The PAN-PACIFIC ENTOMOLOGIST •• Volume 70 January 1994 Number 1 Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES (ISSN 0031-0603) The Pan-Pacific Entomologist EDITORIAL BOARD J. T. Sorensen, Editor R. V. Dowell, Associate Editor R. E. Somerby, Book Review Editor S. M. Sawyer, Editorial Assist. Paul H. Amaud, Jr., Treasurer R. M. Bohart J. T. Doyen J. E. Hafemik, Jr. J. A. Powell Published quarterly in January, April, July, and October with Society Proceed¬ ings usually appearing in the October issue. All communications regarding non¬ receipt of numbers, requests for sample copies, and financial communications should be addressed to: Paul H. Arnaud, Jr., Treasurer, Pacific Coast Entomo¬ logical Society, Dept, of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Application for membership in the Society and changes of address should be addressed to: Stanley E. 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This issue mailed 27 April 1994 The Pan-Pacific Entomologist (ISSN 0031-0603) PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A. © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). The Pacific Coast Entomological Society's portion of page costs for The Pan-Pacific Entomologist , 70 (1), were paid by the society's Henry Clinton Fall Memorial Publication Fund The author's page charges were paid by The Biological Control Program, Integrated Pest Control Branch, California Dept, of Food 8c Agriculture Pacific Coast Entomological Society Statement of Income, Expenditures and Changes in Fund Balances for Years Ending September 30, 1991 and 1990. 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The taxonomic status is changed for E. ( E .) knowltoni braggi Hottes NEW STATUS, E. agilis Hottes NEW SYNONYM, E. claremontiana Hottes NEW SYNONYM, E. cocheta Hottes NEW SYNONYM, E. gillettei Hottes NEW SYNONYM, E. maculata Hottes NEW SYNONYM, E. monelli Hottes NEW SYNONYM, E. oregonensis Hottes NEW SYNONYM, E. palmerae Hottes NEW SYNONYM, E. patchae Hottes NEW SYNONYM, E. pergandi Hottes NEW SYNONYM, E. pineti Hottes NEW SYNONYM, E. robusta Hottes NEW SYNONYM, and E. swaini Hottes NEW SYNONYM. A phylogenetic tree for the genus is reviewed; that estimate, which used Pseudessigella as a outgroup and employs evolutionary quantitative genetic rationales, was produced using discriminant function analysis and a maximum-likelihood net¬ working algorithm, because conventional cladistic characters were inadequate within the genus. The phylogeny is corroborated because it closely reflects the genetic relationships of the aphid’s Pinaceae hosts, and their biogeographic origins. Essigella appear to have evolved with their hosts, or in a resource-tracking fashion, and seem to display instances of character-displacement among closely related species in (or near) sympatry, presumably as a result of competition of their host pines as resources. Key Words. — Insecta, phylogeny, host associations, character displacement, evolutionary quan¬ titative genetics This study addresses the systematics, phylogeny and host associations of Es¬ sigella. The genus, one of three composing the subtribe Eulachnina (Lachninae: Cinarini), is restricted to North America and is the only native Nearctic group of the subtribe. Essigella are linear-bodied and feed on the needles of Pinaceae, chiefly Pinus but also Pseudotsuga and Picea. They are solitary aphids that move quite rapidly when disturbed; several may group facultatively near the base of a needle, where they may be tended by ants. Cage studies of individual aphids (unpublished data) on marked needles indicated that adult virginoparous apterae of Essigella readily wander over pine branches. Essigella is often most abundant early in the season (Burke 1937), and may damage pines (Turpeau & Remaudiere 1990). However, its populations may lower by summer, making the needle yellowing that it causes difficult to diagnose (Brown & Eads 1967). In the southeastern U.S., Essigella density peaks between September and March, and falls to its lowest level during June through August (Patti & Fox 1981a), although outbreaks have occurred in May and June (Hood & Fox 1980). Essigella seem to occur most heavily on young trees, particularly on the lower east side (Patti & Fox 1981b), which has afternoon shading. In the west, Essigella 2 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) occasionally damages Christmas tree plantations (California Department of Food & Agriculture, unpublished data). Sampling (Flood & Fox 1978) and control regimes (Hood & Fox 1980) have been developed for Essigella in southeastern U.S. lumber plantations. Eastop & Hille Ris Lambers (1976) list 21 species in the genus. Lachnus cali- fornicus Essig (1909) was the first described species, but immediately thereafter Del Guercio (1909) described the genus Essigella, with L. californicus Essig as its type species. A second species, E. pini Wilson (1919), was described a decade later, with two more, E. fusca Gillette & Palmer (1924) and E. hoerneri Gillette & Palmer (1924), following shortly thereafter. The genus then sat taxonomically dormant until its synopsis by Hottes (1957), in which 16 new names were created [E. agilis Hottes, E. braggi Hottes, E. claremontiana Hottes, E. cocheta Hottes, E. essigi Hottes, E. gillettei Hottes, E. knowltoni Hottes, E. maculata Hottes, E. monelli Hottes, E. palmerae Hottes, E. patchae Hottes, E. pergandi Hottes, E. pineti Hottes, E. robusta Hottes, E. swaini Hottes, E. wilsoni Hottes]; another name, E. oregonensis Hottes (1958), was added a year later. While working on aphids in California during the 1960s, D. Hille Ris Lambers attempted to treat Essigella. He concluded (unpublished notes) that the only available key (Hottes 1957) to Essigella did not work for numerous reasons, and that the genus needed a major revision using similarly cleared and mounted specimens. In 1978, he advised me (D. Hille Ris Lambers, personal communi¬ cation) that the systematics of Essigella was extremely difficult, and that he ranked the genus as one of the most taxonomically problematic among aphids. Since then, I have analyzed the biological groups in Essigella in relation to their hosts (Sorensen 1983), suggested a phylogeny (Sorensen 1987a), described three new species (Sorensen 1988), analyzed the cladistic placement of the genus among the Eulachnina (Sorensen 1990), assessed phylogenetic changes in shape component variance between Essigella and its sister group (Sorensen 1991), and presented analyses deciphering the biological groupings of Essigella on Pinus contorta Doug¬ las (Sorensen 1992a). The problems of Essigella' s systematics are due to the exceptional reduction of morphological attributes, over that of an already neotinous subtribe (Sorensen 1990). Retained features in the genus are either extremely variable and overlapping among species, represent reductions, or involve pigmentation suites, which often run counter to morphology, and that grade from fully expressed to absent within populations. In addition, several instances of character displacement seem to occur among closely related species under sympatry, or near sympatry (Sorensen 1992a, unpublished data). Discrete characters are unusual within the genus, which shows many internal homoplasies and few conventional autapomorphies or reliable synapomorphies (Sorensen 1987a). In Essigella, many characters have transfor¬ mations that are nebulous and unusually difficult to polarize. Traits that are considered taxonomically meaningful in the genus are often more typical of in¬ traspecific variation in other lachnines, and the converse is also true. Therefore, interspecific variance in Essigella seems to be antipodal to that encountered among many other closely related aphids; traits that might be considered to be indicative of close relationships within the genus, often turn out to show merely superficial resemblance because of homoplasy, intraspecific variance or apparently faulty ontogenic physiognomy. 1994 SORENSEN: A REVISION OF ESSIGELLA 3 Attempts to determine character plasticity in Essigella, by culturing under vari¬ able environmental conditions in the laboratory, failed for several reasons. These aphids, in contrast to others, often could not be successfully transported to the laboratory from field. Those Essigella brought to the laboratory alive were difficult to rear and transfer among host plants. Their laboratory manipulation was ham¬ pered by their solitary, but mobile, habits; to be located, specimens often had to be jarred from the needles of their host, and frequently did not reestablish on the plants; a trait noted by Hottes (1957). Electrophoretic analysis of field collected samples was also problematic: assessing potentially mixed field populations was difficult because morphological differences among species were not yet understood, and isozymic responses were unclear. This revisionary research was based upon newly collected material with proper host associations from throughout most of the range of the genus, following a suggestion from D. Hille Ris Lambers (personal communication). In conjunction with the more traditional approaches to aphid systematics, multivariate analyses were necessary to determine intra- and intersample variation. Ultimately, pre¬ viously existing Essigella material was studied and fit into the derived taxonomic scheme without incident. This approach allowed an unbiased initial view of the genus, which I believe was a prerequisite to its successful revision. Methods and Philosophy Collection and Processing of Specimens. — All potential Essigella hosts, includ¬ ing all conifer genera, were sampled during 1977-1979 for this revision (Sorensen 1983). Collections were made from major geographic populations of the aphids’ hosts throughout western North America, north of Mexico (see Critchfield & Little 1966); of these, over 340 host/sites yielded Essigella. Over 7000 specimens, with an average of over 20 per collection, were collected, processed and studied; ad¬ ditionally, existing material was borrowed from depositories. Elsewhere, I have listed all locations, with host associations, where my sampling found Essigella (Sorensen 1983: appendix Al), and have provided a distribution map of all sam¬ pling locations (Sorensen 1983: figs. 1.1, 1.2). Hottes’ (1957) Essigella specimen processing was poor and resulted in the obscuring of characters or their erroneous interpretation (e.g., Hottes 1957: 108, key couplet 1, “tarsal claws not distinctly bifurcated”). As stressed by D. Hille Ris Lambers (unpublished notes, personal communication), during this project I have processed and mounted all Essigella specimens using standardized clearing and mounting techniques. Preparation followed Hille Ris Lambers (1950), and required sequential boiling in: (a) 95 percent ethanol [5 min], (b) 10 percent potassium hydroxide [4 min], and (c) saturated chlorophenol until the body con¬ tents were translucent [ca. 7-8 minutes]. Treatment times are important for pres¬ ervation and standardization of subtle pigmentation differences. Processed spec¬ imens were mounted immediately (or rarely stored up to 1 week in chlorophenol) in Hille Ris Lambers medium (gum arabic 12 g, concentrated glycerine 6.5 g, chloralhydrate 20 g, distilled water 20 cc). Slides were thick (deep), thus mini¬ mizing common compression artifacts for body width measurements. Immedi¬ ately after mounting, collection numbers were etched into the slides to prevent any mix-up of samples before labeling. 4 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Analytical Methods.— In addition to conventional analytical techniques used in aphid systematics, Sorensen (1983) conducted multivariate analyses to circum¬ scribe the biological groupings of Essigella. Those multivariate analyses were restricted to adult viviparous apterae to limit the influence of seasonal polymor¬ phism and developmental trait variation. Inadequate availability prevented the separate analysis of other morphs. Initially, only Essigella collected from natural stands of native hosts were analyzed to limit potentially confusing environmentally induced variation; later, Essigella from planted stands and nonnative hosts were incorporated without incident. For analysis, individuals from samples were first divided into initial groups by host and geographic location. These initial groups were then circumscribed using ordination analyses to establish common covariant character patterns among both the individuals and groups, and to determine the interrelationships among the initial groups. In the original, exploratory analyses (Sorensen 1983), several subset combinations of up to 66 morphometric characters were employed; these were later reduced to a 26 character subset (see Sorensen 1991: table 1) that was used to circumscribe all final biological groups within the genus. Quantitative analyses of the initial groupings involved the following steps: (1) exploratory delimitation of relationships using principal component analysis [Duncan & Phillips 1980: program PNCOMP] and clustering techniques [Duncan & Phillips 1980: programs GRAPH and CLUST], followed by reassessment of the groupings into biological groups; (2) bivariant plotting of various characters for the deduced biological groups using extended data sets to determine the sim¬ plest character combinations that best allow their separation; (3) corroboration of inter- and intragroup variance, using the deduced biological groups as “knowns” in discriminant function analysis [Nie et al. 1975: SPSS, version 7, program DISCRIMINANT, direct selection mode, Wilks-A criterion]. Sorensen (1992a) details an example of the use of these procedures in deciphering biological group¬ ings in a species complex within Essigella. After the final biological groups were assessed, they were cladistically verified, wherever possible, by establishing con¬ ventional autapomorphies or synapomorphies with reference to out-groups (see Sorensen 1990). Ultimately, a phylogenetic tree was produced for Essigella (see phylogenetic analysis section). With respect to the ordinations used (Sorensen 1983), populations of parthen- ogens can cause problems when assessing their demarcations into groups. Because aphid clones within samples could not be practically ascertained, exemplars were employed in this project. Normally, these were restricted to a single individual per sample. This prevented reduction of effective statistical sample size ( n ); it thus minimized the miscalculation of intragroup genetic variance, because only environmental and error components are left to account for observed intragroup dispersions around centroids as samples become saturated with identical geno¬ types. False low estimates of intragroup genetic variance cause overestimation of intergroup divergence and phyletic anagenic distance (Sorensen 1987b). Operational Species Concepts Employed. — Determining aphid species, in gen¬ eral, is often problematic because of their anholocyclic lineages, which often survive indefinitely in noncontinental climates. I consider an operational aphid species, sensu Doyen & Slobodchikoff (1974), to be those recombinant individuals or parthenogenetic populations that share a unique phyletic lineage. This is pref- 1994 SORENSEN: A REVISION OF ESSIGELLA 5 erably recognized by a conventional autapomorphy. Alternatively, lineages may be deduced from their congruity of conventional diagnostic synapomorphies or, if necessary, plesiomorphies, provided they are genetically stable (not induced by abiotic factors). If nonautapomorphies must be used to deduce a species, its popula¬ tions also must show common distributional and host coincidence; additionally, species must be multivariately circumscribed following the evolutionary quan¬ titative genetic logic outlined in Sorensen & Foottit (1992), using principal com¬ ponent and discriminant function analyses (e.g., Sorensen 1992a: figs. 3 and 6). Here, subspecies are recognized only when they impart ecologically or evolu- tionarily relevant information; they denote divergent or distinct allopatric geo¬ graphic populations (sensu Mayr 1969). This differs from a common usage in aphid systematics, where a subspecies is often named to denote the sympatric variance shown by phenotypically deviant individuals, sometimes on the same host. I consider an operational aphid subspecies to be allopatric and show either: (a) tighter agglomeration in clustering analyses (e.g., Sorensen 1992a: GRAPH analyses), or (b) a more restrictive distribution in the attribute space of ordinations, than do their species within analyses of species-groups (e.g., Sorensen 1992a: fig. 3, as the SNV vs. CAS and RMT distribution in PCA-1). Taxonomic Key Usage. — The key to Essigella species requires adult viviparous apterae (subgenital plate present and entire, gonapophyses and siphunculi present) and, in some instances, their ultimate stadium nymphs (subgenital plate and gonapophyses absent, abdominal dorsum membranous with distinctly demarcated plates at dorsal setal bases). Because Essigella species are exceptionally variable, with overlapping interspecific variation in many traits, calculations of discrimi¬ nant functions (DF), based on several characters, are sometimes required for morphologically based identification. The key appears to have a reliability of at least 90%; host plant information is included for more positive identification. Because of references to subtle pigmentation differences and DF calculations, remounting of some existing slides may be necessary to use the key. Referral to paleness or pigmentation in the keys, diagnoses or discussions throughout this work are to slide mounted material. Slides must be properly cleared, noncom- pressed, and mounted so that the sagittal plane of the aphids is oriented perpen¬ dicular to the slide. Intrapopulational variance in Essigella, or varying slide prep¬ arations, will require that you judge a circumstance in a key couplet to advance by alternative routes. If you are unsure when asked about the degree of specimen pigmentation or slide compression, elect the nonpigmented or compressed slide option. Questions about pigmentation of the body dorsum refer to the background intensity, exclusive of setal bases or muscle attachment plates. Slide compression should be judged conservatively; it is most apparent as a distortion (widening) of the outline of the head, anterad of the eyes, or the rostral base. (Although body widths, especially head width, are important in Essigella, they are generally not used, or are minimized [with warning], in the key due to compression or orien¬ tation faults in many slides.) When asked the number of setae on abdominal tergum VIII, or the number and pattern of dorsal setae on abdominal terga III— IV, compare several specimens and use the mode (the latter can be most easily distinguished on ultimate stadium apterae nymphs, where each seta is on a scleroite). DFs are required for specimens whose trait variance occupies an interspecific 6 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) overlapping range; couplets requiring DFs occur only after those based on non¬ overlapping variation. DF calculation requires (a) the measurement of several characters, (b) the multiplication of each by a given coefficient, (c) the summation of all resulting products, and (d) the addition of a constant [for adjustment]. This grand sum represents a discriminant score (D.S.), whose value represents a thresh¬ old figure for classification of an individual. Where DFs are required, for optimal classification their scores must be calculated to five figures after the decimal, and be based on measurements in mm to three figures after the decimal; the latter may require magnifications of 300-400 x, and error of as little as 5% can result in misclassifications. Presentation of Taxonomic and Distributional Data. — The sequence of species descriptions in this revision reflects their phylogenetic order. Variability and phenotypic similarity among Essigella species preclude their illustration here, except for a schematic topological map of dorsal setal positions on the abdominal terga; my previous Essigella illustrations (i.e., Sorensen 1988: figs. 1-3; 1991: fig. 1) show only that these aphids are relatively linear, varying somewhat in width, or that traits are variable (Sorensen 1991: fig. 2). The keys provided are the most reliable means of identification. I consider all taxonomic names indicated to be new as being legally and orig¬ inally described here; previously, they were considered to be manuscript nomens in Sorensen (1983), a thesis that is unpublished for nomenclatural purposes under nomenclatural rules (ICZN 1985: Articles 8-A1, 8-A3, 8-B, and presently 8-C, 9-2, 9-3, 9-4 and 9-6). Distributional data are listed for each species: use of “JTS” under material examined indicates J. T. Sorensen as the collector. Distributional maps for Es¬ sigella species depict data locales, differentiated as JTS vs. nonJTS collections, superimposed over host ranges; the latter were derived from Critchfield & Little (1966), Little (1971), and Griffin & Critchfield (1972). Abbreviations and Depositories. — My coded references to Essigella taxa else¬ where (Sorensen 1983, 1987a, 1992a, b) are listed at the end of each of the sections for each taxon. The U.S. National Museum of Natural History, Washington, D.C., is represented as NMNH. The Canadian National Collection, Agriculture Canada, Ottawa, Ontario, is represented as CNC. Several of Essig’s types are deposited in the Essig Museum of Entomology, at the Department of Environmental Sciences, Management and Policy, University of California at Berkeley; in 1993, that de¬ partment was created through an amalgamation of several others, including the Department of Entomological Sciences, which formerly housed the Essig Museum. Character Discussion Essigella are extremely variable aphids. Their normal character variation is discussed under each species or subgenus, as are some transformations, synapo- morphies and autapomorphies. This section concentrates on the definitions and phylogenetic transformations of traits; diagnostic autapomorphies are not dis¬ cussed here unless they show intraspecific variation or represent an independent state within a transformation series with multiple states. Aberrations. — Rare Essigella individuals exhibit aberrant traits that are virtually always expressions of plesiomorphic states that should not occur on their species (e.g., the number of dorsal hairs on the abdomen). This probably reflects the 1994 SORENSEN: A REVISION OF ESSIGELLA 7 failure of regulatory genes that normally suppress plesiomorphic phenotypic ex¬ pressions, which is more likely than new creation of a derived state; the suppressed plesiomorphy, already encoded in the genome, requires merely a gene failure for expression. There also appear to be similar suppression failures involving onto- genic phenotypes, where traits of one stage or morph show up erroneously on another (e.g., allometric differences in relative appendage length). Hottes (1957) sometimes erroneously considered such individuals to be new species. Fusion of Terga.— Sorensen (1983 [data used in 1987a], 1988, 1990, 1991, 1992a) previously had misinterpreted the fusion of the meso- and metanota, and demarcation of abdominal tergum I in Essigella. The correct interpretation of fusion of the dorsum in Essigella is: head + pronotum fused, meso + metanota fused, abdominal tergum I free (except E. essigi ), abdominal terga II-VII fused, abdominal tergum VIII free. Previously, I thought the meso + metanotal fusion was solely the mesonotum, abdominal tergum I was the metanotum, and abdom¬ inal terga I-VII, instead of II-VII, were fused. This error was discovered when R. L. Blackman (personal communication) suggested that the autapomorphous tergal fusion in E. ( E.) essigi involved abdominal tergum I, rather than the meta¬ notum. With the exception of species descriptions in Sorensen (1988), corrected here, this reinterpretation does not affect the conclusions of any of those studies; because only the relative definitions of characters, not data, were erroneous. It does mean, however, that the meso + metanotal fusion in Essigella is an additional synapomorphy for the genus, beyond those listed in Sorensen (1990); a meso- metanotal demarcation line is evident in Pseudessigella. The character definitions in error previously are corrected here, as: old ‘number/ [code] definition error ’ > correction ; but {comments} may be injected or substi¬ tuted for full definitions. Sorensen (1983, 1991: table 1): ‘12/[L2THOR] {length of mesonotumY > fused meso + metanota; ‘13/[L3THOR] {length of metano- tum} 9 > abdominal tergum 7; ‘15/[LVABSC],’ ‘16/[NHAB2DT]’ and ‘17/ [NHAB2M]’ {all on abdominal segments II-IV] > III-V. Sorensen (1990: table 1): ‘17/abdominal terga 7-7’ > 2-7. Sorensen (1992a: table 2): ‘ 3/mesothoracic terg. 17 > fused meso + metathoracic ; ‘7 / metathoracic terg. 17 > abd. seg. 7; ‘10/ abdomen {segments 1-7} L, excluding seg. 8’ > abdomen seg. 2-1 L; ‘25/marginal seta L on metathorax ’ and ‘54/W between most-mesal pair of dorsal (spinal) setae on metanotum ’ > abd. terg. 7; ‘28/dorsal (spinal or pleural) seta L on abd. terg. 2’ and ‘29/marginal seta L on abd. terg. 2’ > 3; ‘30/ventral seta L on abd. seg. 2/ ‘41/spiracular plate L on abd. seg. 2’ and ‘57/spiracular plate W on abd. seg. 2’ > 3; ‘ 40/L of presiphuncular abd. {including segment 1}’ > {add} excluding abd. seg. 7; ‘44/sagittal L of largest ventral abd. sclerite on seg. 2-4 7 ‘45/L of dorsal (spinal or pleural) setae between dorsal muscle attachment plates on abd. seg. 2-4, ’ ‘46/N of dorsal (spinal or pleural setae between dorsal muscle attachment plates on abd. seg. 2-4 ’ and ‘58/largest ventral abd. sclerite W on seg. 2-4 ’ > 3-5. Sclerotization.— Sclerotization is not equivalent to pigmentation, although sclerotized areas usually are at least somewhat pigmented. Here, it is the color independent distinctiveness or thickness of a body surface in comparison to an adjacent membranous area, as is evident in Essigella' s tergal fusions. The abdominal dorsum of adult viviparous apterae of Essigella is sclerotized, in contrast to the other genera of the Eulachnina (Sorensen 1990). This scleroti- 8 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) zation is universally present in all adults of that morph, although pigmentation may be quite pale, making it sometimes virtually impossible to detect. When adult viviparous apterae of Pseudessigella and Eulachnus are superficially com¬ pared to Essigella, they resemble the latter’s ultimate nymphal stadia, which lack the dorsal sclerotization. In Essigella, the sclerotized dorsum is what should have been referred to as the “cape” by Hottes (1957); he used the term with reference to pigmentation for more melanic specimens, noting simply its presence or ab¬ sence, despite its actual gradation. The sclerotization of the abdominal dorsum of Essigella seems to have occurred at the inception of the genus, and represents a synapomorphy. It has been strengthened or reduced in various lineages within the genus, and varies homoplasiously from faint to heavy within species groups. Examination of the abdominal dorsum of later stadia nymphs of Essigella, versus adult viviparous apterae, best reveals the latter’s abdominal sclerotization. On adults, scleroites at the base of the dorsal setae of the abdomen have been lost or reduced, probably due to incorporation into the general sclerotic field of the abdominal dorsum; at most, the remnant scleroites appear as indistinctly bordered darker areas at the setal bases (see pigmentation). In contrast, scleroites are usually prominent, with well defined borders on the membranous abdomen of nymphs and alates. Pigmentation. —Pigmentation represents melanization, and, where quantified, is expressed here as a density equivalent to the percentage of solid black in a screen of 52 lines per centimeter (Sorensen 1983). Referral to paleness or pig¬ mentation in the keys, diagnoses or discussions throughout this work are to slide mounted material. Although sometimes aphid taxonomists (in litt.) place little weight on minor or trivial pigmentation patterns, in aphids certain pigmentation suites—denoted here as a covariant series of patterns occurring within phyletic lineages—are more stable within species, over their seasonal polymorphic changes, than are simple length ratios for body segments (D. Hille Ris Lambers, personal communication). Often, major pigmentation suites on viviparous apterae of Es¬ sigella are indicative of one or several species; these are considered to be taxo- nomically important when they have genetic basis and are monophyletic. Unusual environmental conditions can cause variation of pigmentation intensity in aphid species, but not a change in a pigmentation suite. Consequently, reliable pigmen¬ tation is preferred here for identification, where feasible. Unfortunately in Essigella, the expression of a reliable pigmentation suite that is characteristic for a species or group, can vary from strongly pigmented to completely pale within populations; whereas other Essigella species are always pale. Furthermore, to hamper identification, faint pigmentation in slide mounted material can be bleached by excessive clearing or prolonged exposure to sunlight. Geographic variation can also occur in pigmentation suites. For example, many minor pigmentation tendencies (e.g., subtle variations of shade or intensity of melanin) recur as homoplasies within Essigella ; these are usually of little taxo¬ nomic interest, except in regard to intraspecific geographic variation. Two categories of pigmentation suites can be recognized within Essigella for pigmented individuals of adult viviparous apterae: the expression of shading among the tibiae, and of the background of the dorsum of the abdomen or entire body. Reference to body or abdominal pigmentation in the text and keys refers only to background shading, not to darkened muscle attachment sites and setal 1994 SORENSEN: A REVISION OF ESSIGELLA 9 bases. The numerous pigmentation suites are discussed in the descriptions, di¬ agnoses and discussions of species. A general darkening of the body dorsum, as a homoplasious apomorphy, occurs in E. ( L .) eastopi, E. ( E .) essigi, E. (E.) critchfieldi and E. (E.) knowltoni knowltoni. Of these taxa, the dorsum of E. ( L .) eastopi is considered to be an autapomorphy because it shows a unique dorsomedial lightening of the thorax and abdomen (state A). In contrast, the dorsum is evenly dark (state B) in E. (E.) essigi, E. (E.) critchfieldi and some E. ( E .) knowltoni knowltoni (Cascade Range); however, other E. ( E .) knowltoni knowltoni (Rocky Mountains) show a lightening of the frons and head (state C), or of the entire dorsum. The evenly paler dorsum of E. ( E .) knowltoni braggi (state D) is assumed to be a apomorphic reversal from state C. The transformation for the trait is assumed to be A <— B —** C —» D. The most useful tibial pigmentation suite involves the mesotibiae being at least subtly, and usually substantially, paler than both the pro- and metatibiae. This apomorphy is unique to E. (Lambersella), where it is present for pigmented individuals of all morphs. It can be hard to detect on some darkly pigmented E. (L.) eastopi, however, because their legs are quite lightened. Hottes (1957) and Hille Ris Lambers (unpublished notes) erroneously regarded the presence or absence of pigmented spots that often surround the bases of the dorsal setae on the abdomen of adult viviparous apterae to be of taxonomic value. Instead, the spots represent intraspecific or usually intrapopulational variation in most species. They are seldom present, and then only subtly, in E. (Archeoessigella) and E. (E.) wilsoni. Within most species, the spots usually occur only on mod¬ erately pigmented individuals, and I suspect their expression is a remnant of a juvenile factor; they are no doubt homologous with the scleroites that are in¬ variably present in nymphs (see sclerotization). Abdominal Chaetotaxy. — In Essigella, unlike Eulachnus (D. Hille Ris Lambers, personal communication), the number and distributional pattern of setae on the abdominal dorsum appear unaffected by environmental factors. In a principal component analysis of all Essigella taxa (Sorensen 1983), the number of abdom¬ inal setae show variation that is subordinate only to general-size (component 1). These setae can be divided into three categories: (a) dorsal setae on terga III-IV, (b) marginal setae on terga III-IV, and (c) setae on tergum VIII. Within any species, the characteristic state for each of these setal categories is stable among all known morphs. Marginal setae, at least in more apomorphic states, do not appear to show the same type of intraclonal variation as has been found to be dependent upon the number of sequential generations after the fundatrix in other aphids (see Crock & Shanks 1983, Blackman et al. 1987). In this study, such variation would have been detectable as seasonally related variation within natural populations, which would be in contrast to covariant patterns of other diagnostic attributes for taxa; to date no such variation has been found. The dorsal setae of the abdomen are defined here as those setae that occur between (mesad to) the most sagittally-oriented pair of the three muscle attach¬ ment points on each side of the dorsum of each abdominal tergum. The dorsal setae can be further broken into two groups: dorsal major setae, which occur mesally, and dorsal minor setae, which occur more laterally. These subgroupings (Fig. 1) become apparent when tracing the evolutionary transformations of the dorsal setae in the genus. Although the setae may be difficult to see in some cases, 10 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) lateral pair of dorsal muscle /' attachment plates \ anterolateral-most X dorsal minor seta A "0\ / o •/ * triangular set of dorsal major setae Q • B ■ o • ■ ■ o • C D o E F □ □ o © Figure 1. Schematic maps of approximate positions of dorsal setae on an abdominal terga III and IV. See comments under abdominal chaetotaxy in the character discussion section. Maps represent typical relative setal positions, which can vary; the right and left half of each map shows the more common possible positions. Large gray circles = dorsal muscle attachment plates; each: black square = dorsal major seta, black circle = dorsal minor seta; white squares or circles (majors or minors, respectively) show other possible setal positions that are usually absent. A .—Pseudessigella [also see Sorensen (1991: fig. 3)]. B.— E. ( Archeoessigella ). C.—E. (Lambersella ) expression 1 [i.e., most often E. ( L .) hillerislambersi]. D.—E. ( Lambersella ) expression 2 [i.e., most often E. (L.) fusca, E. (L.) eastopi ] (note that a anterolateral-most dorsal minor may move posteromesally). E.—E. ( Essigella ) expression 1 [e.g., E. (E.) essigi, E. (E.) wilsoni, E. (E.) alyeska, E. (E.) critchfieldi, E. (E.) knowltoni]. E.—E. (Essigella) expression 2 (e.g., E. (E.) californica, E. (E.) hoerneri, E. (E.)pini\. Transformation: A -»• B -> C/D —» E —> F. 1994 SORENSEN: A REVISION OF ESSIGELLA 11 they are usually well defined and easily traced in the later stadium nymphs of viviparous apterae (and their adults in Pseudessigella ), where they occur on sclero- ites in the membranous field of the abdominal dorsum. In Pseudessigella (Fig. 1A; Sorensen 1991: fig. 3), the dorsal majors occur as two bilateral, but mesal, groups of three triangularly-arranged setae that flank the mid-line. Each triangular group has one setae to the anterior and two that flank it to the posterior. The dorsal minors occur as one to several smaller setae that are anterolaterad to the dorsal majors. They are posteromesad to the anterad plate of the sagittally-oriented muscle attachment plates on each segment side (i.e., the anterolaterad of the three plates). Apparently at least one dorsal minor seta remains in this position, relative to that muscle attachment plate, throughout the trans¬ formations among the more plesiomorphic Essigella. In E. ( Archeoessigella ) (Fig. IB), the dorsal major setae remain largely un¬ changed, although occasionally an anterad seta in either triangular set may be absent. The dorsal minors, however, usually increase in number over their ex¬ pression in Pseudessigella, and several move posteromesally, coming nearer to the lateral-most dorsal majors. This has the effect of creating what superficially appears to be a band of setae in somewhat irregular positions across the tergum. There is, however, usually a retained association between at least one dorsal minor [the “lateral-most dorsal minor” in descriptions of species here] and the anterolat¬ erad muscle attachment plate. In E. {Lambersella), the next evolutionary step, two transitional expressions are found. In the more plesiomorphic arrangement (Fig. 1C), the dorsal majors remain intact, as do the dorsal minors, although the latter may be reduced in number. In the alternative state (Fig. ID), among the dorsal majors, an anterad seta of either, or both, triangular set(s) may be lost; the dorsal minors may be reduced to two, and the most anterolaterad of these may or may not move posteriorly, away from its formerly associated position near the anterolaterad muscle attachment plate. This can result in the occasional occurrence of an in¬ dividual (or population) with a series of only four dorsal (major + minor) setae occurring in a roughly straight line across the dorsum [a condition that mimics the first transition in E. ( Essigella ) mentioned below]; usually, however, when these setae are reduced to four, one of the laterad dorsal minors retains its an¬ terolaterad position. These two expressions in E. {Lambersella) are not necessarily sequential, and either may occur within populations of any species of that sub¬ genus. Two expressions also occur within E. ( Essigella ), but these derived states are sequential. The first (Fig. 1E) typifies all E. ( Essigella ) with eight (or more) dorsals [i.e., E. ( E .) essigi, E. ( E .) wilsoni, E. (E.) alyeska, and the E. (E.) knowltoni complex]: the anterad seta of each dorsal major set is lost, leaving only two per set (four total). In very darkly pigmented specimens [e.g., E. ( E.) knowltoni knowl¬ toni] this is evident, under high magnification on slide mounted material, as a light spot that is the remnant location of the lost anterad dorsal major. The dorsal minors are also reduced to two on each side, and the lateral-most has moved back into a straight line with the remaining dorsal majors. The second expression (Fig. IF) in E. (Essigella ) occurs as only six (total) dorsals [i.e., E. ( E .) californica, E. (. E .) hoerneri, E. (E.) pini ]. It is similar to that for the eight E. {Essigella) dorsals, except the dorsal minors are reduced to one on each side. Aberrations occur for 12 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) either of these expressions. In either the six or eight state, a suppressed dorsal major may reappear ahead of the line of setae in its normal anterad position. Also, in the eight setae state, one or more unsuppressed dorsal minors may occur, raising the setal count; these may involve a reversion to expression of the anterolat- erad position. The marginal setae are defined here as those setae laterad of the sagittally- oriented pair of muscle attachment points on the dorsum of each abdominal tergum (see Fig. 1; or Sorensen 1991: fig. 3). Like the dorsals, they are most easily seen on later nymphs. Among species, marginals may be expressed in a linear transformation, as states: four to six setae per side, three to five setae per side, or two setae per side. Their numbers are roughly correlated with the number of total dorsals. For example, E. (Archeoessigella ) species, which have the most total dorsals, have four to six marginals per side; in contrast, those E. (Essigella ) species with only six total dorsals have only two marginals per side. Marginal setal patterns show the greatest variation within species in more plesiomorphic states. The marginals of each side may occur in one or two groups, with the latter when their numbers are higher. Setae on abdominal tergum VIII apparently have the same approximate trans¬ formation as the dorsals on segments III-IV. They vary among species in a linear transformation, from: 10-16 setae in two rows; to 8-12 setae in one or two rows; to 6, or occasionally 8 (rarely to 10), setae in one or occasionally two rows. Dorsal Setae on the Metatibiae. —These setae often show considerable inter- and intraspecific variation in Essigella. In a principal component analysis of all Essigella taxa (Sorensen 1983), the length of dorsal setae on the metatibia, along with all other setae, show variation that is subordinate to general-size (component 1) and the number of abdominal setae (component 2). Tentatively, the dorsal setae of the metatibia are considered taxonomically useful only for apterous morphs. Their variation in Essigella contrasts with that of Eulachnus and other Lachninae, where their length appears to be more stable within species. This difference par¬ tially was responsible for the failure of previous attempts to circumscribe and key Essigella species (see Hottes 1957). Hille Ris Lambers (unpublished notes), criticizing Hottes (1957), suggested “constant characters” within Essigella included “the length of tibial setae, but not their being blunt”; he also stated “in the same species more or fewer of the tibial setae may be blunt which accounts for a rather large variation in setal length in some species.” I consider those statements to be erroneous. The retention of incrassate tips regardless of setal length, along with other characters, unifies the E. (E.) knowltoni group. Hottes (1957) erroneously defined his species on narrowly restricted setal length ranges; Hille Ris Lambers thought Hottes’ use of setal length confused blunt versus sharp tipped setae, which although Hille Ris Lambers correctly viewed as a continuum, he unfortunately disregarded as being of any taxonomic value in Essigella. I have studied variation of the tip structure among the dorsal metatibial setae for Essigella, including scanning electron microscopy work, and recognize several degrees of expression of bluntness among tips (un¬ published data); although of some taxonomic merit, this finer level delineation is not presented, because it cannot be used pragmatically to discriminate among most species. Three aspects of the dorsal setae on the metatibiae are recognized and cate- 1994 SORENSEN: A REVISION OF ESS1GELLA 13 gorized here: (a) length dimorphism within individuals and species, (b) variability in the range of setal length within species, and (c) variability in the condition of setal tips within species. Only the characteristic patterns of expression of these setae are considered valid synapomorphies within Essigella. Generally, these setae are long in other Lachninae, and their length is obviously homoplasious within the subfamily. Because of potential confusion concerning reference to particular setae, only those setae on the central one-third of the metatibia are treated here; reference is usually to only the longest of dorsal setae for that section (generally the dorsal metatibial setae are shorter and more incrassate proximally, and longer and sharper distally). The plesiomorphic state (state A) for dorsal setae of the metatibia within Es¬ sigella is short (ca. 0.3-0.7 x tibial diameter) with incrassate tips; as in Pseudes- sigella, E. (Archeoessigella ), and E. (E.) essigi. From this plesiomorphic “short, incrassate” condition, three independent transformation series are hypothesized within Essigella. The first involves elongation of setae to a continuous length range of ca. 0.3-2.0+ x tibial diameter, with the tips of shorter setae incrassate and longer setae sharp. This intermediate state (state B) represents the normal relationship for setal length and tip expression in aphids. Transformation contin¬ ues to an ultimate apomorphic state (state C) for this series, which shows an increase in the range of length variation to ca. 0.1-4.0+ x tibial diameter, and an increase in length variability within populations that I suspect is genetic and consider a weak synapomorphic character. This transformation is A —» B —» C. The second independent transformation involves a single step elongation of the setae (state D) to a continuous length range of ca. 0.3-2.0+ x tibial diameter, but without the development of sharp tips. These setae are always incrassate, regardless of length, and are considered a valid synapomorphy for the E. (E.) knowltoni group. This transformation is: A -* D. The third independent transformation involves setal elongation, to a dimorphic length range spanning ca. 0.3-4.0+ x tibial diameter (state E). This dimorphism can occur as a discrete length difference among different individuals of a popu¬ lation, or may be expressed on single aphids as an abrupt change in setal length on the central part of the metatibiae; it is restricted to E. (Lambersella), and appears to have ecological relevance between species of that subgenus in sympatry as a character displacement (unpublished data). Absence of the dimorphism in E. ( L .) hillerislambersi may be a secondary loss for this accommodation. This transformation is A —* E. Potentially confusing variation occurs in E. ( Essigella ) californica, where a minor increase in length of the dorsal setae is rarely evident on the distad one-third of the metatibiae; this is not considered homologous to the condition in E. {Lambersella). Ventral Abdominal Sclerites. —These sclerites represent attachment plates for muscles on the abdominal venter. Their size and shape may, therefore, have a functional relationship with the degree of sclerotization (not pigmentation) of the abdominal dorsum. Due to the diminished size and irregular variation of these ventral sclerites on more posterad abdominal segments, only segments III-IV are considered for taxonomic (identification) use. The sclerites are measured at their maximal length, along the anteroposterad axis, of the largest such sclerite occurring on abdominal segments III or IV. Unfortunately, the relative shapes and variance of these sclerites must be studied among species to be adequately comprehended. 14 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) As with the dorsal setae on the metatibiae, it is the qualitative pattern of expression of the ventral abdominal sclerites within populations, species, or spe¬ cies groups, that is considered taxonomically meaningful. In Pseudessigella (Sor¬ ensen 1991: figs. 2a-h), the ventral abdominal sclerites can be either uni-, bi-, or tripartite; if broken, the posterad section is relatively large and irregularly ovate, and the more anterad section(s) may be reduced and/or irregularly linear. In most species of less derived Essigella, these sclerites are subcircular to subquadrate [e.g., relative difference, Sorensen (1991: figs. 2e vs. 2f as bottom sclerite only of each, respectively)] (state A); this state occurs in E. (Archeoessigella), E. ( Lam- bersella ) and the more plesiomorphic species of E. {Essigella ). To achieve this shape during evolution, it is unclear if a linear sclerite simply shortens, or if it breaks into multiple subsclerites and loses the more anteriad of these. Alterna¬ tively, in E. (Lambersella) these sclerites can be relatively linear [e.g., Sorensen (1991: figs. 2c-d)], which matches their most linear unbroken development in Pseudessigella, or they can be nearly absent; these alternative expressions are treated as state A here also. Within E. {Essigella), several species show reduction of these sclerites, with expression varying from (at most) irregular, small quasi-stellate shapes through apparent absence (state B). This is considered homoplasious within the subgenus. Within the clade involving the E. {E.) knowltoni group, these sclerites ultimately become relatively enlarged and vary from subcircular or subquadrate to subellip¬ tical [e.g., latter, Sorensen (1991: fig. 2a)] (state D); an intermediate expression (state C) exists for E. (E.) alyeska, however, in which the sclerites vary from between states B and D within populations. The transformation is considered to be A —» B —> C —> D, but it could be independent among the latter three states. Body Widths.