially in groups. In the field, the leaf rolls made by the earlier generations are preferred by young larvae. Laboratory observations indicate that young larvae feed between two leaves and do not make any leafrolls until they are about a week old. Initially, they make tiny leafrolls mostly towards the leaf edge and feed inside. As they grow the leafrolls are enlarged (Fig. 3) and by the time the larval development is complete, one or two leaves may be completely rolled. The density of rolls may be di- rectly related to the population density of leaffolders. However, the ratio of rolls to stages of the insect varies with the date. There are five larval instars. The developmental period of each instar 272 The Pan-Pacific Entomologist Table 1. Duration of the larval instars of the twenty-one grape leaff older larvae reared on grape leaves at 23.9 ± 1° C. Instar Duration in Days Average Maximum Minimum First 3.5 5.5 2 Second 3.9 5.5 2 Third 4 .4 6 3 Fourth 4.3 7.5 3 Fifth 6.1 9 2.5 varies considerably depending upon the temperature. In the laboratory tests at 23.9 ± 1° C, the average developmental period of the first instar larvae was 3.5 days, the second instar 3.9 days, the third instar 4.4 days, the fourth instar 4.3 days, and the fifth instar 6.1 days (Table 1). The length of the larval period varied from 16 to 31 days with an average of 20.8 days. Newly hatched larvae measure about 1.2 to 1.8 mm in length. They are creamy white to pale yellow. The second instar and older larvae are bright green to greenish-yellow. During development, the larvae develop characteristic markings that can be used to distinguish the dif- ferent instars (Barnes 1944). The full grown larvae are light green and measure about 16-22 mm in length (Fig. 4). The larval feeding capacity varies in different instars and in vineyards of different varieties. Barnes (1944) reported that the larvae ate 3.34 square inches of Em- peror grape foliage during their larval development. Out of this, less than 15% of the total foliage was eaten during the first three instars. PREPUPAL AND PUPAL STAGES : After completing development, the larvae become quiescent before pupation. Towards the end of the fifth instar, larvae stop feeding and make a pocket-like case about % to % inch long by cutting the leaves, or gluing the leaves together, and spend their prepupal and pupal stages in these envelopes. In almost all cases, a small piece of a grape leaf is cut on three sides and folded over to form an envelope. In the laboratory, paper towels which were put in rearing cages were similarly cut to form pupal cases. Sometimes, the envelopes are attached to other leaves. In the field, the envelopes fall to the ground with other foilage in late fall, and shelter the dia- pausing pupae until next spring. During the prepupal stage, the larvae shrink considerably and the Vol. 50, No. 3, July 1974 273 Figs. 3-7. Various instars of the grape leaf folder. Fig. 3. A leaf roll made by the grape leaffolder. Fig. 4. Mature larva. Fig. 5. Pre-pupa. Fig. 6. Pupa. Fig. 7. Adult. body color changes from greenish-yellow to pink. Most of the prepupae observed were much fatter and shorter than the fifth instar larvae (Fig. 5). The prepupal period of 42 individuals observed in the laboratory varied from 2 to 6 days with an average of 2.7 days (Table 2). Initially, the pupae of the grape leaffolder are light pink in color, but within a few hours after pupation they become dark brown. They Table 2. The developmental period of different stages of the grape leaffolder reared on grape leaves at 23.9 ± 1° C. Stage Individuals Observed Duration in Days Average Maximum Minimum Egg 202 5.3 9 3.5 Larval 43 20.8 31 16 Pre-pupal 42 2.7 6 2 Pupal 45 11.2 14 8 Egg to adult Males 32 39 54 32 F emales 34 38.2 51 30 Both sexes 66 38.6 54 30 274 The Pan-Pacific Entomologist Table 3. Oviposition and longevity data of 22 to 26 mated grape leaffolder females maintained on sugar solution at 23.9 ± 1°C. All times in days. Pre- Ovi- position Period Ovi- position Period Post-Ovi- position Period Eggs/ Female Adult longevity Males F emales Average 1.5 6.7 1.3 199 6.6 8.8 Maximum 2.5 13.5 2.5 431 12 15 Minimum 0.5 2.5 0 6 2 3 resemble the pupae of other pyralid moths and measure about 1 to 1.5 cm in length (Fig. 6). The pupal period of 45 individuals varied from 8 to 14 days, average 11.2 days (Table 2). ADJJLT STAGE : Before adult emergence, the pupae wiggle vig- orously and free themselves from the pocket-like pupal chamber. Soon after emergence, they move around quite a bit, probably in search of food. Under field conditions they hide in shady places underneath the vines during the daytime and begin their activities after sunset. They are medium sized black moths with patchy white spots on the body and wings (Fig. 7). Adult longevity is dependent upon many factors including tempera- ture and food availability. In the laboratory tests (Table 3), males fed on 10% sugar solution lived 2 to 12 days (average 6.6), while females lived 3 to 15 days (average 8.8). Adults fed on plain water did not survive as long; yet, they lived considerably longer than those provided with no water, which died in two to four days. Field Results The field results were limited to study of adult behavior, daily rhythm and seasonal flight pattern. Field observations indicate that one closed sticky trap with one virgin female as a pheromone source was much more effective than light traps. Results (Fig. 8) show that virgin female traps attracted enormous num- bers of males during a five month season. Although the population of grape leaffolder varies from field to field and from year to year, the comparison of light trap catch data (AliNiazee and Stafford, 1972) with the catches of grape leaffolder in the virgin female traps definitely reflects the remarkable effectiveness of the sex pheromone traps. Also, Vol. 50, No. 3, July 1974 275 Q. re o CL 0) a ra O E <0 4 > E 500 400 300 200 100 M J 1971 Fig. 8. Average number of adult males attracted to virgin female traps per week. the virgin female traps were very effective in detecting low levels of moth activity. Data presented in Figure 8 indicate that there were three popula- tion peaks during 1971. A relatively small peak occurred in October, which probably represented a partial fourth generation. The partial fourth generation may be a suicidal generation because of the lack of food availability and the onset of cold temperature regimes in late fall and early winter. Total numbers of moths collected per trap in each month of the study period indicate that the grape leaffolder population started at a lower level and then built up to a large population by the end of the season. Adult activity during peak summer months, which determines the in- tensity and spread of vineyard infestations, was studied in central Cal- ifornia vineyards. Field observations indicate that the grape leaffolder adults are nocturnal (Fig. 9). They begin their flight activity soon after sunset provided the temperature is not too high (above 32.3°C). However, the early evening flight activity is relatively insignificant. The activity increases with time, peaking about four to six hours after 276 The Pan-Pacific Entomologist