— Body shape differences occur within the genus and primarily involve relative width. Width characters are usually unreliable on most slide mounted material, due to compression artifacts. The standardized mounting tech¬ nique described earlier substantially reduces body distortion. Therefore, width measurements were used in analysis, but are avoided in keys and diagnoses, which must be applied to unstandardized slides. Head width is measured between the most laterad rims of the bases of the antennal sockets. This anterad measurement minimizes the effects of compression that are more likely to occur posteriorly. It also enhances recognition of com¬ pression, because the measurement line is adjacent to the clypeal region and the anterad outline of the frons; distortion of these regions is relatively noticeable when they are compressed. Relatively slight increases in width within species groups are homoplasious in this genus. Only the discrete and statistically significant broad body shown by the E. (E.) knowltoni group and E. (E.) alyeska is considered to be a valid synapo- morphy. Although Moran (1986) warns against using such ecologically influenced traits, it is interesting that E. (E.) hoerneri, a relatively broad Essigella whose width correlates with its pinyon pine host’s needle width, is correctly placed with E. (E.) californica, rather than the Series B E. {Essigella), in several discriminant function analyses (e.g., Sorensen 1992b: figs. 2a-b, 3) of the genus. This dem¬ onstrates the acceptable multivariate use of body width for classification (but not necessarily identification) within Essigella. Lengths and Shapes of Appendage Segments. — Determination of the polarities 1994 SORENSEN: A REVISION OF ESSIGELLA 15 for transformations of antennal segment lengths in Essigella is difficult. The re¬ duction from six to five antennal segments is a synapomorphy for Essigella and Pseudessigella, but is not unique in the Aphididae. In a principal component analysis among all Essigella taxa (Sorensen 1983), antennal segments III, IV and to a lesser extent V, load moderately on the second component. That vector orients largely to setal number on the abdominal dorsum, where polarity is clear. On the vector, however, the antennal segment lengths and abdominal setal numbers load in opposition; consequently, an increase in the relative length of antennal segments probably can be interpreted as apomorphic within the genus, as the abdominal setal number decline (Sorensen 1991, Sorensen & Foottit 1992). Allometric variation in the length of the metatibiae, which is associated with different morphs and stages in Essigella, is confusing. Hottes (1957) and Hille Ris Lambers (unpublished notes) regarded relative tibial lengths as constant among species in the genus. In aphids generally, there is a tendency for alates, because of their generally longer legs, to have relatively longer metatibiae in comparison to body length than do adult viviparous apterae. The converse is often true of later stadia nymphs of viviparous apterae, which generally have relatively shorter metatibiae than do their apterous adults. Variation along this morph factor is discordant in Essigella, however. In some E. (Archeoessigella) and E. ( Essigella ), aberrant adult viviparous apterae exist that retain the relative metatibial length characteristic of the juvenile stages of their species. Several of Hottes’ synonyms can be attributed to this aberrant variation among adult apterae [see the discussion of E. (E) californica\. Contrastingly in E. (Lambersella ), metatibial length is more stable within species; in that subgenus, however, allometric shifts along ontogenic factors, among species, sometimes differentiate the species in sympatry through character displacement. The comparative length of metatarsal segments varies in Essigella. In E. (Ar¬ cheoessigella), the metabasitarsus (first hind tarsus) is relatively short in regression compared to the metadistitarsus (second hind tarsus); a plesiomorphy reflected to a greater degree in Pseudessigella. Apomorphic elongation of the metabasitarsus occurs as a homoplasy in E. (Essigella) and E. (Lambersella), but reliable trans¬ formation of this homoplasious bivariate is difficult to ascertain; see Sorensen (1991) for a discussion of this trait. In lateral view, the shape of the profemur of more primitive Essigella resembles that of Pseudessigella. In the latter, the dorsoproximal base of the femur is strongly swollen and arched. A similar, but less pronounced, swelling is evident in species such as E. (A) kathleenae, and E. (E) pini, among others. In most Essigella, however, the femur usually assumes a more cylindrical shape with elongation in response to increases in body size in various lineages, no doubt as a allometric size transformation. This trait is not employed in identification or phylogenetic assessments, because its variation is inadequately known and is difficult to char¬ acterize; it cannot be measured satisfactorily on slides because it is usually oriented in the dorsoventral axis. Rostral Characters. — The rostrum of Essigella is retractile; consequently, rostral length is measured as the length of the stylets, which are fixed. Unfortunately, on slide mounted specimens the stylets can be withdrawn from the rostrum, and curved, making accurate measurement difficult. Stylets are measured from the sclerotic, basal apophyses in the clypeal region to their unbroken distal tips. 16 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Essigella (E.) hoerneri shows the greatest apomorphic increase in stylet length in response to the exceptional needle fascicle width of its piny on pine hosts. Mea¬ surement of the ultimate rostral segment includes the short, light-colored distal cap and the basal apophyses. Univariate or bivariate use of the ultimate rostral segment has not proven sufficiently reliable for consistent employment. Caudal Protuberance. — Although Hottes (1957) used the median protuberance on the cauda as a diagnostic character, I have avoided this due to its variation and the potential for orientation artifacts on slides where the protuberance is obscured when the cauda points up. The caudal projection shows undoubtedly homoplasious reduction trends, but its transformation and polarity are confusing among species groups. Essigella (E.) pini shows the greatest development of the caudal protuberance, with the protrusion sometimes quite strongly pointed; this undoubtedly accounts for the use of the character as an ultimate, but problematic, diagnostic for that species in Hottes’ (1957) key. Nymphs. — In later stadia nymphs of viviparous apterae (not prealatae), a pair of bilateral sclerotized plates occur that surround the muscle attachment plates on the mesonotum; these may be large or small, depending upon the species. When large (e.g., their diameter approximates the length of the eye), the invasive sclerotizations of these plates extend from the muscle attachment sites to engulf neighboring setal bases. The mesonotal sclerotization on nymphs can be extremely faint, especially in the E. (Archeoessigella ), where the enlarged plates can be difficult to see because of their light pigmentation. This sclerotization probably has a similar history to that of the abdominal dorsum of adult viviparous apterae. Presence of the developed plates may be a synapomorphy unique to Essigella among the Eulachnina. The enlargement of these plates is treated as a plesio- morphy within the genus, however, and secondary losses of the plates are con¬ sidered to be weak synapomorphies. In E. (L.)fusca, where the plates are normally enlarged, one late stadium nymph, within a large and otherwise normal sample, shows the loss of this invasive sclerotization. With this exception, the character appears quite stable among species; therefore, considering the loss state to be plesiomorphic on the basis of that single occurrence (i.e., a suppression failure) would require unacceptably strong homoplasy for the character. Nevertheless, the loss of these invasive sclerotizations, so that neighboring setal bases are free, appears to be a homoplasious apomorphy within E. (Essigella ) for E. ( E.) Cali¬ fornia, E. (E.) hoerneri, E. (E.) wilsoni and E. (E.) alyeska. Alatae. —Essigella alates are poorly known, and in several instances they are unknown. They appear to have few reliable diagnostic characters to identify them beyond species group. Characters often used within other genera, such as the number, shape or arrangement of secondary rhinaria on the antennae, usually show more intra- than interspecific variance in Essigella. Although Essigella alates normally have membranous abdominal terga, aberrants exist that show the normal sclerotic patterns of abdomens for their respective viviparous apterae or oviparae. Because knowledge of alate variation is poor, all statements concerning alate traits are tentative. Although venation is often of quite questionable taxonomic use in aphids (V. Eastop, D. Hille Ris Lambers, personal communications), it may be used in Essigella, with great caution. For instance, in Essigella the medius may have one (or rarely two) furcation(s), or may be single. The furcation may arise on the 1994 SORENSEN: A REVISION OF ESSIGELLA 17 proximal, central or distal one-third of the vein. A single medius is most probably apomorphic; if so, then the only synapomorphy appears to be for E. {E .) pini and E. {E .) essigi. In both species, however, the vein can vary, uncommonly, to having a furcation on its distal one-third. Moreover, the medius is also rarely expressed as a single vein in aberrant alates of the E. (E.) knowltoni group; thus, polarity remains questionable. Other variation of the medius involves the strength of expression of this entire vein system. Where the alates are known in E. (Lambersella), the medius is usually only faintly present along its entire length. This reduction is considered a weak synapomorphy. There are also differences involving the junctions of the anal and cubital veins with the radius. These can be expressed by the distance between the bases of the anal vein and the cubitus, along the radius, or by the truncated angle between them. The character is variable and tentatively considered unreliable as a diagnostic. An aberration displayed by several Essigella species is a darkened band that posteriorly parallels the radius. The epicranial suture may be of use taxonomically. The suture appears to be most prominent and stable in E. (E.) californica and E. (E.) hoerneri, but varies in presence and expression among other species. Oviparae. — Although the oviparae of all Essigella species are not known, among those that are, three conditions exist for the sclerotization/fusion of the abdominal dorsum: (state A) terga II-VII are fused, but I and VIII are free; (state B) abdominal terga II-VI are fused, but I, VII and VIII are free; and (state C) all abdominal terga are free with independently banded sclerotizations. It is unclear whether state A or B is the most plesiomorphic because both occur in E. ( Archeoessigella ); the transformation could be either A-^B^CorA^B— * C. State A occurs in E. ( Archeoessigella) kathleenae, in the E. ( Essigella) knowltoni species-group, and usually in E. {E.) pini. State B occurs in E. {Archeoessigella ) kirki, in E. {Lambersella), and rarely in E. (E.) pini. Therefore, the character is necessarily homoplasious, with state A either as a plesiomorphy, which requires a reversal in E. {Essigella), or as an apomorphy, which requires it to be gained independently in E. {Archeoessigella) and E. {Essigella). The complete loss of tergal fusion in state C, for the E. {E.) calif ornica group, is considered a weak synapomorphy. Uncommon oviparae of species with the banded abdominal terga of state C can show near fusion of terga II-V or II-VI. This suggests an apparent plesiomorphic aberration approaching state B; if so, it appears C is apomorphic to B. No re¬ versions from B or C to A are apparent; however, because E. {E.) pini usually shows fusion of abdominal terga II-VII, but with VI rarely free, this may be evidence for plesiomorphy in state B? Key to the Eulachnina Genera la. Antennae of adult virginoparous apterae 6-segmented. . Eulachnus del Guercio lb. Antennae of adult virginoparous apterae 5-segmented. 2 2a. (lb) Tarsal claws with single, simple tips. Adult apterae with tergum of abdominal segments II-VII membranous. . Pseudessigella Hille Ris Lambers 2b. Tarsal claws incised, with double tips. Adult apterae with tergum of 18 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) abdominal segments II-VII fused, very lightly to heavily sclerotized. . Essigella del Guercio Essigella Del Guercio, 1909 Essigella Del Guercio, 1909, Riv. patol. Veg., Padov, n.s. 3: 329. Lachnus Burmeister, 1835 (in part), Hardbuch der Entomologie, Berlin, 2: 91 (genus attributed to Illiger); Essig, 1909, Pomona J. Entomol., 1: 1-4. Type Species.— Lachnus californicus Essig, 1909, Pomona J. Entomol., 1: 1-4; by monotypy. Viviparous Apterae. — Body elongate, linear to linear-ovate. Antennae 5-segmented; processus ter- minalis short; accessory rhinaria on terminal antennal segment proximad, not directly against primary rhinarium. Head wider than long, fused with pronotum, or nearly so. Eyes without distinct triom- matidia. Rostrum retractile; last rostral segment short, blunt, tip nonfunctionally articulated (if at all), accessory setae absent. Meso- and metanota fused dorsally. Abdominal dorsum lightly to heavily sclerotic; tergum I usually free; terga II-VII fused; tergum VIII free, represented by single, entire sclerotized field not apparently formed from fused lateral sclerites associated with setal bases; pig¬ mentation variable; dorsal setae on segments III—IV in 1 or 2 often irregular rows. Siphunculi rep¬ resented as rimmed pores to short truncated cones, without setae, incorporated into dorsal sclerotic field of abdomen. Cauda rounded, frequently with short, rounded to pointed, median protuberance. Profemora cylindrical-tapering, to dorsoproximad base slightly swollen. Tarsal claws incised, bifid; dorsal tip blunt, ventral tip sometimes slightly projecting, blunt to sharper. Other Morphs.— Known fundatrices and males lacking siphunculi. Known oviparae and males apterous. Known alatae with radial sector short, straight; forewing medius distinct to apparently absent, single or with 1 furcation. Diagnosis.— See the key to the Eulachnina genera and apomorphies section below. Taxonomic Placement. —Essigella, along with Pseudessigella Hille Ris Lam- bers, 1966, and Eulachnus Del Guercio, 1909, comprise the subtribe Eulachnina (Sorensen 1990) of the tribe Cinarini, subfamily Lachninae; the subtribe is con¬ sidered highly derived within the subfamily. The immediate sister-group of Es¬ sigella is Schizolachnus\ which Lampel & Burgener (1987) suggest placing, along with the three eulachnine genera, in a single tribe, the Schizolachnini. Sorensen (1990), however, places Schizolachnus in a separate subtribe, the Schizolachnina. For commentary on the taxonomic relationships of related genera, and previous tribal/subtribal assignments, see Sorensen (1990). Distribution.—Essigella is the only native Nearctic representative within the subtribe Eulachnina, although it has recently been introduced into Europe, in France (Turpeau & Remaudiere 1990) and Spain (Seco Fernandez & Mier Durante 1992). The other eulachnines, Pseudessigella and Eulachnus, are native to the Palaearctic; the former is known only from a single collection in the Himalayas of Pakistan (Sorensen 1991), but the latter has been introduced to the Nearctic where it occurs largely on cultivated Old World pines. Apomorphies. — The three Eulachnina genera share these synapomorphies: body form linear; triommatidia of compound eye undifferentiated; ultimate rostral segment short and blunt, tip nonfunctionally articulated (if at all); accessory setae on ultimate rostral segment absent; setae on siphunculi absent; primary rhinarium without chitinous ring border; and accessory rhinaria on terminal antennal seg- 1994 SORENSEN: A REVISION OF ESSIGELLA 19 ment proximad and not directly against primary rhinarium. Essigella and Pseu- dessigella share these synapomorphies: reduction from six to five antennal seg¬ ments; and abdominal tergum VIII represented by a single, entire sclerotized field that is not apparently formed from fused lateral sclerites that are associated with the dorsal setae bases. Essigella is the most derived genus of the subtribe (Sorensen 1990) and has these autapomorphies: tarsal claws bifid; entire dorsum sclerotized; head and pronotum fused; meso- and metanotum fused dorsally; abdominal tergum I usu¬ ally free (but at least partially fused [laterally] to metanotum as an autapomorphy in one species); abdominal terga II-VII fused. Another tentative autapomorphy for Essigella is a complete loss of the siphunculi in both the fundatrix and male, where these morphs are known; they are yet unknown for Pseudessigella and, therefore, the trait could be synapomorphic at that level. All Essigella have def¬ initely incised tarsal claws, with the resultant presence of an endodontal lobe, despite Hottes’ (1957) comments to the contrary; Hille Ris Lambers (personal communication, unpublished data) correctly interpreted Hottes’ (1957: 108, key couplet la) statement of “Tarsal claws with ends not distinctly bifurcated” as erroneous, and due to over-processing in caustics during slide preparation. Subgenera. — Three Essigella subgenera are recognized and described here; see the phylogenetics section for commentary. Their compelling separation requires discriminant function analysis of morphometric traits because considerable over¬ lap in univariate traits exists among Essigella species (Sorensen 1983). Demar¬ cation of these subgenera was made from an evolutionary perspective (see Sor¬ ensen 1992b) that includes Pseudessigella as an anagenic distance reference. Because many other aphid subgenera can be distinguished by single characters, their dif¬ ferentiation may be under the control of a single, or fairly limited number of, genes. As a result, such univariately defined subgenera probably display less genetic divergence than do the Essigella subgenera, among which realignments have occurred for large suites of genes (Falconer 1981, Sorensen 1991, Sorensen & Foottit 1992) that are responsible for their multivariate divergence. See Sorensen (1991) for a discussion of the multivariate evolution of the shape component among traits between Pseudessigella and Essigella, and among some Essigella groupings. The phylogeny for the genus indicates Archeoessigella, the least derived sub¬ genus, is separated (as Fig. 13: node 1) from Pseudessigella by 18.9 o units (see phylogenetics section). Lambersella is separated from Archeoessigella by 2.08 er units (as Fig. 13: intemode 2-3). Essigella ( Essigella ), the most derived subgenus, is separated from Lambersella by 4.22 a units (as Fig. 13: internode 3-7). All three of these anagenic distances are significant, at a = 0.05, as evolutionary gaps between genus-[or subgenus]-level species assemblages (see phylogenetics section). Also, all three subgenera are phylogenetically convex (sensu Duncan 1980, Es- tabrook 1986), with Lambersella and E. {Essigella) as monophyletic groups. Although the Essigella subgenera are quite valid biologically and evolutionarily, and their status as subgenera gives them the same nomenclatural rights as genera, they are described here with relevance only for intrageneric hierarchy. Because the anagenic distances among the Essigella subgenera are considerably shorter than among the genera in the Eulachnina, I recommend that these subgenera never be elevated to the status of full genera through taxonomic inflation. 20 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Etymology. — The genus was named by Del Guercio (1909) after E. O. Essig, who collected and described its first species as Lachnus californicus Essig. Material Examined.— All taxa proposed here, plus all Eulachnina and Schizolachnina taxa listed in Sorensen (1990: in Phylogenetic Construction sections Ingroup Material Examined and Outgroup Selection). Key to the Subgenera of Essigella This key is intended only for properly cleared, slide-mounted virginoparous apterae, and is meant for populations and species, rather than individuals. Key intraspecific samples with several (preferably n = 10+) individuals to account for variance. Individuals should be keyed using the key to Essigella species. la. Abdominal terga III-IV with dorsal setae in a single [or at most a very slightly staggered] row; lateral-most seta normally absent. Populational mean for number of dorsal setae on abdominal terga III-IV normally 6 or 8, mean number on abdominal tergum VIII normally 6, sometimes 8, never 9 or more [if mean on terga III-IV is 8-10 and the mean for tergum VIII is 8, then : (a) developed pigmentation suite for tibiae described in couplet 2a (below) never occurs in any population; and (b) populational mean for the ratio of length of the metadistitarsus to metabasitarsus is 1.73:1 or less; and either (cj body relatively broad with at least some populations with specimens whose longest dorsal setae on the central one-third of the metatibiae exceed 1.5 x metatibial diameter and remain incrassate regardless of length; or (c 2 ) metanotum and abdominal tergum I fused at least laterally; or (c 3 ) mesonotum of later stadia nymphs of apterae with area immediately surrounding mus¬ cle attachment sites membranous and bases of neighboring setae not on a sclerotized plate contiguous with the muscle attachment sites; or (c 4 ) mean number of marginal setae per side on each of abdominal terga III-IV is 2; or (c 5 ) primary rhinarium on terminal antennal seg¬ ment exceptionally distad with distance from tip of processus terminalis to distal face of rhinarial rim less than 0.5 x diameter of rhinarium, and distal face of rhinarial rim usually perpendicular to longitudinal axis of antennal segment, and rhinarial membrane usually conspicu¬ ously protuberant]. Essigella (Essigella ) del Guercio lb. Abdominal terga III-IV with dorsal setae in a double [or strongly stag¬ gered] row; lateral-most seta normally present. Populational means for number of dorsal setae on abdominal terga III-IV and tergum VIII normally at least 8 or more, never 6 [if means on terga III-IV and tergum VIII are 8-10, then either, (a) any developed pigmentation suite described in couplet 2a (below) may or may not occur in any population; or (b) populational mean for the ratio of length of the metadistitarsus to metabasitarsus is 1.70:1 or greater; but (c) none of conditions Cj-c 5 in couplet la ever exist]. 2 2a. (2a) Populational mean for the ratio of length of the metadistitarsus to metabasitarsus is 1.69:1 or less, but usually under 1.65:1. Intraspecific pigmentation suites ranging from pale (unpigmented) to heavily pig¬ mented, often within populations, but if developed (even subtly) then 1994 SORENSEN: A REVISION OF ESSIGELLA 21 (a) pro-, meso- and metatibiae, respectively, pigmented relatively heavi¬ ly, lightly and heavily [in a dark-light-dark pattern], or (b) body dorsum with darkened pigmentation but with lightened longitudinal stripe in dorsomedial region of thorax and abdomen, or (c) thoracic and ab¬ dominal terga mottled with dorsal setal bases pigmented. In any pop¬ ulation, longest dorsal seta on central one-third of metatibia varying from incrassate and short to long and either sharp or blunt tipped, but if longer than 1.0 x metatibial diameter then they are not incrassate. Populational means for number of dorsal setae: (a) on each of abdom¬ inal terga III-IV usually 10 or less [occasionaly 11], but if mean more than 10 then at least some individuals with 9 or less; and (b) on ab¬ dominal tergum VIII usually 9 or less [occasionally 10], but if mean more than 9 then at least some individuals with 9 or less. . Essigella (Lambersella ) NEW SUBGENUS 2b. Populational mean for the ratio of length of the metadistitarsus to metaba- sitarsus is 1.70:1 or greater, but usually over 1.75:1. Intraspecific pig¬ mentation suites ranging from pale (unpigmented) to very subtly pig¬ mented, but when pigmentation is subtly developed it is generally even and never as in couplet 2a. In any population, longest dorsal seta on central one-third of metatibia always incrassate and less than 1.0 x metatibial diameter. Populational means for number of dorsal setae: (a) on each of abdominal terga III-IV usually 11 or more, but if mean less than 11 then at least some individuals with 13 or more; and (b) on abdominal tergum VIII usually 10 or more, but if mean less than 10 then at least some individuals with 12 or more. . Essigella (Archeoessigella) NEW SUBGENUS Essigella (. Archeoessigella ), NEW SUBGENUS “Essigella (Archoessigella )” Sorensen, 1983: 58 (unpublished manuscript name, note different spelling) Ph.D. Thesis, University of California at Berkeley, Berke¬ ley, California. 605 p. Type Species.—Essigella kathleenae Sorensen, 1988. Viviparous Apterae. —Morphology: Body slender. Meso- and metanota fused dorsally; abdominal tergum I free. Dorsal setae (majors and minors) between muscle attachment plates on abdominal segments III-IV in 2 often irregular rows (see Fig. 1B); lateral-most minor dorsal setae on each side anterad (rarely not) of its immediately mesad neighbor. Abdominal terga III-IV each with 10-16 dorsal (major + minor) and 4-6 (per side) marginal setae; tergum VIII with 10-14, rarely 7-9, setae. Longest dorsal seta on central one-third of metatibiae less than tibial diameter, tips incrassate; these setae of nearly equal length along metatibiae, not dimorphic. Ventral abdominal sclerites on segments III-IV large, subquadrate or subcircular, not rudimentary. Species means for length ratio of metadisti¬ tarsus to metabasitarsus varying between 1.81:1 to 2.05:1. Pigmentation: Body dorsum unicolorously pale; bases of dorsal setae of abdomen concolorous with surrounding terga. All tibiae equally pigmented, usually pale to rarely subtly dusky. Diagnosis.— See the key to the subgenera of Essigella. Discussion.— This plesiomorphic subgenus is paraphyletic, but convex (sensu Duncan 1980, Estabrook 1986); no qualitative synapomorphies exist that uniquely define the group. It characteristically has a high ratio for the metabasitarsus: 22 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70 ( 1 ) metadistitarsus length, relatively many dorsal and marginal setae on abdominal terga III-IV and VIII, metatibial dorsal setae that are short and incrassate, no developed pigmentation suites, and species that are functionally monophagous and restricted to pine species in Pinus (Strobus ), section Strobus, subsection Strobi. Except for some minor differences in placements of certain dorsal setae on the abdominal terga III-IV, intriguingly, all these characteristics are shared by Pseu- dessigella to a great degree. Archeoessigella was named because it differs signifi¬ cantly from Lambersella in several respects, and the two each have distinctly different host associations. Similarities between the Archeoessigella species are relative plesiomorphies. The phylogenetic tree (Fig. 13), based on all 15 available dimensions of discrim¬ inant space, shows E. {A.) kirki to branch from node 1 (distance = 0) as the most primitive Essigella. However, when the multivariate shape-component differences for traits between Essigella and Pseudessigella were analyzed on the more dom¬ inant shape vectors, Sorensen (1991) found E. {A.) kathleenae to be generally more similar to Pseudessigella than to the remaining more derived Essigella, and he found E. (A.) kirki to be intermediate between those groups; he noted each Archeoessigella species was less similar to one another than either was to Pseu¬ dessigella or to the more derived Essigella. The closer proximity between Pseu¬ dessigella and E. (A) kathleenae is also reflected on the second-most dominant minimum selective mortality vector (Fig. 14: DF2). A single, conventional, qual¬ itative trait, the fusion of the abdominal dorsum in oviparae, sheds only vague light on the problem because its transformation and polarity are uncertain [see oviparae under the character discussion section]. Coded References to this Taxon. — Sorensen (1983) referred to this taxon under the manuscript name “ Essigella (Archoessigella ).” Sorensen (1987a) referred to the assemblage that comprise this taxon as group “I” or, with reference to its subcomponents, as “J-K”; in Sorensen (1992b), the latter refers to it. Etymology. — “Archeo-” (Greek) = ancient; the name reflects the old and primitive status of the subgenus; coincidentally, the compounded name includes “-eoessig-” for E. O. Essig. Material Examined.—Essigella (A.) kathleenae, E. (A.) kirki. Essigella {Archeoessigella) kirki Sorensen, 1988 Essigella kirki Sorensen, 1988: 121, Pan-Pacif. Entomol., 64: 121-124. Essigella “hottesi ” Sorensen, 1983: 60 (unpublished manuscript name) Ph.D. Thesis, University of California at Berkeley, Berkeley, California. 605 p. Type Series. — Holotype. vivip. apt.; on slide with 3 paratype vivip. apt., ho- lotype at lower left (8 o’clock position); data: NEW MEXICO. SANTA FE Co.: ca. 30kmNEofSantaFe, hwy475, 3100 m, 10 Aug 1978, J. T. Sorensen (78H55), Pinus flexilis James. Holotype retained in Sorensen collection, eventually to be deposited in The Natural History Museum, London. Paratypes (all same data as holotype): 19 vivip. apt. on 5 slides including holotype slide. Paratype slides deposited: 1 slide in NMNH, Washington, D.C.; 1 slide in CNC, Ottawa, Ontario; 2 slides in Sorensen collection. 1994 SORENSEN: A REVISION OF ESSIGELLA 23 Viviparous Apterae.—Morphology: Body length: 1.73-2.13 (1.92 ± 0.13) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously protuberant. Length of antennal segment V: 95-133 (117 ± 10) p, processus terminalis: 28-45 (37 ± 5) p; IV: 70-91 (82 ± 7) M ; III: 141-188 (157 ± 15) M ; H: 63-73 (68 ± 3) p. Length of longest setae on frons: 10-43 (28 ± 9) p, tips incrassate. Head width: 245-316 (285 ± 19) p. Length of stylets: 530-694 (608 ± 55) p\ ultimate rostral segment: 68-83 (76 ± 5) p, rostral tip reaching abdominal terga I or II in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 367-439 (399 ± 24) p. THORAX: Meso + metanota fused, total length: 296-388 (347 ± 28) p. ABDOMEN: Tergum I free, length: 112-163 (138 ± 15) p\ terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 45-55 (50 ± 4) p; siphunculi nearly flush to truncated conical, protruding to 1.0 x maximal distal width. Ventral abdominal sclerites on segments III-IV subquadrate, subcircular to subelliptical; length: 50-68 (59 ± 6) p, 1.2-2.Ox diameter of metatibiae. Dorsal (major + minor) setae (see Fig. IB) on abdominal terga III-IV: 10-14 (11 ± 1), tips sharp, in 2 irregular rows, lateral-most minor dorsal seta usually in anterad row; marginal setae 4-6 per segment each side. Setae on abdominal tergum VIII: 10-14 (11 ± 1), length: 5-43 (23 ± 11) p, tips incrassate to rarely sharp, in 2 irregular rows. Cauda rounded; caudal protuberance moderately developed to frequently nearly absent; length of longest caudal setae: 70-103 (86 ± 10) p, tips sharp. LEGS: Length of metafemora: 500-663 (578 ± 53) p ; metatibiae: 622-900 (755 ± 70) p; longest dorsal setae on central one-third of metatibiae: 20-30 (24 ± 3) p, 0.1-0.6 x diameter of metatibiae, tips incrassate, approx¬ imately equal or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae: 13-28 (23 ± 4) p, tips sharp. Length of metabasitarsus: 93-118 (104 ± 7) p\ meta- distitarsus: 165-213 (188 ± 13) p. Ratio of metadistitarsus to metabasitarsus averaging 1.81:1, usually less than 1.9:1, rarely reaching 2.0:1 or slightly more. Pigmentation: Color in life: Gray-green, occa¬ sionally pale yellow throughout. Slide-mounted specimens: Background of body dorsum pale (to 10 percent pigment density), unicolorous. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga. Thoracic muscle attachment plates pale, in¬ conspicuous to conspicuous. Dorsal muscle attachment plates of abdomen conspicuous, pale, infre¬ quently dusky. Spiracular plates and ventral abdominal sclerites usually light brown, slightly darker than background of abdominal terga, to pale. Siphunculi concolorous with surrounding terga. Cauda, anal and subgenital plates concolorous with abdominal terga. Antennal segments V and IV slightly to moderately dusky over entire segment, to moderately brown distally; III pale; II and I concolorous with frons. Pro-, meso- and metatibiae usually pale, concolorous, equivalent to body dorsum; fre¬ quently tibiae subtly dusky at distal tip, rarely entire tibiae moderately dusky, slightly darker than body dorsum. Distitarsi usually subtly dusky distally to moderate brown, varying with antennae, infrequently entirely dusky with tibiae. Ultimate Stadium Nymphs of Viviparous Apterae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum with 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; plates usually vague, faintly pigmented, diameter approximately equal¬ ing eye length. Oviparae. -Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae ex¬ cept abdominal terga II-VI fused, lightly to moderately sclerotic, including pleural areas, but VII and VII free; dorsal demarcations of anterad terga not evident; siphunculi usually incorporated into sclerotic dorsum, to free; dorsal abdominal muscle attachment plates pale, unicolorous, except those between terga VI-VII darker. Pseudorhinaria on metatibiae irregular, difficult to distinguish, 7-11. Viviparous Alatae, Males, Fundatrices.— Unknown. Diagnosis. —Essigella (A.) kirki can easily be confused with other pale individ¬ uals of Essigella. It can be separated from all Essigella, except E. (A.) kathleenae, E. (L.) eastopi, E. ( L .) fusca, E. (L.) hillerislambersi, and odd specimens of E. (E.) wilsoni and E. (E.) knowltoni braggi, by having 10 or more dorsal (major + minor) setae on abdominal terga III-IV, in two rows, with the lateral-most minor dorsal seta in the anterad row (e.g., Fig. IB). Essigella (A) kirki lacks the very 24 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Figure 2. Distribution of E. (A.) kirki [dots (JTS samples)], superimposed over the ranges of its hosts, Pinus flexilis [lighter shading] and Pinus strobiformis [darker shading (AZ, NM and south)]. elongate metadistitarsus of E. (A.) kathleenae, having a metadistitarsus to meta- basitarsus ratio of usually less than 1.9:1, but rarely to 2.0:1 [mean: 1.8:1 for E. (A) kirki, 2.05:1 for E. (A.) kathleenae ]. It can be separated from other pale Essigella, however, by having this ratio at over 1.7:1. Essigella (A.) kirki lacks the protuberant, exceptionally distad primary rhinarium of E. {E.) wilsoni. It can be further distinguished from pale E. ( L .) fusca and E. (L.) hillerislambersi, and some pale E. ( L .) eastopi and E. ( E .) knowltoni braggi by having the longest dorsal setae on the central part of the mesotibia less than 0.7 x tibial diameter. All observed E. (E.) knowltoni braggi with 10 or more dorsal (major + minor) setae on abdominal terga III-IV differ from E. {A.) kirki by having the longest dorsal metatibial setae in excess of 1.0 x tibial diameter; however, rare, confusing E. (E.) knowltoni braggi are anticipated, and these could be separated by their broad 1994 SORENSEN: A REVISION OF ESSIGELLA 25 head on noncompressed slides, and by usually longer setae on the frons [see descriptions and E. ( E .) knowltoni diagnosis]. Range. —Rocky Mountains, Montana to Arizona and New Mexico; southern Sierra Nevada (east slope) and White Mountains of California; presumably into Mexico and Canada with its hosts (Fig. 2). Hosts. —Pinus flexilis James and P. strobiformis Engelmann; the latter was pre¬ viously considered to be a variety [as P. flexilis var. reflexa Engelmann] of the former. These pines split the higher elevation niche in the Rocky Mountains, with P. flexilis in the north, P. strobiformis in the south, and some intergradation at their contact in northern New Mexico (Critchfield & Little 1966). The only other Essigella species on these pines is E. (E.) californica, which has secondarily in¬ vaded the niche, opportunistically, and is much less common in it than E. (A.) kirki. It is possible that E. (A.) kirki also occurs on P. ayacahuite Ehrenberg in central Mexico and south, because that pine was formerly considered a variety of P. strobiformis (as P. ayacahuite var. brachyptera Shaw); P. ayacahuite apparently continues the P. flexilis to P. strobiformis morphological and geographic cline (Critchfield & Little 1966), and although discontinuous with the latter, a single isolated stand in western Jalisco, Mexico (Critchfield & Little 1966: map 9) is morphologically intermediate with P. strobiformis (Martinez 1948). Discussion. —Because of previous misinterpretation of meso- and metanotal fusion in Essigella (see character discussion section), the description given here for this species is more accurate than that in Sorensen (1988). Essigella (A.) kirki is a common species that is relatively homogeneous, mor¬ phologically, and always pale, unlike several other Essigella species that can grade from pale to fully pigmented; in these respects it resembles E. {A.) kathleenae. Sorensen (1983) determined that it differs from the latter in bivariate plots of head width, between the lateral rims of the antennal sockets, versus body length, and of metadistitarsus versus metabasitarsus lengths; it also separates under prin¬ cipal component and discriminant function analyses (Sorensen 1983). Coded References to This Taxon.—Essigella (A.) kirki has been referred to previously by: the coding “Sp. K” (Sorensen 1983, 1987a, 1992b) and “HOTT” (Sorensen 1983), and by the manuscript name E. “hottesi ” in Sorensen (1983). Etymology and Common Name.— The species was named for my son, Kirk Hale Sorensen. Common name: Kirk’s limber pine needle aphid. Material Examined.— ARIZONA. APACHE Co.: Lake Harney Rd (hwy 473), nr McNary, 2440 m, 11 Sep 1978, JTS 78114, P. strobiformis, (apt.). COCHISE Co.: nr Rustler Park, Chiricahua Mts, 2500 m, 16 Sep 1978, JTS 78150, P. strobiformis, (apt.). CALIFORNIA. INYO Co.: Lake Sabrina, nr Bishop, 2750 m, 1 Aug 1977, JTS 77H2, P. flexilis, (apt.); Onion Valley Cmpgd, 24 km W of Independence, 2770 m, 4 Aug 1978, JTS 78H13, P. flexilis, (apt.). COLORADO. SAN JUAN Co.: 20 km N of Purgatory, 3020 m, 8 Aug 1978, JTS 78H47, P. flexilis, (apt.). MONTANA. CARBON Co.: Red Lodge, 1770 m, 20 Aug 1978, JTS 78H115, P. flexilis, (apt.). NEVADA. WHITE PINE Co.: Wheeler Peak, 3140 m, 26 Aug 1978, JTS 78H147, P. flexilis, (apt., ovip.). NEW MEXICO. OTERO Co.: 3 km W of Cloudcroft on hwy 82, 2560 m, 13 Sep 1978, JTS 78122, P. strobiformis, (apt., ovip.). SANTA FE Co.: (type series) 30 km NE of Santa Fe on hwy 475, 3100 m, 10 Aug 1978, JTS 78H55, P. flexilis, (apt.). SIERRA Co.: Emory Pass on hwy 90, W of Kingston, 2470 m, 14 Sep 1978, JTS 78134, P. strobiformis, (apt.). UTAH. DUCHESNE Co.: 19 km NE of Castle Gate on hwy 33, 2770 m, 25 Aug 1978, JTS 78H144, P. flexilis, (apt.). WYOMING. ALBANY Co.: 5 km SW of Woods Landing on hwy 230, 2560 m, 15 Aug 1978, JTS 78H92, P. flexilis, (apt.). 26 THE PAN-PACIFIC ENTOMOLOGIST Yol. 70(1) Essigella ( Archeoessigella ) kathleenae Sorensen, 1988 Essigella kathleenae Sorensen, 1988: 115, Pan-Pacif. Entomol., 64: 115-118. Essigella “ kathleeni ” Sorensen, 1988: 124 (lapsus), Pan-Pacif. Entomol., 64: 124. Essigella “ kathleenae ” Sorensen, 1983: 67 (unpublished manuscript name) Ph.D. Thesis, University of California at Berkeley, Berkeley, California. 605 p. Type Series. — Holotype, vivip. apt.; on slide with 3 paratype vivip. apt., ho- lotype at upper left (11 o’clock position); data: CALIFORNIA. SAN BERNAR¬ DINO Co.: 3 km S of jet hwy 38 & Jenks Lake Rd, San Bernardino Mts, 2200 m, 16 Sep 1977, J. T. Sorensen (77138), Pinus lambertiana Douglass. Holotype retained in Sorensen collection, eventually to be deposited in The Natural History Museum, London. Paratypes (all same data as holotype): 30 vivip. apt. on 7 slides including holotype slide. Paratype slides deposited: 1 slide in NMNH, Washing¬ ton, D.C.; 1 slide in CNC, Ottawa, Ontario; 8 slides in Sorensen collection. Viviparous Apterae.—Morphology: Body length: 1.35-2.01 (1.67 ± 0.18) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously protuberant. Length of antennal segment V: 85-113 (102 ± 7) p, processus terminalis: 28-43 (40 ± 4) M ; IV: 60-90 (75 ± 9) p\ III: 98-135 (118 + 11) m; H: 55-68 (62 ± 4) p. Length of longest setae on frons: 8-25 (17 ± 6) p, tips incrassate. Head width: 215-258 (242 ± 11) p. Length of stylets: 428- 653 (581 ± 64) p\ ultimate rostral segment: 55-78 (66 ± 5) p, rostral tip reaching metanotum to abdominal terga III in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 286-377 (334 ± 31) p. THORAX: Meso + metanota fused, total length: 214-306 (280 ±31) p. ABDOMEN: Tergum I free, length: 93-133 (119 ± 12) p; terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 23-38 (32 ± 4) p\ siphunculi flush to truncated conical, protrusion to 0.5 x maximum distal width. Ventral abdominal sclerites on segments III-IV subcircular, subquad¬ rate to subelliptical; length: 36-60 (48 ± 8) p, 1.3-2. lx diameter of metatibiae. Dorsal (major + minor) setae (see Fig. IB) on abdominal terga III-IV: 11-14 (12 ± 1), tips sharp, in 2 irregular rows; marginal setae 4-5 per segment each side. Setae on abdominal tergum VIII: 7-13 (10 ± 2), length: 5-40 (14 ± 10) p, tips incrassate to sharp, in 2 irregular rows. Cauda rounded; caudal protuberance moderately developed, to infrequently nearly absent; length of longest caudal setae: 40-93 (61 ± 16) p, tips sharp. LEGS: Length of metafemora: 316-541 (448 ± 67) p; metatibiae: 428-704 (569 ± 77) p\ longest dorsal setae on central one-third of metatibiae: 5-23 (13 ± 6) p, 0.1-0.8 x diameter of metatibiae, tips incrassate; approximately equal or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae: 10-25(19 ± 5)^, tips sharp. Length of metabasitarsus: 60-95 (79 ± 10) p\ metadistitarsus: 135-180 (162 ± 12) p. Ratio of metadistitarsus to metabasitarsus averaging 2.05:1, greater than 1.9:1, and usually greater than 2.0:1. Pigmentation: Color in life: Pale yellow throughout. Slide-mounted specimens: Background of body dorsum pale (usually to 10, some¬ times to 30, percent pigment density), unicolorous. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen, pale, inconspicuous. Spiracular plates and ventral abdominal sclerites pale. Siphunculi concolorous with surrounding terga. Cauda, anal and subgenital plates pale, concolorous with abdominal terga, to slightly darker. Antennal segments V and IV pale, only very subtly darker than body dorsum; III very pale to distal one-third pale as V and IV; II concolorous with proximal III; I concolorous with frons. Pro-, meso- and metatibiae usually pale, concolorous with body dorsum, to very subtly darker. Distitarsi entirely pale to subtly dusky on distal one-third. Ultimate Stadium Nymphs of Viviparous Apterae. — Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum with 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; plates usually vague, faintly pigmented, diameter approximately equal¬ ing eye length. 1994 SORENSEN: A REVISION OF ESSIGELLA 27 Oviparae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae, ab¬ dominal terga II-VII fused, lightly to moderately sclerotic, including pleural areas, tergum VIII free; dorsal demarcations of anterad terga not evident; siphunculi incorporated into sclerotic dorsum; dorsal abdominal muscle attachment plates pale, unicolorous. Pseudorhinaria on metatibiae irregular, difficult to distinguish, 5-9. Viviparous Alatae, Males, Fundatrices.— Unknown. Diagnosis.—Essigella (A.) kathleenae is consistently pale, and usually can be identified by the unique, exceptionally long metadistitarsus and short metabasi- tarsus. The length ratio of the metadistitarsus to metabasitarsus usually exceeds 2.0:1, and only rarely approaches 1.9:1, the upper value for all other Essigella, except occasional E. (A.) kirki. Range. —California and southwestern Oregon (Fig. 3). Hosts. —Pinus lambertiana Douglass; questionable single occurrences on P. jef- freyi Greville & Balfour, P. sabiniana Douglass and P. monticola Douglass. A single specimen attributed to P. jejfreyi (77166) is probably a beating tray contam¬ ination from a preceding collection (77164) from P. lambertiana, which occurred at dusk. A single specimen from P. sabiniana (77G17) is probably also similarly accidental, following a preceding collection on P. lambertiana (77G16). My col¬ lection from P. monticola (78G7) is a questionable host determination; that col¬ lection is from an isolated, low elevation stand of pines that W. B. Critchfield (personal communication) believes to be P. monticola, but that I believe is possibly P. lambertiana on the basis of its ecological, geographic and elevational circum¬ stances [P. monticola replaces P. lambertiana at higher elevations in the Sierra Nevada, and the P. monticola niche is opportunistically occupied by E. {E.) californica .] Discussion.— Because of previous misinterpretation of meso- and metanotal fusion in Essigella (see character discussion section), the description given here for this species is more accurate than that in Sorensen (1988). Essigella (A.) kathleenae is a common, morphologically homogeneous species. Its elongate metadistitarsus and very shortened metabasitarsus represent a ple- siomorphy within Essigella ; this is shared with Pseudessigella, which has a much higher tarsal ratio and differing metatarsal regression. Essigella (A.) kirki nearly shares the same metatarsal regression with E. (A.) kathleenae, but is displaced along the regression by its slightly longer metabasitarsus. Essigella {A.) kathleenae may have no conventional apomorphies beyond those defining the genus; the confusing polarity for the fused abdominal terga of oviparae, which differs between E. (A) kathleenae and E. {A.) kirki, is discussed in the character discussion section. Coded References to This Taxon.—Essigella (A.) kathleenae has been referred to previously by: the coding “Sp. J” (Sorensen 1983, 1987a, 1992b) and “KATH” (Sorensen 1983), and by the manuscript name E. “kathleenae ” in Sorensen (1983). Etymology and Common Name. — The species is named for my wife, Kathleen Hale Sorensen, who served as my field botanist during this study. Common name: Kathleen’s sugar pine needle aphid. Material Examined. — CALIFORNIA. CALAVERAS Co.: 18 km E of Arnold on hwy 4, 1680 m, 17 Jul 1977, JTS 77G45, P. lambertiana, (apt.). DEL NORTE Co.: Panther Flat Cmpgd, Six Rivers Natl Forest, at Pioneer Rd & hwy 199, E of Gasquet, 4 Jul 1978, JTS 78G7, P. monticola, (apt.). EL DORADO Co.: Lake Tahoe, Emerald Bay, 1980 m, 16 Jul 1977, JTS 77G30, P. lambertiana, (apt.). FRESNO Co.: jet of hwy 180 & Sequoia Lake turnoff, nr Pinehurst, 1710 m, 13 Aug 1977, JTS 28 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Figure 3. Distribution of E. (A.) kathleenae [dots (JTS samples)], superimposed over the range of its host, Pinus lambertiana [shaded]. 77H10, P. lambertiana, (apt.). KERN Co.: Tiger Flat Rd, N of hwy 155, nr Alta Sierra, 1890 m, 20 Sep 1977, JTS 77164, P. lambertiana, (apt.); same but JTS 77166, P. jeffreyi, (apt.). LOS ANGELES Co.: 3 km SE of Big Pines on hwy 2, E of Blue Ridge Summit, 2200 m, 17 Sep 1977, JTS 77148, P. lambertiana, (apt., ovip.). MARIPOSA Co.: Yosemite Natl Park, 13 km W of Crane Flat on hwy 120, 2140 m, 1 Aug 1977, JTS 77H6, P. lambertiana, (apt.). MENDOCINO Co.: Fish Rock Rd, 27 km E of hwy 1, 490 m, 23 Jul 1977, JTS 77G49, P. lambertiana, (apt.). MONTEREY Co.: Cone Peak Rd, 13 km N of jet with Nacimento-Fergusson Rd, Los Pardes Natl Forest, 1310 m, 4 Sep 1977, JTS 77110, P. lambertiana, (apt.). PLACER Co.: 5 km SW of Whitmore on hwy 80, 1430 m, 25 Jun 1977, JTS 77F2, P. lambertiana, (apt.). PLUMAS Co.: hwy 36, 6 km W of jet with hwy 89, 1460 m, 10 Jul 1977, JTS 77G22, P. lambertiana, (apt.); 8 km E of Chester on hwy 36, 1520 m, 4 Jul 1977, JTS 77G16, P. lambertiana, (apt.). RIVERSIDE Co.: South Ridge Rd, nr Idyllwild, 1770 m, 9 Sep 1977, JTS 77121, P. lambertiana, (apt.). SAN BERNARDINO Co.: (type series) San Bernardino Mts, 3 km S of jet of hwy 38 & Jenks Lake Rd, 2200 m, 16 Sep 1977, JTS 77138, P. lambertiana, (apt.); same but 3 km S of Lake Gregory, 1490 m, 17 Sep 1977, JTS 77145, P. lambertiana, (apt.). SISKIYOU Co.: Mt Shasta Ski Bowl Rd, 2450 m, 2 Jul 1977, J. T. Sorensen & D. J. Voegtlin, JTS 77G8, P. lambertiana, (apt.). TEHAMA Co.: Lanes Valley Rd, nr jet with hwy 36, 490 m, 4 Jul 1977, JTS 1994 SORENSEN: A REVISION OF ESSIGELLA 29 77G17, P. sabiniana, (apt.). TRINITY Co.: East County Line Rd, 5 km S of Buckhom Summit on hwy 299, W of Tower, 1530 m, 20 Aug 1977, JTS 77H19, P. lambertiana, (apt.). TUOLUMNE Co.: 2 km E of Groveville on hwy 120, 910 m, 30 Jul 1977, JTS 77G62, P. lambertiana, (apt.); same but JTS 77G63, P. ponderosa, (apt.). VENTURA Co.: Reyes Peak Rd, 10 km E of Pine Mt Summit on hwy 33, 2200 m, 19 Sep 1977, JTS 77158, P. lambertiana, (apt.). OREGON. JACKSON Co.: 15 km S of Union Creek on hwy 62, 850 m, 5 Jul 1978, JTS 78G17, P. lambertiana, (apt.). Essigella ( Lambersella ), NEW SUBGENUS “Essigella ( Lambersella )” Sorensen, 1983: 73 (unpublished manuscript name) Ph.D. Thesis, University of California at Berkeley, Berkeley, California. 605 p. Type Species.—Essigella fusca Gillette & Palmer, 1924, Ann. Entomol. Soc. Am., 17: 6-9. Viviparous Apterae.— Morphology: Body not relatively broad. Meso- and metanota fused dorsally; abdominal tergum I free. Dorsal setae (majors and minors) between muscle attachment plates on abdominal segments III-IV in 2 (rarely 1) often irregular rows (see Figs. 1C-D); lateral-most dorsal minor seta on each side anterad (rarely not) of its immediately mesad neighbor. Abdominal terga III- IV each with 8-12 dorsal (major + minor) and 3-5 (per side) marginal setae; tergum VIII with 8-11, rarely 12, setae. Longest dorsal seta on central one-third of metatibiae to nearly 4x tibial diameter, tips incrassate to sharp, sometimes reflexed; these setae sometimes dimorphic in length or with abrupt length transition, nearly doubling, centrally on the metatibiae. Ventral abdominal sclerites on segments III-IV reduced (rudimentary), irregular stellate to large, subquadrate to sublinear, often broken into linearly separated parts. Species means for length ratio of metadistitarsus to metabasitarsus varying between 1.39:1 to 1.54:1. Pigmentation: Body dorsum variable, pale to dark brown, background unicolorous or not, often mottled; bases of dorsal setae of abdomen concolorous with surrounding terga to substantially darker. Tibiae varying from entirely pale to nearly black, but when darkened mesotibiae at least subtly to usually substantially paler than pro- and metatibiae. Diagnosis. — See the key to the subgenera of Essigella. Discussion.—Lambersella is monophyletic and convex (sensu Duncan 1980, Estabrook 1986), and represents the sister clade of E. (Essigella ). The major nonhomoplasious, qualitative synapomorphy for this subgenus is its “dark-light- dark” tibial pigmentation suite for the pro-, meso- and metatibiae, respectively. Also, unlike any other Essigella, there is also a tendency for the ventral abdominal sclerites on segments III-IV to often be linear; that trait is problematic, however, because it is shared by Pseudessigella (e.g., Sorensen 1991: figs. 2b-e). I consider a length dimorphism of the dorsal setae on the metatibiae [except E. (L.) hilleris- lambersi ] and the usually faint, to absent, forewing medius of alates [morph unknown for E. (L.) eastopi ] to be apomorphies that are unique to Lambersella, but that are not found in, or known from, all its species. Of the Lambersella species, the phylogenetic analysis (Fig. 13) shows E. ( L .) eastopi to be the least derived (closest to the ancestral node 3) and E. ( L .) hillerislambersi the most, in anagenic distance from node 3. Ecologically, Lambersella has invaded the genetically distinct subsection Pon- derosae of the diploxylon pines (subgenus Pinus)\ only E. ( L .) eastopi feeds chiefly on a subsection Sabinianae pine, perhaps reflecting its relatively primitive status in the subgenus. Sorensen (1983: section 2) analyzed the relationships among the taxa within this subgenus; see the discussion under E. (L.) eastopi for a summary. An apparent case of character displacement has occurred under sympatry in California between E. (L.) fusca voegtlini and E. ( L .) hillerislambersi, with respect to bivariant regressions of the length of dorsal setae on the metatibia versus metatibial length (unpublished data); see discussion of E. (L.) fusca voegtlini. This 30 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) suggests that Lambersella species compete for their pine hosts as resources, as apparently do species in the E. ( E .) knowltoni group of E. (Essigella ) (Sorensen 1992a), where a similar form of character displacement also occurs. Coded References to This Taxon. — Sorensen (1983) referred to this taxon under the manuscript name “ Essigella (Lambersella) Sorensen (1987a) referred to the assemblage that comprises this taxon as group “II” or, with reference to its sub¬ components, as “O-N-M-L” [or entire permutations therefore]; in Sorensen (1992b), the latter refers to it. Etymology. — The subgenus is named for Dirk Hille Ris Lambers, who pointed out the tibial pigmentation synapomorphy. Material Examined.—Essigella (L.) eastopi, E. ( L .) fusca fusca, E. (L.) fusca voegtlini, E. (L.) hillerislambersi. Essigella (Lambersella) eastopi, NEW SPECIES Essigella “eastopi ” Sorensen, 1983: 76 (unpublished manuscript name) Ph.D. Thesis, University of California at Berkeley, Berkeley, California. 605 p. Type Series. — Holotype, vivip. apt.; on slide with 1 paratype vivip. apt., ho- lotype on top (12 o’clock position); data: CALIFORNIA. SAN DIEGO Co.: 8 km N of Mt Laguna, hwy SI, 1700 m, 12 Sep 1977, J. T. Sorensen (77132), Pinus coulteri D. Don. Holotype deposited in The Natural History Museum, London. Paratypes (all same data as holotype): 17 vivip. apt. on 5 slides including holotype slide. Paratype slides deposited: 1 slide in NMNH, Washington, D.C.; 1 slide in CNC, Ottawa, Ontario; 2 slides in Sorensen collection. (The type series represents smaller, darker specimens with short setae; these are the more distinctive form of E. (L.) eastopi.) Viviparous Apterae.— Morphology: Body length: 1.65-1.98 (1.84 ± 0.09) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously protuberant. Length of antennal segment V: 105-133 (118 ± 8) g, processus terminalis: 30-40 (34 ± 3) ix- IV: 73-88 (82 ± 5) m; HE 133-183 (146 ± 13) II: 65-73 (69 ± 2) ix. Length of longest setae on frons: 45-66 (52 ± 8) ix, tips incrassate to sharp. Head width: 270-306 (286 ± 11) ix. Length of stylets: 714-836 (766 ± 39) ix: ultimate rostral segment: 65-90 (81 + 8) ix, rostral tip reaching abdominal terga I—III in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 357-428 (385 ± 21) ix. THORAX: Meso + metanota fused, total length: 316-388 (348 ± 21) g. ABDOMEN: Tergum I free, length: 133-184 (158 ± 21) terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 33-45 (40 ± 3) siphunculi truncated conical, protrusion 0.2- 0.6 x maximal distal width. Ventral abdominal sclerites on segments III-IV subquadrate, subelliptical to sublinear; length: 44-59 (52 ± 5) ix, 1.3-2.Ox diameter of metatibiae. Dorsal (major + minor) setae (see Figs. 1C-D) on abdominal terga III-IV: 8-10 (9 ± 1), tips blunt to sharp, in 2 (rarely 1) rows with setae in regular positions, lateral-most minor dorsal seta in anterad row (rarely not); marginal setae 3-4 per segment each side. Setae on abdominal tergum VIII: 8-11 (9 ± 1), length: 45-73 (55 ± 9) ix, tips incrassate to sharp, in 1 or 2 irregular rows. Cauda rounded; caudal protuberance absent to poorly developed; length of longest caudal setae: 75-103 (91 + 9) ix, tips sharp. LEGS: Length of metafemora: 490-704 (578 ± 59) m; metatibiae: 612-908 (699 ± 84) ix\ longest dorsal setae on central one-third of metatibiae: 23-83 (53 ± 17) n, 0.3-2.3 x diameter of metatibiae, tips incrassate to sharp; length variable, either approximately equal to gradually increasing distally, or abruptly doubling in length on central tibiae with setal length dimorphism present; longest ventral setae on metatibiae: 23- 43 (34 ± 7) ix, tips sharp. Length of metabasitarsus: 104-128 (115 ± 7) /x; metadistitarsus: 163-195 (177 ± 11) ix. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.54:1. Pigmentation: Color in life: Yellow throughout to body brown with yellow frons, legs and longitudinal stripe on 1994 SORENSEN: A REVISION OF ESSIGELLA 31 dorsomedial thoracic and abdomen. Dark specimens with tibiae usually yellow, to infrequently pro- and metatibiae as dark as dorsum, mesotibiae yellow; yellow specimens with tibiae concolorous with body to pro- and metatibiae brown, mesotibiae yellow. Slide-mounted specimens: Background of body dorsum variable, unicolorously pale to dark brown with paler frons and longitudinal area on dorso¬ medial terga of thorax and anterad of abdomen; dark pigmentation homogeneous to mottled (to 80 percent pigment density), when latter the pigmentation is density centers around each of the mesal pair of muscle attachment plates on the abdominal dorsum. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga, to very subtly darker. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen pale, vaguely conspicuous, to dark brown, reticulate with well defined borders, conspicuous. Spiracular plates and ventral ab¬ dominal sclerites light to dark brown, conspicuous. Siphunculi concolorous with surrounding terga. Cauda, anal and subgenital plates concolorous with abdominal terga, to slightly darker. Antennal segments V dusky; IV usually dusky on distal one-half or central one-third, frequently entirely dusky; III usually pale, infrequently subtly dusky on distal extreme when IV is entirely dusky; II very pale; I concolorous with frons. Tibiae variable, usually concolorously pale, despite dorsal pigmentation, often to moderate brown with pro- and metatibiae darker than mesotibiae; when metatibiae dark, rarely proximal one-third and ventrodistal tip subtly paler, or rarely pigmentation increasing evenly distally. Distitarsi dusky on distal one-half to three-quarters, when tibiae pale, to entirely brown with tibiae. Ultimate Stadium Nymphs of Viviparous Apterae. — Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distant scleroites. Mesonotum with 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; plates usually heavily, to faintly, pigmented, diameter approximately equaling eye length. Viviparous Alatae, Oviparae, Males, Fundatrices.— Unknown. Diagnosis. —Essigella (L.) eastopi consists of pale to dark brown individuals. Dark specimens usually can be distinguished from other Essigella by having a dark brown body dorsum with a paler, longitudinal area on the dorsomedial region of the thoracic and anterad abdominal dorsum. The tibial pigmentation of E. (L .) eastopi is similar to that of E. (L.) fusca and E. (L.) hillerislambersi, and varies from all tibiae concolorously pale to a pigmentation suite in which the pro- and metatibiae are subtly to substantially darker than the mesotibiae; the latter is less prevalent in those E. ( L .) eastopi with a dark body dorsum. When pale, the three Lambersella species can be difficult to separate. Some E. ( L .) eastopi with short dorsal setae on the metatibiae (less than 1.2 x tibial diameter) and E. (L .) fusca with long setae (greater than 3.0 x tibial diameter) are exclusive. Essigella (L.) hillerislambersi is larger and can be separated from E. ( L .) eastopi if antennal segment III exceeds 0.190 mm. Most morphometric characters overlap in these three species; reliable separation requires application of the discriminant functions in the key to the viviparous apterae [couplets 20 and 21, in that order]. Pale E. ( L .) eastopi also can be confused with most other pale Essigella. They differ from E. (E.) californica, E. (E.) hoerneri and E. ( E .) pini by having eight or more (see Figs. 1C-D), rather than six (Fig. IF), dorsal (major + minor) setae on abdominal terga III-IV. Essigella (L.) eastopi lacks: the abdominal tergum I fusion of E. ( E.) essigi\ the protuberant, unusually distad primary rhinarium of E. (E.) wilsoni; and the exceptionally long metadistitarsus and short metabasi- tarsus of E. (A.) kathleenae. Essigella (L.) eastopi differs from E. (E.) alyeska by having three to five, rather than two, marginal setae on abdominal terga III-IV, and having large invasive, rather than small noninvasive, muscle attachment plates on the mesonotum of later stadia nymphs of apterae. Pale E. (L.) eastopi differ from pale E. ( E.) knowltoni by being narrower, with sometimes sharply 32 THE PAN-PACIFIC ENTOMOLOGIST Yol. 70(1) Figure 4. Distribution of: A. E. (L.) eastopi [dots (JTS samples)], superimposed over the range of its host, Pinus coulteri [shaded]. B. E. ( L .) hillerislambersi [dots (JTS samples), squares (nonJTS samples)], superimposed over the range of its host, Pinus jeffreyi [shaded]. tipped dorsal metatibial setae that frequently have an abrupt increase in length on the central part of the metatibiae. Pale E. (L.) eastopi with short dorsal metatib¬ ial setae (less than 0.7 x tibial diameter) can be especially similar to E. (A.) kirki, but have a metadistitarsus to metabasitarsus ratio of less than 1.70:1. Range. —Coastal ranges of California, south of San Francisco Bay, to Mexico. The geographic range of E. ( L .) eastopi is the most restricted of the genus (Fig. 4 A). Host.—Pinus coulteri D. Don. Essigella ( L .) eastopi, on a subsection Sabinianae pine, is the only E. (Lambersella ) that does not feed primarily on subsection Ponderosae pines, although P. coulteri does hybridize with P. jeffreyi of subsection Ponderosae. Discussion. —Essigella (L.) eastopi is a relatively common species that is fairly 1994 SORENSEN: A REVISION OF ESSIGELLA 33 variable in morphology, especially in the length of the dorsal setae on the metatib¬ iae. There are two semi-discrete, but intergrading, pigmentation morphs for its viviparous apterae. The general darkening of the background of the body dorsum on the darker morph can be considered a homoplasy with that of E. {.Essigella ); however, that morph’s longitudinal, lightened dorsomedial area on the thorax and abdomen is an autapomorphy for E. {L .) eastopi. Essigella ( L .) eastopi is evolutionarily close to E. (L.) fusca, with which it shares several bivariate morphometric regressions. However, it appears to be morpho¬ logically closest to allopatric, rather than sympatric, populations of that species (see below). Dark individuals of E. {L .) eastopi generally have shorter setae than do paler specimens, which can approach E. (L.) fusca in appearance. Bivariant plotting of the longest dorsal seta on the central part of the metatibiae, versus metatibial length (unpublished data), suggests that E. {L.) eastopi may be a di¬ minutive of E. {L .) fusca voegtlini, with respect to that derived regression line; it differs in this respect, however, from allopatric E. (L.) fusca fusca, which has relatively longer metatibiae. The isozymes and nucleic acids of populations of species in E. (Lambersella) should be examined, especially in the Tehachapi, San Gabriel and San Bernardino mountains of southern California; there, one large collection (D. J. Voegtlin 17; Running Springs, San Bernardino Co., on P. coulteri) is troublesome and may obscure clear separation of E. {L .) eastopi from E. {L .) fusca voegtlini. Sorensen (1983) analyzed the relationships among taxa within E. {Lambersella) using principal component and discriminant function analyses on 35 morpho¬ metric traits. The principal component analysis (Sorensen 1983: section 2 PCA- 1) showed that E. {L.) eastopi differed from E. (L.) fusca, as a species, in general- size, as represented as the first vector (which had uniformly high trait loadings and correlations). It was partially displaced from E. (L.) fusca on that vector, which gave the greatest separation to E. (L.) hillerislambersi. Essigella (L.) eastopi differed from sympatric E. (L.) fusca voegtlini on the second principal component vector, which chiefly involved the length of dorsal setae on the metatibiae; how¬ ever, allopatric E. {L.) fusca fusca was intermediate between those two taxa on that vector. Essigella (L.) eastopi chiefly occupied the same size-independent principal component attribute space, defined by vectors 2 and 3 after size removal, as did sympatric E. (L.) hillerislambersi. The discriminant function analysis (Sorensen 1983: section 2 DFA) echoed the findings of the principal component analysis, but with better group resolution and refined intergroup anagenic distances, as expected. Function 1 showed E. (L.) eastopi had greatest separation from E. (L.) hillerislambersi, with sympatric E. {L.) fusca voegtlini intermediate between those two, and allopatric E. (L.) fusca fusca yet intermediate between E. (L.) eastopi and E. (L.) fusca voegtlini. Dis¬ criminant function 2 showed E. ( L .) fusca voegtlini separated from E. (L.) eastopi, E. (L.) fusca fusca and E. (L. ) hillerislambersi, the three of which overlapped. These analyses indicate a form of character displacement occurs among the three E. {Lambersella) species in California under sympatry (unpublished data). Coded References to This Taxon.—Essigella {L.) eastopi has been referred to previously by: the coding “Sp. L” (Sorensen 1983 [but not section 2], 1987a, 1992b), “group B” (Sorensen 1983: section 2) and “EAST” (Sorensen 1983); and by the manuscript name E. “ eastopi ” in Sorensen (1983). 34 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Etymology and Common Name.— The species is named for the aphidologist V. F. Eastop, who introduced me to the use and interpretation of bivariate plotting in aphid taxonomy. Common name: Eastop’s Coulter pine needle aphid. Material Examined. —CALIFORNIA. LOS ANGELES Co.: hwy 2, 7 km NE of jet with Mt Wilson Rd, San Gabriel Mts, 1530 m, 18 Sep 1977, JTS 77151, P. coulteri, (apt.). MONTEREY Co.: Cone Peak Rd, 2 km N of jet with Nacimento-Fergusson Rd, Los Padres Natl Forest, 910 m, 4 Sep 1977, JTS 7719, P. coulteri, (apt.). ORANGE Co.: above Santiago Peak Rd, 10 km N of jet with hwy 74, Cleveland Natl Forest, 1220 m, 10 Sep 1977, JTS 77122, P. coulteri, (apt.). RIVERSIDE Co.: Keen Camp Summit on hwy 74, 3 km N of Mountain Center, San Bernardino Natl Forest, 1500 m, 9 Sep 1977, JTS 77120, P. coulteri, (apt.). SAN BERNARDINO Co.: “view” Picnic Area on hwy 18, W of Rimforest, San Bernardino Natl Forest, 1620 m, 17 Sep 1977, JTS 77144, P. coulteri, (apt.); 7 km W of Barton Flat on hwy 38, 1950 m, 16 Sep 1977, JTS 77136, P. coulteri, (apt.). SAN DIEGO Co.: 5 km S of Julian, Harrison Springs Rd, 1460 m, 12 Sep 1977, JTS 77129, P. coulteri, (apt.); (type series) 8 km N of Mt Laguna on hwy SI, 1700 m, 12 Sep 1977, JTS 77132, P. coulteri, (apt.); Mt Palomar Rd (S6), 3 km S of Mt Palomar, 1530 m, 11 Sep 1977, JTS 77127, P. coulteri, (apt.). SAN LUIS OBISPO Co.: Cuesto Ridge Botanical Area, nr La Cuesta Summit on hwy 101, N of San Luis Obispo, 730 m, 5 Sep 1977, JTS 77114, P. coulteri, (apt.). Essigella ( Lambersella ) fusca fusca Gillette & Palmer 1924, NEW STATUS Essigella fusca Gillette & Palmer, 1924: 6, Ann. Entomol. Soc. Am., 17: 6-9. Essigella agilis Hottes, 1957: 71, Proc. Biol. Soc. Wash., 70: 71-73. NEW SYN¬ ONYM. Essigella palmerae Hottes, 1957: 96, Proc. Biol. Soc. Wash., 70: 96-98. NEW SYNONYM. Primary Types. — Lectotype, vivip. apt., on slide alone; slide data: “Essigella fusca, apt. viv., Holotype, C. P. Gillette & M. A. Palmer, Mt’d. in Damar in xylene/U.S. Nat. Mus., No. 41953/On Pinus ponderosa var. scopulorum, Rocky Mt. Nat. Park (Grags Hill [sic], near Bald Pate Inn) Colo., 7-18-23, Coll. M. A. Palmer, Colo. Agr. Exp. Ac. No. 3422/[on back] lectotype, J. T. Sorensen, 1982.” Lectotype deposited in the U.S. National Museum of Natural History, Washing¬ ton, D.C. There is a problem regarding type designation; Hottes (1957: 88) confusingly mentions both a lectotype and holotype for this species. A slide marked “holotype” exists. In the original description, Gillette & Palmer (1924: 6-9) do not designate a primary type, but later (Gillette & Palmer 1931: 840) state “Types in the U.S. Nat. Mus., Cat. No. 41953; Paratypes in collection of Colo. Agr. Exp. Sta.” Palmer (1952: 15) under the heading “Type” also lists that number. In addition, 2 slides (alio- and morphotypes) bear the number, precluding identification of any indi¬ vidual as lectotype, based on the number alone. Because I cannot tell from Hottes’ publication (1957) that he clearly was designating a lectotype, I presently designate the “holotype” specimen as lectotype, following Hottes’ mention of it. I have added the lectotype label listed above to the back of that slide. Unfortunately, the specimen lacks metalegs and is obscured by debris, but it is recognizable as E. ( L .) fusca. Viviparous Apterae.—Morphology: Body length: 1.79-2.39 (2.09 ± 0.18) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously 1994 SORENSEN: A REVISION OF ESSIGELLA 35 protuberant. Length of antennal segment V: 118-145 (130 ± 7) ju, processus terminalis: 30-48 (37 ± 4) ix\ IV: 73-100 (88 ± 8) n; III: 135-180 (159 ± 12) /u,; II: 65-95 (76 ± 7) n. Length of longest setae on frons: 35-80 (58 ± 11) tips incrassate to sharp. Head width: 245-316 (293 ± 17) ji. Length of stylets: 602-867 (756 ± 62) ultimate rostral segment: 78-98 (90 ± 5) rostral tip reaching abdominal terga I—II, infrequently III, in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 383-479 (424 ± 25) p. THORAX: Meso + metanota fused, total length: 326-459 (400 ± 31) ii. ABDOMEN: Tergum I free, length: 138-184 (162 ± 14) p; terga II—VII fused, VIII free. Maximum distal width of flange on siphunculi: 38-50 (45 ± 3) p\ siphunculi truncated conical, protrusion 0.2-0.6 x maximum distal width. Ventral abdominal sclerites on segments III-IV subquad¬ rate, subelliptical to sublinear, often centrally constricted; length: 48-90 (73 ± 10) p, 1.2-2.6 x diameter of metatibiae. Dorsal (major + minor) setae (see Figs. 1C-D) on abdominal terga III-IV: 8-12 (11 ± 1), tips sharp, in 2 (rarely 1) rows with setae in regular position, lateral-most minor dorsal seta in anterad row; marginal setae 3-5, usually 4 per segment each side. Setae on abdominal tergum VIII: 8-11 (10 ± 1), length: 40-80 (64 ± 9) p, tips incrassate to sharp, in 1-2 rows. Cauda broadly rounded; caudal protuberance absent to poorly developed; length of longest caudal setae: 83-128 (102 ± 13) P, tips sharp. LEGS: Length of metafemora: 581-898 (762 ± 95) p\ metatibiae: 755-1132 (971 ± 104) p\ longest dorsal setae on central one-third of metatibiae: 50 (rarely 15)—120 (76 ± 18) p, 0.5- 3.6, usually 1.2-2.8,x diameter of metatibiae, tips usually incrassate to blunt, occasionally sharp; length variable, either approximately equal along tibiae, gradually increasing distally, or abruptly doubling in length on central tibiae with setal length dimorphism present; longest ventral setae on metatibiae: 29-68 (41 ± 9) p, tips sharp. Length of metabasitarsus: 130-170 (148 ± 12) /u; meta- distitarsus: 170-233 (205 ± 16) p. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.39:1. Pigmentation: Color in life: Head and thorax yellow-brown, abdomen green, pro- and meta¬ tibiae light to dark brown with mesotibiae yellow-brown, dorsal spots brown; or frequently green- yellow to straw yellow, rarely gray throughout. Slide-mounted specimens: Background of body dorsum pale to moderate brown, often mottled, rarely dark brown (usually to 30, rarely to 80, percent pigment density). Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen subtly to substantially darker than surrounding terga. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen varying from moderate to dark brown, conspicuous, often reticulate, sometimes with surrounding tergum more heavily mottled than elsewhere, to pale, inconspicuous. Spiracular plates and ventral abdominal sclerites usually moderate to dark brown, conspicuous, to pale, inconspicuous. Siphunculi concolorous with surrounding terga. Cauda, anal and subgenital plates concolorous with abdominal terga, to slightly darker. Antennal segments V and IV moderate to dark brown, usually concolorous, but frequently paler proximally, infrequently also paler distally; III usually moderate to dark brown on distal one-third, rarely one-half, remainder pale, often entirely pale; II usually subtly darker than proximal III, seldom conspicuously darker, rarely concolorous with proximal III; I usually concolorous with frons, to conspicuously darker. Tibiae variable, usually pro- and metatibiae evenly light to dark brown with mesotibiae substantially paler, sometimes dark pro- and metatibiae paler on proximal and distal tips to one-fourth; commonly all tibiae concolorously pale when body dorsum pale. Distitarsi usually evenly light to dark brown with pro- and metatibiae, to dusky with paler proximal tip when tibiae pale. Ultimate Stadium Nymphs of Viviparous Apterae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum with 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; plates usually distinct, moderately to darkly pigmented, diameter approximately equaling eye length. Viviparous Alatae. - Slide-mounted specimens: Nonmorphometrics as described for viviparous ap¬ terae except lacking body dorsum pigmentation suite; abdominal terga normally membranous, dorsal (major + minor) setae between muscle attachment plates frequently arising from distinct scleroites; antennal segments often as dark as tibiae, except proximal one-fifth of III pale. Antennal segment III with 0-5, IV with 0-1, secondary rhinaria. Epicranial suture absent. Forewing medius with single furcation arising on proximad one-third of vein; cubital base usually arising distad, infrequently proximad, on subcosta with distance between anal and cubital bases on subcosta usually relatively large, ca. 30-40 percent or more of anal vein length; usually medius, sometimes cubitus and anal veins faint, vague to absent. Abdominal terga frequently with irregular sclerites that engulf or join muscle attachment plates and dorsal (major + minor) setal bases or scleroites. 36 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Oviparae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae ex¬ cept abdominal terga II-VI fused, lightly to moderately sclerotic, including pleural areas, but VII and VIII free; dorsal demarcations of anterad terga rarely evident; siphunculi usually incorporated into sclerotic dorsum, to free; dorsal abdominal muscle attachment plates unicolorous. Pseudorhinaria on metatibiae irregular, 9-27. Males. — Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except body slightly smaller, with slightly longer antennae and tibiae; dorsal demarcations of abdominal terga evident. Antennal segment III with 13-15, IV with 8-10, secondary rhinaria. Fundatrices. — Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except siphunculi absent; longest dorsal setae on central part of metatibiae 0.8-1. Ox tibial diameter. Diagnosis. — Pigmented Essigella (X.) fusca can be distinguished from all Es- sigella, except E. (L.) hillerislambersi and E. (X.) eastopi, by the tibial pigmentation suite in which the pro- and metatibiae are often substantially darker than the mesotibiae. Essigella (X.) fusca lack the completely developed pigmentation for the body dorsum shown by some E. ( L .) eastopi, but cannot be separated reliably from E. ( L .) hillerislambersi by pigmentation. Pale E. (X.) fusca, E. (X.) eastopi and E. (X.) hillerislambersi can be separated from other pale Essigella by the diagnostics given for pale E. (X.) eastopi. Pale E. ( L .) fusca have longer dorsal setae on the metatibiae (greater than 1.2 x tibial diameter) than some E. ( L .) eastopi, and a shorter antennal segment III than E. (X.) hillerislambersi ; but these differences are indiscrete. Reliable separation of these Lambersella species requires application of the discriminant functions in the key to the viviparous apterae [couplets 20 and 21, in that order]. As subspecies, E. (X.) fusca fusca and E. (X.) fusca voegtlini are morphologically indiscrete, with clinal univariate characters; their separation is locality dependent, but they can be classified using the discriminant function in the key to the vivip¬ arous apterae [couplet 22]. In Essigella (X.) fusca fusca, the metatibiae and an¬ tennal segment V are generally longer, and the dorsal setae on the metatibiae are generally shorter, than in E. (X.) fusca voegtlini. Synonyms. —Essigella agilis Hottes, NEW SYNONYM: holotype, vivip. apt., on slide with 4 other specimens, holotype shown by arrow (7-8 o’clock position); data: COLORADO. MESA Co.: Glade Park, 26 Jun 1956, F. C. Hottes, Pinus ponderosa Lawson. Essigella agilis holotype deposited in NMNH. Essigella palmerae Hottes, NEW SYNONYM: holotype, vivip. alat., on slide with morphotype vivip. apt.; data: ARIZONA. PIMA Co.: Summerhaven, 13 Jun 1954, F. C. Hottes, Pinus ponderosa. Essigella palmerae holotype deposited in NMNH. Range.— Southern British Columbia, south: (in the east) through the Rocky Mountains to Arizona, New Mexico and into Mexico; (in the west) to northern and eastern Oregon, but not California or southwestern Oregon (Fig. 5). [For species, see E. (X.) f. voegtlini also.] Hosts. — Subsection Ponderosae pines, principally Pinus ponderosa Lawson, but also P. ponderosa var. arizonica Engelmann, P. engelmannii Carriere, and P. leiophylla Schiede & Deppe (latter, subsection Leiophyllae). Assuming the iden¬ tification is correct, a reputed collection from Callitris drummondii Betham & Hooker f. ex F. Mueller (Cupressaceae), listed in Blackman & Eastop (in press) as “BMNH colln, leg. H. G. Walker” [R. L. Blackman, personal communication], and in Walker et al. (1978: 588) under that host as “31/1/71 Moderate (VFE),” is undoubtedly from a nonresident host in one of Walker’s many Los Angeles 1994 SORENSEN: A REVISION OF ESSIGELLA 37 Arboretum samples; most identifications of Essigella to species that are listed in Walker et al. (1978) are questionable, because only Hottes’ (1957) key was avail¬ able at the time. [For species, see E. (L.) f voegtlini also.] Discussion.—Essigella (L.) fusca is a common and morphologically variable species. Sorensen (1983) analyzed the E. (Lambersella) species; see the discussion of E. (L.) eastopi for a brief summary. That study also analyzed E. ( L .) fusca itself, after breaking it further into geographic subunits for other principal com¬ ponent analyses (Sorensen 1983: section 2 PCA-2, PCA-3). Within E. (L.) fusca, general-size variance (Sorensen 1983: section 2 PCA-2, vector 1) dominated any difference among populations. There was, however, a general east-west gradient (Sorensen 1983: section 2 PCA-2, vector 2) with longer dorsal setae on the metatib¬ iae, and shorter metatibiae occurring in the west [E. ( L.) f voegtlini ], and the opposite combination in the more eastern portions of the range [E. (L.) f. fusca]. The next most dominant vector (Sorensen 1983: section 2 PCA-2, vector 3) in that analysis suggested a very rough north-south morphocline among non- Californian populations [E. (L .) f fusca], which the Californian material [E. (L .) f voegtlini] spanned. When material from California was omitted from those analyses to improve resolution further, general-size variance (Sorensen 1983: section 2 PCA-3, vector 1) still dominated interpopulational differences among the nonCalifomian pop¬ ulations. However, subordinate to that, nonCalifomian populations [E. ( L .) f fusca] showed a general north-south gradient (Sorensen 1983: section 2 PCA-3, vector 2) that involved the length of dorsal setae on the metatibiae and lateral setae on the body, plus the number of dorsal (major + minor) setae on the abdomen; this vector, in the absence of interference from E. ( L .) f voegtlini, oriented to, and improved the resolution of, the variance revealed in the second vector of the previous analysis (Sorensen 1983: section 2 PCA-2, vector 3). The third vector (Sorensen 1983: section 2 PCA-3, vector 3) for the nonCalifomian populations showed mostly intrapopulational variance. With respect to qualitative traits, E. (L.) fusca fusca populations from Arizona and New Mexico frequently are slightly paler, with slightly darker metatibiae that sometimes show both their distal and proximal ends to be paler. This southwestern material, however, is not considered sufficiently distinct to warrant recognition with subspecific status on the basis of either quantitative or qualitative traits. Coded References to This Taxon.—Essigella (L.) fusca fusca has been referred to previously by: the coding “Sp. M” (Sorensen 1983 [but not section 2], 1987a, 1992b), “group D” (Sorensen 1983: section 2) and “FUSC” (Sorensen 1983); and by the name E. fusca fusca in Sorensen (1983). Etymology and Common Name.—“Fusca,” from the Latin “ fuscus ,” meaning “dusky,” “dark” or “swarthy” (Brown 1978); apparently with reference to “. . . having dorsum of abdomen dark in color in apterous virgogenia” (Gillette & Palmer 1924: 8). Common name: the dusky ponderosa pine needle aphid; although Palmer (1952:14) refers to this species as “The Brown and Green Pine needle Aphid,” the common name indicated here is more appropriate and less confusing because other Essigella are brown and green. Material Examined. — [E. (L.) fusca fusca only :] ARIZONA. APACHE Co.: 10 km N of Lupton on hwy 12 (= 166), 2070 m, 11 Sep 1978, JTS 78118, P. ponderosa, (apt.). COCHISE Co.: nr Rustler 38 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Figure 5. Distribution of E. (L.) fusca fusca [black dots (JTS samples), black squares (nonJTS samples)] and E. ( L.) fusca voegtlini [white triangles (JTS samples), white squares (nonJTS samples)], superimposed over the range of its principal host, Pinus ponderosa [shaded]. Park, Chiricahua Mts, 2500 m, 16 Sep 1978, JTS 78147, P. ponderosa, (apt.). COCONINO Co.: 9 km W of Williams on hwy 66, 2070 m, 9 Sep 1978, JTS 7815, P. ponderosa, (apt.). GILA Co.: Pine, 17 May 1978, C. F. & C. S. Smith, CFS 78-31, Pinus sp., (apt.). GRAHAM Co.: SW of Stafford on hwy 366, 1830 m, 15 Sep 1978, JTS 78136, P. leiophylla, (apt.); same but 1980 m, JTS 78137, P. ponderosa var. arizonica, (apt.). NAVAJO Co.: Mogollon Rim Rd, 8 km SW of Showlow, 2070 m, 10 Sep 1978, JTS 78113, P. ponderosa, (apt.). PIMA Co./Summerhaven, 13 Jun 1954, F. C. Hottes, (alat.). COUNTY UNCERTAIN: Sitgreaves Natl Forest, 18 Jun 1969, D. T. Jennings, P. ponderosa, (alat.). COLORADO. ARCHULETA Co.: 25 km W of Pagosa Springs on hwy 160, 2140 m, 8 Aug 1978, JTS 78H50, P. ponderosa, (apt.). GUNNISON Co.: 16 km NW of Kebler Pass, 2440 m, 13 Aug 1978, JTS 78H75, P. ponderosa, (apt.). LARIMER Co.: (lectotype) Bald Pate Inn, nr Craig’s Hill, Rocky Mt Natl Park, 12 Jul 1923, M. A. Palmer, CAES 3420, P. ponderosa, (apt.); (paratype) Craigs, Estes Park, 27 Jul 1923, M. A. Palmer, CAES 3430, P. ponderosa, (alat.); (type) Craigs, nr twin Sisters Mt, 27 Jul 1923, M. A. Palmer, CAES 3430/USNM 41953, P. ponderosa, (alat.); Estes Park, 1 Sep 1922, F. C. Hottes, CAES 3312/USNM 41953, P. ponderosa, (ovip., male); same but 24 Jul 1921, C. P. Gillette, CAES 2804, (apt.). MESA Co.: Glade Park, 26 Jun 1956, F. C. Hottes, P. ponderosa, (apt.); Carson Hole, 3/8 Aug 1956, (apt.). SAN MIGUEL Co.: 6 km NE of Placerville on hwy 62, 2320 m, 7 Aug 1978, 1994 SORENSEN: A REVISION OF ESSIGELLA 39 JTS 78H43, P. ponderosa, (apt.). IDAHO. IDAHO Co.: Deep Creek, nr Old Warrior’s Face, Bitteroot Natl Forest, 16 Aug 1979, D. J. Voegtlin, DJV 691, P. ponderosa, (apt.). VALLEY Co.: McCall, 23 Sep 1956, M. J. Forsell, P. ponderosa, (apt., alat., ovip., male). MONTANA. MISSOULA Co.: Big Larch Cmpgd, Seeley Lake, Lola Natl Forest, NE of Missoula, 20 Aug 1979, D. J. Voegtlin, DJV 713, P. ponderosa, (apt.). NEVADA. WHITE PINE Co.: Wheeler Peak, 2750 m, 26 Aug 1978, JTS 78H153, P. ponderosa, (apt.). NEW MEXICO. BERNALILLO Co.: 2 km NW of San Antinito on hwy 44, 2290 m, 12 Sep 1978, JTS 78120, P. ponderosa, (apt.). OTERO Co.: Cloudcroft, hwy 82, 2710 m, 13 Sep 1978, JTS 78125, P. ponderosa, (apt.). SANTA FE Co.: 20 km NE of Santa Fe on hwy 475, 2680 m, 10 Aug 1978, JTS 78H59, P. ponderosa, (apt.). SIERRA Co.: 3 km W of Kingston on hwy 90, 2140 m, 14 Sep 1978, JTS 78132, P. ponderosa, (apt., alat.). COUNTY UNCERTAIN: Gila Natl Forest, 19 Jul 1965, H. G. Kinzer, P. ponderosa, (alat.); same but 1 Nov 1967, (apt.). OREGON. BAKER Co.: 11 km W of Unity on hwy 26, 20 Jul 1978, JTS 78G112, P. ponderosa, (alat.). HARNEY Co.: 20 km N of Bums on hwy 395, 20 Jul 1978, JTS 78G117, P. ponderosa, (apt.). SOUTH DAKOTA. LAW¬ RENCE Co.: 20 km S of Deadwood on hwy 385, 1650 m, 18 Aug 1978, JTS 78H98, P. ponderosa, (apt.). UTAH. DAGGETT Co.: 21 km S of Manila on hwy 44, 2350 m, 24 Aug 1978, JTS 78H135, P. ponderosa, (apt.). KANE Co.: 50 km SE of Cedar City on hwy 14, 2560 m, 5 Aug 1978, JTS 78H28, P. ponderosa, (apt.). WYOMING. ALBANY Co.: hwy 287, 2 km N of state border, 15 Aug 1978, JTS 78H93, P. ponderosa, (apt.). CROOK Co.: 6 km W of Devil’s Tower Jet on hwy 14, 1100 m, 19 Aug 1978, JTS 78H104, P. ponderosa, (apt.). CANADA. BRITISH COLUMBIA: Fairmont Hotsprings, hwy 93, 17 Jul 1978, JTS 78G91, P. ponderosa, (apt.). MEXICO. PUEBLA: Puebla, km 43 Corn Fed., 11 Jun 1983, A. L. Munuz, 267, Pinus sp., (apt.). STATE UNCERTAIN: Sierra Largo, at El Passo, 12 Jun 1966, Eads & Rood, Pinus sp., (apt.). Essigella (Lambersella) fusca voegtlini, NEW SUBSPECIES Essigella “ fusca voegtlinr Sorensen, 1983: 89 (unpublished manuscript name) Ph.D. Thesis, University of California at Berkeley, Berkeley, California. 605 p. Type Series. — Holotype, vivip. apt., on slide with 2 paratype vivip. apt., the holotype is only complete specimen on the slide, at top (1 o’clock position); data: CALIFORNIA. FRESNO Co.: jet hwys 180 & 245, 1620 m, 13 Aug 1977, J. T. Sorensen (77H9), Pinus ponderosa. Holotype deposited in The Natural History Museum, London. Paratypes (all same data as holotype): 18 vivip. apt., on 5 slides including holotype slide. Paratype slides deposited: 1 slide in NMNH, Washington, D.C.; 1 slide in CNC, Ottawa, Ontario; 2 slides in Sorensen collec¬ tion. Viviparous Apterae.— Morphology: As E. (L.) fusca fusca, except as follows: Body length: 1.88-2.21 (2.04 ± 0.09) mm. HEAD: Length of antennal segment V: 105-135 (121 ± 8) p, processus terminalis: 30-40 (36 ± 3) m; IV: 70-100 (85 ± 7) p\ III: 128-178 (147 ± 14) p; II: 65-85 (73 ± 5) p. Length of longest setae on frons: 44-88 (59 ± 12) p. Head width: 275-311 (289 ± 10) p. Length of stylets: 551— 857 (732 ± 66) p; ultimate rostral segment: 84-95 (89 ± 3) p. Total length of fused head + pronotum: 377-449 (419 ± 16) p. THORAX: Total length of fused meso + metanota: 347-428 (389 ± 19) p. ABDOMEN: Tergum I length: 138-179 (157 ± 10) p. Maximum distal width of flange on siphunculi: 30-48 (39 ± 5) p. Ventral abdominal sclerite length: 44-80 (69 ± 9) p. Dorsal (major + minor) setae (see Figs. 1C-D) on abdominal terga III-IV: 8-12 (10 ± 1); marginal setae 4-5 per segment each side. Setae on abdominal tergum VIII: 8-10 (8 ± 1), length: 45-100 (69 ± 14) p. Length of longest caudal setae: 80-120 (98 ± 9) p. LEGS: Length of metafemora: 612-831 (732 ± 62) p\ metatibiae: 806-1061 (941 ± 75) p\ longest dorsal setae on central one-third of metatibiae: 58-135 (103 ± 19) p\ longest ventral setae on metatibiae: 38-83 (52 ± 11) p. Length of metabasitarsus: 128-155 (141 ± 9) p\ metadistitarsus: 173-223 (198 ± 11) p. Mean ratio of metadistitarsus to metabasitarsus: 1.40:1. Pigmentation: As E. (L.) fusca fusca. Diagnosis.—See the E. (L.) fusca fusca diagnosis, and couplet 22 in the key to the viviparous apterae. 40 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Range.— California, southwestern Oregon, extreme western Nevada (Fig. 5). [For species, see E. ( L .) f fusca also.] Hosts. — Subsection Ponderosae pines, principally Pinus ponderosa Lawson, but also P. jeffreyi Greville & Balfour and P. coulteri D. Don (latter, subsection Sabinianae); single collections from P. sabiniana Douglass (77F15), P. monophylla Torrey & Fremont (77H3), P. contorta murrayana Greville & Balfour (R. Luck sample) and P. quadrifolia Parlatore (77119) are probably not resident. [For spe¬ cies, see E. ( L .) f. fusca also.] Discussion. —Essigella (L.) fusca voegtlini, although a gradient subspecies [see discussion under E. ( L .) f. fusca], is named to recognize the morphometric prob¬ lems of E. (L.) fusca in sympatry with E. (L.) eastopi, and especially E. (L.) hillerislambersi. Essigella (.L.)f. voegtlini [and E. (L.) f. fusca to a very much lesser extent] shows a common dimorphism of length for the dorsal setae on the meta¬ tibiae. The dimorphism is evident in either of two forms: (a) on a given individual, as an abrupt transition from shorter to longer setae along the central portion of the dorsum of the metatibia; or (b) among various individuals in, or among, populations as the length of the longest setae on the central portion of the dorsum of the metatibia. This dimorphism causes a gap along a regression line of length of these setae when plotted against metatibial length (Sorensen 1983). Interestingly, the length of the dorsal setae of the metatibiae on E. ( L.) hillerislambersi correspond to this gap, indicating a character displacement in sympatry. These setal lengths do not appear to be influenced by host on E. ( L.) f voegtlini. Coded References to This Taxon. —Essigella (L.)fusca voegtlinih.?iS been referred to previously by: the coding “Sp. N” (Sorensen 1983 [but not section 2], 1987a, 1992b), “group C” (Sorensen 1983: section 2) and “VOEG” (Sorensen 1983); and by the manuscript name E. ‘ fusca voegtlini ” in Sorensen (1983). Etymology and Common Name.— The California subspecies is named for aphidologist D. J. Voegtlin, and his ever-present beard. Common name: Voegtlin’s dusky ponderosa pine needle aphid. Material Examined. — [E. (L.) fusca voegtlini only :] CALIFORNIA. BUTTE Co.: Feather River Cyn, 5 km NE of jet of hwy 70 & Cherokee Rd, 26 Jun 1977, JTS 77F15, P. sabiniana, (apt.). CALAVERAS Co.: 18 km E of Arnold on hwy 4, 1680 m, 17 Jul 1977, JTS 77G46, P. ponderosa, (apt.); 2 km NE of Murphys on hwy 4, 670 m, 17 Jul 1977, JTS 77G47, P. ponderosa, (apt.). EL DORADO Co.: Georgetown, 29 May 1977, J. T. Sorensen, P. ponderosa, (apt.). FRESNO Co.: (type species) jet of hwys 180 & 245, 1620 m, 13 Aug 1977, JTS 77H9, P. ponderosa, (apt.). INYO Co.: jet of Lake Sabrina Rd & Southern Calif. Edison Plant 2 Rd, nr Bishop, 2130 m, 1 Aug 1977, JTS 77H3, P. monophylla, (apt.). KERN Co.: Tehachapi Mtn Park, S of Tehachapi, 1980 m, 19 Sep 1977, JTS 77160, P. ponderosa, (apt., ovip.); same but JTS 77161, P. jeffreyi, (apt.); Tiger Flat Rd, N of hwy 155, nr Alta Sierra, 1890 m, 20 Sep 1977, JTS 77165, P. ponderosa, (apt.). LAKE Co.: 5 km S of Lake Pillsbury, Elk Mt Rd, 640 m, 24 Jul 1977, JTS 77G56, P. jeffreyi, (apt.). LASSEN Co.: 1 km SW of Susanville on hwy 36, 1460 m, 4 Jul 1977, JTS 77G13, P. jeffreyi, (apt.). LOS ANGELES Co.: Camp Baldy, 5 Dec 1956, J. MacSwain, “on fir,” (apt.); hwy 2, 7 km NE of jet with Mt Wilson Rd, San Gabriel Mts, 1530 m, 18 Sep 1977, JTS 77151, P. coulteri, (apt.). MONTEREY Co.: Plaskett Ridge Rd, Los Padres Natl Forest, 1040 m, 4 Sep 1977, JTS 77111, P. ponderosa, (apt.). PLUMAS Co.: 2 km SE of Graeagle on hwy 89, 1310 m, 26 Jun 1977, JTS 77F10, P. ponderosa, (apt.); Halsted Cmpgd, Plumas Natl Forest, 19 km NE of Beldon on hwy 70, 790 m, 26 Jun 1977, JTS 77F13, P. ponderosa, (apt.). RIVERSIDE Co.: 2 km N of Paradise Valley on hwy 74, 1500 m, 9 Sep 1977, JTS 77119, P. quadrifolia, (apt.). SAN BERNARDINO Co.: 7 km W of Barton Flat on hwy 38, 1950 m, 16 Sep 1977, JTS 77136, P. coulteri, (apt.); San Bernardino Natl Forest, “view” Picnic Area on hwy 18, W of Rimforest, 1610 m, 17 Sep 1977, JTS 77143, P. ponderosa, (apt.); same but 1620 m, JTS 77144, 1994 SORENSEN: A REVISION OF ESSIGELLA 41 P. coulteri, (apt., alat.); same but Barton Flat, 29 Aug 1972, D. J. Voegtlin, DJV 67, (apt.); same but Camp Angeles, 29 Aug 1972, D. J. Voegtlin, DJV 66, P. ponderosa, (apt.); same but Dogwood, 28 Aug 1972, DJV 25, (apt.); same but Snow Valley, DJV 69, P.jejfreyi, (apt.); same but Running Springs, 4 Aug 1973, DJV 77, P. coulteri, (apt.); San Bernardino Mts, nr jet of Jenks Lake Rd & hwy 38, 2010 m, 16 Sep 1977, JTS 77135, P. ponderosa, (apt.); same but 2 km S of jet of hwy 38 & Jenks Lake Rd, 2200 m, JTS 77139, P. jeffreyi, (apt.). SAN DIEGO Co.: 2 km E of Mt Palomar on hwy S6, 1650 m, 11 Sep 1977, JTS 77128, P. attenuata, (apt.); 5 km S of Julian, Harrison Springs Rd, 1460 m, 12 Sep 1977, JTS 77129, P. coulteri, (apt.); lake Cuyamaca, nr Cuyamaca State Park, 1800 m, 12 Sep 1977, JTS 77130, P. ponderosa, (apt.); Mt Palomar Rd (S6), 3 km S of Mt Palomar, 1530 m, 11 Sep 1977, JTS 77127, P. coulteri, (apt.). SHASTA Co.: 2 km W of Fall River Mills on hwy 299, 21 Jul 1978, JTS 78G123, P. ponderosa, (apt.); Hat Creek, 24 Jun 1955, E. O. Essig, P. ponderosa, (apt.). SISKIYOU Co.: Edson Creek access Rd, Shasta Natl Forest, 8 km W of Bartel on hwy 89, 1160 m, 3 Jul 1977, JTS 77G10, P. jeffreyi, (apt.). TUOLUMNE Co.: 2 km E of Groveville on hwy 120, 910 m, 30 Jul 1977, JTS 77G63, P. ponderosa, (apt.); Yosemite Natl Park, 17 May 1938, E. O. Essig, P. ponderosa, (fund.). VENTURA Co.: Mt Pinos Summit, 2680 m, 18 Sep 1977, JTS 77155, P. jeffreyi, (apt.); Reyes Peak Rd, 10 km E of Pine Summit on hwy 33, 2200 m, 19 Sep 1977, JTS 77159, P. jeffreyi, (apt.). COUNTY UNCERTAIN: Lake Tahoe, 16 Jul 1969, R. Luck, P. contorta murrayana, (apt.); same but 17 Jul 1969, P. jeffreyi, (apt.). NEVADA. CLARK Co.: Charleston Mts, Lee Canyon Ski Area, 2590 m, 4 Aug 1978, JTS 78H18, P. ponderosa, (apt.). OREGON. JACKSON Co.: 21 km S of Union Creek on hwy 62, 5 Jul 1978, JTS 78G15, P. ponderosa, (apt., alat.). JOSEPHINE Co.: Grant’s Pass, 2 Sep 1914, H.F.W., P. ponderosa, (apt.). LAKE Co.: 28 km N of Lakeview on hwy 395, 20 Jul 1978, JTS 78G119, P. ponderosa, (apt.). Essigella (Lambersella) hillerislambersi, NEW SPECIES Essigella “ hillerislambersi ” Sorensen, 1983: 99 (unpublished manuscript name) Ph.D. Thesis, University of California at Berkeley, Berkeley, California. 605 p. Type Series. — Holotype, vivip. apt.; on slide with 1 paratype vivip. apt., ho- lotype at bottom (6 o’clock position) and mounted inverted; data: CALIFORNIA. PLUMAS Co.: jet hwys 36 & 89, 1340 m, 19 Jul 1977, J. T. Sorensen (77G23), Pinus jeffreyi. Holotype deposited in The Natural History Museum, London. Para- types (all same data as holotype): 8 vivip. apt. on 6 slides including holotype slide; 4 paratype slides with 1 adult vivip. apt. and 1 nymph, only the adults are paratypes. Paratype slides deposited: 1 slide in NMNH, Washington, D.C.; 1 slide in CNC, Ottawa, Ontario; 3 slides in Sorensen collection. Viviparous Apterae.— Morphology: Body length: 2.10-2.64 (2.29 ± 0.14) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously protuberant. Length of antennal segment V: 113-145 (132 ± 8) p, processus terminalis: 35-45 (38 ± 3) p\ IV: 90-139 (116 ± 16) p; III: 183-230 (201 ± 13) p\ II: 83-90 (87 ± 2) p. Length of longest setae on frons: 48-88 (71 ± 10) p, tips incrassate to sharp. Head width: 265-357 (329 ± 20) p. Length of stylets: 622-969 (821 ± 81) p; ultimate rostral segment: 88-108 (101 ± 5) p , rostral tip reaching abdominal terga I—II in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 428-500 (470 ± 20) p. THORAX: Meso + metanota fused, total length: 400-510 (446 ± 32) p. ABDOMEN: Tergum I free, length: 158-204 (183 ± 14) m; terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 40-60 (49 ± 5) p\ siphunculi truncated conical, protruding 0.3- 0.6 x maximal distal width. Ventral abdominal sclerites on segments III-IV subquadrate to sublinear, often centrally constricted; moderate to large, length: 50-90 (67 ± 11) p, 1.2-2.3x diameter of metatibiae. Dorsal (major + minor) setae (see Fig. 1C) on abdominal terga III-IV: 8-11 (10 ± 1), tips sharp, in 2 rows with regular positions, lateral-most minor dorsal seta in anterad row; marginal setae 4-6 each side. Setae on abdominal tergum VIII: 8-11 (9 ± 1), length: 55-98 (70 ± 12) p, tips incrassate to sharp, in 1-2 rows. Cauda broadly rounded; caudal protuberance absent to poorly developed; length of longest caudal setae: 93-130 (108 ± \2) p, tips sharp. LEGS: Length of meta- 42 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) femora: 836-1142 (948 ± 87) /x; metatibiae: 1061-1561 (1276 ± 130) u', longest dorsal setae on central one-third of metatibiae: 60-113 (92 ± 16) n, 1.5-2.8x diameter of metatibiae, tips blunt to sharp; approximately equal or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae: 33-68 (49 ± 10) n, tips sharp. Length of metabasitarsus: 140-200 (168 ± 15) /z; metadistitarsus: 208-275 (240 ± 22) p. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.43:1. Pigmentation: Color in life: Body straw yellow, frequently with dark spots; head con- colorous to orange-brown; tibiae variable, entirely concolorous yellow, to pro- and metatibiae nearly black, mesotibiae yellow. Slide-mounted specimens: Background of body dorsum very pale to rarely light brown (usually less than 10, rarely to 30 percent pigment density), unicolorous. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga, to subtly darker. Thoracic muscle attachment plates light to moderate brown, often spotted, conspic¬ uous. Dorsal muscle attachment plates of abdomen pale to dark brown, conspicuous. Spiracular plates and ventral abdominal sclerites light to dark brown, conspicuous, rarely pale, inconspicuous. Siphun- culi concolorous with surrounding terga, to subtly darker. Cauda, anal and subgenital plates pale, concolorous with abdominal tergum, to substantially darker. Antennal segments V and IV dark brown, concolorous, to V and distal one-half of IV dusky; III entirely pale to distal one-third dark brown, remainder pale; II pale; I concolorous with frons, to subtly darker. Tibiae variable, pro- and metatibiae usually uniformly light to dark brown, often nearly black, mesotibiae pale; commonly all tibiae concolorously pale. Distal four-fifths of distitarsi dusky to nearly black with tibiae. Ultimate Stadium Nymphs of Viviparous Apterae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum with 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; plates usually distinct, faintly to darkly pigmented, diameter approx¬ imately equaling eye length. Viviparous Alatae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous ap¬ terae except lacking body dorsum pigmentation suite; abdominal terga normally membranous, dorsal (major + minor) setae between muscle attachment plates infrequently arising from distinct scleroites; antennal segments often as dark as tibiae, except proximal one-fifth of III pale. Antennal segment III with 0-2, IV with 0-1, secondary rhinaria. Epicranial suture absent. Forewing medius with single furcation arising on proximad one-third of vein; cubital base usually arising distad, infrequently proximad, on subcosta with distance between anal and cubital bases on subcosta usually relatively large, ca. 30-40 percent or more of anal vein length; usually medius, sometimes cubitus and anal veins faint, vague to absent. Abdominal terga infrequently with irregular sclerites that engulf or join muscle attachment plates and dorsal (major + minor) setal bases or scleroites. Oviparae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae ex¬ cept abdominal terga II-VI fused, lightly to moderately sclerotic, including pleural areas, but VII and VIII free (rarely VII not free); dorsal demarcations of anterad terga not evident; siphunculi usually incorporated into sclerotic dorsum, to free; dorsal abdominal muscle attachment plates unicolorous. Pseudorhinaria on metatibiae irregular, 5-19. Males, Fundatrices. — Unknown. Diagnosis.— Essigella ( L .) hillerislambersi and E. (L.)fusca are difficult to dis¬ tinguish. Darker individuals of both can be separated from other Essigella, except E. (L.) eastopi, by their tibial pigmentation suite [see E. ( L .) eastopi diagnosis]. Dark E. (L.) hillerislambersi usually have a paler background on the body dorsum than, and lack the developed body dorsum pigmentation of, E. (L.) eastopi. All three Lambersella species grade into completely pale individuals that can be differentiated from other pale Essigella by the diagnostics given for E. (L.) eastopi. Although E. (L.) hillerislambersi is a larger species, with an often longer antennal segment III, than either E. ( L .) fusca or E. ( L .) eastopi, overlapping morphometric variation in these species requires that reliable separation use the discriminant function in the key to the viviparous apterae [couplet 20]. Range. —California, southwestern Oregon, extreme western Nevada (Fig. 4B). Hosts. —Pinus jejfreyi Greville & Balfour; a single occurrence on P. attenuata 1994 SORENSEN: A REVISION OF ESSIGELLA 43 Lemmon is a questionable host determination; that sample (77142), which in¬ cluded oviparae, was from a host tree that was very small and lacked cones, but was among mature P. attenuata. Discussion. —Essigella ( L.) hillerislambersi is the largest Essigella, and is a rea¬ sonable common species. It is the most multivariately divergent within the sub¬ genus, and appears to be involved in a character displacement phenomenon with the other E. (Lambersella ) taxa in sympatry. See the discussions of E. (L.) eastopi, E. (L.) fusca fusca and E. (L.) fused voegtlini for its relationships, unrepeated here. Coded References to This Tdxon.—Essigelld (L.) hillerisldmbersi has been re¬ ferred to previously by: the coding “Sp. O” (Sorensen 1983 [but not section 2], 1987a, 1992b), “group A” (Sorensen 1983: section 2)and“HRL” (Sorensen 1983); and by the manuscript name E. “hillerisldmbersi ” in Sorensen (1983). Etymology dnd Common Nome. — This species is named for the aphidologist Dirk Hille Ris Lambers, who served as a mentor during my early aphid taxonomy studies. Common name: Hille Ris Lambers’ Jelfrey pine needle aphid. Material Examined.— CALIFORNIA. ALPINE Co.: Upper Cascade Creek, E side of Ebbett’s Pass on hwy 4, 5 km E of summit, 2350 m, 17 Jul 1977, JTS 77G40, P. jeffreyi, (apt.); W side of Monitor Pass on hwy 89, 2 km E of jet with hwy 4, 1830 m, 17 Jul 1977, JTS 77G38, P. jeffreyi, (apt.). EL DORADO Co.: Lake Tahoe, Meek’s Bay, 1980 m, 16 Jul 1977, JTS 77G29, P. jeffreyi, (apt.); South Lake Tahoe, 1950 m, 16 Jul 1977, JTS 77G32, P. jeffreyi, (apt.). INYO Co.: jet of Lake Sabrina Rd & Southern Calif. Edison Plant 2 Rd, nr Bishop, 2130 m, 1 Aug 1977, JTS 77H4, P. jeffreyi, (apt.). LOS ANGELES Co.: 3 km SE of Big Pines on hwy 2, E of Blue Ridge Summit, 2200 m, 17 Sep 1977, JTS 77149, P. jeffreyi, (apt., ovip.). MONO Co.: Deadman Summit on hwy 395, nr Crestview, 2440 m, 31 Jul 1977, JTS 77G72, P. jeffreyi, (apt., alat.); E side of Monitor Pass on hwy 89, 2070 m, 17 Jul 1977, JTS 77G37, P. jeffreyi, (apt.). NEVADA Co.: Prosser Lake Recreation Area, hwy 89, 25 Jun 1977, JTS 77F5, P. jeffreyi, (apt.). PLUMAS Co.: hwy 36, 8 km W of jet with hwy 89, 1460 m, 10 Jul 1977, JTS 77G25, P. jeffreyi, (apt.); (type series) jet of hwys 36 & 89, 1340 m, 10 Jul 1977, JTS 77G23, P. jeffreyi, (apt.). RIVERSIDE Co.: 2 km N of Paradise Valley on hwy 74, 1500 m, 9 Sep 1977, JTS 77118, P. jeffreyi, (apt.). SAN BERNARDINO Co.: San Bernardino Natl Forest, Heart Bar, 30 Aug 1972, D. J. Voegtlin, DJV 73, P. jeffreyi, (apt.); same but Keller Peak Cmpgd, 2200 m, 17 Sep 1977, JTS 77142, P. attenuata, (apt., ovip.); same but JTS 77141, P. jeffreyi, (apt., ovip.); San Bernardino Mts, 2 km S of jet of hwy 38 & Jenks Lake Rd, 2200 m, 16 Sep 1977, JTS 77139, P. jeffreyi, (apt.). SAN DIEGO Co.: Pioneer Mail Trail Picnic Area, Cleveland Natl Forest, 3 km N of Mt Laguna on hwy SI, 1740 m, 12 Sep 1977, JTS 77131, P. jeffreyi, (apt.). SIERRA Co.: 18 km S of Sierraville on hwy 89, 26 Jun 1977, JTS 77F7, P. jeffreyi, (apt.). TEHAMA Co.: 5 km E of Childs Meadows on hwy 89, 1460 m, 10 Jul 1977, JTS 77G21, P. jeffreyi, (apt.). TULARE Co.: E of Big Meadows Cmpgd, Sierra Natl Forest, 2320 m, 13 Aug 1977, JTS 77H13, P. jeffreyi, (apt., alat.). TUOLUMNE Co.: Yosemite Natl Park, nr Porcupine Rat-Porcupine Creek, 2500 m, 30 Jul 1977, JTS 77G67, P. jeffreyi, (apt., alat.). VENTURA Co.: Mt Pinos Summit, 2680 m, 18 Sep 1977, JTS 77155, P. jeffreyi, (apt.); Reyes Peak Rd, 10 km E of Pine Summit of hwy 33, 2200 m, 19 Sep 1977, JTS 77159, P. jeffreyi, (apt.). COUNTY UNCERTAIN: Eagle Peak, Stanislaus Natl Forest, 8 Jul 1979, D. J. Voegtlin, DJV 558, P. jeffreyi, (apt.); Lake Tahoe, 17 Jul 1969, R. Luck, P. jeffreyi, (apt.). NEVADA. ORMSBY Co.: E side of Spooner Summit on hwy 50, 1770 m, 16 Jul 1977, JTS 77G33, P. jeffreyi, (apt.). Essigella ( Essigella ) Del Guercio, 1909, NEW STATUS Ldchnus Burmeister, 1835 (in part), Handbuch der Entomologie, Berlin, 2: 91 (genus attributed to Illiger); Essig, 1909, Pomona J. Entomol., 1: 1-4. “Essigella (.Essigella )” Sorensen, 1983: 73 (unpublished manuscript name) Ph.D. Thesis, University of California at Berkeley, Berkeley, California. 605 p. Type Species. — Ldchnus cdlifornicus Essig, 1909, Pomona J. Entomol., 1: 1-4. 44 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Viviparous Apterae. —Morphology: Body relatively broad to narrow. Meso- and metanota fused dorsally; abdominal tergum I usually free but may be fused with meso + metanota, especially laterally. Dorsal setae (majors and minors) between muscle attachment plates on abdominal segments III-IV in 1 rarely irregular row with mesad (spinal) setae occasionally slightly anterad or posterad of others (see Figs. 1E-F); lateral-most dorsal minor seta on each side not anterad (very rarely so) of the immediately mesad neighbor. Abdominal terga III-IV each with 5-10, rarely to 12, dorsal (major + minor) and 2-5 (per side) marginal setae; tergum VIII with usually 6, frequently to 8, rarely to 10, setae. Longest dorsal seta on central one-third of metatibiae to nearly 4 x tibial diameter, tips incrassate to sharp; these setae not dimorphic in length but sometimes exceptionally variable among specimens, with nearly equal length, or gradually increasing, along metatibiae. Ventral abdominal sclerites on segments III-IV reduced (rudimentary), irregular stellate to large, subquadrate, subcircular or subel¬ liptical. Species means for length ratio of metadistitarsus to metabasitarsus varying between 1.47:1 to 1.69:1. Pigmentation: Body dorsum variable, pale to nearly black, unicolorous or variable, but not strongly mottled; bases of dorsal setae of abdomen concolorous with surrounding terga to substantially darker. Tibiae varying from entirely pale to nearly black; when darkened, all concolorous or pro- and mesotibiae paler than metatibiae. Diagnosis.— See the key to the subgenera of Essigella. Discussion. — This clade has ecologically transferred to diploxylon pines of the subgenus Pinus (Sorensen 1987a). The transfer is exclusive of subsection Pon- derosae pines, although E. (E.) californica, which is relatively polyphagous within Pinus, feeds on that subsection also. Some E. (.Essigella ) taxa have moved to Pinaceace hosts other than Pinus [i.e., E. (E.) wilsoni, E. (E.) alyeska]. Some have partially reinvaded haploxylon pines in the subgenus Strobus, through their rel¬ atively polyphagous feeding habits [i.e., E. (E.) californica, E. (E.) pini\. Others [i.e., E. ( E .) hoerneri ] have entirely reinvaded subgenus Strobus, in the unoccupied niches of section Parrya subsection Cembroides. Sorensen (1983) examined the relationships among the E. (Essigella ) species with discriminant function and principal component analyses, using morphometric data, and with principal co¬ ordinate analysis, multidimensional scaling and various UPGMA and single¬ linkage clustering algorithms, using coded quantitative and qualitative data. The phylogenetic analyses here indicate that Essigella (. Essigella ) can be divided into two series with historical biogeographic relevance: series A, which contains E. (E.) californica, E. (E.) essigi, E. (E.) hoerneri, E. (E.) pini, and E. (E.) wilsoni; and series B, which contains E. (E.) alyeska, E. (E.) critchfieldi, E. (E.) knowltoni braggi, and E. (E.) knowltoni knowltoni. Series A is paraphyletic, and shares hosts suspected of having an austral origin during the Tertiary [i.e., Madro-Tertiary geoflora] (Axelrod 1958, 1967; Raven & Axelrod 1978). Essigella (E.) californica, and E. (E.) hoerneri, however, clearly form a monophyletic species group within this series. Essigella ( E .) essigi is the least derived species in the subgenus, and its immediate ancestral node (Fig. 13: node 7) on the phylogenetic tree is shared by all other E. (Essigella ), making it the functional sister-group for the remainder of the subgenus. Series B is monophyletic, with a synapomorphy as a relatively broad head width [see discussion of E. ( E .) alyeska ]. Members of Series B occur on Pinus contorta Douglass ex Loudon, Pinus banksiana Lambert and Picea glauca (Moench) Voss; hosts that have relatively northern distributions in North America (Critchfield & Little 1966, Little 1971), and are of boreal origin during the Tertiary [i.e., Arcto- Tertiary geoflora]. Sorensen (1992a) has analyzed the biological groupings and host associations within the E. ( E .) knowltoni complex, and has found its species, subspecies and populations to closely overlay the geographic and terpene variance 1994 SORENSEN: A REVISION OF ESSIGELLA 45 in Pinus contorta and its subspecies [see the discussions under E. (E.) critchfieldi and E. (E.) knowltoni]. He also notes the presence of character displacement in both qualitative and multivariate quantitative traits among the taxa in this species group. Note that Sorensen (1983) reversed the letters for series A and B, as manuscript references; in that unpublished work, series A and B were paraphyletic and mono- phyletic groups, respectively. Coded References to This Taxon. —Sorensen (1983) referred to this taxon under the manuscript name “ Essigella (. Essigella ).” Sorensen (1987a) referred to the assemblage that comprise this taxon as group “III” or, with reference to its sub¬ components, as “I-H-B-A-C-D-E-F-G” [or entire permutations therefore]; in Sor¬ ensen (1992b), the latter refers to it. Material Examined.—Essigella (E.) alyeska, E. (E.) calif or nica, E. (E.) critchfieldi, E. (E.) essigi, E. (E.) hoerneri, E. ( E .) knowltoni braggi, E. (E.) knowltoni knowltoni, E. (E.) pini, E. (E.) wilsoni. Series A Essigella ( Essigella) essigi Hottes, 1957 Essigella essigi Hottes, 1957: 84, Proc. Biol. Soc. Wash., 70: 84-85. Primary Type. — Holotype, vivip. alat., on slide with 5 other alat. and 7 apt., holotype shown by arrow (near center position among all specimens, 9 o’clock among alat.); slide data: “ Pinus radiata, Redwood City, California, June 10, 1939, L. Blanc/Paratype, Essigella holotype essigi F. C. Hottes, Essig.” (Redwood City is in San Mateo Co.). Holotype deposited in the Essig Museum of Entomology, University of California at Berkeley, Berkeley, California. Viviparous Apterae.— Morphology: Body length: 1.33-1.93 (1.62 ± 0.18) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously protuberant. Length of antennal segment V: 90-125 (109 ± 11) p, processus terminalis: 23-43 (36 ± 5) m; IV: 60-93 (78 ± 10) p\ III: 110-153 (133 ± 14) p- II: 56-68 (63 ± 4) p. Length of longest setae on frons: 13-65 (29 ± 13) p, tips incrassate. Head width: 228-275 (249 ± 13) p. Length of stylets: 541-755 (621 ± 56) p\ ultimate rostral segment: 50-78 (63 ± 7) p, rostral tip reaching abdominal terga I—III in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 245-388 (324 ± 39) p. THORAX: Meso + metanota fused, combined total length when dorsally demarcated from abdominal tergum I: 250-377 (312 ± 36) p. ABDOMEN: Tergum I fused with metanotum, completely so across dorsum (pale individuals) to fused laterally only (dark individuals), length when dorsally demarcated: 92-143 (123 ± 19) p; terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 15-29 (21 ±4) p\ siphunculi flush with tergum. Ventral abdominal sclerites on segments III-IV subquadrate, subcircular to subelliptical; length: 35-60 (50 ± 7) p, 1.1- 2.0 x diameter of metatibiae. Dorsal (major + minor) setae (see Fig. IE) on abdominal terga III-IV: 8, very rarely 7 or 9, tips sharp, in 1 row, infrequently with mesad pair of setae posterad; marginal setae 3, infrequently 2, per segment each side. Setae on abdominal tergum VIII: 6 to rarely 8, length: 5-48 (23 ± 14) p, tips incrassate to sharp, in 1 row. Cauda rounded to broadly rounded; caudal protuberance moderately developed to nearly absent; length of longest caudal setae: 20-92 (60 ± 19) p, tips sharp. LEGS: Length of metafemora: 367-581 (476 ± 66) p\ metatibiae: 418-694 (567 ± 87) p; longest dorsal setae on central one-third of metatibiae: 3-43 (18 ± 11) p, 0.1-1.3 x diameter of metatibiae, tips incrassate; approximately equal or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae: 5-35 (24 ± 9) p, tips sharp. Length of metabasitarsus: 65-98 (85 ± 11) p\ metadistitarsus: 118—163 (144 ± 15) p. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.69:1. Pigmentation: Color in life: Black to green throughout or green with 46 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Figure 6. Distribution of E. ( E .) essigi [dots (JTS samples), squares (nonJTS samples)], superim¬ posed over the range of its hosts, Pinus attenuata [lighter shading] and Pinus radiata [darker shading (coastal Santa Cruz, Monterey and San Luis Obispo Counties)]. yellow-green head; frequently with dark dorsal spots when body not dark. Slide-mounted specimens: Background of body dorsum pale to dark brown or nearly black (to nearly 100 percent pigment density), unicolorous. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga, to subtly darker. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen pale, inconspicuous to dark brown, conspicuous. Spiracular plates and ventral abdominal sclerites pale to nearly black. Siphunculi concolorous with surrounding terga. Cauda, anal and subgenital plates subtly to substantially darker than abdominal terga. Antennal segments V and IV dusky to moderate brown, concolorous; III entirely pale to dusky on distal one- third, remainder pale; II concolorous with proximal III, to subtly darker; I concolorous with frons. Pro-, meso- and metatibiae concolorous, and usually equivalent to abdominal terga, sometimes subtly lighter (dark individuals), rarely darker. Distitarsi evenly moderate brown, sometimes subtly paler at proximal tip. Ultimate Stadium Nymphs of Viviparous Apterae. - Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga 1994 SORENSEN: A REVISION OF ESSIGELLA 47 membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites; meso- and metathorax not fused. Mesonotum with 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; plates usually heavily, to faintly, pigmented, diameter approximately equaling eye length. Viviparous Alatae. -Slide-mounted specimens: Nonmorphometrics as described for viviparous ap- terae except lacking body dorsum pigmentation suite, and meso- and metathorax not fused; abdominal terga normally membranous, dorsal (major + minor) setae between muscle attachment plates fre¬ quently arising from distinct scleroites; antennal segments often as dark as tibiae, except proximal one-fifth of III pale. Antennal segment III with 2-3, IV with 0-1, secondary rhinaria. Epicranial suture usually strongly developed, to absent. Forewing medius usually single, infrequently single furcation arising on distad one-third of vein; cubital base usually arising distad, infrequently proximad, on subcosta with distance between anal and cubital bases on subcosta usually relatively large, ca. 30-40 percent or more of anal vein length; medius, especially cubitus and anal veins usually distinct, except infrequently proximad 10-15 percent vague. Abdominal terga infrequently with irregular sclerites that engulf or join muscle attachment plates and dorsal (major + minor) setal bases or scleroites. Oviparae, Males, Fundatrices.— Unknown. Diagnosis. — Adult viviparous apterae of E. (E.) essigi can be identified by the unique additional fusion of abdominal tergum I to fused meso + metanota. They vary from entirely pale to dark brown [like E. ( E.) critchfleldi and some E. ( E .) knowltoni knowltoni], and the fusion of abdominal tergum I varies inversely with pigmentation: it extends entirely across the dorsum in pale individuals, but is restricted to the lateral edges of the dorsum in dark specimens. The forewing medius of alates usually is single, or occasionally 1-branched with the furcation distad and closer to the posterad margin of the wing than to the subcosta. This alate character is similar in E. (E.) pini [see that diagnosis] and potentially can be confused in E. (E.) knowltoni knowltoni and E. (E.) alyeska. Range. —California and southwestern Oregon (Fig. 6). Hosts.—Pinus radiata D. Don and P. attenuata Lemmon, both subsection Oocarpae pines that hybridize (W. Libby, personal communication). Although E. (E.) essigi is commonly found on closed-cone pines, it has not been found on P. muricata D. Don, despite extensive collecting; its appearance on that pine would not be surprising, however. Also, one collection (77G12) exists from P. ponderosa, at a site about 50 mi east, by air, of the nearest stand of P. attenuata, near the southern Modoc-Siskiyou county border (Critchfield & Little 1966: map 58) in northwest California. Another collection (7718) exists from P. sabiniana, possibly a contaminant, at the same location and immediately after a collection (7717) from P. attenuata. I suspect that the locations where E. (E.) essigi can be found in California have probably increased substantially in recent years because of the extensive landscape planting of Pinus radiata. Discussion.—Essigella (E.) essigi is relatively homogeneous in morphology, although specimens from southwest Oregon and the northern Californian coast tend to be very slightly more linear than those near the San Francisco Bay area and south. The autapomorphic fusion of abdominal tergum I in this species is incomplete dorsally on darker specimens, probably due to an increased scleroti- zation of the body dorsum; the break may be necessary for articulation (?) of the more rigid tergum on those individuals. Subtle differences between coastal pop¬ ulations on P. radiata, and inland populations on P. attenuata should be examined in more detail with isozyme or nucleic acid techniques. Hottes (1957: 85) stated “I am sure that specimens of this species [E. (E.) essigi ] were part of the original material from which Essig described Lachnus californicus, 48 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) although there is no actual proof in the remaining cotype material.” He based this supposition on Essig’s illustration (Essig 1909: fig. 2), which failed to indicate long tibial setae. Unfortunately, Hotte’s reliance upon length of the dorsal setae on the tibia was entirely misguided. I have found no evidence to support his supposition. The phylogenetic position of E. ( E.) essigi within the subgenus is problematic. Clustering analyses on qualitative coded characters (Sorensen 1983) suggest it is probably primitive within E. {Essigella) because its similarities to other taxa are mostly plesiomorphies at the level of the subgenus. Ordinations on morphometric data indicate that it is close to E. ( E .) pini and E. (E.) wilsoni in ordinant space; the evolutionary aspects of this are discussed in Sorensen (1992b). Cladistic anal¬ ysis of coded data (unpublished data) suggest that E. (E.) essigi is either para- phyletically one of the more primitive E. (Essigella ), along with E. (E.) pini, or that it potentially forms a monophyletic subgroup, which is the sister-group to the remainder of the subgenus, with E. (E.) pini. Intuitively, I consider E. (E.) essigi to be closest to E. ( E .) pini, but I am uncertain of the exact relationship between them. Both share reduction of the alate medius to a single vein, but I am reluctant to accept the trait as a convincing, nonhomo- plasious synapomorphy. I also suspect the genetic compatibility of subsection Oocarpae and Australes pines [the latter chiefly hosts E. (E.) pini ] suggests a potentially common biogeographic origin; see the section on ecological collabo¬ ration of phylogenetic hypotheses for comments. Coded References to This Taxon. —Essigella (E.) essigi has been referred to previously by: the coding “Sp. I” (Sorensen 1983, 1987, 1992b) and “ESSG” (Sorensen 1983), and by the name E. essigi in Sorensen (1983). Etymology and Common Name. — Hottes (1957) apparently named this species for the aphidologist E. O. Essig, presumably because he described the first Essi¬ gella, E. ( E.) calif or nica, albeit as a Lachnus. Common name: Essig’s closed-cone pine needle aphid [see etymology for E. (E.) californica ]. Material Examined.— CALIFORNIA. ALAMEDA Co.: Berkeley, 10 Nov 1935, E. O. Essig, P. radiata, (apt.); same but 15 Feb 1982, J. T. Sorensen, P. radiata, (apt.). DEL NORTE Co.: 16 air km NWW of Crescent City on hwy 199, Six Rivers Natl Forest, 4 Jul 1978, JTS 78G6, P. attenuata, (apt., alat.). LAKE Co.: 21 km N of Upper Lake, Elk Mt Rd, 1030 m, 24 Jul 1977, JTS 77G60, P. attenuata, (apt., alat.); same but 23 km N of Upper Lake, 1160 m, JTS 77G59, (apt.); same but 26 km N of Upper Lake, JTS 77G58, (apt.). MODOC Co.: E side of Cedar Pass, 29 km E of Alturas, 1890 m, 3 Jul 1977, JTS 77G12, P. ponderosa, (apt.). MONTEREY Co.: 3 km N of Point Lobos State Park Reserve on hwy 1, 29 Dec 1978, JTS 78L1, P. radiata, (apt.); Carmel, 16 Jun 1973, D. J. Voegtlin, DJV 25, P. radiata, (apt.); Lockwood-San Ardo Rd, 13 km SW of jet with Paris Valley Rd, 550 m, 4 Sep 1977, JTS 7717, P. attenuata, (apt.); same but JTS 7718, P. sabiniana, (apt.); Monterey, 18 Jun 1973, D. J. Voegtlin, DJV 24, P. radiata, (apt.); same but 19 Feb 1974, T. Kono, CDFA 79B20-10- 2, (apt.). NAPA Co.: 3 km N of Angwin, Howell Mt Rd, 4 Feb 1978, JTS 79B2, P. attenuata, (apt.). PLACER Co.: 6 km W of Dutch Flat on hwy 80, 2 Aug 1978, JTS 78H2, P. attenuata, (apt.). SAN LUIS OBISPO Co.: Cambria Pines, 5 Sep 1977, JTS 77112, P. radiata, (apt.). SAN MATEO Co.: (paratype) Redwood City, 10 Jun 1939, L. Blanc, P. radiata, (apt., alat.). SANTA CLARA Co.: (paratype) Palo Alto, Stanford Univ., 30 Mar 1938, E. O. Essig, P. radiata, (apt.,); (paratype) same but 25 Apr 1930, P.S.B., (apt.); Morgan Hill, 14 Oct 1942, Bell, CDFA 42J10, P. attenuata, (apt.). SISKIYOU Co.: Snowman Hill Summit on hwy 89, 8 km E of jet with hwy 5, 1360 m, 2 Jul 1977, JTS 77G3, P. attenuata, (apt.). TRINITY Co.: Junction City, 11 km of Weaverville on hwy 299, 430 m, 20 Aug 1977, JTS 77H23, P. attenuata, (apt.). OREGON. JOSEPHINE Co.: 2 km N of O’brien on hwy 199, 4 Jul 1978, JTS 78G11, P. attenuata, (apt.). 1994 SORENSEN: A REVISION OF ESSIGELLA 49 Essigella ( Essigella) pini Wilson, 1919 Essigella pini Wilson, 1919: 2, Entomol. News, 30: 2-3. Essigella patchae Hottes, 1957: 98, Proc. Biol. Soc. Wash., 70: 98-100. NEW SYNONYM. Primary Type. — Lectotype, vivip. alat., on slide alone; data: “(Lectotype)/82- 14/11, Essigella pini Wilson, Cotype, Pinus virginiana, Plummer’s Is., Md., May 27, 1914.” Lectotype deposited in the Granovsky Collection, Department of Entomology, Fisheries & Wildlife, University of Minnesota, St. Paul, Minnesota. Hottes (1957: 103) designated a vivip. alat., from the Granovsky collection as lectotype. A slide that is so labeled does exist, although I doubt that Hottes personally labeled it because the printing does not match his, and he did not label other lectotypes in this genus; I have seen this specimen and consider it the lectotype. Viviparous Apterae.— Morphology: Body length: 1.57-2.03 (1.79 ± 0.15) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously protuberant. Length of antennal segment V: 98-133 (115 ± 9) p, processus terminalis: 25-38 (31 ± 4) m; IV: 73-100 (84 ± 8) HI: 110-158 (136 ± 16) p\ II: 63-79 (68 ± 4) p. Length of longest setae on frons: 15-73 (29 ± 16) p, tips incrassate. Head width: 245-316 (276 ± 19) p. Length of stylets: 541-663 (572 ± 34) p\ ultimate rostral segment: 55-75 (67 ± 5) p, rostral tip reaching abdominal terga I—II in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 326-454 (400 ± 36 ) /x. THORAX: Meso + metanota fused, total length: 265-367 (327 ± 33) p. ABDOMEN: Tergum I free, length: 112-163 (134 ± 15) p\ terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 15-35 (28 ± 5) p; siphunculi usually flush to slightly protruding to 0.3 x maximal distal width. Ventral abdominal sclerites on segments III-IV usually subcircular, subquadrate to subelliptical, sometimes irregular, asterisk-shaped, or constricted anteriorly; length: 40-75 (54 ± 8) p, 1.0-2.Ox diameter of metatibiae. Dorsal (major + minor) setae (see Fig. IF) on abdominal terga III-IV: 6, infrequently 7, tips sharp, in 1 row; marginal setae 2 per segment, each side. Setae on abdominal tergum VIII: 6, length: 8-25 (13 ± 5) p, tips incrassate to sharp, in 1 row. Cauda rounded; caudal protuberance well developed, often pointed, to moderately developed; length of longest caudal setae: 43-120 (75 ± 20) p, tips sharp. LEGS: Length of metafemora: 398-581 (486 ± 57) p ; metatibiae: 561-831 (675 ± 79) p\ longest dorsal setae on central one-third of metatibiae: 11-33 (17 ± 6) p, 0.3-1.Ox diameter of metatibiae, tips usually incrassate, infrequently sharp; ap¬ proximately equal or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae: 23-68 (35 ± 11) p, tips sharp. Length of metabasitarsus: 79-103 (89 ± 6) p\ metadistitarsus: 138-163 (147 ± 9) p. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.65:1. Pigmentation: Color in life: Green with yellow-orange to red-orange head (from notes on C. F. Smith slides). Slide-mounted specimens: Background of body dorsum pale (to 10 percent pigment density), unicolorous. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen pale, inconspicuous. Spiracular plates and ventral abdominal sclerites pale, to moderate brown, conspicuous. Siphunculi concolorous with surrounding terga. Cauda, anal and subgenital plates pale, concolorous with, to subtly dusky and slightly darker than abdominal terga. Antennal segments V and IV pale to light brown, concolorous; III entirely pale to distal one-third concolorous with V and IV; II concolorous with proximal III; I concolorous with frons. Pro-, meso- and metatibiae usually pale, concolorous and equivalent to body dorsum, infrequently entire tibiae slightly dusky, subtly darker than dorsum. Distitarsi entirely pale to subtly dusky on distal one-third. Ultimate Stadium Nymphs of Viviparous Apterae. — Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum with, very rarely lacking, 2 sclerotized plates extending from muscle 50 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) attachment sites to engulf neighboring setal bases; plates faintly to heavily pigmented, diameter ap¬ proximately equaling eye length. Viviparous Alatae. -Slide-mounted specimens: Nonmorphometrics as described for viviparous ap- terae except lacking body dorsum pigmentation suite; abdominal terga normally membranous, dorsal (major + minor) setae between muscle attachment plates occasionally arising from distinct scleroites; antennal segments often as dark as tibiae, except proximal one-fifth of III pale. Antennal segment III with 2-3, IV with 0, secondary rhinaria. Epicranial suture vaguely to strongly developed. Forewing medius usually single, infrequently single furcation arising on distad one-third of vein; cubital base usually arising distad on subcosta with distance between anal and cubital bases on subcosta usually relatively large, ca. 30-40 percent or more of anal vein length; medius, especially cubitus and anal veins distinct, except infrequently proximad 10-15 percent vague. Abdominal terga infrequently with irregular sclerites that engulf or join muscle attachment plates and dorsal (major + minor) setal bases or scleroites. Oviparae. -Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae, ab¬ dominal terga II-VII fused, moderately sclerotic, including pleural areas, tergum VIII free (uncom¬ monly tergum VII free, demarcations of II-VI evident laterally); dorsal demarcations of anterad terga sometimes evident; siphunculi incorporated into sclerotic field, to free; dorsal abdominal muscle attachment plates unicolorous. Pseudorhinaria on metatibiae irregular, 8-15. Males, Funda.trices. — Unknown. Diagnosis. — This species is pale and may be confused with other pale Essigella-, it requires the combination of several characters for identification. Essigella (E.) pini can be separated from all Essigella, except E. (E.) californica and E. (E.) hoerneri, by having six (Fig. IF), instead or eight or more, dorsal (major + minor) setae on abdominal terga III-IV. It differs from E. (E.) calif ornica and E. ( E .) hoerneri by having usually large subcircular-subquadrate, rather than usually small irregular, ventral abdominal sclerites on abdominal segments III-IV, and by hav¬ ing usually large invasive, instead of small noninvasive, mesonotal muscle at¬ tachment plates on later stadia nymphs of apterae. The mean lengths of antennal segments III and the metatibiae are shorter, but overlapping, in proportion to body length in E. (E.) pini than in either E. (E.) calif ornica or E. ( E .) hoerneri. The caudal protuberance of E. ( E .) pini also is often abnormally long and pointed, but the trait is not an entirely satisfactory discriminator. Alates of E. ( E.) pini differ from other species, except E. (E.) essigi, in having the medius usually single, or infrequently 1-branched with the furcation exceptionally distad; but E. (E.) knowltoni knowltoni has also rarely shown this condition. Although the relative stability of the trend to a single medius for E. {E.) pini and E. (E.) essigi seems a reasonable partial diagnostic for those species, unknown alate morphs and interspecific variance in several Essigella species make discriminatory use of venation potentially questionable. Synonyms. —Essigella patchae Hottes, NEW SYNONYM: holotype, vivip. alat., as a single fragmented specimen on the slide; data: MAINE. PENOBSCOT Co.: Stillwater, 4 Jul 1909, Pinus strobus L. Essigella patchae holotype deposited in the NMNH. Range. —Eastern U.S.; one record from southern Quebec (Fig. 7). Hosts. —Notably Pinus virginiana P. Miller, P. taeda L., P. strobus L.; presum¬ ably many species of Pinus subsection Australes; subsection Sylvestres pines also recorded as hosts. Note that because E. (E.) pini is the only Essigella that I have not personally collected, during extensive sampling of the western Nearctic (Sor¬ ensen 1983), I cannot attest to the accuracy of determination of its hosts, as with other Essigella. Indeed, because the aphid occurs only in the eastern U.S., and it 1994 SORENSEN: A REVISION OF ESSIGELLA 51 Figure 7. Distribution of E. ( E .) pini [squares (nonJTS samples)], superimposed over the ranges of its hosts, Pinus strobus [darker + moderate shading] and an amalgamation of subsection Australes plus eastern subsection Contortae pines [lighter + moderate shading] (moderate shading indicates the distributional overlap of hosts). could not often be successfully identified until now (see discussion), many western pine species attributed as its hosts are in error. Discussion.—Essigella ( E .) pini, known from the eastern Nearctic only, has been the most confused Essigella with regard to misidentifications. This is, no doubt, due in large part to Hottes’ (1957) erroneous key. Using that key, it is conceivable that at least some individuals (albeit, small and pale in several in¬ stances) of all species, except E. ( L .) hillerislambersi, might key out to Hottes’ “is. pint” References to Essigella “pini” from the western U.S. and Canada (i.e., Knowlton 1930, Gillette & Palmer 1931, Palmer 1952, Smith & Parron 1978) are in error and clearly do not represent that species. I cannot determine exactly to what they correctly refer, because adequate diagnostics are not mentioned. Such references evoke potential confusion with E. ( E .) californica, E. (is.) hoerneri, E. (is.) wilsoni, E. (L.) fusca fusca, E. (is.) knowltoni knowltoni and E. (A.) kirki, due to the geography involved and the earlier erroneous diagnostic fixation on the length of the dorsal setae of the metatibiae. A single alate from Quebec, captured in a 52 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Malaise trap (F. W. Quednau, personal communication), appears to be the only Canadian record of this species. Patti & Fox (1981a, b) discuss the seasonal occurrence and intratree distribution of E. (E .) pini in South Carolina. Essigella ( E .) pini appears to be relatively homogeneous morphologically. Be¬ cause it was not extensively sampled over its range during this study, however, knowledge of its variation is drawn only from collections by others, which have relatively limited geographic spread and continuity. It shows infrequent variability in reduction of the ventral abdominal sclerites on segments III-IV, but this trait is paralleled within other species. Placement of E. (E.) pini within the subgenus was confusing during analysis; for comments, see the discussion of E. ( E .) essigi. It shows several qualitative loss-reduction apomorphies that are considered to be homoplasious with E. (E.) californica and E. (E.) hoerneri, and a few others with E. ( E .) wilsoni and E. ( E.) alyeska. The fusion of abdominal terga on its oviparae is also problematic [see character discussion section]. Coded References to This Taxon. —Essigella (E.) pini has been referred to pre¬ viously by: the coding “Sp. H” (Sorensen 1983, 1987a, 1992b) and “PINI” (Sor¬ ensen 1983), and by the name E. pini in Sorensen (1983). Etymology and Common Name. —Wilson (1919) presumably named this spe¬ cies with reference to the host genus, Pinus. Common name: the eastern pine needle aphid; although Palmer (1952: 16) refers to this species as “The Speckled Pine Needle Aphid,” the common name indicated here is more appropriate and less confusing because many Essigella are speckled. Material Examined. — ALABAMA. BARBOUR Co.: Eufaula, 10 Feb 1876, Pinus sp., (apt.). FLOR¬ IDA. ALACHUA Co.: Gainesville, 23 Mar 1928, A. Tissot, ANT F323-28, P. taeda, (apt., alat.); same but no date, ANT F886-32, (apt., alat.). COLUMBIA Co.: 22 Mar 1973, G. Hertel, P. elliottii En- gelmann, (apt.); 26 May 1972, G. Hertel, P. elliottii, (alat.). SAINT JOHNS Co.: St. Augustine, 10 May 1945, D. & B. Darry, P. taeda, (apt.); same but 24 May 1945, D. & B. Darry, (alat.). SEMINOLE Co.: Sanford, 23 Feb 1929, A. Tissot, ANT F480-29, P. taeda, (alat.). MAINE. PENOBSCOT Co.: Stillwater, 4 Jul 1909, E. M. Patch, MAES 46-09, P. strobus, (alat.). MARYLAND. BALTIMORE Co.: Sheppard Pratt, 3 Aug 1974, A. Scarbough, (alat.). PRINCE GEORGES Co.: Beltsville, 19 Jun 1978, W of Cantelo, yellow pan trap, (alat.). COUNTY UNCERTAIN: (lectoype) Plummer’s Island, 27 Apr 1914, P. virginiana, (apt., alat.). NORTH CAROLINA. ALLEGANY Co.: Gladesville, 17 Jun 1959, D. A. Young, Pinus sp., (apt.). BANCOMBE Co.: Twin Tunnels, (Blue Ridge) Parkway, 29 Jul 1958, C. F. Smith, CFS 58-347, Pinus sp., (apt.). CHEROKEE Co.: N of Andrews, 24 Jul 1958, C. F. Smith, CFS 58-309, Pinus sp., (apt., alat.). DURHAM Co.: Durham, 10 Jan 1978, D. Whitman, P. lambertiana, (apt.); same but 22 Oct 1959, S.S.T.,P. taeda, (apt.); same but 8 Jan 1979, J. Richmond, (apt.). MACON Co.: Highlands, Mt Satulah, 29 Sep 1970, C. F. Smith C. S. Smith & C. Sullivan, P. rigida P. Miller (alat.). MOORE Co.: West End, 30 Oct 1958, S.S.T., Pinus sp., (apt., alat.). RICH¬ MOND Co.: Norman, 30 Oct 1958, S.S.T., Pinus sp., (apt.). WAKE Co.: McCullers, 18 May 1967, C. F. Smith, CFS 67-28b, P. taeda, (apt.); Umstead Park, 30 May 1960, C. F. Smith, CFS 60-303, P. taeda, (apt., alat.). WASHINGTON Co.: Roper, 10 Feb 1975, C. G. Livingston, P. taeda, (apt.). WILKES Co.: McGrady, 14 Oct 1963, C. F. & C. S. Smith, CFS 63-166, (alat.). YANCEY Co.: 2.4 km (1.5 mi) E of Mt Mitchell State Park entrance, 23 Jul 1970, G. Fedde, P. pungens Lambert (apt., alat.). Crabtree Meadows, (Blue Ridge) Parkway, 12 Oct 1958, S.O.T., P. strobus, (apt., ovip.); same but Pinus sp., (apt.). OKLAHOMA. LATIMER Co.: Robber’s Cave State Park, 19 Sep 1957, Van Cleave, P. echinata P. Miller (apt., alat.). McCURTAIN Co.: Broken Bow, 13 Sep 1960, Van Cleave, P. echinata, (apt.). PENNSYLVANIA. CENTRE Co.: State College, 3 Oct 1959, J. Pepper, P. sylvestris L., P. resinosa Aiton, P. strobus, (apt., alat.). SOUTH CAROLINA. OCONEE Co.: Seneca, 26 May 1962, R. Eikenbarry, Pinus sp., (apt.). PICKENS Co.: Clemson, 4 Apr 1973, K. Griffith, Pinus sp., (apt., alat.); same but 6 Jun 1977, W of Hood, P. taeda, (alat.). VIRGINIA. MONTGOMERY Co.: Blacksburg, 6 Feb 1967, W. A. Allen, P. taeda, (apt.). CANADA. QUEBEC: Mt St. Hilaire, Cte Rouville, Meteo, 5 Jul 1979, R. Roy, Malaise trap (alat.). 1994 SORENSEN: A REVISION OF ESSIGELLA 53 Essigella ( Essigella) californica (Essig), 1909 Lachnus californicus Essig, 1909: 1, Pomona J. Entomol., 1: 1-4. Essigella californica Del Guercio, 1909: 328, Rivista di Patologia Vegetale, Anno III Num. 20-21: 328-329. Essigella claremontiana Hottes, 1957: 79, Proc. Biol. Soc. Wash., 70: 79-81. NEW SYNONYM. Essigella cocheta Hottes, 1957: 82, Proc. Biol. Soc. Wash., 70: 82-84. NEW SYNONYM. Essigella monelli Hottes, 1957: 95, Proc. Biol. Soc. Wash., 70: 95-96. NEW SYNONYM. Essigella pineti Hottes, 1957: 101, Proc. Biol. Soc. Wash., 70: 101-102. NEW SYNONYM. Essigella swaini Hottes, 1957: 105, Proc. Biol. Soc. Wash., 70: 105-106. NEW SYNONYM. Primary Type. — Lectotype, vivip. alat., on slide alone; data: “Monterey pine, Claremont, Cal., Feb. 14, 1909, E.O.E./type/Cotype, Lachnus californicus Essig, Essig/[on back] Lectotype of Hottes, J. T. Sorensen ‘82” (Claremont is in Los Angeles Co.). Lectotype deposited in the Essig Museum of Entomology, University of California at Berkeley, Berkeley, California. The extent of the original type series is somewhat confused; but the [1909] series must have involved an ovipara (see discussion). The series was scattered and possibly adulterated by the addition of material collected in 1911 (“47” slides), which may be involved with Essig’s (1912) redescription (see Hottes 1957: 78). Hottes (1957: 78) mentions a lectotype and describes the slide as thick, but the apparent slide merely bears a small label stating “type” below the coverslip. The label is asymmetrically placed, and could have had a prefix “lecto” removed. There are also, however, two other slides, deposited in the NMNH, bearing “type” in blue ink on the right-hand label that are part of the 1911 “47” series. I deduced what must be Hottes’ lectotype based upon his description of the slide and its location, and I have labeled the slide as lectotype. Viviparous Apterae.— Morphology: Body length: 1.30-2.38 (1.90 ± 0.24) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously protuberant. Length of antennal segment V: 103-155 (128 ± 11) /x, processus terminalis: 30-48 (37 ± 5) p\ IV: 88-138 (109 ± 14) p; III: 135-250 (194 ± 31) u; II: 63-88 (77 ± 8) p. Length of longest setae on frons: 6-73 (39 ± 18) p, tips incrassate to sharp. Head width: 214-347 (270 ± 26) p. Length of stylets: 551-806 (688 ± 71) /u.; ultimate rostral segment: 70-98 (83 ± 8) p, rostral tip reaching abdominal terga I—II, occasionally III, in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 316-510 (393 ± 42) p. THORAX: Meso + metanota fused, total length: 245-571 (365 ± 57) p. ABDOMEN: Tergum I free, length: 82-194 (131 ± 23) p; terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 25-45 (36 ± 5) p\ siphunculi flush to truncated conical, protrusion to 0.7 x maximal distal width. Ventral abdominal sclerites on segments III-IV usually irregular, to subcircular when small (length less than 0.6 x metatibial diameter), subquadrate when large (length greater than 1.0 x metatibial diameter); length: 8-63 (23 ± 14) p, 0.3-1 . 1 x diameter of metatibiae. Dorsal (major + minor) setae (see Fig. IF) on abdominal terga III-IV: 6, rarely 7, tips incrassate to sharp, in 1 row; marginal setae 2 each side. Setae on abdominal tergum VIII: 6, occasionally 7, infrequently 8, length: 8-85 (36 ± 20) p, tips incrassate to sharp, in 1 row. Cauda rounded; caudal protuberance moderately to poorly developed, sometimes absent; length of longest caudal setae: 48- 103 (75 ± 12) p, tips sharp. LEGS: Length of metafemora: 469-938 (718 ± 126) p\ metatibiae: 653- 54 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) 1397 (1042 ± 195) fx; longest dorsal setae on central one-third of metatibiae: 8-118 (58 ± 33) n, tips 0.1-3.0 x diameter of metatibiae, tips incrassate to sharp; approximately equal or very gradually increasing distally, no setal length dimorphism, but very rarely with dorsal setae breaking on proximad one-third of metatibiae to a ca. 30-50 percent increase in length; longest ventral setae on metatibiae: 14-73 (35 ± 14) n, tips sharp. Length of metabasitarsus: 84-148 (122 ± 16) u; metadistitarsus: 148— 230(191 ± 20) /i. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.57:1. Pigmentation: Color in life: Thorax gray-green, abdomen lime green, legs light to dark brown, or straw yellow throughout; dorsal spots absent to brown. Slide-mounted specimens: Background of body dorsum pale to moderate brown (usually to 20, infrequently to 40 percent pigment density), unicolorous. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga, to conspicuously darker. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen, spiracular plates and ventral abdominal sclerites pale, inconspicuous, to dark brown, conspicuous. Siphunculi concolorous with surrounding terga, to substantially darker. Cauda, anal and subgenital plates concolorous with abdominal terga, to substantially darker. Antennal segments V and IY subtly dusky distally, pale proximally, to entirely dusky; III entirely very pale to subtly dusky on distal one-third, remainder pale; II concolorous with proximal III, to subtly darker; I concolorous with frons. Pro-, meso- and metatibiae usually concolorous, evenly pale or pale with subtly dusky distal tip, to evenly dark brown; or metatibiae subtly to substantially darker than pro- and mesotibiae. Distitarsi dusky on distal one-third, pale proximally, to evenly dark brown with tibiae. Ultimate Stadium Nymphs of Viviparous Apterae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum lacking 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; area surrounding muscle attachment sites membranous. Viviparous Alatae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous ap¬ terae except lacking body dorsum pigmentation suite; abdominal terga normally membranous, dorsal (major + minor) setae between muscle attachment plates occasionally arising from distinct scleroites; antennal segments often as dark as tibiae, except proximal one-fifth of III pale. Antennal segment III with 0-5, IV with 0-3, secondary rhinaria. Epicranial suture strongly developed. Forewing medius with single furcation arising on proximad, very rarely on central, one-third of vein; cubital base arising proximad, rarely distad, on subcosta with distance between anal and cubital bases on subcosta usually relatively small, ca. 20-30 percent or less of anal vein length; medius, especially cubitus and anal veins usually distinct, except infrequently proximad 10-15 percent vague. Abdominal terga lacking irregular sclerites that engulf or join muscle attachment plates and dorsal (major + minor) setal bases or scleroites. Oviparae. — Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae ex¬ cept abdominal dorsum membranous with irregular transverse sclerites containing dorsal (major + minor) setae, occasionally dorsal muscle attachment plates, on each tergum; marginal setae usually on separate scleroites, occasionally engulfed by dorsal sclerites on posterad terga; siphuncular cones sclerotized, regular, separated from other dorsal sclerotic fields; dorsal abdominal muscle attachment plates unicolorous. Rarely with abdominal II-VI sclerotic/fused, terga VII and VIII free; dorsal de¬ marcations of anterad terga then evident and siphuncular cones surrounded closely by, sometimes engulfed by, sclerotic fields. Pseudorhinaria on metatibiae irregular, 12-21; also on procoxa and pro- and metafemora (Essig 1909: figs. 2a-b, 2d). Males.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except body slightly smaller, with slightly longer antennae and tibiae; dorsal demarcations of abdominal terga evident. Antennal segment III with 13-15, IV with 8-10, secondary rhinaria. Fundatrices. -Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except siphunculi absent; longest dorsal setae on central part of metatibiae 0.5-1.5x tibial diameter. Diagnosis. —Essigella (E. ) californica requires the combination of several char¬ acters for identification; individuals usually are pale and may be confused with other pale Essigella. Essigella (E. ) californica can be separated from all Essigella, except E. (E .) hoerneri and E. ( E .) pini by having six (Fig. IF), instead of eight or more, dorsal (major + minor) setae on abdominal terga III-IV. Essigella (E.) californica differs from E. (E.) pini by having usually small and irregular, rather 1994 SORENSEN: A REVISION OF ESSIGELLA 55 than large, ventral abdominal sclerites on abdominal segments III—IV, and also small and noninvasive, rather than large and'invasive, muscle attachment plates on the mesonotum of later stadia nymphs of apterae. Alates differ from those of E. (E.) pini by having the forewing medius always 1-branched with the furcation proximad, near the subcosta, instead of usually unbranched or occasionally 1-branched but with the furcation no more distad than half way between the subcosta and the posterad margin of the wing, as in E. (E.) pini. Oviparae of E. ( E .) californica and E. (E.) pini differ in the sclerotic pattern of the abdominal dorsum; in E. ( E .) pini, usually all terga, except terga I and VIII, are fused; in E. (. E .) californica, usually all terga are separate (as independent sclerotic bands), or infrequently segments II-VI are (sometimes only partially) united, with evident segmental demarcations, but terga VII and VIII remain independent [E. ( E .) pini rarely shows the latter condition]. Essigella (E.) californica and E. (E.) hoerneri are difficult to distinguish; the qualitative characters listed above for alates, oviparae and nymphs are identical between them. Although E. (E.) californica has a shorter rostrum, narrower head and longer antennal segment IV than does E. ( E .) hoerneri, these differences are indiscrete and reliable separation requires application of the discriminant function in the key to the viviparous apterae [couplets 27 or 28, see 26]. Synonyms. —Essigella claremontiana Hottes, NEW SYNONYM: holotype, vi- vip. apt., on slide with 5 other apt., holotype shown by arrow (11 o’clock position); data: CALIFORNIA. LOS ANGELES Co.: Claremont, 14 Feb 1909, Pinus ra- diata. Essigella claremontiana holotype deposited in the NMNH. Essigella cocheta Hottes, NEW SYNONYM: holotype, vivip. apt., on slide with 9 other apt. (including the holotype of E. monelli), E. cocheta holotype shown by circle (7 o’clock position); data: CALIFORNIA. MENDOCINO Co.: Fort Bragg, 8 May 1936, E.O.E[ssig]., Pinus “ tuberculata ” [= P. muricata]. Essigella cocheta holotype is deposited in the Essig Museum of Entomology, University of Cali¬ fornia at Berkeley, Berkeley, California. Essigella monelli Hottes, NEW SYNONYM: holotype, vivip. apt., on same slide as holotype of E. cocheta (see above), E. monelli holotype shown by circle (12 o’clock position). Essigella monelli holotype data and depository same as E. cocheta, above. Essigella pineti Hottes, NEW SYNONYM: holotype vivip. alat., on slide with fundatrix of E. (L.) fusca voegtlini; data: CALIFORNIA. MARIPOSA Co.: Yo- semite, 1218 m (4000 ft), 17 May 1938, E.O.E.[ssig]., Pinus ponderosa. Essigella pineti holotype is deposited in the Essig Museum of Entomology, University of California at Berkeley, Berkeley, California. Essigella swaini Hottes, NEW SYNONYM: holotype, vivip. alat., on slide with 6 other specimens, holotype shown by circle (12 o’clock position); data: CALI¬ FORNIA. LAKE Co.: Kelseyville, 12 Apr 1936, P. Schulthess, Pinus sabiniana. Essigella swaini holotype is deposited in the Essig Museum of Entomology, Uni¬ versity of California at Berkeley, Berkeley, California. Range. — Southern British Columbia and Alberta, throughout the western U.S. (exclusive of Alaska), to southern Mexico (Fig. 8); extensive sampling (Sorensen 1983) has most commonly collected it west of the Cascade-Sierra Nevada ranges and through Arizona and New Mexico. One confirmed record from Miami, Flor¬ ida suggests it may occur in the Caribbean and have a Pan-Mexican distribution. 56 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Figure 8. Distribution of E. (E.) californica [dots (JTS samples), squares (nonJTS samples)], su- perimposd over an amalgamation of the ranges of its hosts. Collections (not shown) also exist in southern Florida, central Mexico, France and Spain. Recently (Turpeau & Remaudiere 1990), I have identified it as having been introduced into France, in Pinus radiata plantations; since then, it has also been found in Spain (Seco Fernandez & Mier Durante 1992). Hosts. —Pinus and Pseudotsuga. Frequently found on: Pinus albicaulis Engel- mann, P. monticola, P. flexilis, P. leiophylla, P. ponderosa, P. ponderosa var. arizonica, P. jeffreyi, P. engelmannii, P. sabiniana, P. coulteri, P. torreyana Parry, P. radiata, P. attenuata, P. muricata\ infrequently found on: Pinus strobiformis, P. Iambertiana, P. contorta latifolia Engelmann ex S. Watson, P. washoensis Mason & Stockwell, Pseudotsuga menziesii (Mirbel) Franco, Pseudotsuga macrocarpa (Vasey) Mayr; not found on: Pinus cembroides Zuccagni, P. edulis Engelmann, P. 1994 SORENSEN: A REVISION OF ESSIGELLA 57 monophylla, P. quadrifolia, P. balfouriana Greville & Balfour, P. aristata Engel- mann, P. contorta contorta, P. contorta murrayana, P. contorta bolanderi (Sor¬ ensen 1983). During a recent introduction in France, Turpeau & Remaudiere (1990) report P. rigida, P. strobus, P. taeda, P. virginiana and P. griffithi McClelland as additional hosts. Essigella (E.) californica is frequently abundant on nonnative pines in the central valley of California, and also on native pines, especially subsection Sabinianae, in the surrounding foothills and in southern California. Although it is quite po- lyphagous within Pinus, it is notably absent on piny on pines [see discussion of E. ( E .) hoerneri ]. It occasionally feeds on Pseudotsuga, but apparently not in the presence of E. (E.) wilsoni. Although E. (E.) calif ornica may occur on hosts that are occupied by other, more restricted Essigella species, it is generally less nu¬ merous (i.e., relative abundance on an individual tree) on such hosts than are the species that are restricted to that niche; this is especially true when both occur on an individual tree. Note that with reference to “Sp. A” [= E. (. E .) calif ornica ], Sorensen (1987a: 255, lines 27, 28) mentioned Pseudotsuga as a “(secondary host capture)”; this unfortunate wording refers to opportunistic capture of a host species as a primary host, not a secondary (i.e., summer) host in the sense of the life cycle of the aphidines. Discussion. — This species, together with E. ( E .) hoerneri, forms the E. ( E .) calif ornica complex. Essigella (E.) calif ornica is the commonest Essigella in the western Nearctic and appears to be the species [followed closely by E. (E.) hoerneri] that is most prone to produce alates, as determined by their frequency among and within collection samples. This species is operationally defined and may actually represent a series of sibling entities on different host groups; however, I am comfortable with considering it to be a single taxon, because the range of its morphological variance does not appear to significantly exceed that shown by other, less polyphagous Essigella species. Further taxonomic division, beyond the current operational level, seems unwarranted unless biological and genetic anal¬ yses are carried out. Essigella (E.) calif ornica and E. (E.) hoerneri share several nonexclusive apo- morphies, although I have no doubt that they form a monophyletic group: re¬ duction of the dorsal (major + minor) setae to six [shared with E. (E.) pini] and the marginal setae to two [with E. (E.) pini, E. (E.) wilsoni and E. ( E .) alyeska] on abdominal terga III-IV; reduction of the ventral abdominal sclerites on seg¬ ments XII-IV to small, irregular plates [with E. (E.) wilsoni and, in part, E. ( E .) alyeska, E. (L.) fusca, E. (L.) hillerislambersi and (rarely) E. (E.) pini]; and the reduction of the mesonotal muscle attachment plates on the latter stadia apterae nymphs to noninvasive [with E. ( E .) wilsoni, E. ( E .) alyeska, and (very rarely) E. ( E .) pini]. I have analyzed character variation among and within populations of the E. ( E.) calif ornica complex, in comparison to the E. ( L.) fusca and E. (E.) knowltoni complexes, using principal component analyses (unpublished data). In those anal¬ yses, the body setal lengths of the former generally loaded uniformly and heavily on the second principal component vector, with the first vector representing general-size. The extent of that setal loading was generally much more uniform for the E. ( E .) calif ornica complex, than it was for these other complexes, indicating 58 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) that setal lengths vary more, in unison, in the E. (E.) californica complex than in either of the others. In other analyses (unpublished data), when only E. (E.) calif ornica and E. (E.) hoerneri were subjected to principal component analysis, where general-size was again represented by vector 1, they diverged on combi¬ nations of vectors 2 versus 3. The loadings of those vectors indicated that their separation was chiefly on the basis of body widths and stylet length. In contrast, Hottes (1957: 109, key couplet 11) used 0.030 mm as a threshold value for the length of the dorsal setae on the metatibia (his “hairs on the mid region of outer margin of metathoracic tibiae”) to separate his E. “ calif ornica” and E. “ hoerneri ”; clearly Hottes’ approach was erroneous. Essigella (E.) calif ornica and E. ( E .) hoerneri show extreme variation of the length of the setae on the frons and dorsal setae on the metatibiae. This population attribute could be considered synapomorphic, as a mutation that allows greater phenotypic plasticity of setal length in response to environmental conditions, but the mechanism of expression is not understood. Essigella (E.) wilsoni also shows a similar tendency, but to a lesser degree; the number of incidences of extreme setal length reduction (i.e. , < 0.3 x tibial diameter) is much less in it. Other variation shared by both E. {E.) calif ornica and E. ( E .) hoerneri entails the relative length of the metatibiae on adult viviparous apterae. On occasion, aberrant individual viviparous apterae retain the relatively shorter metatibiae that is characteristic of the ultimate stadium of their nymph (i.e., the holotype of E. “cocheta ”). This trait also occurs in other Essigella species, but it seems to be less frequently expressed than in the E. (E.) calif ornica complex. I suspect that such aberrantly short tibiae result from the failure of a regulatory gene that controls physiognomic transitions between nymphal versus adult (or alate versus aptera) allometries. With the exception of E. ( E.) pini, E. ( E .) californica has caused the greatest confusion in the genus. Hottes did not have adequate samples to reflect its vari¬ ability, and used poor preparatory techniques. Among his synonyms, E. “monelli ” and E. “ cocheta ” were stated to lack distinctly bifid tarsal claws, but their types actually have them; excessive clearing in potassium hydroxide obscures this char¬ acter when viewed through only low power magnification. Other synonyms, E. “ claremontiana” and E. “ swainif were previously separated on the basis of setal lengths, which vary continuously. In fact, based upon setal length, Hottes (1957: 85) believed that the series from which Essig originally described “Lachnus cal- ifornicus ” must have had specimens of E. “ essigi ” in it [see discussion of that species], Wilson’s (1919: 1) (re)description of “Essigella californica (Essig),” from material on Pseudotsuga “ douglassi ” [= P. menziesii], “Pinus ponderosaT ’ and specimens sent to him by Essig, most probably incorporates species other than E. ( E.) californica ; if so, E. ( E .) wilsoni and E. ( L.) fusca might presumably be involved, but I cannot determine this. Ironically, even Essig (1909) had some trouble recognizing the different morphs when originally describing E. ( E .) californica. For example, Essig’s illustration of an aptera of the species (Essig 1909: fig. 2a-b, 2d) clearly shows the legs with sensoria, which he mentions in the description, but the description is labeled “Apterous Vivivarous Female”; these are pseudorhinaria of the ovipara morph, however. Coded References to This Taxon.—Essigella (E.) californica has been referred 1994 SORENSEN: A REVISION OF ESSIGELLA 59 to previously by: the coding “Sp. A” (Sorensen 1983, 1987a, 1992b) and “CALF” (Sorensen 1983), and by the name E. californica in Sorensen (1983). Etymology and Common Name.— Essig named this species for California, the state in which its original collection occurred, and where it is most commonly encountered (e.g., Fig. 8). Common name: the Californian pine needle aphid; although Essig (1936: 229), Doane et al. (1936: 360), Palmer (1952: 14), and Furniss & Carolin (1977: 99) refer to this species as the “Monterey Pine Aphid,” the common name indicated here is more appropriate and less confusing because Pinus radiata, Monterey pine, as a niche is occupied much more representatively by E. (E.) essigi, whereas E. (E.) californica is quite polyphagous within Pinus. Material Examined. — ARIZONA. COCHISE Co.: Amer. Mus. Nat. Hist. Southwest Research Station, Chiricahua Mts, 1700 m, 16 Sep 1978, JTS 78143, P. engelmannii, (apt.); Carr Canyon Rd, Huachuca Mts, 2070 m, 17 Sep 1978, JTS 78155, P. engelmannii, (apt.); Miller Canyon Rd, Huachuca Mts, 1700 m, 17 Sep 1978, JTS 78152, P. leiophylla, (apt.); nr Steward Camp, Chiricahua Mts, 1530 m, 16 Sep 1978, JTS 78142, P. leiophylla, (apt., alat.). COCONINO Co.: 9 km W of Williams on hwy 66, 2070 m, 9 Sep 1978, JTS 7815, P. ponderosa, (apt.). GILA Co.: 16 km E of Kohles Ranch on hwy 260, 1700 m, 9 Sep 1978, JTS 78111, P. ponderosa, (apt.). GRAHAM Co.: SW of Stafford on hwy 366, 1830 m, 15 Sep 1978, JTS 78136, P. leiophylla, (apt.); same but 1980 m, JTS 78137, P. ponderosa var. arizonica, (apt.). MARICOPA Co.: Phoenix, 13 Jan 1972, D. Carver, P. canariensis, (apt.); Sun City, 27 Jan 1972, D. Carver, P. taeda, (apt.). NA VAJO Co.: Mogollon Rim Rd, 8 km SW of Showlow, 2070 m, 10 Sep 1978, JTS 78113, P. ponderosa, (apt.). PIMA Co.: Bear Canyon Picnic Area, Santa Catalina Mts, 1830 m, 18 Sep 1978, JTS 78157, P. leiophylla, (apt., alat.); same but JTS 78160, P. ponderosa var. arizonica, (apt.). COUNTY UNCERTAIN: Sitgreaves Natl Forest, 19 Jun 1969, D. T. Jennings, P. ponderosa, (apt.). CALIFORNIA. ALAMEDA Co.: Berkeley, 23 Apr 1947, E. O. Essig, P. radiata, (apt., alat.); same but 10 Nov 1935, (apt.); same but 28 Oct 1952, trap pan, (alat.). ALPINE Co.: E. side of Ebbett’s Pass on hwy 4, 3 km E of summit, 2440 m, 17 Jul 1977, JTS 77G41, P. monticola, (apt.); same but W side, 5 km W of summit, 2500 m, JTS 7 7 G4 3, P. jeffreyi, (gcpi). AMADOR Co.: 13 km N of Plymouth, 29 May 1977, J. T. Sorensen, P. sabiniana, (apt., alat.). BUTTE Co.: Chico, 27 Oct 1949, H. T. Osborn, CDFA 40-K-5, P. yunnanensis Franchet [?], (apt., alat.); Feather River Cyn, 22 km NE of jet of hwy 70 & Cherokee Rd, 26 Jun 1977, JTS 77F14, P. ponderosa, (apt.). CALAVERAS Co.: 18 km E of Arnold on hwy 4, 1680 m, 17 Jul 1977, JTS 77G46, P. ponderosa, (apt.); 2 km NE of Murphys on hwy 4, 670 m, 17 Jul 1977, JTS 77G47, P. ponderosa, (apt.); 7 km NE of Angel’s Camp on hwy 4, 460 m, 17 Jul 1977, JTS 77G48, P. sabiniana, (apt., alat.). COLUSA Co.: W of William on hwy 20, 18 Apr 1979, T. Kono & P. Crane, CDFA 79D19-35, P. sabiniana, (apt.). CONTRA COSTA Co.: Mt Diablo, 23 Apr 1939, E. O. Essig, P. sabiniana, (alat.); Orinda, 29 Sep 1961, E. I. Schlinger, EIS 61-9-30b, Pinus sp., (apt., alat.). DEL NORTE Co.: Gasquet, 21 Sep 1966, P. Allen, CDFA 66-116-14, Pseudotsuga menziesii, (alat.). EL DORADO Co.: Blodgett Exper¬ imental Forest (Univ. Calif.), E of Georgetown, 26 Jul 1973, D. J. Voegtlin, DJV 55, P. ponderosa, (apt., alat.); same but 28/29 May 1977, J. T. Sorensen, P. lambertiana, P. ponderosa, (alat.); George¬ town, 29 May 1977, J. T. Sorensen, P. sabiniana, (apt., alat.); Lake Tahoe, Meek’s Bay, 1980 m, 16 Jul 1977, JTS 77G29, P. jeffreyi, (apt.); Mutton Cyn, 3 Oct 1961, T. Kono, Pinus sp., (alat.); S Fork of American River, 5 Jul 1973, D. J. Voegtlin, DJV 35, P. sabiniana, (apt.). FRESNO Co.: 22.4 km (14 mi) W of Coalinga on hwy 145, 25 Apr 1979, D. Taylor, CDFA 79D27-8, P. sabiniana, (apt., alat.); Clovis, 7 Apr 1965, Dunnegan, CDFA 65D9-21, P. canariensis, (apt.); Trimmer, Pine Flat Lake, 13 Aug 1977, JTS 77H8, P. sabiniana, (apt.); jet of hwys 180 & 245, 1620 m, 13 Aug 1977, JTS 77H9, P. ponderosa, (apt.). HUMBOLT Co.: nr Little River State Beach, 17 km N of Areata on hwy 101, 4 Jul 1978, JTS 78G3, P. muricata, (apt., alat.). KERN Co.: Caliente-Bodhsh Rd, S of Bodhsh, 820 m, 20 Sep 1977, JTS 77167, P. sabiniana, (apt.); Heritage Park, 19 Jun 1967, K. Hench, P. canariensis C. Smith, (apt.); Keene, 760 m, 20 Sep 1977, JTS 77162, P. sabiniana, (apt.); Kemville, 22 May 1978, C. F. & C. S. Smith, CFS 78-76, Pinus sp., (apt., alat.); Lebec, 25 Mar 1958, E. I. Schlinger, EIS 58-3-259, P. sabiniana, (apt.); Tehachapi Mtn Park, S of Tehachapi, 1980 m, 19 Sep 1977, JTS 77160, P. ponderosa, (apt.); same but JTS 77161, P. jeffreyi, (apt.); Tiger Flat Rd, N of hwy 155, nr Alta Sierra, 1890 m, 20 Sep 1977, JTS 77164, P. lambertiana, (ovip.); same but JTS 77166, P. jeffreyi, (apt.). LAKE Co.: 10 km S of Lake Pillsbury, Elk Mt Rd, 930 m, 24 Jul 1977, JTS 77G57, 60 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) P. ponderosa, (apt., alat.); same but 5 km S, 640 m, JTS 77G56, P. jejfreyi, (apt.); 21 km N of Upper Lake, Elk Mt Rd, 1030 m, 24 Jul 1977, JTS 77G60, P. attenuata, (alat.); Kelseyville, 12 Apr 1936, P. Schulthess, P. sabiniana, (apt.); same but 15 Jul 1935, E. Doybell, (alat.); W of Lake Pillsbury, Eel River Rd, 340 m, 24 Jul 1977, JTS 77G54, P. sabiniana, (apt., alat.). LASSEN Co.: 7 km SW of Susanville on hwy 36, 1460 m, 4 Jul 1977, JTS 77G13, P. jejfreyi, (apt., alat.); Lassen Natl Park, summit area, 2440 m, 10 Jul 1977, JTS 77G18, P. monticola, (apt.); hwy 89 (nr Lassen Natl Park), 6 km N of jet with hwy 36, 2013 m, 10 Jul 1977, JTS 77G20, P. monticola,(apt.). LOS ANGELES Co.: (lectotype) Claremont, 14/18 Feb 1909, E. O. Essig, P. rndiata, (apt., alat.); Lake Hughes, 22 May 1959, E. I. Schlinger, EIS 59-5-23k, P. ponderosa, (alat.); same but 2 km NE on hwy N2, 1000 m, 18 Sep 1977, JTS 77152, P. sabiniana, (apt.); 3 km SE of Big Pines on hwy 2, E of Blue Ridge Summit, 2200 m, 17 Sep 1977, JTS 77149, P. jejfreyi, (alat.); Azuza, 4 Nov 1969, McHom & Weber, CDFA 69K7-8, P. canariensis, (apt.); Camp Baldy, 5 Dec 1956, J. MacSwain, “on fir,” (apt., alat.); hwy 2, 7 km NE of jet with Mt Wilson Rd, San Gabriel Mts, 1530 m, 18 Sep 1977, JTS 77151, P. coulteri, (apt.). MARIN Co.: San Rafael, 18 May 1967, C. Schmid, P. radiata, (apt.). MARIPOSA Co.: Feliciana Mt, 25 Jul 1946, H. Chandler, P. “ tuberculata ” [= muricata!], (apt.); Yosemite Natl Park, Camp Foresta, 1340 m, 30 Jul 1977, JTS 77G64, P. ponderosa, (apt., alat.); same but JTS 77G65, P. lambertiana, (apt.). MENDOCINO Co.: 10 km W of Laytonville, Branscomb Rd, 580 m, 24 Jul 1977, JTS 77G53, P. ponderosa, (apt.); Albion-Little River Rd, 5 km E of hwy 1, 210 m, 23 Jul 1977, JTS 77G52, P. muricata, (apt., alat.); Fish Rock Rd, 7 km E of Anchor Bay, 23 Jul 1977, JTS 77G51, P. muricata, (apt.); Fort Bragg, 8 May 1936, E. O. Essig, P. “ tuberculata ” [= muricata!], (apt.). MODOC Co.: 1 km W of Adin Pass on hwy 299, 21 Jul 1978, JTS 78G120, P. ponderosa, (apt.); E side of Cedar Pass, 29 km E of Alturas, 1890 m, 3 Jul 1977, JTS 77G12, P. ponderosa, (apt.). MONO Co.: Deadman Summit on hwy 395, nr Crestview, 2440 m, 31 Jul 1977, JTS 77G72, P. jejfreyi, (apt.); Saddlebag Lake, 3050 m, 31 Jul 1977, JTS 77G69, P. albicaulis, (apt., fund.). MONTEREY Co.: Cone Mt, 9 Aug 1962, E. I. Schlinger, EIS 62-8-9a, Pinus sp., (apt.); Cone Peak Rd, 2 km N of jet with Nacimento-Fergusson Rd, Los Padres Natl Forest, 910 m, 4 Sep 1977, JTS 7719, P. coulteri, (apt.); Lockwood-San Ardo Rd, 13 km SW of jet with Paris Valley Rd, 550 m, 4 Sep 1977, JTS 7717, P. attenuata, (apt.); same but JTS 7718, P. sabiniana, (apt.); Monterey, 16 Feb 1979, T. Kono, CDFA 79B20-10-3, P. radiata, (apt.); same but 18 Jun 1973, D. J. Voegtlin, DJV 24, P. radiata, (apt., alat.). NAPA Co.: 16 km NE of Angwin, jet of Howell Mt Rd & Pope Canyon Rd, 4 Feb 1978, JTS 79B1, P. sabiniana, (apt.). ORANGE Co.: Anaheim, 9 Feb 1965, J. Seapy, CDFA 65B15-26-2, P. radiata, (apt.); above Santiago Peak Rd, 10 km N of jet with hwy 74, Cleveland Natl Forest, 1220 m, 10 Sep 1977, JTS 77122, P. coulteri, (apt.); same but JTS 77123, P. attenuata, (apt.). PLACER Co.: 5 km SW of Whitmore on hwy 80, 1430 m, 25 Jun 1977, JTS 77F1, P. ponderosa, (apt.); 6 km W of Dutch Flat on hwy 80, 2 Aug 1978, JTS 78H2, P. attenuata, (apt.); same but 27 Aug 1978, JTS 78H160, (apt.). PLUMAS Co.: 13 km E of Chester on hwy 36, 1520 m, 4 Jul 1977, JTS 77G15, P. jejfreyi, (apt., alat.); hwy 36, 6 k W of jet with hwy 89, 1460 m, 10 Jul 1977, JTS 77G22, P. lambertiana, (apt.); same but JTS 77G25, P. jejfreyi, (apt.); Jackson Creek Cmpgd, Plumas Natl Forest, 2 km SE of Cromberg on hwy 70/89, 1280 m, 26 Jun 1977, JTS 77F12, P. ponderosa, (apt.); Keddie, 18 Oct 1966, Swanson, CDFA 66J24-27, Pseudotsuga menziesii, (apt.). RIVERSIDE Co.: Idyllwild, 1 Jun 1940, C. Michner, P. ponderosa, (apt., alat.); Keen Camp Summit on hwy 74, 3 km N of Mountain Center, San Bernardino Natl Forest, 1500 m, 9 Sep 1977, JTS 77120, P. coulteri, (apt.); Riverside, 28 Nov 1961, C. Lagace, EIS 61-2-28a, P. canariensis, (apt., alat.); same but 9 Mar 1960, E. I. Schlinger, EIS 60-3-9a, P. canariensis, (apt., alat.). SACRAMENTO Co.: Wm. Land Park, Sacramento, 26 Aug 1961, T. Kono, Pinus sp., (apt., alat.). SAN BENITO Co.: Clear Creek Rd, 10 km SE of jet with Coalinga Rd, 1000 m, 3 Sep 1977, JTS 7714, P. coulteri, (apt.); same but 14 km SE of that jet, 1370 m, JTS 7715, (apt., alat.); same but Clear Creek Recreation Area entrance, 2600 m, JTS 7716, P. sabiniana, (apt.); Coalinga Rd, 2 km SE of jet with hwy 25, 550 m, 3 Sep 1977, JTS 7713, P. sabiniana, (apt.); Gloria-Bickmore Rd, 14 km W of jet with hwy 25, 580 m, 3 Sep 1977, JTS 7711, P. sabiniana, (apt.); same but JTS 7712, P. coulteri, (apt.); Pinnacles Natl Monument, 24 Apr 1948, J. MacSwain, Pinus sp., (apt.). SAN BERNARDINO Co.: 7 km W of Barton Rat on hwy 38, 1950 m, 16 Sep 1977, JTS 77136, P. coulteri, (apt., alat.); San Bernardino Natl Forest, Keller Peak Cmpgd, 2200 m, 17 Sep 1977, JTS 77142, P. attenuata, (apt.); same but Dogwood, 28 Aug 1972, D. J. Voegtlin, DJV 72, P. ponderosa, (alat.); same but Running Springs, 4 Aug 1973, DJV 77, P. coulteri, (alat.); same but Snow Valley, 28 Aug 1972, DJV 69, P. jejfreyi, (alat.); Redlands, 22 Dec 1978, CDFA 78L26-28, P. radiata, (apt.). SAN DIEGO Co.: 2 km E of Mt Palomar on hwy S6, 1650 m, 11 Sep 1977, JTS 77128, P. attenuata, (apt.); Mt Palomar Rd (hwy S6), 5 km S of Mt Palomar, 1370 m, 11 Sep 1977, JTS 77126, Pseudotsuga macrocarpa, (apt.); 5 km S of Julian, Harrison Springs Rd, 1460 m, 12 Sep 1977, JTS 1994 SORENSEN: A REVISION OF ESSIGELLA 61 77129, P. coulteri, (apt.); La Jolla, Univ. Calif, campus, 11 Sep 1977, JTS 77125, P. torreyana, (apt., alat.); San Diego, 10 May 1967, R. Roberson, CDFA 67E15-68, P. radiata, (apt.); same but 29 Jun 1961, O. Beck, CDFA 61F29-53, (apt., alat.); same but 11 Dec 1957, W. Radcliffe, P. canariensis, (apt., alat.); Torrey Pines State Reserve, 10 Sep 1977, JTS 77124, P. torreyana, (apt.); Valley Center, 18 Apr 1975, G. Gordun, CDFA 75D24-34, Pinus sp., (apt.). SAN FRANCISCO Co.: San Francisco, 28 Apr 1967, M. Stufflebeam, CDFA 67E1-10, P. radiata, (apt., alat.). SAN LUIS OBISPO Co.: 2 km E of Santa Margarita on hwy 58, 300 m, 5 Sep 1977, JTS 77113, P. sabiniana, (apt.); Cuesta Ridge Botanical Area, nr La Cuesta Summit on hwy 101, N of San Luis Obispo, 730 m, 5 Sep 1977, JTS 77114, P. coulteri, (apt., alat.); Ragged Point, 21 Jul 1973, D. J. Voegtlin, DJV 56, P. radiata, (alat.). SANTA BARBARA Co.: Happy Canyon Rd, 16 km NE of jet with hwy 154, Los Padres Natl Forest, 370 m, 6 Sept 1977, JTS 77116, P. sabiniana, (apt., alat.); Purissima Hills, 10 km N of jet of hwys 1 & 246, 6 Sept 1977, JTS 77115, P. muricata, (apt., alat.); San Marcos Pass, 740 m, 14 Apr 1960, E. I. Schlinger & J. Hall, EIS 60-4-15c, P. “ monticola’' [?], (apt., alat.); Santa Barbara, 1 May 1939, G. Woodham, Pinus sp., (apt., alat.); Santa Cruz Island, Prisoner’s Harbor, 25 Sep 1978, JTS 78164, P. muricata, (apt.); Santa Ynez, 23 Apr 1975, B. Jarvis, CDFA 75D24-39, Pinus sp., (apt.); Tequepis Cyn, 17 May 1957, M. Cravens, CDFA 57E21-14, P. radiata, (apt.). SANTA CLARA Co.: Campbell, 19 Apr 1967, G. Prole, CDFA 67E5-34, Pinus sp., (apt.); Palo Alto, Stanford Univ., 25 Apr 1930, P.S.B., P. radiata, (apt.); same but 7 Apr 1912, H. Morrison, P. “maritima ” [?], (apt., alat.). SANTA CRUZ Co.: Santa Cruz, 20 Jul 1966, J. Bauer, CDFA 66G26-3, Pinus sp., (alat.). SHASTA Co.: 24 km (15 mi) E of Redding, nr Bella Vista, 29 Mar 1979, D. Henry, CDFA 79C29-19, P. sabiniana, (apt.); 16 km S of Castella on hwy 5, 400 m, 2 Jul 1977, JTS 77G1, P. sabiniana, (apt.); 2 km W of Fall River Mills on hwy 299, 21 Jul 1978, JTS 78G121, P. sabiniana, (apt., alat.); same but JTS 78G123, P. ponderosa, (apt.); 3 km N of hwy 299 on Rock Creek Rd, W of Redding, 300 m, 20 Aug 1977, JTS 77H14, P. attenuata, (apt.); Whiskeytown Lake, 370 m, 20 Aug 1977, JTS 77H15, P. attenuata, (apt.); same but JTS 77H16, P. sabiniana, (apt., alat.). SISKIYOU Co.: Edson Creek access Rd, Shasta Natl Forest, 8 km W of Bartel on hwy 89, 1160 m, 3 Jul 1977, JTS 77G10, P. jeffreyi, (apt., alat.); Mt Shasta Ski Bowl Rd, 2450 m, 2 Jul 1977, J. T. Sorensen & D. J. Voegtlin, JTS 77G8, P. lambertiana, (apt.); same but JTS 77G6, P. ponderosa, (apt., alat.); same but JTS 77G4, P. albicaulis, (apt., fund.); Snowman Hill Summit on hwy 89, 8 km E of jet with hwy 5, 1360 m, 2 Jul 1977, JTS 77G2, P. ponderosa, (apt., alat.). SOLANO Co.: Green Valley, 29 Oct 1939, N of Frazier, (alat.). SONOMA Co.: hwy 101, at Sonoma-Mendocino Co. line, 3 Jul 1978, JTS 78G1, P. sabiniana, (apt., alat.). TEHAMA Co.: 29 km E of Dales on hwy 36, 910 m, 10 Jul 1977, JTS 77G27, P. sabiniana, (apt., alat.); same but 970 m, JTS 77G28, P. ponderosa, (apt., alat.); Lanes Valley Rd, nr jet with hwy 36, 490 m, 4 Jul 1977, JTS 77G17, P. sabiniana, (apt.). TRINITY Co.: Big Flat, 1 Jun 1978, C. F. Smith, P. sabiniana, (apt.); Buckhom Summit on hwy 299, W of Tower House, 980 m, 20 Aug 1977, JTS 77H17, P. ponderosa, (apt., alat.); Weaverville, 550 m, 20 Aug 1977, JTS 77H20, P. sabiniana, (apt., alat.); same but JTS 77H21, P. ponderosa, (apt., alat.). TULARE Co.: E of Big Meadows Cmpgd, Sierra Natl Forest, 2320 m, 13 Aug 1977, JTS 77H13, P. jeffreyi, (apt.); same but JTS 77H12, P. monticola, (apt.); Visalia, 9 Apr 1971, J. Gilley, CDFA 71D12-11, Pinus sp., (apt., alat.). TUOLUMNE Co.: 2 km E of Groveville on hwy 120, 910 m, 30 Jul 1977, JTS 77G63, P. ponderosa, (apt., alat.); 7 km W of Big Oak Flat on hwy 120, 550 m, 30 Jul 1977, JTS 77G61, P. sabiniana, (apt.); Kennedy Meadows, 12 Jul 1951, W. Lange, P. ponderosa, (alat.); Mocassin, 14 Jun 1973, D. J. Voegtlin, DJV 37, P. sabiniana, (apt., alat.); Strawberry, 26 Apr 1951, J. MacSwain, Pinus sp., (alat.); Yosemite Natl Park, 1330 m, 17 May 1938, E. O. Essig, P. ponderosa, (alat.). VENTURA Co.: 4.8 km (3 mi) S of Pine Mt Summit, 16 May 1961, R. Van den Bosch, RVdB 61 -V-19j, P. sabiniana, (apt., alat.); Mt Pinos Summit, 2684 m, 18 Sep 1977, JTS 77154, P. flexilis, (apt., ovip., male); Santa Paula, 26 Jun 1911, E. O. Essig (USNM type 16243, P. radiata, (apt., alat.). YOLO Co.: Davis, 1 Mar 1979, R. Harris, CDFA 79C2-1-2, P. sabiniana, (apt., alat.); Davis, 19 May 1979, T. & C. Kono, CDFA 79E21- 42, P. sabiniana, (apt.). COLORADO. LARIMER Co.: Stove Prairie Hill, nr Bellvue, 16 Jun 1922, M. A. Palmer, CAES 3118, P. “ murraryana" [= contorta latifoliaP. ], (alat.); Estes Park, 22 Jul 1922, M. A. Palmer, CAES 3152, P. flexilis, (apt.). SAN JUAN Co.: 20 km N of Purgatory, 3020 m, 8 Aug 1978, JTS 78H47, P. flexilis, (apt.). FLORIDA. DADE Co.: Opa Locka, 29 Feb 1956, C. Shepard & L. Daigle, Pinus sp., (apt.). IDAHO. BONNER Co.: 6 km E of Colburn on hwy 95, 18 Jul 1978, JTS 78G102, P. monticola, (apt.); 6 km S of Cocolalla on hwy 95, 18 Jul 1978, JTS 78G105, P. ponderosa, (apt., alat.). CLEAR WATER Co.: 5 km EofOrofino on hwy 12, 18 Jul 1978, JTS 1&G10&, P. ponderosa, (apt.). MONTANA. CARBON Co.: E side of Beartooth pass on hwy 212, 2780 m, 20 Aug 1978, JTS 78H118, P. albicaulis, (apt., ovip.). FLATHEAD Co.: 5 km W of MacGregor Lake on hwy 2, E of Happy Inn, 18 Jul 1978, JTS 78G101, P. ponderosa, (apt., alat.); hwy 93, nr Olney, 17 Jul 1978, JTS 62 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) 78G99, P. monticola, (apt.). NEBRASKA. THOMAS Co.: Halsey, 14 Sep 1958, R. Henzlik, (alat.). NEVADA. CLARK Co.: Charleston Mts, Lee Canyon Ski Area, 2590 m, 4 Aug 1978, JTS 78H16, P. flexilis, (apt., alat.). WASHOE Co.: Mt Rose Summit Cmpgd, Toiyabe Natl Forest, 2 Aug 1978, JTS 78H8, P. albicaulis, (apt., fund.); same but JTS 78H9, P. monticola, (apt.); Mt Rose, Slide Mountain Ski Area, 2 Aug 1978, JTS 78H6, P. washoensis, (apt.). NEW MEXICO. BERNALILLO Co.: 2 km NW of San Antinito on hwy 44, 2290 m, 12 Sep 1978, JTS 78120, P. ponderosa, (apt.). CATRON Co.: Apache Natl Forest, 11 May 1978, C. F. & C. S. Smith, CFS 78-24, (apt., alat.). OREGON. BAKER Co.: 11 km W of Unity on hwy 26, 20 Jul 1978, JTS 78G112, P. ponderosa, (apt., alat.). CLACKAMAS Co.: Timberline Lodge, Mt Hood, 1770 m, 6 Jul 1978, JTS 78G35, P. albicaulis, (fund.). DESCHUTES Co.: Sisters, 23 May 1968, C. F. Smith & B. Zak, CFS 68-115, P. ponderosa, (apt.). HARNEY Co.: 20 km N of Bums on hwy 395, 20 Jul 1978, JTS 78G117, P. ponderosa, (apt., alat.). JACKSON Co.: 21 km S of Union Creek on hwy 62, 5 Jul 1978, JTS 78G15, P. ponderosa, (apt., alat.); same but 3 km E, 110 m, JTS 78G18, (apt). JOSEPHINE Co.: O’brien, 4 Jul 1978, JTS 78G9, P. jeffreyi, (apt.). KL AMATH Co.: 16 km S of LaPine on hwy 97, 5 Jul 1978, JTS 78G26, P. ponderosa, (apt., alat.). LAKE Co.: 28 km N of Lakeview on hwy 395, 20 Jul 1978, JTS 78G119, P. ponderosa, (apt.). WASCO Co.: 21 km SE of Government Camp on hwy 26, 970 m, 6 Jul 1978, JTS 78G34, P. monticola, (apt.); same but 46 km SE, 670 m, JTS 78G32, P. c. murrayana, (apt., alat.); jet of Mills Creek & hwy 26, 35 km NW of Madras, 6 Jul 1978, JTS 78G29, P. ponderosa, (apt., alat.). UTAH. CACHE Co.: Bearlake Summit on hwy 89, 8 km W of Garden City, 2350 m, 24 Aug 1978, JTS 78H132, P. flexilis, (apt.). DUCHESNE Co.: 19 km NE of Castle Gate on hwy 33, 2770 m, 25 Aug 1978, JTS 78H144, P. flexilis, (apt.); W of Duchesne, 29 Jun 1958, G. F. Knowlton, (alat.). IRON Co.: 32 km SE of Cedar City on hwy 14, 3020 m, 5 Aug 1978, JTS 78H27, P. flexilis, (apt.). PIUTE Co.: Marysville Cyn, 11 Jun 1943, G. F. Knowlton, P. ponderosa, (alat.). WASHINGTON. CHELAN Co.: 8 km SW of Chelan on hwy 97, 12 Jul 1978, JTS 78G68, P. ponderosa, (apt., alat.); Washington Pass on hwy 20, 1700 m, 12 Jul 1978, JTS 78G75, P. albicaulis, (apt., fund.). CLALLAM Co.: Olympic Natl Park, Hurricane Ridge, 9 Jul 1978, JTS 78G51, P. monticola, (apt., fund.). GRAYS HARBOR Co.: 16 km W of Amanda Park on hwy 101, 10 Jul 1978, JTS 78G54, P. monticola, (apt.). KING Co.: Seattle, 16 Jun/20 Oct 1955, M. Forsell, Pinus sp., (apt.). KITSAP Co.: 8 km S of Hood Canal bridge on hwy 3, 9 Jul 1978, JTS 78G49, P. monticola, (apt.). OKANOGAN Co.: 17 km NW of Winthrop on hwy 20, 550 m, 12 Jul 1978, JTS 78G71, P. ponderosa, (apt., alat.). WHITMAN Co.: Pullman, 26 Sep 1956, F. C. Hottes, P. ponderosa, (alat.). YAKIMA Co.: 16 km W of Naches on hwy 410, 11 Jul 1978, JTS 78G65, P. ponderosa, (apt.); E side of Chinook Pass on hwy 410, 1310 m, 11 Jul 1978, JTS 78G63, P. monticola, (apt.); Union Gap, 22/26 Sep 1952, E. Davies, trap pan, (alat.). WYOMING. CROOK Co.: 6 km W of Devil’s Tower Jet on hwy 14, 1100 m, 19 Aug 1978, JTS 78H104, P. ponderosa, (apt.). PLATTE Co.: S of Glendo on hwy 25, 1920 m, 17 Aug 1978, JTS 78H94, P. ponderosa, (apt.). TETON Co.: Hoback Jet, 19 km S of Jackson on hwy 89, 1860 m, 23 Aug 1978, JTS 78H128, P. flexilis, (apt., male); nr Togwotee Pass on hwy 287, 2800 m, 23 Aug 1978, JTS 78H125, P. albicaulis, (apt., fund., ovip.). WASHAKIE Co.: 19 km NE of Tensleep on hwy 16, 2350 m, 19 Aug 1978, JTS 78H107, P. flexilis, (apt.); Tensleep Cyn, Bighorn Mts, 1580 m, 20 Aug 1978, JTS 78H111, P. ponderosa, (apt.). CANADA. BRITISH COLUMBIA: 21 km N of Cache Creek on hwy 97, 13 Jul 1978, JTS 78G81, P. ponderosa, (apt.); 21 km S of Lytton on hwy 1, 13 Jul 1978, JTS 78G78, P. ponderosa, (apt.); 5 km N ofSpuzzum on hwy 1, 13 Jul 1978, JTS 78G77, P. monticola, (apt., male); 7 km S of Canal Flats on hwy 93, 17 Jul 1978, JTS 78G95, P. c. latifolia, (apt.); Fairmont Hotsprings, hwy 93, 17 Jul 1978, JTS 78G91, P. ponderosa, (apt.). MEXICO. DISTRITO FEDERAL: Ajusco, 2800 m, 2 Apr 1981, G. Remaudiere, Pinus sp., (apt., alat.). STATE UNCERTAIN: Chapingo, 27 Oct 1980, G. Remaudiere, Pinus sp., (apt., alat.). FRANCE. PROVINCE UNCERTAIN: Rennesle Rheu, 30 Jun 1989, “R 42,” piege, (alat.); Landemeau Finistere, 6 Sep 1989, E. Turpeau, 16580, “P. radiatalfl (apt.). Essigella ( Essigella ) hoerneri Gillette & Palmer, 1924 Essigella hoerneri Gillette & Palmer, 1924: 5, Ann. Entomol. Soc. Am., 17: 5-6. Essigella gillettei Hottes, 1957: 88, Proc. Biol. Soc. Wash., 70: 88-90. NEW SYNONYM. Essigella maculata Hottes, 1957: 93, Proc. Biol. Soc. Wash., 70: 93-95. NEW SYNONYM. 1994 SORENSEN: A REVISION OF ESSIGELLA 63 Primary Type. — Lectotype, vivip. apt., on slide with 6 other specimens, lec- totype in upper right; slide data: “ Essigella hoerneri n. sp., lectotype (vivip. apt.) (type—others paratypes) C. P. Gillette & M. A. Palmer/U.S. Nat. Mus. No. 41952/ On Pinus edulis, 9-25-21, Owl Canon [sic] Larimer Co., Colo., Coll. J. L. Hoemer, Colo. Agr. Exp. Sta. Ac. 2894/[on back] lectotype, Essigella hoerneri Gillette & Palmer, J. T. Sorensen 1981.” Lectotype deposited in the U.S. National Museum of Natural History, Washington, D.C. There is confusion concerning the lectotype designation. In the original de¬ scription, Gillette & Palmer (1924: 5-6) list no primary type, but later (Gillette & Palmer 1931: 841) state “Types in U.S. Nat. Mus., Cat. No. 41952; Paratypes in collection of Colo. Agr. Exp. Sta.” Palmer (1952:16) again refers to that museum number under the heading Type. There is a second “type” slide, containing one ovipara with four other specimens, that also bears the U.S. Nat. Mus. number 41952. Consequently, the slide cannot be identified from that number alone. Hottes (1957: 92) mentions a lectotype and gives the slide data (as above), but does not tell the position of the designated individual on the slide. Although there is a “map” of position of the “type” on the slide, I am uncertain that this represents the lectotype mentioned by Hottes. Because the “type” individual on that slide was incomplete, I have designated a different, intact specimen from the slide (upper right corner, 2 o’clock position) as lectotype. Viviparous Apterae.— Morphology: Body length: 1.49-2.36 (1.86 ± 0.22) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously protuberant. Length of antennal segment V: 110-148 (130 ± 10) p, processus terminalis: 30-43 (37 ± 4) p- IV: 75-120 (101 ± 12) p\ III: 118-238 (175 ± 29) p\ II: 59-88 (70 ± 7) p. Length of longest setae on frons: 13-88 (48 ± 24) p, tips incrassate to sharp. Head width: 245-329 (291 ± 22) p. Length of stylets: 714-1130 (860 ± 107) p\ ultimate rostral segment: 58-100 (80 ± 9) p, rostral tip reaching abdominal terga III-V in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 316-459 (385 ± 37) p. THORAX: Meso + metanota fused, total length: 275-439 (371 ± 46) p. ABDOMEN: Tergum I free, length: 102-148 (123 ± 13) p\ terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 28-45 (35 ± 5) p; siphunculi flush to truncated conical, protrusion to 0.7 x maximal distal width. Ventral abdominal sclerites on segments III-IV usually irregular, subcircular or sublinear when small (length less than 0.6 x metatibial diameter), subquadrate when large (length greater than 1.0 x metatibial diameter); length: 5-51 (22 ± 11) p, 0.3-1.1 x diameter of metatibiae. Dorsal (major + minor) setae (see Fig. IF) on abdominal terga III-IV: 6, very rarely 7, tips sharp, in 1 row; marginal setae 2 per segment, each side. Setae on abdominal tergum VIII: 6, rarely 7, length: 13-60 (31 ± 15) p, tips incrassate to sharp, in 1 row. Cauda rounded; caudal pro¬ tuberance moderately to poorly developed, occasionally absent; length of longest caudal setae: 43- 105 (78 ± 17) p, tips sharp. LEGS: Length of metafemora: 388-857 (596 ± 106) p\ metatibiae: 561- 1275 (908 ± 179) p\ longest dorsal setae on central one-third of metatibiae: 10-113 (38 ± 26) p, 0.1- 2.9 x diameter of metatibiae, tips incrassate to sharp; approximately equal or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae: 18-50 (34 ± 10) p, tips sharp. Length of metabasitarsus: 93-148 (118 ± 15) p\ metadistitarsus: 148-223 (187 ± 18) p. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.58:1. Pigmentation: Color in life: Thorax gray-green, abdomen lime green, legs yellow-brown. Slide-mounted specimens: Background of body dorsum usually pale to rarely light brown (to 20 percent pigment density), usually unicolorous, oc¬ casionally abdominal terga dorsomedially dusky to entire abdomen evenly moderate brown (to 50 percent pigment density) in contrast to pale head and thorax. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga, to rarely subtly darker. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen usually pale, inconspicuous, to rarely light brown, vaguely evident. Spiracular plates and ventral abdominal sclerites pale, inconspicuous, to moderate brown, conspicuous. Siphunculi concolorous with surrounding terga. 64 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Cauda, anal, and subgenital plates concolorous with abdominal terga to subtly darker. Antennal segments Y and IV subtly dusky distally, pale proximally, to entirely dusky, infrequently to moderate brown; III entirely very pale to subtly dusky on distal one-third, remainder pale; II as pale as proximal III to subtly darker; I concolorous with frons. Pro-, meso- and metatibiae concolorous, usually evenly pale, equivalent to thoracic tergum or slightly darker, to subtly dusky on distal and occasionally proximal tip, rarely entirely dusky. Distitarsi usually dusky on distal one-half, pale proximally, to entirely dusky with distal tip of tibiae. Ultimate Stadium Nymphs of Viviparous Apterae. — Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum lacking 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; area surrounding muscle attachment sites membranous. Viviparous Alatae. - Slide-mounted specimens: Nonmorphometrics as described for viviparous ap¬ terae except lacking body dorsum pigmentation suite; abdominal terga normally membranous, dorsal setae between muscle attachment plates very rarely arising from distinct scleroites; antennal segments often as dark as tibiae, except proximal one-fifth of III pale. Antennal segment III with 0-5, IV with 0-3, secondary rhinaria. Epicranial suture strongly developed. Forewing medius with single furcation arising on proximad one-third of vein; cubital base arising proximad on subcosta with distance between anal and cubital bases on subcosta relatively small, ca. 20-30 percent or less of anal vein length; medius, especially cubitus and anal veins usually distinct, except infrequently proximad 10-15 percent vague. Abdominal terga lacking irregular sclerites that engulf or join muscle attachment plates and dorsal (major + minor) setal bases or scleroites. Oviparae. — Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae ex¬ cept abdominal dorsum membranous with irregular transverse sclerites containing dorsal (major + minor) setae on each tergum; marginal setae usually on separate scleroites; siphuncular cones scler¬ otized, regular, separated from other dorsal sclerotic fields; dorsal abdominal muscle attachment plates unicolorous. Rarely with abdominal terga II-VI sclerotic/fused, terga VII and VIII free; then dorsal demarcations of anterad terga evident and siphuncular cones surrounded closely by, sometimes en¬ gulfed by, sclerotic fields. Pseudorhinaria on metatibiae irregular, 11-23. Males, Fundatrices.— Unknown. Diagnosis. —Essigella (E.) hoerneri requires the combination of several char¬ acters for identification; individuals usually are pale and may be confused with other pale Essigella. Essigella (E.) hoerneri can be separated from all Essigella, except E. (E.) californica and E. (E.) pini by having six (Fig. IF), instead of eight or more, dorsal (major + minor) setae on abdominal terga III-IV. Diagnostics for all morphs of E. ( E .) hoerneri that separate it from E. ( E .) pini are the same as for E. (E.) californica [see that diagnosis]. Essigella (E.) hoerneri and E. (E.) californica are difficult to distinguish. Al¬ though E. ( E .) hoerneri has a longer rostrum, wider head and shorter antennal segment IV than does E. ( E .) californica, these differences are indiscrete, and reliable separation requires application of the discriminant function in the key to the viviparous apterae [couplets 27 or 28, see 26]. Synonyms.—Essigella gillettei Flottes, NEW SYNONYM: holotype, vivip. alat., on slide alone; data: COLORADO. LARIMER Co.: Stove Prairie Hill, Bellevue, 16 Jun 1922, M. A. Palmer, P. murrayana [= P. contorta latifolial ]. Essigella gillettei holotype deposited in the NMNH. Essigella maculata Hottes, NEW SYNONYM: holotype, vivip. alat., on slide alone; data: COLORADO. MESA Co.: Grand Junction, 2 Sep 1956, Pinus edulis. Essigella maculata holotype deposited in the NMNH. Range. — Great Basin, from the Sierra Nevada to the Rocky Mountains, south of Idaho and Wyoming to Arizona, New Mexico and southern California; pre¬ sumably into Mexico following its hosts (Fig. 9). 1994 SORENSEN: A REVISION OF ESSIGELLA 65 Figure 9. Distribution of is. ( E .) hoerneri [dots (JTS samples), squares (nonJTS samples)], super¬ imposed over the range of its hosts, Pinus monophylla [darker shading] and Pinus edulis [lighter shading (UT, AZ and east)]. Hosts. —Pinus section Parrya, subsection Cembroides: P. edulis Engelmann, P. monophylla Torrey & Fremont, P. cembroides Zuccagni and P. quadrifolia Par- latore [see discussion]. Essigella ( E .) hoerneri is the only Essigella regularly on piny on pines [E. (L .) fusca has rarely been taken on pinyons, but is considered nonresident]. References to E. (E.) hoerneri on P. flexilis and P. ponderosa are probably erroneous, or at least nonresident; although I have not yet seen the slides, I suspect they represent E. (E .) californica, or in the case of ponderosa pine possibly E. (. L .) fusca. Discussion. —Essigella (E.) hoerneri is closely related to E. (E.) californica\ see the discussion of that species, where most comments apply equally to E. (E.) hoerneri ]. The long rostrum and styli of E. (E.) hoerneri are autapomorphic; although, within other Essigella species complexes, some species may have these features slightly lengthened in comparison to their close relatives, that lengthening is not in the same class as here. In E. ( E .) hoerneri, this appears to be an adaptation to feeding on pinyons, whose needles are relatively thick. Their needle thickness results from a failure to split into the multiple needles (Mirov 1967) that normally arise from a needle fascicle (e.g., Pinus monophylla). There is an east-west gradient for stylet length in this aphid, which appears to roughly reflect the needle diameter of the hosts. The more eastern populations of E. (. E.) hoerneri (Colorado, New Mexico) have a relatively shorter rostrum, and the rostrum reaches maximal 66 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) relative length in the populations of the Owens Valley area of California, and western Nevada. This reflects their host pine geography and needle diameter: Pinus monophylla, a single-needled pinyon with a large diameter needle, occurs west of the Nevada-Utah border; whereas P. edulis, a double-needled pinyon with needles of proportionately less diameter, occurs to the east. The exact species status of these two pines may be questionable (W. B. Critchfield, personal com¬ munication). Essigella (E.) hoerneri has apparently split (Fig. 13: node 10) from the E. (E.) californica lineage to ecologically reinvade Pinus (Strobus ), on subsection Cem- broides pines. These section Parrya pine niches are unoccupied by other Essigella. The Archeoessigella species feed monophagously within Pinus (Strobus), but in section Strobus, subsection Strobi; E. (E.) californica and E. (E.) pini also feed in that subsection in polyphagy; further, E. (E.) californica occurs on P. albicaulis, the sole Nearctic representative of Pinus (Strobus) section Strobus, subsection Cembrae. Interestingly, the other section Parrya subsections are not occupied by Essigella : subsection Gerardianae is Asian, but the Nearctic subsection Balfour- ianae probably predates Essigella (see Ecological Corroboration of tl). Coded References to This Taxon. —Essigella (E.) hoerneri has been referred to previously by: the coding “Sp. B” (Sorensen 1983, 1987a, 1992b) and “HOER” in (Sorensen 1983), and by the name E. hoerneri in Sorensen (1983). Etymology and Common Name.— This species was apparently named after J. L. Hoerner, who collected the series upon which the original description was based (Gillette & Palmer 1924: 5). Common name: Hoemer’s pinyon pine needle aphid; although Palmer (1952: 15) refers to this species as “The Immaculate Pine Needle Aphid,” the common name indicated here is more appropriate and less confusing because other Essigella are immaculate, in the sense of lacking “speckles” [see etymology for E. (E.) pini]. Material Examined.— ARIZONA. APACHE Co.: 5 km W of Eagar on hwy 273, 2140 m, 11 Sep 1978, JTS 78115, P. edulis, (apt.); 6 km N of Lupton on hwy 12 (= 166), 1980 m, 11 Sep 1978, JTS 78116, P. edulis, (apt., alat.). COCHISE Co.: Miller Canyon Rd, Huachuca Mts, 1700 m, 17 Sep 1978, JTS 78153, P. cembroides, (apt.). COCONINO Co.: 22 km N of Williams on hwy 64, 2070 m, 9 Sep 1978, JTS 7817, P. edulis, (apt.); 32 km S of Grand Canyon Village on hwy 180, 2070 m, 9 Sep 1978, JTS 7819, P. edulis, (apt.); 8 km W of Grand Canyon Caverns on hwy 60, 1700 m, 9 Sep 1978, JTS 7813, P. edulis, (apt.). CALIFORNIA. INYO Co.: Bristlecone Pine Forest entrance on hwy 168, Inyo Natl Forest, W of Westgard Pass, 2230 m, 31 Jul 1977, JTS 77G74, P. monophylla, (apt.); jet of Lake Sabrina Rd & Southern Calif. Edison Plant 2 Rd, nr Bishop, 2130 m, 1 Aug 1977, JTS 77H3, P. monophylla, (apt., alat.). KERN Co.: Valle Vista Cmpgd, 13 km W of Apache Saddle Ranger Station, 1500 m, 18 Sep 1977, JTS 77156, P. monophylla, (apt.); W side of Walker Pass on hwy 178, 26 km E of Oyx, 1530 m, 20 Sep 1977, JTS 77163, P. monophylla, (apt., alat.). MONO Co.: Cedar Flat, nr White Mt, 15 Jul 1961, E. I. Schlinger, EIS 61-7-15h, “Pinon pine,” (apt.); E side of Monitor Pass on hwy 89, 2070 m, 17 Jul 1977, JTS 77G36, P. monophylla, (apt., alat.); Lee Vining Cmpgd, Inyo Natl Forest, W of Tioga Pass on hwy 120, 2290 m, 31 Jul 1977, JTS 77G70, P. monophylla, (apt.); Sherwin Summit, 17 Jul 1972, D. J. Voegtlin, DJV 47, P. monophylla, (apt.); Topaz Lake, 1680 m, 5 Jul 1979, S. Paulaitis, DJV 541, P. monophylla, (apt.). RIVERSIDE Co.: 2 km N of Paradise Valley on hwy 74, 1500 m, 9 Sep 1977, JTS 77119, P. quadrifolia, (apt.); Alpine Village, 21 km S of Palm Desert on hwy 74, 1160 m, 9 Sep 1977, JTS 77117, P. monophylla, (apt.); Joshua Tree Natl Monument, Key’s View, 1530 m, 12 Sep 1977, JTS 77133, P. monophylla, (apt.). SAN BERNARDINO Co.: 16 km W of Barton Flat on hwy 38, 2140 m, 16 Sep 1977, JTS 77140, P. monophylla, (apt.); Pipes Cyn, NW of Yucca Valley & Pioneer Town, 8 km NW of jet of Pioneertown Rd & Pipes Canyon Rd, 1530 m, 15 Sep 1977, JTS 77134, P. monophylla, (apt., alat.); Sheep Cyn, 2 km NW of Mountain Top Jet on hwy 138, 1525 m, 17 Sep 1977, JTS 77147, P. monophylla, (apt.); nr Ivanpah, New York Mts, 1994 SORENSEN: A REVISION OF ESSIGELLA 67 1600 m, 8 Sep 1978, JTS 7812, P. monophylla, (apt.); same but 1700 m, JTS 7811, P. edulis, (apt.). VENTURA Co.: Cuyama Valley, 22 May 1959, E. I. Schlinger, EIS 59-5-23i, P. “ cembroides ” [?], (apt., alat.); Lake of the Woods, 10 km W of Tejon Pass, 1556 m, 18 Sep 1977, JTS 77153, P. monophylla, (apt., alat.). COUNTY UNCERTAIN: Frazier Park, 22 May 1959, E. I. Schlinger, EIS 59-5-23d, P. “ cembroides ” [?], (apt., alat.) (slide is labeled “Los Angeles” Co., but Frazer Park is in Kern Co.; however, main roads in NW L.A. Co. are less than ~5 km away and Ventura Co. is also immediately adjacent). COLORADO. CHAFFE Co.: Poncha Springs, 2440 m, 12 Aug 1978, JTS 78FI69, P. edulis, (apt.). LARIMER Co.: (lectotype) Owl Cyn, 25 Sep 1921, C. P. Gillette & J. Hoemer, CAES 2894, P. edulis, (apt.); (paratype) same but 10 Oct 1921, C. L. Corkins, CAES 3028/USNM 41952, (apt., ovip.); (paratype) same but 6 Nov 1921, J. Hoemer, CAES 3035, (ovip.). MESA Co.: Grand Junction, 2 Sep 1956, P. edulis, (alat.); same but 3 Oct 1956, F. C. Hottes, (alat.). SAN MIGUEL Co.: 6 km NE of Placerville on hwy 62, 2320 m, 7 Aug 1978, JTS 78H41, P. edulis, (apt.). COUNTY UNCERTAIN: Refe, 17 Aug 1956, P. edulis, (apt.). NEVADA. CLARK Co.: Charleston Mts, Lee Canyon Rd (hwy 52), 2290 m, 4 Aug 1978, JTS 78H14, P. monophylla, (apt.); same but Lee Canyon Ski Area, 2590 m, JTS 78H18, P. ponderosa, (alat.); W of Las Vegas, 20 Apr 1978, C. F. Smith, CFS 78-56, P. monophylla, (apt., alat.). DOUGLAS Co.: hwy 395, 16 km NW of jet with hwy 3, Pine Nut Mts, 1650 m, 16 Jul 1977, JTS 77G35, P. monophylla, (apt.). LANDER Co.: Scott Summit on hwy 50, 11 km E of Austin, 2230 m, 26 Aug 1978, JTS 78H158, P. monophylla, (apt., alat.). WASHOE Co.: Mt Rose, Slide Mountain Ski Area, 2 Aug 1978, JTS 78H6, P. washoensis, (alat.). WHITE PINE Co.: Leyland Cave Natl Monument, 2074 m, 26 Aug 1978, JTS 78H156, P. monophylla, (apt.); Little Antelope Summit on hwy 50, 56 km E of Eureka, 2260 m, 26 Aug 1978, JTS 78H157, P. monophylla, (apt.). NEW MEXICO. BERNALILLO Co.: Crest View, hwy 14, 2280 m, 12 Sep 1978, JTS 78121, P. edulis, (apt., alat., ovip.). RIO ARRIBA Co.: 8 km S of Tierra Amarilla on hwy 84, 2410 m, 8 Aug 1978, JTS 78H52, P. edulis, (apt., alat.). SANTA FE Co.: 20 km NE of Santa Fe on hwy 475, 2680 m, 10 Aug 1978, JTS 78H57, P. edulis, (apt.). SIERRA Co.: 2 km E of Kingston on hwy 90, 1860 m, 14 Sep 1978, JTS 78131, P. edulis, (apt.). OKLAHOMA. CIMARRON Co.: Kenton, 16 May 1961, Van Cleave, “Pinon pine,” (apt., alat.). UTAH. DAGGETT Co.: Flaming Gorge Dam, Dutch John, 22 Jun 1978, C. S. Smith, CFS 78-238, “pinyon pine,” (apt., alat.) (slides of this series marked Butch John and Dutch John, Wyoming). DUCHESNE Co.: Starvation lake, hwy 40, 1800 m, 25 Aug 1978, JTS 78H140, P. edulis, (apt.). GARFIELD Co.: hwy 20, 5 km W of jet with hwy 89, 2040 m, 6 Aug 1978, JTS 78H32, P. edulis, (apt.). SEVIER Co.: 35 km E of Salina on hwy 70, 2130 m, 6 Aug 1978, JTS 78H33, P. edulis, (apt., alat.). WASHINGTON Co.: 43 km SW of Cedar City on hwy 15, 1220 m, 5 Aug 1978, JTS 78H22, P. monophylla, (apt.). WAYNE Co.: 2 km NE of La Sal on hwy 46, 2230 m, 6 Aug 1978, JTS 78H36, P. edulis, (apt.). Essigella (Essigella) wilsoni Hottes, 1957 Essigella wilsoni Hottes, 1957: 106, Proc. Biol. Soc. Wash., 70: 106-108. Essigella pergandei Hottes, 1957: 100, Proc. Biol. Soc. Wash., 70: 100. NEW SYNONYM. Essigella oregonensis Hottes, 1958: 155, Proc. Biol. Soc. Wash., 71: 155-156. NEW SYNONYM. Primary Type. — Holotype, vivip. apt., on slide with 6 other specimens, holotype shown by arrows (12 o’clock position); data: “Douglas Fir, Whitby Isd., Seattle, Wash., Aug 29, 1955, M. J. Forsell, Coll./Holotype, Essigella wilsoni F.C. Hottes” (reference to “Whitby Isd.” may be to “Whidbey Island” in Island Co.; Seattle is in King Co.). Holotype deposited in the Essig Museum of Entomology, Uni¬ versity of California at Berkeley, Berkeley, California. Viviparous Apterae.—Morphology: Body length: 1.24-2.03 (1.49 ± 0.22) mm. HEAD: Primary rhinarium on terminal antennal segment (V) exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim less than 0.5, usually 0.3, x diameter of rhinarium; distal face of rhinarial rim usually perpendicular to longitudinal axis of antennal segment; rhinarial membrane usually conspicuously protuberant. Length of antennal segment V: 83-113 (101 ± 8) n, processus terminalis: 20-40 (31 ± 5) ju; IV: 66-95 (82 ± 9) n\ III: 83-183 (133 ± 23) n\ II: 55-80 (66 ± 6) fi. 68 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Figure 10. Distribution of E. ( E .) wilsoni [dots (JTS samples), squares (nonJTS samples)], super¬ imposed over the range of its hosts, Pseudotsuga menzeisii [darker shading] and Pseudotsuga macro¬ carp a [lighter shading (Santa Barbara Co. and south in CA)]. Length of longest setae on frons: 13-35 (22 ± 6) g, tips incrassate to sharp. Head width: 225-286 (248 ± 14) g. Length of stylets: 408-653 (525 ± 64) /x; ultimate rostral segment: 43-73 (64 ± 7) g, rostral tip reaching abdominal terga I—II, infrequently III, in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 281-428 (332 ± 37) g. THORAX: Meso + metanota fused, total length: 204-357 (265 ± 35) g. ABDOMEN: Tergum I free, length: 71-153 (99 ± 20) g\ terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 31-48 (38 ± 5) g; siphunculi flush to truncated conical, protruding to 0.5 x maximal distal width. Ventral abdominal sclerites on segments III-IV usually irregular, subcircular when small (length less than 0.6 x metatibial diameter), to subquadrate when large (length greater than 1.0 x metatibial diameter); length: 5-53 (27 ± 15) m, 0.1-1.3 x diameter of metatibiae. Dorsal (major + minor) setae (see Fig. IE) on abdominal terga III-IV: usually 8, occasionally 10, very rarely 12, tips sharp, when 8 setae then in 1 row, infrequently single mesad or lateral-most minor dorsal seta anterad, when 10 or more then in 2 irregular rows, usually with lateral-most minor dorsal seta on each side anterad to its next mesad neighbor; marginal setae 2 per segment, each side. Setae on abdominal tergum VIII: 6, rarely to 8, length: 8-28 (12 ± 5) g, tips incrassate to sharp, in 1 row. Cauda rounded; caudal protuberance 1994 SORENSEN: A REVISION OF ESSIGELLA 69 moderately to poorly developed, infrequently absent; length of longest caudal setae: 45-88 (58 ± 11) li, tips sharp. LEGS: Length of metafemora: 393-592 (466 ± 52) metatibiae: 561-836 (682 ± 78) m; longest dorsal setae on central one-third of metatibiae: 10-35 (18 ± 7) u, 0.05-1.5 x diameter of metatibiae, tips incrassate to sharp; approximately equal or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae: 15-38 (24 ± 6) p, tips sharp. Length of metabasitarsus: 79-110 (94 ± 8) ju; metadistitarsus: 125-180 (155 ± 16) p. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.65:1. Pigmentation: Color in life: Lime green throughout. Slide-mounted specimens: Background of body dorsum pale (to 10 percent pigment density), unico- lorous. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen pale, inconspicuous. Spiracular plates and ventral abdominal sclerites pale to infrequently light brown. Siphunculi concolorous with surrounding terga. Cauda, anal and subgenital plates pale, concolorous with abdominal terga, to subtly darker. Antennal segments V and IV dusky, concolorous; III entirely pale, to distal one-half dusky, remainder pale; II concolorous with proximal III; I con¬ colorous with frons, to subtly darker. Pro-, meso- and metatibiae usually pale, concolorous and equivalent to body dorsum, infrequently entire tibiae slightly dusky, subtly darker than body dorsum. Distitarsi entirely pale to subtly dusky on distal one-third. Ultimate Stadium Nymphs of Viviparous Apterae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum lacking 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; area surrounding muscle attachment sites membranous. Viviparous Alatae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous ap¬ terae except lacking body dorsum pigmentation suite; abdominal terga normally membranous, dorsal (major + minor) setae between muscle attachment plates occasionally arising from distinct scleroites; antennal segments often as dark as tibiae, except proximal one-fifth of III pale. Antennal segment III with 1, IV with 0, secondary rhinaria. Epicranial suture absent to vaguely developed. Forewing medius with single furcation arising on proximad one-third of vein; cubital base usually arising distad, infre¬ quently proximad, on subcosta with distance between anal and cubital bases on subcosta usually relatively large, ca. 30-40 percent or more of anal vein length; medius, especially cubitus and anal veins distinct, except infrequently proximad 10-15 percent vague. Abdominal terga lacking irregular sclerites that engulf or join muscle attachment plates and dorsal (major + minor) setal bases or scleroites. Oviparae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae ex¬ cept abdominal dorsum membranous with faint, irregular transverse sclerites containing dorsal (major + minor) setae on each tergum; marginal setae usually on separate faint scleroites; siphuncular cones sclerotized, regular, separated from other dorsal sclerotization fields; dorsal abdominal muscle at¬ tachment plates faint, unicolorous. Pseudorhinaria on metatibiae irregular, 9-15. Males, Fundatrices.— Unknown. Diagnosis. —Essigella (E.) wilsoni can be identified by the unique primary rhi- narium that is unusually protuberant and exceptionally close to the tip of the antennal segment V. This species is pale. Synonyms. —Essigella oregonensis Hottes, NEW SYNONYM: holotype, vivip. apt., on slide with 1 male; data: OREGON. CLACKAMAS Co.: Government Camp, 17 Aug 1957, Pinus albicaulis. Essigella oregonensis holotype deposited in the NMNH. Essigella pergandei Hottes, NEW SYNONYM: holotype, vivip. apt., several specimens on slide, holotype circled; data: WASHINGTON. KING Co.: Seattle, 17 Jul 1955, M. J. Forsell, “ Abies concolor ” (Gordon) Lindberg [reference to “ Abies concolor ” presumably is a mistaken identification of Pseudotsuga men- ziesii, see host discussion below]. Essigella pergandei holotype deposited in the NMNH. Range. — Southern British Columbia and Alberta, throughout the western U.S. (exclusive of Alaska), presumably south into Mexico as far as its hosts (Fig. 10). 70 THE PAN-PACIFIC ENTOMOLOGIST Yol. 70(1) Hosts. —Pseudotsuga menziesii (Mirbel) Franco and Pseudotsuga macrocarpa (Vasey) Mayr. Many museum slides (other collectors), are labeled Abies concolor (e.g., E. pergandei holotype), or simply “fir.” Also, there is one record ( E. ore- gonensis holotype) from Pinus albicaulis, which is probably opportunistic. Essi- gella (E.) wilsoni is a commonly collected species that has transferred exclusively to a host other than Pinus. Numerous records from Abies are most probably in error; my extensive sampling (Sorensen 1983) on Abies did not yield any Essigella. Whenever I sampled E. ( E .) wilsoni from what I thought to be an Abies, there was invariably an adjacent Pseudotsuga with a branch intermingled that proved to be the host. If Abies is a host, it is very much less commonly used than Pseudotsuga. Discussion.—Essigella (E.) wilsoni is common and morphologically homoge¬ neous. The condition of the primary rhinarium on antennal segment V is an autapomorphy. Its other apomorphies are listed in the discussion of E. (E.) Cal¬ ifornia. The length of dorsal setae on the metatibiae varies somewhat similarly to, but not to the extent of, E. (E.) californica. The male of E. (E.) wilsoni is unknown. The morphotype male [synonym E. pergandei ] is too poorly mounted and positioned to determine its species; its primary rhinarium is not characteristic of E. (E.) wilsoni, and it may be a male E. (E.) calif ornica. The rhinarial difference may be a sexual character difference or preparation artifact, however? The phylogenetic placement of E. (. E.) wilsoni is confusing; see alternative analyses in the phylogenetics section. Ordinations (Sorensen 1992b) place it closest to E. (. E .) pini and E. ( E.) essigi in discriminant space, but conventional (coded data) cladistic analyses suggest it forms a trichotomy with the E. ( E .) californica clade and series B (unpublished data). On the basis of many bivariant regression plottings (unpublished data), I feel E. (E.) wilsoni shows the closest relationship to the E. (E.) californica complex. Many traits that I discount as homoplasies suggest a relationship to E. ( E .) alyeska; however, I believe the broad body of the latter is a strong synapomorphy linking it to the E. ( E .) knowltoni complex, as is its host association. Coded References to This Taxon.—Essigella (E.) wilsoni has been referred to previously by: the coding “Sp. C” (Sorensen 1983, 1987a, 1992b) and “WILS” (Sorensen 1983), and by the name E. wilsoni in Sorensen (1983). Etymology and Common Name. — Hottes (1957: 107) named this species after aphidologist H. F. Wilson, who described the second named Essigella species, E. ( E.) pini. Hottes apparently attributed the naming to Wilson’s (1919: 1) mention of “j E. californica (Essig)” from Pseudotsuga “douglasii ” [= P. menziesii]. He probably (correctly) deduced, from mention of that host, that his own “ E. wilsoni ” was involved, because Hottes undoubtedly could not make such a deduction from the description or characters mentioned by Wilson. Common name: Wilson’s Douglas fir needle aphid. Material Examined.—ARIZONA. COCHISE Co.: nr Rustler park, Chiricahua Mts, 2500 m, 16 Sep 1978, JTS 78151, Pseudotsuga menziesii, (apt.). GILA Co.: 32 km E of Kohles Ranch on hwy 260, 10 Sep 1978, JTS 78112, Pseudotsuga menziesii, (apt.). GRAHAM Co.: 34 km SW of Stafford 1994 SORENSEN: A REVISION OF ESSIGELLA 71 on hwy 366, 2170 m, 15 Sep 1978, JTS 78141, Pseudotsuga menziesii, (apt.). CALIFORNIA. EL DORADO Co.: Blodgett Experimental Forest (Univ. Calif.), E of Georgetown, 26 Jul 1973, D. J. Voegtlin, DJV 57, Pseudotsuga menziesii, (apt.); same but 28 Apr 1977, J. T. Sorensen, (apt.). HUM- BOLT Co.: Lord Ellis Summit on hwy 299, W of Willow Creek, 670 m, 21 Aug 1977, JTS 77H25, Pseudotsuga menziesii, (apt.). LAKE Co.: 18 km W of Lake Pillsbury, Eel River Rd, 490 m, 24 Jul 1977, JTS 77G55, Pseudotsuga menziesii, (apt.). LOS ANGELES Co.: hwy 2, 15 km NE of jet with hwy 39, San Gabriel Mts, 2290 m, 17 Sep 1977, JTS 77150, Pseudotsuga macrocarpa, (apt.). MARIN Co.: Alpine Lake, 25 Mar 1977, J. T. Sorensen, Pseudotsuga menziesii, (apt.); Muir Woods, 27 Mar 1964, C. F. Smith & Graham, CFS 64-27, “ Abies ” [assumed erroneous], (apt.). MENDOCINO Co.: Fish Rock Rd, 27 km E of hwy 1, 490 m, 23 Jul 1977, JTS 77G50, Pseudotsuga menziesii, (apt.); Nature Conservancy Coastal Redwood Preserve, 8 km N of Branscomb, 13 May 1978, JTS 78E103, Pseudotsuga menziesii, (apt.). PLUMAS Co.: Jackson Creek Cmpgd, Plumas Natl Forest, 2 km SE of Cromberg on hwy 70/89, 1280 m, 26 Jun 1977, JTS 77F11, Pseudotsuga menziesii, (apt.). SAN BERNARDINO Co.: 3 km NE of Lake Gregory Village, San Bernardino Mts, 1310 m, 17 Sep 1977, JTS 77146, Pseudotsuga macrocarpa, (apt.); 8 km W of Barton Flat on hwy 38, 1920 m, 16 Sep 1977, JTS 77137, Pseudotsuga macrocarpa, (apt.). SAN DIEGO Co.: Mt Palomar Rd (S6), 5 km S of Mt Palomar, 1370m, 11 Sep 1977, JTS 77126, Pseudotsuga macrocarpa, (apt.). SISKIYOU Co.: Deadhorse Summit on hwy 89, 10 km SE of Bartel, 1370 m, 3 Jul 1977, JTS 77G11, Pseudotsuga menziesii, (apt.); Mt Shasta Ski Bowl Rd, 2450 m, 2 Jul 1977, JTS 77G7, Pseudotsuga menziesii, (apt.). TEHAMA Co.: 45 km E of Dales on hwy 36, 1460 m, 10 Jul 1977, JTS 77G26, Pseudotsuga menziesii, (apt.). TRINITY Co.: 3 km W of Weaverville on hwy 299, 730 m, 20 Aug 1977, JTS 77H22, Pseudotsuga menziesii, (apt.); Buckhorn Summit on hwy 299, W of Tower House, 980 m, 20 Aug 1977, JTS 77H18, Pseudotsuga menziesii, (apt.); Ironside Mt Lookout Rd, W of Junction City, 1070 m, 21 Aug 1977, JTS 77H24, Pseudotsuga menziesii, (apt.). TUOLUMNE Co.: Yosemite Natl Park, hwy 120 entrance, 1700 m, 1 Aug 1977, JTS 77H7, Pseudotsuga menziesii, (apt.). VENTURA Co.: 5 km NNE of Pine Mt Summit on hwy 33, 1340 m, 19 Sep 1977, JTS 77157, Pseudotsuga macrocarpa, (apt.); Pine Mt Summit, 16 May 1961, R. Van den Bosch & J. Hall, RVdB 61-V-19L, “white fir” [assumed erroneous], (apt.). COLORADO. GUNNISON Co.: 16 km NW of Kebler Pass, 2440 m, 13 Aug 1978, JTS 78H76, Pseudotsuga menziesii, (apt.). SAN MIGUEL Co.: 6 km NE of Placerville on hwy 62, 2320 m, 7 Aug 1978, JTS 78H42, Pseudotsuga menziesii, (apt.). IDAHO. BONNER Co.: 6 km S of Cocolalla on hwy 95, 18 Jul 1978, JTS 78G106, Pseudotsuga menziesii, (apt.). MONTANA. GALLATIN Co.: Battle Ridge Pass, Bridger Mts, S of Bozeman, 19 Aug 1979, D. J. Voegtlin, DJV 702, Pseudotsuga menziesii, (apt.). LINCOLN Co.: 4 km S of Stryker on hwy 93, nr Flathead Co. line, 17 Jul 1978, JTS 78G96, Pseudotsuga menziesii, (apt.). NEW MEXICO. OTERO Co.: 3 km W of Cloudcroft on hwy 82, 2560 m, 13 Sep 1978, JTS 78124, Pseudotsuga menziesii, (apt.). SANTA FE Co.: 30 km NE of Santa Fe on hwy 475, 3110 m, 10 Aug 1978, JTS 78H54, Pseudotsuga menziesii, (apt.). OREGON. BENTON Co.: Corvallis, 25 Jan 1915, L. Childs, Pseudotsuga menziesii, (apt.). CLACKAMAS Co.: Government Camp, 17 Aug 1958, P. albicaulis, (ovip.). JOSEPHINE Co.: O’brien, 4 Jul 1978, JTS 78G10, Pseudotsuga menziesii, (apt.). POLK Co.: 6 km W of Grand Ronde on hwy 18, 7 Jul 1978, JTS 78G41, Pseudotsuga menziesii, (apt.). WASCO Co.: 46 km SE of Government Camp on hwy 26, 670 m, 6 Jul 1978, JTS 78G30, Pseudotsuga menziesii, (apt.). WASHINGTON Co.: 21 km W of Manning on hwy 26, 7 Jul 1978, JTS 78G48, Pseudotsuga menziesii, (apt.). UTAH. DUCHESNE Co.: 19 km NE of Castle Lake on hwy 33, 2780 m, 25 Aug 1978, JTS 78H146, Pseudotsuga menziesii, (apt.). IRON Co.: 16 km SE of Cedar City on hwy 14, 2170 m, 5 Aug 1978, JTS 78H23, Pseudotsuga menziesii, (apt.). SEVIER Co.: 66 km E of Salina on hwy 70, 2227 m, 6 Aug 1978, JTS 78H34, Pseudotsuga menziesii, (apt.). WASHINGTON. KING Co.: Seattle, 17 Jul 1955, J. W. Forsell, “Abies concolor" [assumed erroneous], (apt.); same but 25 Apr 1973, D. Pike, Pseudotsuga menziesii, (apt.). KITSAP Co.: 8 km S of Hood Canal bridge on hwy 3, 9 Jul 1978, JTS 78G50, Pseudotsuga menziesii, (apt.). OKANOGAN Co.: 17 km NW of Winthrop on hwy 20, 550 m, 12 Jul 1978, JTS 78G72, Pseudotsuga menziesii, (apt.); Loup Loup Pass, Okanogan Natl Forest, 19 Sep 1979, D. J. Voegtlin, DJV 759, Pseudotsuga menziesii, (apt.). PIERCE Co.: hwy 706, nr Ashford, 11 Jul 1978, JTS 7%G51, Pseudotsuga menziesii, (apt., alat.). COUNTY UNCERTAIN: [see primary type paragraph] (type) “Whitby Island,” “Seattle,” 29 Aug 1955, M. J. Forsell, (apt.). WYOMING. TETON Co.: 35 km SE of Jackson on hwy 187, 1950 m, 23 Aug 1978, JTS 78H130, Pseudotsuga menziesii, (apt.). CANADA. BRITISH CO¬ LUMBIANS km SofRadium Hot Springs on hwy 93, 17 Jul 1978, JTS 78G90, Pseudotsuga menziesii, (apt.). 72 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Series B Essigella ( Essigella ) alyeska Sorensen, 1988 Essigella alyeska Sorensen, 1988: 118, Pan-Pacif. Entomol., 64: 118-121. Essigella “ alyeska ” Sorensen, 1983: 112 (unpublished manuscript name) Ph.D. Thesis, University of California at Berkeley, Berkeley, California. 605 p. Type Series. — Holotype, vivip. apt., on slide with 1 paratype vivip. apt., ho- lotype on top (12 o’clock position); data: ALASKA. FAIRBANKS NORTH STAR BOROUGH: College (Univ. Alaska campus), nr Fairbanks, 24 Jun 1979, J. T. Sorensen (79F1), Picea glauca. Holotype retained in Sorensen collection, even¬ tually to be deposited in The Natural History Museum, London. Paratypes (all same data as holotype): 25 vivip. apt. on 13 slides including holotype slide, Paratype slides deposited: 1 slide in NMNH, Washington, D.C.; 1 slide in CNC, Ottawa, Ontario; 11 slides in Sorensen collection. Viviparous Apterae.— Morphology: Body length: 1.42-1.65 (1.51 ± 0.07) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminals to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously protuberant. Length of antennal segment V: 100-120 (108 ± 8) p, processus terminalis: 28-38 (34 ± 4) m; IV: 83-98 (86 ± 5) p- HI: 138-170 (151 ± 11) IE 63-73 (67 ± 3) p. Length of longest setae on frons: 33-53 (41 ± 7) p, tips incrassate, rarely sharp. Head width: 286-301 (292 ± 301) p. Length of stylets: 561-775 (600 ± 69) p\ ultimate rostral segment: 63-85 (74 ± 8) p, rostral tip reaching abdominal terga I—II in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 337-388 (361 ± 16) p. THORAX: Meso + metanota fused, total length: 265-316 (298 ± 17) p. ABDOMEN: Tergum I free, length: 102-118 (108 ± 7) p\ terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 43-48 (46 ± 2) p\ siphunculi strongly protuberant, protruding 0.7-1.1 x maximal distal width. Ventral abdominal sclerites on segments III-IV irregular, to subcircular when large; length: 26-40 (35 ± 5) p, 0.8-1.4x diameter of metatibiae. Dorsal (major + minor) setae (see Fig. IE) on abdominal terga III-IV: 7-9, usually 8, tips sharp, in 1 row; marginal setae 2 each side, per segment. Setae on abdominal tergum VIII: 6-8, length: 15-45 (36 ± 10) p, tips incrassate to sharp, in 1 row. Cauda broadly rounded; caudal protuberance poorly developed to absent; length of longest caudal setae: 83-100 (91 ± 7) p, tips sharp. LEGS: Length of metafemora: 428-520 (488 ± 33) p\ metatibiae: 663-785 (731 ± 44) p; longest dorsal setae on central one-third of metatibiae: 30-45 (38 ± 5) p, 0.7-1.5 x diameter of metatibiae, tips incrassate, rarely sharp; approximately equal or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae: 23-33 (28 ± 7) p, tips sharp. Length of metabasitarsus: 95-103 (99 ± 2) p; metadistitarsus: 135-158 (147 ± 8) p. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.48:1. Pigmentation: Color in life: Body gray-green, head yellow-orange. Slide-mounted specimens: Background of body dorsum pale to light brown (to 20 percent pigment density), unicolorous. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen pale, inconspicuous, to moderate brown, conspicuous. Spiracular plates and ventral abdominal sclerites pale, to dark brown, conspicuous. Siphunculi concolorous with surrounding terga, to subtly darker, especially distally near flange. Cauda, anal and subgenital plates light to moderate brown, subtly to substantially darker than abdominal terga. Antennal segments V and IV light to moderate brown, IV sometimes proximally pale; III pale if proximal IV pale, to dusky on distal one-half, if IV entirely darker; II subtly darker than proximal III: I as dark as V, or nearly so, and subtly darker than frons. Pro-, meso- and metatibiae usually concolorous, pale, equivalent to body dorsum, sometimes slightly dusky on distal tip, entire tibiae infrequently slightly darker. Distitarsi entirely dusky. Ultimate Stadium Nymphs of Viviparous Apterae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum lacking 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; area surrounding muscle attachment sites membranous. Viviparous Alatae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous ap- 1994 SORENSEN: A REVISION OF ESSIGELLA 73 terae except lacking body dorsum pigmentation suite; abdominal terga normally membranous, dorsal (major + minor) setae between muscle attachment plates sometimes arising from distinct scleroites; antennal segments often as dark as tibiae, except proximal one-fifth of III pale. Antennal segment III with 0-2, IV with 0, secondary rhinaria. Epicranial suture absent to weakly developed. Forewing medius with furcation arising on central one-third of vein; cubital base usually arising distad, uncom¬ monly proximad, on subcosta with distance between anal and cubital bases on subcosta usually relatively large, ca. 20-40 percent or more of anal vein length; medius, especially cubitus and anal veins usually distinct, except infrequently proximad 10-15 percent vague. Abdominal terga lacking irregular sclerites that engulf or join muscle attachment plates and dorsal (major + minor) setal bases or scleroites. Oviparae, Males, Fundatrices.— Unknown. Diagnosis.— Essigella (E.) alyeska requires the combination of several char¬ acters for identification, because it may be confused with other pale Essigella. Essigella (E.) alyeska can be separated from E. (E.) californica, E. (E.) hoerneri and E. (E.) pini by having eight (Fig. IE), rather than six, dorsal (major + minor) setae on abdominal terga III-IV. It can be distinguished from E. (A) kathleenae, E. (A.) kirki, E. (L.) eastopi, E. (L.) fusca and E. ( L .) hillerislambersi by having two, instead of three or usually more, marginal setae on abdominal terga III-IV, and having small and noninvasive, rather than large and invasive, mesonotal muscle attachment plates on later stadia nymphs of apterae. Essigella (E.) alyeska lacks the thoracic fusion of E. (. E .) essigi, and the protuberant, exceptionally distad primary rhinarium of E. (E.) wilsoni. Some individuals of E. ( E .) alyeska are particularly similar to small, pale E. ( E .) critchfieldi and E. ( E .) knowltoni [es¬ pecially E. ( E.) knowltoni braggi\, but differ from these by often having: often small, instead of always large, ventral abdominal sclerites on segments III-IV; two, instead of three or four, marginal setae on abdominal terga III-IV; and small and noninvasive, rather than large and invasive, mesonotal muscle attachment plates on later stadia nymphs of apterae. Range. —Interior of Alaska, Ontario and Quebec (Fig. 11) [known only from the type series and three other collections]. I anticipate that E. (E.) alyeska will be found in the northern Rocky Mountains in the U.S., and across Canada, wherever the hosts occur. Hosts. —Picea glauca (Moench) Voss, Pinus banksiana Lambert. Collections of E. (E.) alyeska are too few to reliably suggest which host is usual. My extensive sampling of Picea and all other conifers in Alaska, beyond the northern limits of Pinus, yielded E. ( E .) alyeska in only two locations; it was not found during extensive samplings of all conifers in the western U.S. and western Canada (Sor¬ ensen 1983). Collections from Quebec and Ontario list the host as Pinus bank- SldYlQ . Discussion. —Because of previous misinterpretation of meso- and metanotal fusion in Essigella (see character discussion section), the description given here for this species is more accurate than that in Sorensen (1988). Essigella (E.) alyeska is apparently uncommon; the limited collections of it preclude an adequate understanding of its morphological variation over its range. It is a broad-shaped species, but use of its body width characteristic, and that of the E. (E.) knowltoni complex, are not suggested for diagnostics because of the measurement error often associated with nonstandardized (compressed) slides that have been made by others. In contrast, I have attempted to standardize my Essigella slides for noncompression (Sorensen 1983) and, thus, have been able to use body width as an attribute in classification. This allowed the monophyletic 74 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Figure 11. Distribution of E. ( E .) alyeska [dots (JTS samples), squares (nonJTS samples)], super¬ imposed over the range of its hosts, Picea glauca [lighter shading (inclusive of darker)] and Pinus banksiana [darker shading]. grouping of E. (E.) alyeska with the E. ( E .) knowltoni complex, on the basis of width as a nonhomoplasious synapomorphy. In the absence of use of that trait, several other homoplasies would have indicated a closer relationship to E. ( E .) wilsoni (see discussion of that species). Essigella (E.) alyeska has no autapomor- phies, or nonhomoplasious synapomorphies beyond its broad body width; most characters separating it from the E. ( E .) knowltoni complex are reductions or losses. Biology and biogeography also indicate its relationship to the E. (E.) knowltoni complex. The latter feed on the western members of Pinus {Pinus) Subsection Contortae, of which P. banksiana is an eastern member (Little & Critchfield 1969). Essigella (E.) alyeska superficially resembles very pale E. (E.) critchfieldi, despite several finer level differences. That resemblance, and E. {E.) alyeska' s central Alaskan distribution, which is very close to the potential Alaskan panhandle distribution of E. (E.) critchfieldi on Pinus contorta contorta, leads to the suspicion that these two species may be divergent sisters. That hypothesis is furthered by the proximity of these species on the phylogenetic tree derived from discriminant analysis (see the phylogenetics section). Essigella {E.) alyeska may have arisen from the progenitor of the E. (E.) knowltoni complex after the host capture of Picea, or when P. contorta and P. banksiana probably were separated during glaciations. In either event, these species, as members of the series B clade (Figs. 13-15), must have had an origin in the Arcto-Tertiary geoflora, unlike other Essigella, whose hosts had an origin in the Madro-Tertiary geoflora (Sorensen 1992a). Coded References to This Taxon.—Essigella (E.) alyeska has been referred to previously by: the codings “Sp. D” (Sorensen 1983, 1987a, 1992b) and “ALYE” (Sorensen 1983), and by the manuscript name E. “alyeska ” (Sorensen 1983). 1994 SORENSEN: A REVISION OF ESSIGELLA 75 Etymology and Common Name. — The aphid’s species name is the Athabascan Indian term for “Alaska.” Common name: the Alaskan conifer needle aphid. Material Examined. -ALASKA. FAIRBANKS NORTH STAR BOROUGH: (type series) College, (Univ. Alaska Campus), nr Fairbanks, 24 Jun 1979, JTS 79F1, Picea glauca, (apt.). BOROUGH UNCERTAIN: 20 km NE of entrance Mt McKinley Natl Park, 15 Jul 1979, JTS 79G1, Picea glauca, (apt.). CANADA. ONTARIO: Perrault Falls, 17 Jul 1963, G. A. Bradley 63-147-O-APV, Pinus banksiana. QUEBEC: St. Bruno, Lac St. Jean, 10 Aug 1985, A. St. Hilaire, Pinus banksiana. Essigella ( Essigella ) critchfieldi, NEW SPECIES Essigella “ critchfieldi ” Sorensen, 1983: 112 (unpublished manuscript name) Ph.D. Thesis, University of California at Berkeley, Berkeley, California. 605 p. Type Series. — Holotype, vivip. apt., on slide with 4 paratype vivip. apt., ho- lotype in lower right (5 o’clock position); data: WASHINGTON. GRAYS HAR¬ BOR Co.: 16 km W of Amanda Park, hwy 101, 10 Jul 1978, J. T. Sorensen (78G56), Pinus contorta contorta. Holotype deposited in the Natural History Museum, London. Paratypes (all same data as holotype): 20 vivip. apt., on 5 slides including holotype slide. Paratype slides deposited: 1 slide in NMNH, Washington, D.C.; 1 slide in CNC, Ottawa, Ontario; 1 slide in Sorensen collection. Viviparous Apterae.— Morphology: Body length: 1.65-1.88 (1.78 ± 0.08) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously protuberant. Length of antennal segment V: 100-163 (132 ± 16) g, processus terminals: 30-43 (34 ± 4) u; IV: 95-113 (102 ± 6) g; HI: 138-160 (147 ± 7) g\ II: 65-73 (70 ± 2) g. Length of longest setae on frons: 28-55 (41 ±7) g, tips incrassate. Head width: 296-325 (308 ± 10) g. Length of stylets: 510-653 (597 ± 41) g\ ultimate rostral segment: 65-78 (74 ± 4) g, rostral tip reaching abdominal terga I—II, infrequently III, in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 326-418 (372 ± 39) g. THORAX: Meso + metanota fused, total length: 316-357 (340 ± 17) ix. ABDOMEN: Tergum I free, length: 122-143 (132 ± 6) g\ terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 39-50 (45 ± 4) g\ siphunculi protuberant, protrusion 0.5-0.8x maximal distal width. Ventral abdominal sclerites on segments III-IV subquadrate to sub- circular; length: 38-48 (43 ± 3) ix , 1.0-1.4x diameter of metatibiae. Dorsal (major -l- minor) setae (see Fig. IE) on abdominal terga III-IV: 8-9, tips sharp, in 1 row; marginal setae 3-4 each side, per segment. Setae on abdominal tergum VIII: usually 6, infrequently 7, anticipated rarely to 8, length: 23-40 (29 ± 5) ix, tips incrassate to sharp, in 1 row. Cauda broadly rounded; caudal protuberance moderately developed to absent; length of longest caudal setae: 50-100 (78 ± 15) ix, tips sharp. LEGS: Length of metafemora: 490-581 (529 ± 32) g\ metatibiae: 683-826 (760 ± 50) g\ longest dorsal setae on central one-third of metatibiae: 14-28 (22 ± 4) /x, 0.3-1.2 x diameter of metatibiae, tips incrassate; approximately equal or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae: 25-58 (39 ± 9) g, tips sharp. Length of metabasitarsus: 98-110 (103 ± 3) g\ metadistitarsus: 153-178(162 ± 8)/u. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.57:1. Pigmentation: Color in life: Black to very dark brown. Slide-mounted specimens: Background of body dorsum dark brown to nearly black (to nearly 100 percent pigment density), rarely pale, unicolorous. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen conclorous with surrounding terga; on dark individuals, dorsal (major + minor) setal sockets transparent, resem¬ bling pinholes. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen moderate to dark brown, vaguely evident (dark individuals) to conspicuous (pale individuals). Spi- racular plates and ventral abdominal sclerites moderate brown (light individuals) to nearly black (dark individuals). Siphunculi concolorous with surrounding terga. Cauda, anal and subgenital plates subtly to substantially darker than abdominal terga. Antennal segments V and IV concolorously dusky to moderate brown, paler than I and subtly paler than abdominal terga (dark individuals), rarely darker than I and abdominal terga (light individuals); III distal one-fifth to one-third dusky to moderate brown, remainder pale; II at least subtly darker than proximal III; I at least subtly darker than II and frons (all individuals) and substantially darker than V and IV (dark individuals). Pro-, meso- and 76 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) metatibiae all concolorous, as dark as (dark individuals) body tergum, to subtly darker (pale individ¬ uals). Distitarsi entirely moderately brown, to proximal tip sometimes subtly paler. Ultimate Stadium Nymphs of Viviparous Apterae. — Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum with 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; plates distinct, darkly pigmented, diameter approximately equaling eye length. Viviparous Alatae, Oviparae, Males, Fundatrices.— Unknown. Diagnosis. —Essigella (E.) critchfieldi is usually dark brown to nearly black, but infrequently nonteneral specimens are moderately brown. When dark, it is easily confused with E. ( E .) essigi and E. ( E .) knowltoni knowltoni. It lacks the abdominal tergum I fusion of E. (E.) essigi. It differs from E. ( E .) knowltoni knowltoni by: having the darkest pigmentation of antennal segments IV and V at least subtly lighter, instead of subtly darker, than antennal segment I and the abdominal dorsum; usually having a wider maximal distal width of the siphunculi (more than, versus less, than 0.040 mm, although this is an indiscrete difference); and having the body dorsum unicolorous, rather than the frons and sometimes the head and anterad abdomen at least subtly lighter than the abdominal dorsum. The last character can be troublesome for separating E. ( E.) knowltoni knowltoni from the Cascade range and southwestern regions of Oregon, where some indi¬ viduals have the frons concolorously as dark as the abdomen. Essigella ( E .) critchfieldi might also be confused with dark E. (L.) eastopi or aberrant, dark E. (L.) fusca, but differs from these by: having eight dorsal (major + minor) setae on abdominal terga III-IV in a single row (Fig. IE), rather than double (or rarely single) row with the lateral-most minor dorsal seta in the anterad row (e.g., Fig. ID); and having six, instead of eight, setae on abdominal tergum VIII. Addition¬ ally, the pigmentation patterns of E. ( L.) eastopi and E. (L.) fusca differ [see their diagnoses]. Uncommon, pale E. (E.) critchfieldi individuals require the combination of several characters for identification. In particular, E. {E.) alyeska may be confused with these [see diagnosis: E. (E.) alyeska]. Pale E. (E.) critchfieldi can be diagnosed by their chaetotaxy pattern and the number of setae on abdominal terga III-IV and VIII (see above); by their ventral abdominal sclerites on abdominal segments III-IV always being large and subquadrate to subcircular; by the mesonotal muscle attachment plates on their later stadia nymphs of apterae being large and invasive; and by the longest dorsal setae on the central part of their mesotibiae being 0.5- 1.5 x tibial diameter, with tips always incrassate. Range. — Coastal in: Washington, Oregon, northern California, and presumably British Columbia and the Alaskan panhandle (Fig. 12A). Host.—Pinus contorta contorta Douglass ex Loudon; one collection (78G61) from P. contorta latifolia Englemann ex S. Watson shows E. ( E.) critchfieldi and E. (E.) knowltoni knowltoni occur discretely in sympatry. Discussion.—Essigella (E.) critchfieldi is the most divergent member of the E. (. E .) knowltoni complex, and it was generally difficult to procure when sampling its host. It is relatively homogeneous in morphology, and is nearly always very dark when nonteneral. Its dark body dorsum is homoplasious within E. (. Essigella ), and causes confusion with evenly dark E. (E .) knowltoni knowltoni (e.g., Cascades) and the more distantly related E. (E.) essigi. The species’ incrassate tips of the setae on the frons and dorsal setae on the metatibiae, regardless of their length, 1994 SORENSEN: A REVISION OF ESSIGELLA 77 Figure 12. Distribution of: A. E. ( E .) critchfieldi [dots (JTS samples)], superimposed over the range of its host, Pinus contorta contorta [shaded]; B. E. (E.) knowltoni knowltoni [black dots (JTS samples), black squares (nonJTS samples)] and E. (E.) knowltoni braggi [white triangles (JTS samples), white squares (nonJTS samples)], superimposed over the range of their hosts, Pinus contorta latifolia [lighter shading] and Pinus contorta murrayana [darker shading (CA and cascade OR)]. is a synapomorphy for the E. (. E.) knowltoni complex, as is its broad body. On E. (. E .) critchfieldi, the length of dorsal setae on the central part of the metatibiae appears less variable and generally somewhat shorter than on E. (E.) knowltoni, but these setae are generally longer and more variable than on E. (E.) essigi. The lighter antennal pigmentation in E. (E .) critchfieldi, in contrast to the dark body dorsum, is an autapomorphy. Sorensen (1992a) analyzed the relationships within the E. (E .) knowltoni com¬ plex, and in that study assigned populations of E. ( E .) critchfieldi to the coastal group (Sorensen 1992a: COA). There [see discussion under E. (E.) knowltoni knowltoni], composite clusterings indicate that E. (E.) critchfieldi is the most divergent entity of the complex, and remained distinct until the ultimate clustering 78 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) level. Its relative distinction was confirmed by principal component and discrim¬ inant function analyses, both of which indicate that it is less like either of the E. (. E .) knowltoni subspecies than they are to themselves. The analyses showed that E. (E.) critchfieldi separates from the most geographically proximal populations of E. ( E .) knowltoni [the equally and evenly dark, Cascade E. ( E .) knowltoni knowltoni] by the relative difference, albeit subtle, in pigmentation of its antennal segments, the distal width of its extended siphuncular flange, and its smaller general size. It separates easily from E. ( E .) knowltoni braggi because the latter has much paler general pigmentation. Character displacement involving quali¬ tative and qualitative traits occurs within this aphid complex [ see discussion of E. ( E .) knowltoni ]. Coded References to This Taxon.—Essigella (E.) critchfieldi has been referred to previously by the codings: “Sp. E” (Sorensen 1983, 1987a, 1992b), “CRIT” (Sorensen 1983), and “COA” (Sorensen 1992a). Sorensen (1983) referred to this taxon under the manuscript name E. “ critchfieldi .” Etymology and Common Name.— The species is named for the botanist and plant geneticist W. B. Critchfield, who provided much of the information on pines and their relatedness that was necessary for this aphid revision. Common name: Critchfield’s shore pine needle aphid. Material Examined. —CALIFORNIA. DEL NORTE Co.: Crescent City, 4 Jul 1978, JTS 78G5, P. c. contorta, (apt.). HUMBOLT Co.: Manila, 7 km W of Areata on hwy 255, 3 Jul 1978, JTS 78G2, P. c. contorta, (apt.). OREGON. CLATSOP Co.: Seaside, 7 Jul 1978, JTS 78G46, P. c. contorta, (apt.). TILAMOOKCo.: Pacific City, 7 Jul 1978, JTS 78G44, P. c. contorta, (apt.). WASHINGTON. GRAYS HARBOR Co.: (type series) 16 km W of Amanda Park on hwy 101, 10 Jul 1978, JTS 78G56, P. c. contorta, (apt.). YAKIMA Co.: E side of Chinook Pass on hwy 410, 1310 m, 11 Jul 1978, JTS 78G61, P. c. latifolia, (apt.). Essigella {Essigella) knowltoni knowltoni Hottes, 1957, NEW STATUS Essigella knowltoni Hottes, 1957: 92, Proc. Biol. Soc. Wash., 70: 92-93. Primary Type. — Lectotype, vivip. apt., on slide with 3 other apt., lectotype in lower right comer; slide data: “Colo. Aphids, Host Pinus contorta var. muriana, Essigella fusca G. & P., Pink-gree Park, Color., Date 23 Aug 1935, G. F. Knowlton- collector/Holotype Essigella knowltoni F. C. Hottes (over)/[on back] Essigella knowltoni knowltoni Hottes, lectotype, designated J. T. Sorensen, 1981/[specimen position map on slide label] Lectotype Sorensen 1981, as seen from this side.” Lectotype deposited in the U.S. National Museum of Natural History, Washing¬ ton, D.C. There is confusion concerning the type designation. Hottes (1957: 93) designated a holotype (data as above). The slide has “holotype” written on it, and a circle drawn to indicate the individual, but there is no specimen subtending, or even near, the designation circle. I have selected one of the same morph on the slide as technical lectotype, because no single individual was identifiable as the “ho¬ lotype.” Viviparous Apterae.— Morphology: Body length: 1.60-2.32 (1.99 ± 0.21) mm. HEAD: Primary rhinarium on terminal antennal segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial rim greater than 0.5 x diameter of rhinarium; distal face of rhinarial rim usually oblique to longitudinal axis of antennal segment; rhinarial membrane not conspicuously 1994 SORENSEN: A REVISION OF ESSIGELLA 79 protuberant. Length of antennal segment V: 108-150 (132 ± 12) /x, processus terminalis: 28-50 (39 ± 5) iu; IV: 78-115 (100 ± 9) /x; III: 148-218 (184 ± 20) /x; II: 63-95 (75 ± 6) ix. Length of longest setae on frons: 20-73 (44 ± 14) g, tips incrassate. Head width: 296-377 (333 ± 23) ix- Length of stylets: 520-836 (658 ± 74) fx\ ultimate rostral segment: 69-90 (80 ± 5) u, rostral tip reaching abdominal terga I—III in dorsal view through slide-mounted specimens. Head + pronotum fused, total length: 367-479 (420 ± 33) ix. THORAX: Meso + metanota fused, total length: 306-449 (373 ± 42) ix. ABDOMEN: Tergum I free, length: 102-173 (142 ± 18) /u; terga II-VII fused, VIII free. Maximum distal width of flange on siphunculi: 28-44 (36 ± 3) m; siphunculi nearly flush to truncated conical, protruding to 0.6 x maximum distal width. Ventral abdominal sclerites on segments III-IV subcircular to less commonly subelliptical, length: 40-68 (54 ± 7) /x , 1.0-1.6 x diameter of metatibiae. Dorsal (major + minor) setae (see Fig. IE) on abdominal terga III-IV: 8-10, tips sharp, in 1 row, often irregular with setae next to most mesad slightly anterad, or most mesad pair slightly posterad; marginal setae 3-4 each side. Setae on abdominal tergum VIII: 6, infrequently 7, very rarely to 8 (anticipated), length: 20-63 (39 ± 12) /x, tips incrassate to sharp, in 1 row. Cauda broadly rounded; caudal protu¬ berance usually absent to poorly developed, sometimes to moderately developed; length of longest caudal setae: 70-123 (96 ± 16) ix, tips sharp. LEGS: Length of metafemora: 479-775 (655 ± 77) ix; metatibiae: 669-1102 (939 ± 127) m; longest dorsal setae on central one-third of metatibiae: 10-78 (39 ± 18) ix, 0.3-2.3 x diameter of metatibiae, tips incrassate; approximately equal or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae: 25-50 (38 ± 8) ix, tips sharp. Length of metabasitarsus: 105-163 (131 ± 14) ju; metadistitarsus: 150-230 (195 ± 18) ix. Ratio of metadistitarsus to metabasitarsus less than 1.9:1, mean 1.49:1. Pigmentation: Color in life: Body usually dark brown to black, infrequently gray-green or gray; when dark, frons usually paler, yellow. Slide-mounted specimens: Background of body dorsum variable, usually moderately brown to often nearly black (to nearly 100 percent pigment density), occasionally moderately pale to light brown; when dark, frons and sometimes anterad of thorax usually paler than abdominal dorsum; dorsum rarely slightly mottled, or abdominal dorsum rarely darkened more dorsomedially. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga; on dark individuals, dorsal (major + minor) setal sockets transparent, resembling pinholes; occasionally on paler specimens pigmentation of setal bases on abdominal terga subtly darkened and laterally expanded to form a nearly complete, vague band on each terga. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen conspicuous, slightly darker than (pale indi¬ viduals) to as dark as (dark individuals) abdominal terga. Spiracular plates and ventral abdominal sclerites conspicuous, usually dark brown to nearly black (dark individuals), rarely pale (pale individ¬ uals). Siphunculi concolorous with surrounding terga. Cauda, anal and subgenital plates slightly darker than (pale individuals) to as dark as (dark individuals) abdominal terga. Antennal segments V and IV moderate to very dark brown, often distal one-half paler, infrequently pale individuals with proximal one-third of IV paler; III usually moderate to dark brown on distal one-half, remainder pale, infre¬ quently entirely pale (pale individuals), rarely proximal one-half moderate brown and substantially darker distally (dark individuals); II concolorous with proximal III; I concolorous with frons, always lighter than darkest part of V and IV. Pro-, meso- and metatibiae concolorous but variable, usually equivalent with (dark individuals) abdominal terga, often paler, infrequently slightly darker (pale individuals); infrequently tibiae dusky at both tips, paler centrally. Distitarsi variable with tibiae, entirely dark to dusky on distal one-third, proximally paler. Ultimate Stadium Nymphs of Viviparous Apterae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous apterae except lacking body dorsum pigmentation suite; abdominal terga membranous with dorsal (major + minor) setae, between muscle attachment plates, arising from distinct scleroites. Mesonotum with 2 sclerotized plates extending from muscle attachment sites to engulf neighboring setal bases; plates usually distinct, faintly to heavily pigmented, diameter approx¬ imately equaling eye length. Viviparous Alatae.— Slide-mounted specimens: Nonmorphometrics as described for viviparous ap¬ terae except lacking body dorsum pigmentation suite; abdominal terga normally membranous, dorsal (major + minor) setae between muscle attachment plates frequently arising from distinct scleroites; antennal segments often as dark as tibiae, except proximal one-fifth of III pale. Antennal segment III with 0-5, IV with 0, secondary rhinaria. Epicranial suture usually absent, to vaguely developed. Forewing medius with single furcation, very rarely 2 or rarely medius single, (if 2, then first) furcation point usually arising on proximad, infrequently on central or distad, one-third of vein; cubital base usually arising distad, infrequently proximad, on subcosta with distance between anal and cubital bases on subcosta usually relatively large, ca. 30-40 percent or more of anal vein length; medius, 80 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) especially cubitus and anal veins distinct, except infrequently proximad 10-15 percent vague. Ab¬ dominal terga frequently with irregular sclerites that engulf or join muscle attachment plates and dorsal (major + minor) setal bases or scleroites. Oviparae.— Slide-mounted specimens: Nonmorphometrics as described for viviparious apterae, abdominal terga II-VII fused, moderately to heavily sclerotic, including pleural areas, tergum VIII free; dorsal demarcations of anterad terga not evident; siphunculi incorporated into sclerotic dorsum; dorsal abdominal muscle attachment plates unicolorous. Pseudorhinaria on metatibiae irregular, 8-9. Males, Fundatrices.— Unknown. Diagnosis.—Essigella (E.) knowltoni can be difficult to distinguish because it varies in body pigmentation from pale to nearly black. These are relatively wide (broad) aphids, for Essigella, but that trait is shared with E. (E.) critchfieldi and E. (E .) alyeska, and is very easily distorted by slide compression [see discussion of E. (E.) alyeska]-, therefore, it is not recommended. Although dark individuals can be confused with E. (E.) essigi and E. (E.) critchfieldi, E. (E.) knowltoni lacks the abdominal tergum I fusion of E. (E.) essigi, and differs, most reliably, from E. (E.) critchfieldi in having antennal segments IV and V at least subtly darker, rather than lighter, than antennal segment I. Essigella ( E .) knowltoni also may be confused with dark E. (L. ) eastopi or dark, aberrant E. (L.) fusca, but differs from these as E. (E.) critchfieldi does [see diagnosis: E. (E.) critchfieldi]. Pale E. (E.) knowltoni individuals can be confused with most pale Essigella, and require the combination of several characters for identification. They differ from (my limited samples of) E. (E.) alyeska by having three to four, rather than two, marginal setae on abdominal terga III-IV, and always large and invasive, rather than small and noninvasive, muscle attachment plates on the mesonotum of later stadia nymphs of apterae. They differ from E. (E.) californica, E. (E.) hoerneri and E. ( E.) pini by having eight to infrequently 10 (Fig. IE), rather than six, dorsal (major + minor) setae on abdominal terga III-IV. They lack the protuberant, excep¬ tionally distad primary rhinarium of E. (. E .) wilsoni, and the exceptionally long metadistitarus and short metabasitarsus of E. (A.) kathleenae. Many pale E. ( L.) eastopi, E. ( L .) fusca and E. (L.) hillerislambersi may be confused with pale E. ( E .) knowltoni-, individuals with sharp dorsal setae on the metatibiae can be distinguished from E. (is.) knowltoni, which always have these setae incrassate, regardless of length; other E. (is.) knowltoni with fewer than eight setae on ab¬ dominal tergum VIII can be separated from these three species, which always have eight or more such setae. Problems arise in separating pale E. (E.) knowltoni with eight or more setae on abdominal tergum VIII from pale E. ( L.) eastopi, E. (L.) fusca and E. (L.) hillerislambersi with incrassate or blunt dorsal setae on the metatibiae; such E. (L.) fusca, E. ( L .) hillerislambersi and (usually) E. ( L .) eastopi have dorsal (major + minor) setae on abdominal terga III-IV in two rows with the lateral-most minor dorsal seta in the anterad row (e.g., Fig. 1C); although rare E. (. E .) knowltoni braggi may approach this condition, they usually resemble Fig. ID. In E. (L.) fusca and E. (L.) hillerislambersi, the ventral abdominal sclerites on segments III-IV vary from small to large and sublinear, but not large and subcircular-subelliptical, as is always the case for E. (E.) knowltoni. Odd, pale E. ( E .) knowltoni [usually is. (E.) k. braggi] with 10 setae on abdominal tergum VIII, and short dorsal setae on the metatibiae, might be potentially confused with E. (A.) kirki [see diagnosis: E. (A.) kirki]. Separation of the E. ( E .) knowltoni subspecies depends chiefly on pigmentation differences, because univariate traits overlap to a large degree. However, pig- 1994 SORENSEN: A REVISION OF ESSIGELLA 81 mentation differences between E. ( E .) knowltoni knowltoni and E. (E.) knowltoni braggi can be subtle for paler specimens. Essigella (E.) k. knowltoni are usually dark, ranging to nearly black, and usually have the frons, and sometimes the entire head and anterad of the thorax, paler than the abdominal terga; some specimens from the Cascade Range and southwest Oregon have the frons as dark (to black) as the rest of the body dorsum. Most E. ( E .) k. braggi are pale, rarely moderately dark, but then always with the frons concolorous with the body dorsum; they have six to often eight, rarely 10, setae on abdominal tergum VIII, versus the six, to rarely eight for E. (E.) k. knowltoni. The degree of sclerotization of the terga, and the subgenital and anal plates, for E. (E.) k. braggi is less than for equivalently pigmented E. (E.) k. knowltoni. Essigella (E.) k. braggi individuals have indis- cretely longer metatibiae for their body length, than do E. (E.) k. knowltoni. See couplet 16 in the key to the viviparous apterae for separation of these subspecies. Range. — Interiors of Oregon, Washington and British Columbia; south through the Rocky Mountains to central Utah and southern Colorado (Fig. 12B). [For species, see E. (E.) knowltoni braggi also.] Hosts.—Pinus contorta latifolia Engelmann ex S. Watson; P. contorta murray- ana Greville & Balfour (only to southern Oregon). [For species, see E. ( E .) knowl¬ toni braggi also.]. Discussion. — This species [including E. (E.) k. braggi as a subspecies], and E. (E.) critchfieldi, comprise the E. ( E.) knowltoni complex. A unique, qualitative synapomorphy for this complex is that the tips of the setae on the frons and dorsum of the central part of the metatibiae are always incrassate, regardless of the length of those setae. The complex also has a broad body, which is a syna¬ pomorphy with E. (E.) alyeska, denoting a clade on subsection Contortae pines within E. (Essigella ) [see discussion of E. (E.) alyeska]. The fusion of the abdom¬ inal terga of the oviparae for E. ( E .) knowltoni is problematic [see the character discussion section]. Essigella (. E .) knowltoni shares the Pinus contorta niche with E. (E.) critchfieldi, but occurs only on the interior (noncoastal) morphocline of P. contorta latifolia- murrayana. Sorensen (1992a) analyzed and discusses the taxonomic and host relationships within the complex, as coded groups of populations. In those anal¬ yses, E. (E.) k. knowltoni and E. (E.) k. braggi represent the populations assigned to the Cascade-Rocky Mountain (Sorensen 1992a: CAS + RMT) and the Sierra Nevada (Sorensen 1992a: SNV) groups, respectively. The analyses, using exem¬ plars from populations, combined character information from both coded qual¬ itative traits and factor loading scores derived from principal component analyses. The composite data was then clustered, and showed that the groupings of pop¬ ulations that make up E. (E.) k. knowltoni and E. (E.) k. braggi are best circum¬ scribed separately. These groups of populations show somewhat divergent trends in their covariance distributions of morphometric traits in the attribute space defined by principal component analysis, but both differ markedly from the pop¬ ulations comprising E. ( E .) critchfieldi (Sorensen 1992a: fig. 5). The closer rela¬ tionship of the E. (E.) knowltoni subspecies to one another, in comparison with either to E. ( E.) critchfieldi, was also confirmed by discriminant function analysis (Sorensen 1992a: fig. 6). The subspecies of E. (E.) knowltoni break in the Klamath-Siskiyou region (California-Oregon border) along the Pinus contorta latifolia-murrayana mor- 82 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) phocline, which arcs around the Great Basin and Columbia Plateau. Interestingly, this schism of aphid populations does not occur slightly further north, at the Columbia river (Oregon-Washington border), where Critchfield (1957) states the morphological break in the P. c. latifolia-murrayana morphocline, itself, occurs. The Klamath-Siskiyou area marks a steep terpene gradient within Pinus contorta murrayana, which separates the California Sierra Nevada populations of the tree from those of the Cascades (Forrest 1980). Thus, the aphid relationships within the E. ( E .) knowltoni complex show excellent geographic congruence with the genic and biochemical diversity in the Pinus contorta complex (Wheeler & Guries 1982a, b; Wheeler et al. 1983). Essigella (E. ) k. knowltoni is the most variable of the two subspecies, grading from completely pale to nearly black within populations; Sorensen (1992a: figs. 2a-d) shows maps depicting qualitative character variance over geography. Rocky Mountain populations have the highest incidence of pale individuals, show the greatest size variation, and generally have the longest (but quite variable) dorsal setae on the metatibiae. Populations from British Columbia are similar to those of the Rocky Mountains, except darker individuals show greater development of the paleness of the frons and usually the entire head and anterad of the thorax. Dark individuals from Oregon cascade populations often show the frons to be unicolorously as dark as the rest of the body dorsum, as does E. ( E .) critchfieldi. Populations of E. (E.) k. braggi are pale to seldom moderately brown, but then always have the entire body dorsum unicolorous. Pigmentation suites within the E. (E.) knowltoni complex do not appear influ¬ enced by host or environment. For example, a collection of specimens (77G61) of E. (E.) k. braggi spuriously from Pinus monticola maintained their characteristic pigmentation suite, despite occurring opportunistically on that haploxylon pine. Analysis of 25 environmental variables (unpublished data) from sample locations, in relation to qualitative characteristics of individuals from those samples, did not appear to indicate relatedness. Within the E. (E.) knowltoni complex, the pigmentation suite is involved in a character displacement phenomenon (Sorensen 1992a) that also involves general- size. Among the most geographically proximal populations of E. (E.) knowltoni knowltoni, E. (E.) knowltoni braggi and E. (E.) critchfieldi, where these taxa are relatively adjacent in southern Oregon and northern California, those sharing the most similar pigmentation differ the most markedly in covariance relationships among morphometric traits and general-size, and vice versa. When the adjacent populations of these taxa are similarly colored, they differ in size, but when similar in size, they differ in color. The body is relatively unicolorously dark or pale in these geographic areas, but becomes differentiated, as gradiently bicolored, in E. (E.) knowltoni knowltoni in the Rocky Mountains, at a maximal distance from the zone of contact (Sorensen 1992a: figs. 2a-c). Although I consider the populations that comprise E. ( E .) k. knowltoni and E. ( E .) k. braggi to be subspecific, based upon the relative anagenic distance between them when compared to E. (E.) critchfieldi (Fig. 13; Sorensen 1992a: fig. 6), they are definitely more distinct (nonclinal) than the subspecies of E. (L.) fusca. The subspecific status of E. (E.) k. braggi is assigned here to reflect the relative inter¬ taxon distances shown by Sorensen (1992a); however, it would be more appro¬ priate to elevate E. (E.) k. braggi to full species status, rather than synonymize 1994 SORENSEN: A REVISION OF ESSIGELLA 83 it. Hottes (1957) apparently did not recognize the relationship between his E. “ knowltoni ” and E. “braggi ” [or E. “robusta ”], which he believed to be separate species; although he does contrast his E. “braggi ” with ft. “robusta,” suggesting an awareness of similarity between them. He considered his E. “knowltoni ” [here £. (if.) knowltoni knowltoni ] to be “perhaps most closely allied to E. essigi citing the dorsal darkness of the body. Toward the beginning of this study, Dirk Hille Ris Lambers (personal communication [1980]), also citing the dark body dorsum, but without close examination, indicated that he thought my collections of E. ( E .) knowltoni knowltoni, E. (E.) critchfieldi and E. (E.) essigi were conspecific, whereas E. (E.) knowltoni braggi was distinct. Several analyses (Sorensen 1983, 1987a, 1992a, b, unpublished data) indicate that the melanic dorsum trait, which varies to pale within many of the populations of all those species that display it, is homoplasious within E. ( Essigella ). Coded References to This Taxon. —Essigella (E.) knowltoni knowltoni has been referred to previously by the codings: “Sp. F” (Sorensen 1983, 1987a, 1992b), “KNOW” (Sorensen 1983), and “CAS + RMT” (Sorensen 1992a). Sorensen (1983) referred to this taxon under the name E. knowltoni knowltoni. Etymology and Common Name. — This species was named for G. F. Knowlton, who collected the holotype (Hottes 1957: 93), and had a long time friendship with F. C. Hottes that began when they were students under A. A. Granovsky (G. F. Knowlton, personal communication). Common name: Knowlton’s lodgepole pine needle aphid. Material Examined. —[E. ( E .) knowltoni knowltoni only:] COLORADO. CLEAR CREEK Co.: Empire, 2530 m, 14 Aug 1978, JTS 78H86, P. ponderosa, (apt.). GRAND Co.: 24 km NW of Grandby on hwy 125, 2530 m, 15 Aug 1978, JTS 78H90, P. c. latifolia, (apt., ovip.). GUNNISON Co.: W side of Monarch Pass on hwy 50, 2870 m, 13 Aug 1978, JTS 78H74, P. c. latifolia, (apt.). HUERFANO Co.: North La Veta Pass Summit on hwy 160, 2870 m, 12 Aug 1978, JTS 78H63, P. c. latifolia, (apt.). LAKE Co.: 11 km W of Twin Lakes on hwy 82, 3050 m, 14 Aug 1978, JTS 78H77, P. c. latifolia, (apt., ovip.). LARIMER Co.: (paratype) Cameron Pass, 18 Aug 1940, G. F. Knowlton, P. c. latifolia, (apt.); (lectotype) Pingree Park, 23 Aug 1935, G. F. Knowlton, P. contorta var. “ muriana (apt.); (paratype) same but 19 Aug 1935, P. c. latifolia, (apt.). IDAHO. ADAMS Co.: Tamarack, 18 Jul 1978, JTS 78G110, P. c. latifolia, (apt.). BONNER Co.: 6 km S of Cocolalla on hwy 95, 18 Jul 1978, JTS 78G104, P. c. latifolia, (apt.). VALLEY Co.: McCall, 5 Jun 1978, C. S. Smith, CFS 78-170, P. c. latifolia, (apt.). MONTANA. FLATHEAD Co.: 16 km S of Stryker on hwy 93, 17 Jul 1978, JTS 78G97, P. c. latifolia, (apt.). PARK Co.: Silver Gate, hwy 212, 2170 m, 21 Aug 1978, JTS 78H122, P. c. latifolia, (apt.). RAVALLI Co.: Chief Joseph Pass on hwy 13 [93?] on continental divide, 17 Jul 1979, D. J. Voegtlin, DJV 692, P. c. latifolia, (apt.). OREGON. BAKER Co.: Blue Mt Summit on hwy 26, 20 Jul 1978, JTS 78G113, P. c. latifolia, (apt.). GRANT Co.: Canyon Meadows Cmpgd, Malheur Natl Forest, nr John Day, 21 Aug 1979, D. J. Voegtlin, DJV 613, P. c. murrayana, (apt.). JACKSON Co.: 3 km E of Union Creek on hwy 62, 1100 m, 5 Jul 1978, JTS 78G19, P. c. murrayana, (apt.). KLAMATH Co.: 16 km S of LaPine on hwy 97, 5 Jul 1978, JTS 78G22, P. c. murrayana, (apt., alat.). WASCO Co.: 46 km SE of Government Camp on hwy 26, 670 m, 6 Jul 1978, JTS 78G32, P. c. murrayana, (apt.). UTAH. CACHE Co.: 11 km W of Garden City on hwy 89, 2350 m, 24 Aug 1978, JTS 78H133, P. c. latifolia, (apt.); Beaver Creek, Logan Cyn, 25 Jul 1929 Aug 1937, C. F. & C. S. Smith, P. c. latifolia, (apt.). DAGGETT Co.: 32 km S of Manila on hwy 44, 2400 m, 24 Aug 1978, JTS 78H139, P. c. latifolia, (apt.). WASHINGTON. FERRY Co.: Sherman Pass, Colville Natl Forest, 14 Sep 1979, D. J. Voegtlin, DJV 751, P. c. latifolia, (alat.). KING Co.: Arboretum, Seattle, 12 Aug 1956, M. J. Forsell, P. contorta, (apt.). WHATCOM Co.: NE of Newhalem on hwy 20, 300 m, 12 Jul 1978, JTS 78G76, P. c. latifolia, (apt.). YAKIMA Co.: E side of Chinook Pass on hwy 410, 1310 m, 11 Jul 1978, JTS 78G61, P. c. latifolia, (apt.). WYOMING. JOHNSON Co.: 26 km W of Buffalo on hwy 16, 2290 m, 19 Aug 1978, JTS 78H105, P. c. latifolia, (apt.). TETON Co. /Huckleberry Hotsprings, hwy 287, between Yellowstone & Grand Teton Natl Parks, 2010 m, 23 Aug 1978, JTS 84 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) 78H124, P. c. latifolia, (apt., alat., ovip.). CANADA. ALBERTA: 3 km (2 mi) S of Cypress Hills, 18 Jul 1966, P. Rauch, RVdB CL66-VII-18B, P. c. latifolia, (apt.). BRITISH COLUMBIA. 21 km S of 100 Mile House on hwy 97, 910 m, 13 Jul 1978, JTS 78G82, P. c. latifolia, (apt.); 40 km E of Prince George on hwy 16, 14 Jul 1978, JTS 78G85, P. c. latifolia, (apt.); 5 km N of Spuzzum on hwy 1, 13 Jul 1978, JTS 78G77, P. monticola, (apt.); 7 km S of Canal Flats on hwy 93, 17 Jul 1978, JTS 78G95, P. c. latifolia, (apt.); Mt Robson Prov Park, 15 Jul 1978, JTS 78G88, P. c. latifolia, (apt.); nr Clearwater, 9 Sep 1979, D. J. Voegtlin, DJV 744, P. c. latifolia, (apt.). Essigella ( Essigella ) knowltoni braggi Hottes, 1957, NEW STATUS Essigella braggi Hottes, 1957: 73, Proc. Biol. Soc. Wash., 70: 73-75. Essigella robusta Hottes, 1957: 103, Proc. Biol. Soc. Wash., 70: 103-105. NEW SYNONYM. Primary Type. — Holotype, vivip. apt., on slide with 4 other apt., holotype shown by arrow (upper right); slide data: “ Pinus contorta, Tuolumne Meadows, Calif., VIII-22-1955, J. W. MacSwain/Holotype, Essigella braggi F. C. Hottes” (Tuol¬ umne Meadows is in Tuolumne Co., in Yosemite National Park, west of Tioga Pass). Holotype deposited in the Essig Museum of Entomology, University of California at Berkeley, Berkeley, California. Viviparous Apterae.—Morphology: As E. (E.) knowltoni knowltoni, except as follows. Body length: 1.67-2.39 (2.04 ± 0.21) mm. HEAD: Length of antennal segment V: 125-153 (141 ± 9) p, processus terminalis: 33-63 (41 ± 8) p- IV: 95-118 (107 ± 7) p; III: 168-215 (190 ± 17) p ; II: 70-88 (77 ± 4) p. Length of longest setae on frons: 28-78 (52 ± 12) p. Head width: 316-398 (349 ± 22) p. Length of stylets: 632-816 (718 ± 60) p m , ultimate rostral segment: 68-88 (81 ± 6) p, rostral tip reaching abdominal terga I—II in dorsal view through slide-mounted specimens. Total length of fused head + pronotum: 393-490 (438 ± 29) p. THORAX: Total length of fused meso + metanota: 296-439 (381 ± 39) p. ABDOMEN: Tergum I length: 112-173 (149 ± 20) p. Maximum distal width of flange on siphunculi: 25-45 (37 ± 6) p. Ventral abdominal sclerite length: 48-65 (56 ± 6) p. Dorsal (major + minor) setae (see Fig. IE) on abdominal terga III-IV: 8-10, rarely to 12, when 12 the lateral-most minor dorsal seta usually anterad of its next mesad neighbor (i.e., Fig. ID). Setae on abdominal tergum VIII: 6-8, rarely to 10, length: 23-88 (52 ± 16) p, in 1 row, to 2 when 12. Length of longest caudal setae: 70-125 (93 ± 15) p. LEGS: Length of metafemora: 622-842 (724 ± 61) p; metatibiae: 928- 1219 (1048 ± 76) p; longest dorsal setae on central one-third of metatibiae: 25-55 (39 ± 9) p\ longest ventral setae on metatibiae: 23-48 (34 ± 7) p. Length of metabasitarsus: 125-158 (140 ± 11) p; metadistitarsus: 190-225 (206 ± 12) p. Mean ratio of metadistitarsus to metabasitarus: 1.47:1. Pig¬ mentation: As E. (E.) knowltoni knowltoni, except as follows. Color in life: Gray-green or gray to light brown, throughout. Slide-mounted specimens: Background of body dorsum pale to light brown (to 40 percent pigment density), unicolorous. Terga at bases of setae on frons and dorsal (major + minor) setae on abdomen concolorous with surrounding terga, infrequently subtly darker. Thoracic muscle attachment plates and dorsal muscle attachment plates of abdomen usually substantially darker than body dorsum, often only subtly darker. Spiracular plates and ventral abdominal sclerites conspicuous, usually dark brown, infrequently light brown, but always darker than body dorsum. Cauda, anal and subgenital plates usually substantially darker than abdominal terga, often only slightly darker. Antennal segments with darkest areas usually moderate brown, sometimes lighter; proximal base of III never moderate brown; II usually concolorous with proximal one-half of III, but infrequently darker. Pro-, meso- and metatibiae usually pale to frequently moderately brown, often substantially darker than abdominal dorsum. Diagnosis. — For separation of the E. (E.) knowltoni subspecies, see the diagnosis of E. (E. ) knowltoni knowltoni, and couplet 16 in the key to the viviparous apterae. Synonyms. —Essigella robusta Hottes, NEW SYNONYM: lectotype (formerly “holotype”), vivip. apt., on slide with 7 other specimens (lectotype in 10 o’clock position); data: CALIFORNIA. EL DORADO Co.: Upper Echo Lake, 2285 m 1994 SORENSEN: A REVISION OF ESSIGELLA 85 (7500 ft), 6 Aug 1937, E.O.E[ssig]., Pinus contorta murrayana. Essigella robusta lectotype deposited in the Essig Museum of Entomology, University of California at Berkeley, Berkeley, California. Although Hottes (1957: 104-105) designated a “holotype” for E. robusta, his designation circle on the slide encompasses 2 adult and 1 nymphal vivip. apt.; I have selected the center specimen within the des¬ ignation circle as technical lectotype, because no single individual was clearly identifiable as “holotype.” Range. — Sierra Nevada and Cascades of California (Fig. 12B). [For species, see E. (E.) knowltoni knowltoni also.] Host.—Pinus contorta murrayana Greville & Balfour (south of the Oregon- Califomia border only) (see discussion). [For species, see E. (E.) knowltoni knowl¬ toni also.] Discussion. —See E. ( E .) knowltoni knowltoni. Coded References to This Taxon.—Essigella (E.) knowltoni braggi has been referred to previously by the codings: “Sp. G” (Sorensen 1983, 1987a, 1992b), “BRAG” (Sorensen 1983), and “SNV” (Sorensen 1992a). Sorensen (1983) referred to this taxon under the manuscript name E. “knowltoni braggi .” Etymology and Common Name. — Hottes (1957:) named “ Essigella braggi” for L. C. Bragg, presumably because he collected many aphids early in this century; 1 cannot find reference, however, to his association with “ Essigella braggi ” in particular. Common name: Bragg’s lodgepole pine needle aphid. Material Examined. — [E. (E.) knowltoni braggi only:] CALIFORNIA. ALPINE Co.: E side of Ebbett’s Pass on hwy 4, 3 km E of summit, 2400 m, 17 Jul 1977, JTS 77G42, P. c. murrayana, (apt.); same but JTS 77G41, P. monticola, (apt.); Upper Cascade Creek, E side of Ebbett’s Pass on hwy 4, 2 km E of summit, 2350 m, 17 Jul 1977, JTS 77G39, P. c. murrayana, (apt., alat.); W side of Ebbett’s Pass on hwy 4, 18 km W of summit, 2470 m, 17 Jul 1977, JTS 77G44, P. c. murrayana, (apt.). EL DORADO Co.: South Lake Tahoe, 1950 m, 16 Jul 1977, JTS 77G31, P. c. murrayana, (apt., alat.); Upper Echo Lake, 2400 m, 6 Aug 1937, E. O. Essig. P. c. murrayana, (apt.); Wright’s Lake, 850 m, 28 Sep 1969, C. Lagace, Pinus sp., (ovip.). INYO Co.: Bishop, 15 Sep 1969, T. Kono & M. Wasbauer, CDFA 69-J30-32, P. c. murrayana, (apt.); Lake Sabrina, nr Bishop, 2750 m, 1 Aug 1977, JTS 77H1, P. c. murrayana, (apt., alat.). MONO Co.: 1 km S of Crestview on hwy 395, 1 Aug 1977, JTS 77H5, P. c. murrayana, (apt.); Deadman Summit on hwy 395, nr Crestview, 2440 m, 31 Jul 1977, JTS 77G71, P. c. murrayana, (apt.). NEVADA Co.: Prosser Lake Recreation Area, hwy 89, 25 Jun 1977, JTS 77F6, P. c. murrayana, (apt.). PLUMAS Co.: 13 km E of Chester on hwy 36, 1520 m, 4 Jul 1977, JTS 77G14, P. c. murrayana, (apt., alat.); hwy 36, 8 km W of jet with hwy 89, 1460 m, 10 Jul 1977, JTS 77G24, P. c. murrayana, (apt.). SIERRA Co.: 18 km S of Sierraville on hwy 89, 26 Jun 1977, JTS 77F8, P. c. murrayana, (apt.); Donner Summit on hwy 80, 2200 m, 27 Aug 1978, JTS 78H159, P. c. murrayana, (apt.); same but 2290 m, 25 Jun 1977, JTS 77F3, P. c. murrayana, (apt.). SISKIYOU Co.: Edson Creek access Rd, Shasta Natl Forest, 8 km W of Bartel on hwy 89, 1160 m, 3 Jul 1977, JTS 77G9, P. c. murrayana, (apt.). TULARE Co.: 8 km NW of Stoney Creek Cmpgd, Sierra Natl Forest, 2380 m, 13 Aug 1977, JTS 77H11, P. c. murrayana, (apt., alat.). TUOLUMNE Co.: Yosemite Natl Park, Tuolumne Meadows, 22 Aug 1955, J. MacSwain, P. c. murrayana, (apt.); same but nr Porcupine Flat-Porcupine Creek, 2500 m, 30 Jul 1977, JTS 77G66, (apt.). COUNTY UNCERTAIN: Lake Tahoe, 16/17/21 Jul 1969, R. Luck, P. c. murrayana, (apt., alat.). Key to the Viviparous Apterae of Essigella Before using this key, see the commentary under taxonomic key usage in the methods section; also see the character discussion section. la. Abdominal terga III-IV each with 6 dorsal (major + minor) setae be¬ tween muscle attachment plates (e.g., Fig. IF). 23 86 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) lb. Abdominal terga III-IV each with 7 or more dorsal (major + minor) setae between muscle attachment plates. 2 2a. (lb) Pro- and metatibiae subtly to conspicuously darker than mesotibiae, with mesotibial pigmentation approximately that of abdominal terga, or paler. Abdominal tergum VIII with 8 or more setae. Lateral-most minor dorsal seta on each side, between muscle attachment plates on abdominal terga III-IV, usually conspicuously anterad of immediately mesad neighbor (e.g., Figs. 1C-D). ... [pigmented E. (Lambersella)] 17 2b. All tibiae concolorous, or metatibiae darker than pro- and mesotibiae. Setae on abdominal tergum VIII and arrangement of dorsal (major + minor) setae on abdominal segments III-IV variable (e.g., Figs. 1B-E). 3 3a. (2b) Abdominal tergum I fused to the amalgamated meso + metanota; this fusion at least along their lateral contacts, but may be dorsally complete in more heavily pigmented specimens. .. E. ( E .) essigi Hottes (Host: P. radiata, P. attenuata ) 3b. Abdominal tergum I always completely free. 4 4a. (3b) Metadistitarsal length usually > 2.0 (rarely to 1.9) x length of meta- basitarsus (if 1.9-2.0 x, then: color in life of vivip. apt. is pale yellow, oviparae with abdominal terga II-VII fused). Slide-mounted speci¬ mens always concolorously pale. E. (A.) kathleenae Sorensen (Host: P. lambertiana ) 4b. Metadistitarsal length at most 1.95 x, usually less, length of metaba- sitarus (if 1.9-2.Ox, then: color in life of vivip. apt. is usually not pale yellow, oviparae with abdominal terga II-VI fused but VII free, host P. flexilis or P. strobiformis). Slide-mounted specimens variable, pale to developed pigmentation of some sort. 5 5 a. (4b) Primary rhinarium on antennal segment V close to tip; distance between distal portion of rim of primary rhinarium and tip < 0.5, usually < 0.3, x diameter of rhinarium. Distal face of rim of primary rhinarium extending perpendicularly to longitudinal axis of segment V. Membrane of primary rhinarium, if extended, often reaching tip of segment. E. (E.) wilsoni Hottes (Host: Pseudotsuga menziesii, Pseudotsuga macrocarpa) 5b. Distal rim of primary rhinarium and tip of antennal segment V more distant than 0.5 x diameter of primary rhinarium. Distal face of rim of primary rhinarium usually extending obliquely from antennal seg¬ ment. Membrane of primary rhinarium, when protruding, not reach¬ ing tip of segment. 6 6a. (5b) Abdominal tergum VIII bearing 6, occasionally 7, setae. 14 6b. Abdominal tergum VIII with 8 or more setae. 7 7a. (6b) Metadistitarsal length > 1.70 x length of metabasitarsus (if 1.9- 2.0 x, see couplet 4b). Slide-mounted specimens always concolorously pale. E. (A.) kirki Sorensen (Host: P. flexilis, P. strobiformis ) 7b. Metadistitarsal length < 1.70 x length of metabasitarsus. Side-mounted specimens with variable pigmentation, pale to developed. 8 8a. (7b) Abdominal terga III-IV each with 8 dorsal (major + minor) setae 1994 SORENSEN: A REVISION OF ESSIGELLA 87 in 1 roughly linear row (but occasionally with the pair of setae im¬ mediately lateral to the most mesal pair displaced anterad) so that the lateral-most minor dorsal seta on each side of these terga is not con¬ spicuously anterad to its immediately mesad neighbor (e.g., Fig. IE). 9 8b. Abdominal terga III-IV each with 8 or more dorsal (major + minor) setae in 2 rows or staggered, so that the lateral-most minor dorsal seta on each side is conspicuously anterad to its immediately mesad neigh¬ bor (e.g., Figs. 1B-C). 10 9a. (8a) Adult apterae with 2 marginal setae on abdominal segments III- IV. Mesonotum of later stadia nymphs of apterae with area imme¬ diately surrounding muscle attachment sites membranous; bases of neighboring setae not on a sclerotized plate contiguous with the muscle attachment sites. E. [E.) alyeska Sorensen (Host: Picea glauca, Pinus banksiana ) 9b. Adult apterae with 3 or more marginal setae on abdominal segments III-IV. Mesonotum of later stadia nymphs of apterae with scleroti- zation (light to dark pigmentation) extending from muscle attachment sites to form a pair of contiguous plates (approximately the diameter of the eye length) that engulf some neighboring setal bases. 10 10a. (8b, 9b) Ventral abdominal sclerites on segments III-IV linear to sub- linear (when not folded), the length (anteroposterior axis) of longest sclerite > 2.0 x width (mesolateral axis). . [some £. ( Lambersella)\ 17 10b. Ventral abdominal sclerites on segments III-IV circular to subcircular (when not folded), the length of longest sclerite, < 2.0, usually < 1.5 x width. 11 1 la. (10b) At least some setae on frons and some dorsal setae on central one- third of metafemora and metatibiae with sharp tips. .[some E. [Lambersella )] 17 lib. Tips of all setae on frons and all dorsal setae on central one-third of metafemora and metatibia incrassate. 12 12a. (lib) Dorsal setae on metatibiae with an abrupt transition in length (nearly doubling) about midway along segment. .[some E. [Lambersella)} 17 12b. Dorsal setae on metatibiae approximately equal in length or gradually increasing distally. 13 13a. (12b) Body dorsum dark with a longitudinal paler region on dorsomedial region of thoracic and abdominal terga. .. [some E. [Lambersella)} 17 13b. Body dorsum pale to dark, but if dark then either concolorous or frontal area of head is paler than abdominal dorsum. 14 14a. (6a, 13b) Mesonotum of later stadia nymphs of apterae with a pair of light to dark pigmented sclerotizations that extend from muscle at¬ tachment sites to form contiguous plates that engulf some neighboring setal bases; the diameter of these plates approximates the eye length. Adult apterae with pigmentation of body dorsum variable, often tho¬ racic and abdominal terga are dark brown to black; if pale [nonteneral ) then : (a) ventral abdominal sclerites on segments III and IV conspic¬ uous, circular or subcircular (when not folded), and large (minimum 88 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) diameter at least 0.6 x metatibial diameter); (b) antennal segment III length usually > 0.160 mm; (c) maximal distal diameter of extended siphuncular flange usually < 0.040 mm; and (d) 3-4 marginal setae on abdominal terga III-IV. [E. (E.) knowltoni complex] 15 14b. Mesonotum of later stadia nymphs of apterae with only membranous areas immediately surrounding muscle attachment sites; bases of neighboring setae not on contiguous sclerotized plates with the muscle attachment sites. Adult apterae with thoracic and abdominal terga pale and: (a) ventral abdominal sclerites segments III and IV variable, frequently small with minimum diameter < 0.6 x metatibial diam¬ eter; (b) antennal segment III length < 0.160 mm; (c) maximal distal diameter of extended siphuncular flange usually > 0.040 mm; and (d) abdominal terga III-IV with 2 marginal setae. .. E. (E.) alyeska Sorensen (Host: Picea glauca, Pinus banksiana ) 15a. (14a) Darkest pigmentation (nonteneral) of antennal segments III, IV, and V subtly to substantially lighter than antennal segment I. Antennal segment III pale except distal one-quarter frequently slightly pig¬ mented. Body dorsum, including frons, uniformly dark brown to black. Longest dorsal seta on central one-third of metatibiae usually <0.8, rarely > 1.1, x metatibial diameter. Maximum distal diameter of extended siphuncular flange usually > 0.040 mm. . E.(E.) critchfieldi NEW SPECIES (Host: P. contorta contorta ) 15b. Darkest pigmentation (nonteneral) of antennal segments III, IV, and V darker than antennal segment I. Distal one-third to one-half of an¬ tennal segment III dark. Body dorsum variable; if dark, frons often lighter than abdominal dorsum. Longest dorsal seta on central one- third of metatibiae variable, often > 1.1 x metatibial diameter. Max¬ imum distal diameter of extended siphuncular flange usually < 0.040 mm. [E. (E.) knowltoni] 16 16a. (15b) Body dorsum usually moderately to extremely dark, occasionally pale; when dark, frons often lighter than abdominal dorsum; when pale, the ventrolateral border of abdominal tergum, anteroventral border of frons, and posterad border of subgenital plate well defined and demarcated from adjacent membranous regions. Abdominal ter¬ gum VIII with 6, rarely to 8, setae. E. ( E .) knowltoni knowltoni Hottes (Host: P. contorta latifolia; P. contorta murrayana [Oregon]) 16b. Body dorsum usually pale to infrequently moderately dark; not extreme¬ ly dark. Darker specimens with frons concolorous with abdominal dorsum; and with either the ventrolateral border of abdominal tergum, anteroventral border of frons, or posterad border of subgenital plate usually poorly defined compared to adjacent membranous regions. Abdominal tergum VIII with 5-8, rarely to 10, setae. . E. (E.) knowltoni braggi Hottes (Host: P. contorta murrayana [California]) 17a. (2a, 10a, 11a, 12a, 13a) Background of body dorsum darker than tibiae, with a paler longitudinal area on dorsomedial region of the thoracic and abdominal terga. E. (L.) eastopi NEW SPECIES (Host: P. coulteri) 1994 SORENSEN: A REVISION OF ESSIGELLA 89 17b. Background of body dorsum variable from uniformly pale to dark, or mottled, but not as described in couplet 17a; if dark then lacking a paler longitudinal area on dorsomedial region of thoracic and abdom¬ inal terga, and pro- and metatibiae substantially pigmented. 18 18a. (17b) Length of dorsal setae on central one-third of metatibiae > 0.100 mm, and metatibial length < 0.905 mm. Tips of these setae sharp, often reflexed. 22 18b. Length of dorsal setae on central one-third of metatibiae < 0.100 mm, or metatibial length > 0.905 mm. Tips of these setae variable, in- crassate to sharp, but not reflexed. 19 19a. (18b) Metatibial length >1.30 mm, and antennal segment III length > 0.200 mm, and head width (noncompressed slide) measured at lateral bases of antennae > 0.330 mm E. (L.) hillerislambersi NEW SPECIES (Host: P. jejfreyi ) 19b. Metatibial length < 1.30 mm, or antennal segment III length < 0.200 mm, or head width (noncompressed slide) measured at lateral bases of antennae < 0.330 mm [default here if unsure of degree of slide compression]. 20 20a. (19b) Discriminant score (D.S.) < —1.2769, where D.S. - [(antennal segment III length in mm)x (—41.1157)] + [(antennal segment IV length in mm) x (—71.1238)] + [(antennal segment V in mm)x (50.8637)] + [(eye length in mm) x (—58.8556)] + [(number of dorsal {major + minor} setae on abdominal tergum III)x (0.5209)] + (9.81618). E. (L.) hillerislambersi NEW SPECIES (Host: P. jejfreyi ) 20b. D.S. (couplet 20a) > —1.2769. 21 21a. (20b) Discriminant score (D.S.) > 1.3945, where D.S. — [(metatibial length in mm)x (—8.3479)] + [(metabasitarsal length in mm) x (—63.4133)] + [(antennal segment III length in mm)x (65.4496)] + [(dorsomedial length of head + pronotum in mm)x (—29.4826)] + [(dorsomedial length of abdominal tergum I in mm)x (38.7739)] + (12.4544). E. (L.) eastopi NEW SPECIES (Host: P. coulteri) 21b. D.S. (couplet 21a) < 1.3945. [E. (L.)fusca] 22 22a. (18a, 21b) Discriminant score (D.S.) < —0.0803, where D.S. [(met a has i tarsal length in mm)x (71.9890)] + [(length of longest dorsal seta on central part of metatibiae in mm) x (—51.6627)] T- [(number of dorsal {major + minor} setae on abdominal tergum VIII) x (0.9549)] + [(length from anterior of eye to posterolateral comer of protho¬ rax) x (-28.9019)] + [(length of longest seta on antennal segment II) x (—98.3813)] 90 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) + (-6.11263). E. (L.) fusca voegtlini NEW SUBSPECIES (Host: P. ponderosa, P. jejfreyi, P. coulteri) 22b. D.S. (couplet 22a) > —0.0803.. E. (L.) fusca fusca G & P (Host: P. ponderosa, P. ponderosa var. arizonica, P. engelmannii, P. leiophylla) 23a. (la) Mesonotum of later stadia nymphs of apterae with area immediately surrounding muscle attachment sites membranous; bases of neigh¬ boring setae not on a pair of contiguous, sclerotized plates (light to dark pigmentation) extending from muscle attachment sites. Western Nearctic in distribution [default here if specimen is from the Rocky Mountains, Black Hills or westward, Mexico or not North American]. . [E. (E.) californica complex] 23b. Mesonotum of later stadia nymphs of apterae with sclerotization (light to dark pigmentation) extending from muscle attachment sites to form a pair of contiguous plates (approximately the diameter of the eye length) that engulf some neighboring setal bases. Eastern Nearctic in distribution [default here if specimen is from east of the Rocky Moun¬ tains or Black Hills and not Mexico]. E. (E.) pini Wilson (Host: most eastern Pinus sp.) 24a. (23a) Siphunculi conspicuously darker than surrounding abdominal ter- ga, or dorsal (major + minor) setae between muscle attachment plates on abdominal terga II-VI, on dark basal scleroites with well defined borders (nonnymph).. E. (E.) californica (Essig) (Host: most Pinus sp. [except pinyons]) 24b. Siphuncular pigmentation approximately equivalent to surrounding ab¬ dominal terga. Dorsal (major + minor) setae between muscle attach¬ ment plates on abdominal terga II-VI arising from undifferentiated areas of abdominal terga, or from areas that are subtly darker, but have only vaguely defined borders. 25 25a. (24b) Abdominal terga (excluding tergum immediately adjacent to setal bases) subtly to conspicuously darker than thoracic terga and head dorsum. E. (E.) hoerneri G & P (Host: P. monophylla, P. edulis, P. cembroides, P. quadrifolia ) 25b. Abdominal terga (excluding tergum immediately adjacent to setal bases) concolorous with thoracic and head terga; if body dorsum pigmented, abdominal terga not contrasting with thoracic terga and head dor¬ sum. 26 26a. (25b) Slide compressed or sagittal plane of aphid rolled from vertical axis; body width measurements potentially distorted by compression or perspective artifacts of slide preparation [default here if uncer¬ tain]. 27 26b. Slide not compressed and sagittal plane of aphid not rolled; body width measurements not distorted by compression or perspective artifacts of slide preparation. 28 27a. (26a) Discriminant score (D.S.) > 0.3991, where D.S. = [(antennal segment IV length in mm)x (0.0093)] + [(antennal segment II length in mm) x (-0.1345)] 1994 SORENSEN: A REVISION OF ESSIGELLA 91 + [(stylet length in mm) x (0.0134)] + (0.39912). E.(E.) hoerneri G & P (Host: P. monophylla, P. edulis, P. cembroides, P. quadrifolia) 27b. D.S. (couplet 27a) < 0.3991. E. ( E .) californica (Essig) (Host: most Pinus sp. [except pinyons]) 28a. (26b) Discriminant score (D.S.) < —0.4847, where D.S. = [(antennal segment IV length in mm)x (0.0059)] + [(antennal segment II length in mm)x (0.2023)] + [(head width at lateral base of antennae in mm)x (—0.0456)] + [(stylet length in mm)x (-0.0083)] + (3.18802). E.(E.) hoerneri G & P (Host: P. monophylla, P. edulis, P. cembroides, P. quadrifolia ) 28b. D.S. (couplet 28a) > —0.4847. E. (E.) californica (Essig) (Host: most Pinus sp. [except pinyons]) Phylogenetic Analyses: tl During this project, the need for development of an operational method to estimate phylogeny using noncoded, morphometric attributes became apparent, because Essigella have few morphological traits that could be treated or coded objectively using the then existing numerical cladistic procedures that were based upon discrete-state data. While analyzing the genus (Sorensen 1983), I developed a phylogenetic procedure for morphometric data that employed discriminant function analysis to reveal unshared variance among groups, and then linked the group centroids to yield a phylogenetic network. That procedure was later modified (Sorensen 1987a) to yield the currently accepted phylogeny for the genus. The method estimates minimum selective mortality indices (sensu Lande 1979) that account for divergence resulting from past selection (Pimentel 1992). Sorensen & Foottit (1992) present the quantitative genetic rationales for the procedure, and Sorensen (1992b) discusses its operational limitations. The phylogeny developed for Essigella, as tl of Sorensen & Foottit (1992), Sorensen (1992b) and here [= 0 of Sorensen (1987a)], is based on adult virgi- noparous apterae only. Sorensen (1983) used 26 morphometric traits (see methods section) to derive tl and circumscribe all Essigella taxa; these are listed elsewhere (see Sorensen 1991: table 1). For Essigella, tl was generated using discriminant function analysis (Nie et al. 1975: SPSS, version 7, program DISCRIMINANT, direct selection mode, Wilks-X criterion) to derive group centroids, as mean group phenotypes (z, sensu Lande 1979) for taxa; this was followed by their linkage using a maximum-likelihood cladistic algorithm (Felsenstein 1984: PHYLIP, ver¬ sion 2.5, program CONTML, c-option); the analysis included Pseudessigella, as an outgroup (Sorensen 1990), for rooting. Because this phylogenetic estimate is probability based, as a maximum-likelihood network, confidence intervals for each phyletic segment (intemode) were generated; these are listed in Sorensen (1987a: table 1). In Fig. 1 3,11 is shown with the length of its intemodes, which represent evolved apomorphic anagenic distance, scaled proportionally to their divergence. Figure 14 shows this phylogenetic pathway as it navigates through the 3-dimensional 92 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(1) Evolutionary Distance 1 a unit * = anagenic distance significant at a =0.05(1.96 a units) to Pseudessigella Figure 13. Anagenic distance preserving portrayal of the phylogenetic estimate, tl, for Essigella (after Sorensen 1987a). White nodes are taxa (as group centroids); black nodes (numbered in square brackets) are ancestors; intemode lengths are proportionate to their anagenic distances (indicated) in pooled standard deviation units (o-); intemodes that are significant (as > 1.96 a for a = 0.05) for the genus/subgenus level are shown in bold numbers with an asterisk. Subgenera are indicated by shading. 2 . 08 * discriminant space that is represented by the dominant three minimum selective mortality vectors (Sorensen & Foottit 1992, Sorensen 1992b) occurring over the given evolutionary episode. Although preserving the furcation patterns in that space, Fig. 14 necessarily distorts the intemodal distances, which are derived from the full 15 dimensions (= Groups — 1) of the data matrix. The cladistic relation¬ ships among Essigella species should be obvious from these figures; for discussion, see Sorensen (1992b) or Sorensen (1987a). Justification for Delimitation of Subgenera.— Because Pseudessigella was nec¬ essarily included in the analysis for polarity, and because it represents the nearest separate genus (Sorensen 1990), the analytical perspective (sensu Sorensen 1992b) used in generating tl allows inference of reasonable, quantitatively determined subgeneric demarcations within Essigella. This is possible, and logically called for, because, for the general analytical procedure: (1) All, and only, unshared variance (apomorphy, sensu Sorensen & Foottit 1992) among the taxa is used in network construction, so that the derived inter- 1994 SORENSEN: A REVISION OF ESSIGELLA 93 Figure 14. Spatial-based portrayal for tl (after Sorensen 1992b). Here, tl transverses the evolu¬ tionary space defined by the dominant three minimum selective mortality vectors (sensu Lande 1979), represented as discriminant functions (DF) (see Sorensen & Foottit 1992, Sorensen 1992b). Taxa (as group centroids) are abbreviated by their first two letters; the subgenera and Pseudessigella are cir¬ cumscribed by dashed lines that indicate the maximal spatial distributions of their contained indi¬ viduals. The first two vectors (DF1, DF2) are shown, and the third (DF3) is implied by the relative size and color of the dot, and size of letters, for each taxon: larger (white) dots are forward of, intermediate (gray) dots are on, and smaller (black) dots are backward from, the plane of the page. Phyletic internodes here are not proportionate to their anagenic distances (as in Fig. 13), but spatially demonstrate the furcation events in this evolutionary space. node values appropriately portray a maximum-likelihood representation of solely apomorphic anagenic distance among the network nodes. (2) The intemodal lengths are in Mahalonobis’ distance, as standard deviation units, o [= SD units of Sorensen (1987a, 1992b)], that are parsimoniously pooled across all incorporated groups (as taxa). Thus, these 0.05) (Table 2). This suggests a positive correlation between the duration and number of copulations. All females mated in the laboratory experiment were exposed to three additional 1994 ICHINOSE: PARATRECHINA MATING BEHAVIOR 185 Table 1. The number and duration (seconds) of copulations and days from the last copulation to the death for males of P. flavipes. Male Duration Days 1 Number First b Second Third ml 3 92 360 205 0.5 m2 2 235 1210 — 1.5 m3 2 60 722 — 0.5 m4 2 287 408 — 1.5 m5 2 40 140 — 1.5 m6 2 128 972 — 0.5 Mean ± SD 140.3 ± 99.5 632.0 ± 410.9 — 1.00 ± 0.50 ml 1 100 — — 2.5 m8 1 195 — — 2.0 m9 1 223 — — 1.0 mlO 1 719 — — 1.0 ml 1 1 322 — — 2.0 ml2 1 160 — — 2.0 m 13 1 531 — — 2.0 ml4 1 282 — — 1.5 ml5 0 — — — 2.0 Mean ± SD 316.5 ± 208.4 1.75 ± 0.50 a Significantly different, t = 2.78, P < 0.02. b Significantly different, t = 2.24, P < 0.05, using log-transformation. males after copulation. They did not mate again and eventually shed their wings. They laid eggs in July, which became workers in August, indicating that they were successfully inseminated. Discussion The field observations of P. flavipes nuptial flights and previous data (Ichinose 1987) indicate that females of this ant mate once. The female that was observed mating twice on 25 Jun 1986 seems to be exceptional. Both of her copulations lasted longer than those of other females. The long durations may be due to her copulating with previously mated males. This inference is supported by the lab¬ oratory experiment where the duration of a second or third copulation was sig¬ nificantly longer than those of a first copulation. Some females used in the laboratory could have mated before collection. This seems improbable, however, because females observed in the field removed their T able 2. A two-way AN OVA test comparing the duration of first and second copulations in multiple mating males. Variables are transformed logarithmically. Source of variation df ss MS Fs Significance level Mating 1 1.265 1.265 27.978 P < 0.01 Individuals 5 0.901 0.180 3.987 ns P > 0.05 Error 5 0.226 0.045 Total 11 2.393 186 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) wings less than 10 min after the end of copulation. Laboratory experiments were started at least 1 h after the collection and all the females had their wings. Multiple mating of male ants is known or suggested to occur in several ant species (Scherba 1961, Kannowski 1963, Ito & Imamura 1974). However, P. flavipes is the first species in which the number of male matings has been experimentally shown to be limited, although Hdlldobler (1976) has suggested that the number of male matings is limited in Pogonomyrmex spp. Multiple matings in some P. flavipes males indicates that their genitalia are not lost at copulation as occurs in F. rufa (Marikovsky 1961) or A. mellifera (Wilson 1971). The reason that copulation of the male is limited may exist in the amount of their sperm, not in the structure of their genitalia. If P. flavipes males cannot produce sperm after eclosion as in other ants (Holldobler & Bartz 1985), the male should have less sperm at the second copulation than at the first. Thus, the ANOVA test in Table 2 suggests a negative correlation between sperm amount and copulation duration in the males. If so, the sperm amount in the male decreases in the order: the multiple mating male at the first copulation; the single mating male; and the multiple mating male at the second copulation. Because survival after the last copulation is shorter in multiple mating males than in single mating ones, sperm usage may promote the death of P. flavipes males. Acknowledgment I thank S. F. Sakagami and M. J. Toda for their comments; K. Isigaki, of the Tomakomai Experiment Forest, facilitated this research in the forest. Literature Cited Bartz, H. S. 1982. On the evolution of male workers in Hymenoptera. Behav. Ecol. Sociobiol., 11: 223-228. Cole, B. J. 1983. Multiple mating and the evolution of social behavior in the Hymenoptera. Behav. Ecol. Sociobiol., 12: 191-201. Corso, C. R. & A. Serzedello. 1981. A study of multiple mating habit in Atta laevigata based on the DNA content. Comp. Biochem. Physiol., 69B: 901-902. Crozier, R. H. & R. E. Page. 1985. On being the right size: male contributions and multiple mating in social Hymenoptera. Behav. Ecol. Sociobiol., 18: 105-115. Glancey, B. M. & C. S. Lofgren. 1985. Spermatozoan counts in males and inseminated queens of the imported fire ants, Solenopsis invicta and Solenopsis richieri (Hymenoptera: Formicidae). Fla. Entomol., 68: 162-168. Holldobler, B. 1976. The behavioral ecology of mating i n harvester ants (Hymenoptera: Formicidae: Pogonomyrmex). Behav. Ecol. Sociobiol., 1: 405-423. Holldobler, B., &H. S. Bartz. 1985. Sociobiology of reproduction in ants. pp. 237-257. In Holldobler, B. & M. Lindauer (eds.). Experimental behavioral ecology and sociobiology (Fortschritte Der Zoologies, Bd. 31). Sinauer Associates, Sunderland, Massachusetts. Ichinose, K. 1987. Annual life cycle of Paratrechina flavipes (Hymenoptera, Formicidae) in the Tomakomai Experiment Forest, southern Hokkaido. Kontyu, 55: 9-20. Ito, M. & S. Imamura. 1974. Observations on the nuptial flight and intemidal relationship in a polydomous ant, Formica ( Formica ) vessensis Forel. J. Fac. Sci. Hokkaido Univ. VI Zool., 19: 681-694. Kannowski, P. B. 1963. The flight activities of formicine ants. Symp. Genet. Biol. Ita., 12: 74-102. Marikovsky, P. I. 1961. Material on sexual biology of the ant Formica rufa L. Ins. Soc., 8: 23-30. Page, R. E. Jr. & R. A. Metcalf. 1982. Multiple mating, sperm utilization, and social evolution. Am. Nat., 119: 263-281. Scherba, G. 1961. Nest structure and reproduction in the mound-building ant Formica opaciventris Emery in Wyoming. J. N. Y. Entomol. Soc., 69: 71-87. 1994 ICHINOSE: PARATRECHINA MATING BEHAVIOR 187 Talbot, M. 1959. Flight activities of two species of the ants of the genus Formica. Am. Midi. Nat., 61: 124-132. Wilson, E. O. 1971. The insect societies. Belknap Press of Harvard University Press, Cambridge, Massachusetts. Yamauchi, K., T. Furukawa, K. Kinomura, H. Takamine & K. Tsuji. 1991. Secondary polygyny by inbred wingless sexuals in the dolichoderine ant Technomyrmex albipes. Behav. Ecol. Sociobiol., 29:313-319. PAN-PACIFIC ENTOMOLOGIST 70(3): 188-205, (1994) A BIOGRAPHICAL ACCOUNT OF HAROLD COMPERE (1896-1978), BIOLOGICAL CONTROL FOREIGN EXPLORER Gordon Gordh 1 Department of Entomology, University of California, Riverside, California 92521 Abstract.— An historical account is provided for Harold Compere, a biological control foreign explorer for the University of California (1923-1963). Compere’s early involvement in aviation and its application to entomology are discussed. Some of his foreign exploration exploits are reported and success with control of citrophilous mealybug is noted. His taxonomic and mor¬ phological interests in parasitic Hymenoptera are discussed. A bibliography of Harold Compere’s scientific publications is provided. Key Words. — George Compere, biological control, foreign exploration, citrophilous mealybug [.Pseudococcus calceolariae ], black scale [Saissetia oleae ], California red scale [Aonidiella au- rantii ], parasitic Hymenoptera, taxonomy, morphology Biographical notes and observations provide a point of reference which time and memory otherwise diminish. This account reviews high points in the life of Harold Compere. He had no children or direct descendants; he was not a sub¬ scribing or participating member of any scientific society and he did not supervise graduate students. Consequently his passing has gone largely unrecognized. For more than 40 years Harold Compere served the University of California and State of California as a foreign explorer and research entomologist. His contribution to biological control of citrus pests was substantial and his contribution to the sys- tematics of parasitic Hymenoptera was noteworthy. The following notes serve to document some of his accomplishments. Early Years Harold Compere was bom on 17 Jan 1896 on Washington Street, Los Angeles, California (Fig. 1). His mother, Amy Caypless Compere, was bom in Bridgeport, Connecticut. His father, George Compere (1858-1928), was bom in Davenport, Iowa. The elder Compere was a prominent figure in southern California agriculture and an early foreign explorer for biological control. George Compere was at least indirectly responsible for Harold’s involvement with biological control. The elder Compere moved to California in 1874. During 1878 George was in charge of the Vejar Orchard which was seriously infested with black scale. In 1891 he became a horticulture inspector for Los Angeles County. George was a pioneer in foreign exploration and in 1898 he accepted a position in Hawaii to search for beneficial insects, first in Australia and Hong Kong. In 1901 he was employed by the Western Australia Department of Agriculture to search for natural enemies; in 1904 he was jointly employed by the State of California and Western Australia to search for natural enemies of injurious insects. George held the post of Entomologist to the Bureau of Agriculture of Western Australia until 1910. In 1908 he went to China and took parasites of red scale to Perth. When he returned to California, 1 Present address: Entomology Department, University of Queensland, St. Lucia 4072, Queensland, Australia. 1994 GORDH: HAROLD COMPERE HISTORY 189 Figure 1. Harold Compere in a photograph taken about 1901. George accepted a position as a port inspector in San Francisco. For an account of George Compere’s entomological activities see Howard (1930), Essig (1931) and an anonymous obituary (1935). Harold Compere lacked a formal education but succeeded in an academic environment despite this apparent handicap. By his own account he repeatedly dropped out of elementary school but was forced to return by his mother. His last classroom training was at Lincoln High School in Santa Monica, where he finally quit the 9th grade in 1912. Harold Compere thought he would like to become a fisherman. After leaving school he went to work as a deck hand on a pleasure boat, the McKinley, which operated off “Frazer’s Million Dollar Pier” at Ocean Park. He was paid eight dollars a week, but the job did not last long. He told the writer on more than one occasion that the fishing business involved too much physical labor. However, 1 Jan 1913 found him an apprentice gardener at Golden Gate Park in San Francisco where the labor was just as intensive and his pay was $1.50 per day. At Golden Gate Park, Harold Compere was noticed by the park’s founder, John McLaren. Apparently McLaren was impressed by Compere and his abilities. After a short while at the park, Compere was put in charge of insect control because 190 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) he apparently displayed a flair for this type of work. This aspect of his employment lasted from 1914-1915. He was called “The Professor” and to quote Compere he “. . . had the social status of a foreman but the pay of an apprentice gardener.” His terminal pay at the park was $3.00 per day. Compere’s professional development in entomology began to solidify during 1915. San Francisco officials announced the establishment the position of County Horticultural Commissioner. Competition for the position was open and required an examination. Compere wanted the position because it carried a salary of $6.00 per day, an office and an expense account. He was granted a leave of absence from Golden Gate Park by McLaren and studied for the exam. McLaren told Compere that if he passed the exam, the job would be his. McLaren was a very influential person in San Francisco at the time and agreed to use his influence with the Chairman of the Board of Supervisors who made the appointment. Compere achieved the highest score on the exam but he was not appointed to the position because he was not 21 years old. Instead, Compere became a Lab¬ oratory Assistant with the California State Commission of Horticulture on 16 Sep 1915. Compere received this offer from Harry Scott Smith (1883-1957), then Superintendent of the State Insectary at Capitol Park in Sacramento. Smith served as administrator for the County Horticultural Commissioner exam and was im¬ pressed with Compere’s performance on the written and oral examination. In effect, Compere became an assistant to Smith and served him for more than 35 years. The initial phase of the relationship was temporarily interrupted by World War I. A few details about Smith are relevant to Compere’s story. Smith was bom in Aurora, Illinois and attended the University of Nebraska (A.B. 1907; M.S. 1908; honorary Ph.D. 1953). Smith was hired by L. O. Howard (1857-1950) in 1908 to study natural enemies of the boll weevil. Smith subsequently moved to the Melrose Highlands Laboratory in Massachusetts where he worked on natural enemies imported to control gypsy moth. Smith left federal employment to try ranching in Wyoming, but returned to entomology when his efforts at livestock management failed. Smith’s first foreign exploration involved searching for natural enemies of the alfalfa weevil in Europe; one of the species he imported was useful in control of alfalfa weevil. Upon return to the United States, Smith was hired by Albert John Cook, California State Horticultural Commissioner, to serve as superintendent of the State Insectary. Smith went on to become the “father” of biological control in California and the person generally credited with coining the term “biological control.” Harold Compere saw service as a 2nd Lieutenant in the U.S. Army Air Service during World War I. Compere was granted a leave of absence from the State Commission of Horticulture during 1917, and he enlisted as a private in the Aviation Section, Signal Enlisted Reserve Corps. His entrance into flight training was unusual because college training was a requirement, and most officers in the Army were college graduates. The Army Air Corps was even more selective. For a high-school dropout to be accepted into this prestigious program was remarkable. Compere entered the School of Military Aeronautics, University of California at Berkeley on 27 Oct 1917 and completed the course eight weeks later. Compere, with a ninth grade education, passed all examinations. He once confided that he studied after “lights out” in the latrine. His studies paid off; most students with college training failed to complete the course. 1994 GORDH: HAROLD COMPERE HISTORY 191 Compere received his flight training at Rockwell Field, North Island, San Diego. As part of his training he made a cross-country flight to Allesandro, Riverside County, California and became the first pilot to land at what eventually became March Air Force Base (See March Field Story, 50th Anniversary 1918-1968, p. 12). This Strategic Air Command base continues as an important military in¬ stallation, and airplanes continue to fly over Compere’s home in Riverside and the experiment station where he worked. After completion of training, Compere was assigned to duty as a flight instructor at Ellington Field in Houston, Texas. Compere later stated that he desired an instructorship because duty in the war was dangerous. (The life of a pilot at the front was measured in days or weeks.) Compere’s life as a pilot was not dull. Sport among stateside pilots included cloud flying and racing railroad trains. Compere participated in cloud flying until pilots were killed in accidents and he realized the dangers associated with this activity. Racing trains seemed less dan¬ gerous to Compere, but the custom was strictly prohibited by the military. Nev¬ ertheless, Compere raced trains until the day an Air Patrol spotted him. To elude authorities Compere became lost in the landing traffic, taxied his plane into a hanger and fled the scene. Compere was responsible for the pioneering effort to use the airplane as an applied entomological tool. The story was briefly mentioned, but has gone gen¬ erally unnoticed. [See “The Airplane as a Farm Scout” 29 Mar 1919, Literary Digest, 60 (13): 133-134.] During the fall of 1918 he discussed the possibilities of using the airplane in survey work with Walter D. Hunter (1875-1925), then a member of the Federal Horticulture Board and responsible for control of pink bollworm, Pectinophora gossypiella (Saunders), in Texas. On 23 Nov 1918 Com¬ pere forwarded a letter to the Commanding Officer of Ellington Field, requesting that permission be granted to use a military aircraft for the survey of cotton fields near Ellington. That request was disapproved by the Executive Officer, 1 st Lt. J. H. Sullivan. Subsequently, Compere wrote to L. O. Howard and indicated that the work was feasible. Howard was impressed with the possibilities of the aircraft being used for entomological purposes. On 9 Dec 1918 he forwarded Compere’s letter to the Division of Military Aeronautics with an endorsement of the plan. In response, Lt. Colonel John Sullivan, Chief of the Photographic Branch and acting under orders of Major General Kenly, directed the Commanding Officer of El¬ lington Field to undertake the work outlined by Compere and submit a record of the work to the War Department in Washington. Sullivan’s letter is dated 13 Dec 1918. In a letter to Howard, Compere summarized the survey program. “Dr. Hunter assigned a very capable man to this work. He will be in charge and direct the pilot. A ship has been equipped with speaking tubes so that any pilot will be capable of carrying on this work when directed by a quarantine inspector. The inspector makes the charts and is responsible for the success of the work. Several days ago we made our first flight. It was a success. A storm prevented us from continuing the survey.” “I have just returned from a visit to California where I met Mr. Smith who is holding a position open for me. He urged that I immediately return to Sacramento. On returning from my visit, I found my discharge papers and also the orders to undertake the boll worm survey. I had accepted my discharge and on Tuesday 192 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) will return to California. I have made arrangements for another pilot to conduct this work. He will be under the direction of Dr. Hunter’s quarantine inspector. In this work a pilot only acts as an aerial chauffeur, so that I do not feel my personal services will be of any great advantage.” (Compere to Howard, 17 Jan 1919.) The quarantine inspector selected for the work was Carl Heinrich (1880-1955) (Wade 1955). The pilot was 2nd Lt. William H. Tillisch, Harold Compere’s roommate. Heinrich and Tillisch made two flights on the afternoon of 21 Jan 1919 from Ellington Field. A report of the trip was prepared by Heinrich and forwarded to Washington. Tillisch had been selected for the entomological work personally by Compere. Tillisch had intended to remain in the Army but he was killed in a scouting accident on 7 Aug 1919. Harold Compere was discharged from the Army on 18 Jan 1919 and returned to California. The role of the airplane expanded in service of entomology. From a commercial standpoint, aerial application of pesticides on cotton was initiated about 1922 when equipment for application of dust was developed. Sprays were used from airplanes in 1930 for the control of mosquitoes. Widespread usage of the aircraft for mosquito control was established during World War II. Curiously, Compere dropped flying completely after discharge from military service. During the remainder of his life he rode as a passenger on airplanes only a few times out of medical necessity. All of his foreign exploration was conducted by ship. Compere married Joan Tillman on 25 Aug 1919 in San Francisco. The marriage was the second for Joan, and the only marriage for Harold. Joan was bom 27 Oct 1898 at Brownswood, Texas, and died during Jan 1974 at Riverside. Compere returned to employment with Smith in California under the title of Junior En¬ tomologist, California State Department of Agriculture (1919-1921). During 1921- 1923 Compere was employed as an Assistant Entomologist. The Division of Beneficial Insect Investigations was established as a part of the Citrus Experiment Station (CES) in 1923. Several years earlier, the state legislature was convinced by progressive agriculturists that a research program for citrus and subtropical crops should be located in the southern part of the state. For an account of the Citrus Experiment Station see Boyce (1969). Riverside was selected as the site of the CES. The working group in beneficial insects represented a reorgani¬ zation of the California State Commission of Horticulture. Biological control as an organized unit had been established in 1907 with construction of the State Insectary in Sacramento. The first superintendent of the Insectary was E. K. Cames, assisted by E. J. Branigan. Cames was succeeded as superintendent by H. S. Smith in 1913. The research program on beneficial insects became part of the Citrus Experiment Station with its CSCH staff transferred from Sacramento to Riverside. Smith was Head of the Division. Original members of the Division were Harold Compere, as foreign explorer and A. J. Basinger (1886-1984) as secretary. Philip H. Timberlake (1884-1981) was hired in 1924 to serve as a taxonomic specialist. Stanley Flanders (1894-1984) joined the group in 1929 and later served as the quarantine officer. Foreign Exploration Compere’s major contribution to entomology was as a foreign explorer for natural enemies of pests associated with subtropical agriculture. During a twenty 1994 GORDH: HAROLD COMPERE HISTORY 193 year period Compere made trips to South America, Africa, Australia and the Orient. He assumed a position similar to one held by George Compere with the state of California intermittently during the years 1899-1910. To understand why someone would be employed to wander the globe searching for insects, we must understand the agricultural conditions and views of growers regarding insect pest control. Biological control was viewed optimistically by many citrus growers. Their enthusiasm stemmed directly from the spectacular success achieved over the cottony cushion scale, Iceryia purchasi (Masked) (CCS). CCS was the most serious insect pest of citrus during the 1880s and threatened elimination of the crop from California. The pest originated in Australia. Beneficial insects were found in Australia during 1888 and imported into California. These beneficial insects included a predaceous ladybird beetle, Rodolia cardinalis (Mul- sant) and a parasitic fly Cryptochaetum iceryae (Williston). These insects became established and were responsible for complete control of CCS (Doutt 1958). The exploration work was undertaken by Albert Koebele (1852-1924); George Com¬ pere was not associated with the spectacular success, but he did witness it. In part because growers were highly receptive to this kind of work, the State constructed the insectary in Sacramento. Harry Smith reinstituted foreign exploration after he became superintendent of the State Insectary in 1913. During that year he searched in Japan and the Philippines for natural enemies of mealybugs and scale insects attacking citrus. The work of others followed. The 1920s saw Smith increasingly involved in administrative duties with a continuing strong need for foreign exploration. The exploration work was conducted by Harold Compere. During his career, Compere imported natural enemies for numerous pests of citrus, most notably the black scale, California red scale and citrophilous mealybug. 1926-1927 Australia, New Zealand (Citrophilous Mealybug).— After control of CCS, the most serious pests of citrus in California were mealybugs, including citrophilous mealybug [Pseudococcus calceolariae (Maskell)], citrus mealybug [Planococcus citri (Risso)] and long-tailed mealybug [Pseudococcus longispinus (Targioni-Tozzetti)]. Before WWI biological control work on these pests had been limited to use of the coccinellid Cryptolaemus montrouzieri Mulsant which had been imported from western Australia by Koebele during 1892. The beetle was a general predator of mealybug whose success was limited. Considerable work was invested in collecting adults and transporting them elsewhere in California where pest outbreaks occurred. The problem was one of demand surpassing sup¬ ply. The beetles could not be produced in large numbers because their mealybug prey could not be mass produced in the laboratory. The citrophilous mealybug (CM) was first detected by C. P. Clausen (1893— 1975) at Upland (near Riverside) during 1913. By the mid 1920s, this insect was recognized as the most significant pest, infesting more than 100,000 acres of citrus. Cryptolaemus montrouzieri was an established predator but it was not effective. Therefore, Compere was sent to Australia by Smith to search for natural enemies of the CM. In Sydney during March 1928 Compere recovered the parasitic wasps Coccophagus gurneyi Compere and Tetracnemus pretiosus Timberlake from a mulberry tree infested with CM. Compere returned from Australia during 1929 with these parasites and by 1931 C. gurneyi completely controlled this pest in California. The instance stands second only to the Vedalia beetle in importance in the annals of applied biological control. The trip cost the state of California 194 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) $1700 but saved the citrus growers millions of dollars. The loss in revenue and cost of control of this pest would have been astronomically large had it not been controlled over the past 60 years. 1929-1930 Japan, China, India, Ceylon, Eritrea (Black Scale).— Black scale (BS) [Saissetia oleae (Oliver)] was first noted in California about 1880 and peri¬ odically had been a serious pest on citrus and olives. In the early years of southern California citrus, BS was regarded as the most serious pest. Africa has been generally regarded as the native home of BS. In 1917 E. J. Vossler (1890-1918) imported the encyrtid Metaphycus lounsburyi (Howard) from Australia, but this internal parasite was not consistent in its effect. Other natural enemies were imported earlier with inconsequential results. For the 1929 trip Compere traveled via steamer and carried Wardian cages containing potted plants infested with BS. Several parasites of minor importance were collected at various localities. In earlier work on BS the celebrated Italian entomologist Filippo Silvestri (1873-1949) had reported parasites ofBS in Eritrea. While in Eritrea, Compere exposed parasitic wasps to the scale insects and cultures were established. However, in Egypt the customs officials put all of his material in a warehouse where the temperature exceeded 120° F. All of his work was lost due to excessive heat. Compere continued to search for BS parasites throughout his career. In spite the efforts of Compere, and many other entomologists, BS remains a sporadic problem. Richards & Morse (1992) provide an excellent review of BS in southern California, including a list of natural enemies collected and imported by Compere. 1932-1933 Hong Kong, China, Pakistan, India, Ceylon (California Red Scale).— The California red scale (CRS) [Aonidiella aurantii (Maskell)] was first considered a serious problem in 1877. The pest was noticed by Thomas A. Garey and L. M. Holt. Garey was a member of the Southern California Horticultural Society, dealer in nursery stock and founding member of Pomona, California. Holt was secretary of the Horticultural Society and editor of the Society’s journal. The CRS was found in Garey’s stock during 1878, but not mentioned. Subsequent histories note that CRS was first discovered in 1879, without association to Garey. Ac¬ cording to Compere, Garey was more concerned with the damage that knowledge of CRS would do to his business, than damage the scale would do to his plants (The Riverside Press, Tuesday, 26 Dec 1961). In 1892 natural enemies of CRS had been sent to California from Australia by Koebele. From 1900-1906 George Compere sent shipments intermittently from Australia to California. Silvestri searched in China during 1924-1925. Harold Compere’s objective in the orient was to find effective natural enemies of CRS. A basic tenet of classical biological control states that the most effective natural enemies of a pest are located in the center of endemicity of the pest. At that time CRS occurred virtually everywhere citrus was grown commmercially. The origin of CRS was obscured by movement of many plants by commerce. If citrus was the original host of CRS, then logic dictated an oriental origin because that is where citrus originated. However, CRS was known to thrive and reproduce on several host plants and there was no firm reason to believe that CRS necessarily originated on citrus in its native home. A notable achievement during this trip was that Compere discovered that pre¬ ceding entomologists had misidentified yellow scale [then called Chrysomphalus citrinus (Coquillett)] for the California red scale [then called Chrysomphalus au- 1994 GORDH: HAROLD COMPERE HISTORY 195 Figure 2. Harold Compere in Pakistan during 1932. rantii] . The discovery was important because these scale insects had been regarded as taxonomically indistinguishable. This discovery permitted entomologists to correctly sort parasite-host, host plant and distribution records. We have little documentation of this trip (Fig. 2). When reviewing his impor¬ tation records, we find that Compere did not significantly increase the number of species of parasitic Hymenoptera available for control of CRS. In fact, he later speculated that he had deliberately excluded species oiAphytis because he believed they were A. chrysomphali, a species ineffective in the control of CRS and known to exist in California. Later he recognized that a complex of species operated under this name (Compere 1961). 1934-1935 South America (California Red Scale). — Compere’s objective in South America was to obtain natural enemies of citrus pests, principally parasites of the CRS. The trip was subsidized by Sunkist (formerly the California Citrus Growers Exchange). Compere’s first port-of-call was Cartagena, Colombia on 25 Jul 1934. The excursion was brief. Only door-yard citrus was examined and no parasites collected. Next, he arrived at Puerto Rico on 27 Jul. Subsequent stops were made at Mayaguez, Port-of-Spain and Georgetown, but nothing was ob¬ tained. Compere arrived at Rio de Janeiro on 18 Aug 1934. Brazil was the focal point of this trip because of its extensive citrus plantings. In Brazil, he encountered several obstacles. The Brazilian government prohibited removal of any botanical or zoological material unless similar material existed in the Ministry of Agriculture or National Museum. In Rio Compere’s microscope was confiscated by Customs Officials. He was compelled to recover it at the American Embassy. [The writer had the same problem in Rio with the same microscope during 1978.] Exploration in Brazil was less than spectacular. Citrus had been in Brazil since 196 THE PAN-PACIFIC ENTOMOLOGIST VoL 70(3) the early days of Portuguese colonization. During the early part of the 20th century the citrus industry fell on hard times locally and many growers were shifting their agricultural efforts to cotton. Compere spent the last of August through November exploring Rio de Janeiro, Sao Paulo and Bahia. In Sao Paulo Compere worked with Edson Hambleton (1902-1980), and met Adolph Hempel, an American ex¬ patriot from Ohio who had worked on coccoids in Brazil since 1895. Hambleton lived in South America from 1929-1943 and was Professor and Head of the Escola Superior de Agricultura e Veterinaria in Vicosa and working on the mealy¬ bug genus Pseudococcus [For an obituary of Hambleton see Russell (1981)]. The following comments in a letter from Compere to Smith (11 Oct 1934) state rather well some aspects of foreign exploration. “The trip to Alagoinhas as a guest of the Government was not a great success. The train was scheduled to make the trip in 2 x h hours. After traveling 2 x k hours we were informed the train was 2 x /i hours late. Arrangements had been made for a return at 4:00 P.M. Sunday night. At noon we learned that the schedule had been changed and our train had left at 4:00 A.M. Caldeira, the plant inspector escort, and the interpreter that I borrowed from the Consulate had to be in Bahia Monday morning. They approached me with an offer to pay their part if I would hire a car to take us to Bahia. It was not their funeral so I paid the entire cost. First we found a motor car driver who agreed to make the trip for 180 milreis. It rained as usual. Thick gumbo mud churned by cattle in the hilly country mired us down. For a distance of several miles horses towed and countrymen pushed. For at least once in my life I looked the part of a foreign explorer. Covered with mud, on a horse in a mud hole I had my picture taken for the benefit of my critics. It was rather enjoyable except for the uncertainty of whether we were to spend the night out in rural Brazil. The principal part of the entertainment at the Citrus Experiment Station was the Director performing difficult maneuvers on a new tractor for my benefit.” The collections of Brazil for CRS were disappointing and the only species of promise were parasites of black scale. In Nictheroy, Compere found promising parasites of BS, including an undescribed species of Coccophagus. Compere was excited about this find because he felt that it offered a possibility of controlling BS in coastal California plantings where these parasites were probably too small for hyperparasites to attack. The collections in other countries of South America were not productive of parasites of CRS. 1936-1937 South and Central Africa (Black Scale, California Red Scale). — The trip to Africa was funded by Sunkist. This trip resulted in Compere collecting, importing and describing Metaphycus helvolus (Compere), now a dominant par¬ asite of BS in California. Earlier, BS was kept under some biological control by M. lounsburyi. Over the years ecological conditions changed and the pest became more pernicious. When M. lounsburyi was imported into California, BS attacked many species of trees, including false pepper, citrus and olive. The scale occurred in so-called “uneven broods.” That is, a tree could harbor all stages of BS at all times. Female M. lounsburyi attacked the “rubber-stage” which occurred im¬ mediately before adulthood. Over time, the impact of M. lounsburyi was such that the parasite induced the host to become even-brooded, or development was shifted such that long periods lapsed when the suitable host stage was not available. As a consequence, parasite populations declined. 1994 GORDH: HAROLD COMPERE HISTORY 197 Olive and citrus infested with BS became more widespread in the even-brood. False pepper continued to produce “uneven brooded” BS which in turn supported low populations of M. lounsburyi. The explanation given for this condition was that false pepper continued to produce new growth which favored scale devel¬ opment. Control of BS was still regarded as good because before the importation of M. lounsburyi, BS would frequently kill the trees. Importation of M. helvolus resulted in good control of BS, particularly along the coast of California, although the success was not as spectacular as that witnessed with the CCS or the citro- philous mealybug. Control of BS with M. helvolus was such that the pest become a problem only through pesticide upset. In the interior regions of California where the scale was even-brooded, the scale was completely controlled. Some insectaries shifted their activities from production of citrophilous mealybug parasites and Cryptolaemus to the production of M. helvolus. Many other parasites were im¬ ported by Compere during this trip including M. stanleyi Compere, Coccophagus rusti Compere and C. cowperi Girault. 1947-1948 South Africa, East Africa, Zanzibar (Codling Moth, Grape Mealy¬ bug, Citrus Red Mite).— Foreign exploration was impossible during World War II. Compere was too old for military service and he remained in Riverside at the Experiment Station working on entomological projects (Fig. 3). Following the War, Smith sent Compere to South Africa in search of natural enemies for several citrus pests, including CRS, citrus red mite, long-tailed mealybug, and Baker’s mealybug. During the war years, mealybugs had become a problem in Orange and Los Angeles Counties, red scale was a perennial problem and citrus red mite was becoming a more serious pest. This was Compere’s last foreign exploration trip. He arrived in Capetown 21 Mar 1947. The trip to Africa had been authorized nearly two years earlier and was confounded by delays. Compere wanted a car for exploration in South Africa. His experience from the earlier trip to South Africa led him to feel this would give him the mobility necessary to work effectively. During the period following WW II, new cars were difficult to secure and it took Compere nearly eight months to obtain a 1946 Chevrolet. After he purchased the car another eight months passed before he secured passage on a steamer for himself, Joan and the car. The first month in Capetown was spent with the police, traffic control, insurance, immigration and related activities. Moving around within South Africa was difficult due to police and immigration regulations. The housing problem was acute in Capetown. In large part, housing accommodations dictated collecting plans. He would not relinquish occupancy of one lodging until another had been secured. If Compere had not brought a car, then living conditions would have been considerably worse. The cost of living had skyrocketed. He was required to pay for meals missed at boarding houses and hotels. In a letter to Smith, Compere estimated that the cost of the visit doubled while the productivity was reduced. The difficulties were partially com¬ pensated by the generous loan of a fully equipped laboratory at the Low Tem¬ perature Laboratory on Portswood Road, Capetown by Mr. Rees-Davies. Compere’s objectives in the Cape involved the codling moth [Cydia pomonella (Linneaus)], predatory coccinellids and the grape mealybug (GM) [Pseudococcus maritimus (Ehrhorn)]. Smith was anxious to obtain codling moth parasites reared 198 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) / / i I Figure 3. Harold Compere in Riverside during 1940. at the Western Province Research Institute. When Compere arrived in Stellen¬ bosch during Apr 1947 he found none to spare for export to California. Conse¬ quently he shifted his attention to other objectives. He had seen a coccinellid feeding on CRS at Clan william in 1936 and hoped to locate it again. Also in the Cape the so-called GM was known as a pest of grapes only. During 1936 he could 1994 GORDH: HAROLD COMPERE HISTORY 199 not find this pest on citrus, ornamentals or deciduous fruit. Compere speculated that the pest on grape in the Cape was not the same as that in southern California. Compere visited Clanwilliam 16-17 Apr 1947. The CRS picture had changed considerably from his earlier visit. In 1947 he noted citrus trees being killed by CRS infesting the branches. The situation was so dramatic that in one grove more than 10,000 dead citrus trees had just been removed. Compere could not find the coccinellid he had seen earlier and the only natural enemies located were Chilo- corus lophanthae (Blaisdell) and an Aphytis, presumably chrysomphali (Mercet). During May 1947 he visited the Letaba Estates of northeastern Transvaal near the Murchison Range. The estate was owned by a Dr. Merensky, a German geologist and mining engineer who arrived in Southwest Africa during 1908 and explored Namaqualand. Returning to Germany in 1909, Merensky wrote a paper which predicted the existence of diamonds in the alluvial deposits not connected with the Kimberley Mines. Merensky’s paper went unnoticed so he put together enough money to explore the area in 1926. He discovered diamonds in Nama¬ qualand as predicted, filed mining rights to 20 claims, sold these in 1928 and went into agriculture. Merensky was Compere’s host at Letaba. The Estate consisted of about 200,000 acres planted half in Valencia and half in Navel oranges. CRS was as abundant as in southern California. As in California, the navels were more heavily infested and more prone to attack by CRS than Valencias. Compere noted that the trees at Letaba sustained heavier infestation and recovered more rapidly than trees planted in California. Letaba provided a superb natural laboratory for the study of CRS as it affected citrus. Nowhere in his travels had he encountered a problem of this magnitude under the control of one grower. The climatic conditions were comparable with California but the summer was slightly warmer than Orange County. May was considered winter in Transvaal but he recorded Aphytis chrysomphali and coccinellid predators as abundant. The distribution of CRS was not even and the distribution of parasites was not straightforward. Although this visit was informative, it was not productive in terms of new natural enemies. Compere reluctantly visited Zanzibar during October and November 1947. Bureaucratic entanglements with his car and the problems of securing living ac¬ commodations enhanced Compere’s reluctance to voluntarily move. Smith sec¬ onded Compere to the Pacific Science Board to search for Scolia ruficornis (Fabr.), a wasp parasitic on the beetle Oryctes rhinocerus (L.), then a serious pest of coconut palm in the South Pacific. Compere had been in Nairobi during July thru Sep¬ tember. Smith finally ordered Compere to undertake the work, an action which both men found unpleasant. In more than 25 years of association Smith never ordered Compere to do anything. Smith always relied on Compere’s judgement to act independently in the field. F. X. Williams of the Hawaiian Sugar Planters Association replaced Compere in this work. Compere returned to Nairobi and was operated on 9 Jan 1948 for an hernia. Ten weeks later the hernia reappeared. Medical attention was provided at the European Hospital in Nairobi, the same hospital in which Joan was resident three months with a broken leg. During April 1948 he was particularly interested in working with parasites of long-tailed mealybug in Kenya. He recovered a species of an Anagyrus near A. maritimus which had been sent to California earlier. THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) 200 Compere later described the parasite as Anagyrus kivuensis. Compere felt that if the parasite did succeed that it would justify the trip to Africa. In Kenya this parasite was rated as the most important parasite of coffee mealybug, Planococcus kenyae (LePelly). At this time S. E. Flanders made requests for Chilocorus wahl- bergi Mulsant. Compere located this coccinellid in navel orange groves at Thika which were heavily infested with California and Florida red scale, purple scale, russet mite and red spider mite. The occurrence was particularly noteworthy because Compere had standing orders to secure natural enemies of RSM. He located staphylinid larvae feeding on this mite on avocado, but apparently other insects were not observed feeding on this pest. By June 1948 Compere was looking forward to returning to the United States. The medical problems encountered by Harold and Joan had been significant. This, coupled with the poor collecting conditions in Kenya, hastened his attempt to depart for home. Unfortunately, the political situation complicated matters. His Chevrolet, according to Kenya law, had to be sold at a price fixed by the Controller of Motor Vehicles. Payment was to be in East African shillings and could not be converted into U.S. dollars legally. To further complicate matters removal of large sums (including shillings) from East Africa was prohibited. Domestic Exploration 1951 (Western Grape Leaf Skeletonizer).— Compere’s last expedition for natural enemies was to the Atlantic and Central States for parasites of Harrisina americana (Guerin) which might be used in California against H. brillians (GLS) Barnes & McDunnough. The trip (13 Jul-26 Sep 1951) involved 17 shipments. At the time GLS was a significant pest of grapes in California. Smith had retired shortly before Compere’s trip, but Smith directed the project. He believed that beneficial insects could be obtained from the east. Compere selected Columbus, Ohio as the starting point owing to its central location and because H. americana had been recorded from there, but had not achieved pest status in commercial plantings of grapes. The trip was made by car and he engaged in virtually no collecting along the way to Columbus because of the lateness of the season and prevailing weather con¬ ditions. Compere visited grape plantings in Ohio which revealed nothing. He shifted to New York due to the reported outbreak of H. americana in 1949 at Cayuga County. Unfortunately, Compere could not locate the site of the reported infes¬ tation. Plantings of grapes were infested with another skeletonizer, not H. amer¬ icana. P. J. Hartzell at the Geneva Experiment Station suggested Compere try the Hudson Valley but this exploration was unproductive. A subsequent visit to New Jersey revealed nothing. Compere next tried Pennsylvania because H. americana was considered a pest there. The Lepidopterist S. W. Frost took Compere to Shingletown Gap where the first collection of H. americana larvae was made and shipped to Riverside on 9 Aug. A second shipment was sent from material taken at Morganstown, West Virginia on 13 Aug 1951. Compere had been plagued with asthma and the condition had become intol¬ erable. Compere went to Florida for relief and where there was some promise of finding natural enemies. A shipment of larvae was taken near Jacksonville and sent to Riverside on 17 Aug. Additional collections were taken at Jacksonville which revealed parasites, and another collection made at Titusville. On 1 Sep he 1994 GORDH: HAROLD COMPERE HISTORY 201 left the state and next made a small collection in Tennessee. The trip yielded nothing of substance and must be regarded as a failure. Collecting Techniques Harold Compere was among the most successful collectors in terms of number of species obtained abroad, the number of establishments in California and bi¬ ological control achievements. His methodology probably influenced his perfor¬ mance record. Compere’s approach to foreign exploration remained essentially constant throughout his career and is best summarized in his own words in a letter to S. E. Flanders dated 9 Jun 1947. “I am more than ever of the opinion that sidewalk exploring in the big cities is far more profitable than orchard ex¬ ploration. However, sidewalk exploring is [sic] nean and dirty work. Orchard exploring is a grand life, especially in Africa where the total citrus plantings are about equal to those of Tulare County. ... In the past I have been criticized for not getting out into the orchards.” Someone with foreign exploration experience can appreciate the wisdom in Compere’s words. The threat to personal safety has always been a real and persistent problem. Dogs were Compere’s particular nem¬ esis and he always carried a pair of oversize gardener’s shears for self protection on sidewalk exploration. Systematic Studies of Parasitic Hymenoptera From the standpoint of applied biological control, parasitic Hymenoptera are among the most important beneficial insects. Many of the insects which he col¬ lected were new to science. Taxonomy of the parasitic Hymenoptera was in a confused state and there were too few taxonomists working on these wasps to provide rapid and correct identifications. P. H. Timberlake was hired to work on the taxonomy of parasitic Hymenoptera but he shifted interests and worked on the taxonomy of bees. These facts made Compere’s involvement in taxonomy imperative. Between foreign exploration trips, Compere focused his attention on the taxonomy of parasitic Chalcidoidea, mostly Aphelinidae and Encyrtidae. These families are numerically large, taxonomically difficult and represented the bulk of the material collected by Compere. The descriptions of his species may be found in the publications listed below. Compere was a self-educated taxonomist whose work with aphelinids is note¬ worthy. Compere (1931) provided the first modem revision of Coccophagus, a cosmopolitan genus consisting of about 200 species. Members of the genus dem¬ onstrate complex biological habits and may serve as primary or secondary par¬ asites. Compere (1955) also developed the first revisionary study of the genus Aphytis. Species of Aphytis are the primary parasites of armored scale insects and are important in control of diaspidids (Rosen & DeBach 1978). The history of Aphytis in biological control makes a compelling story. Aspects of that history were provided by Compere (1961) in his classic paper on the California red scale. Before Compere’s work, Aphytis contained only a few species. He described several species and drew attention to several species which were morphologically similar and biologically useful. His taxonomic studies served as the foundation for the work of Paul DeBach (1913-1992) in applied biological control of scale insects. DeBach, with graduate students and colleagues, pointed to the existence of nu¬ merous sibling species complexes within Aphytis. Presently we recognize more 202 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) than 200 species of Aphytis with many species remaining undescribed (Rosen & DeBach 1979). Compere’s work in the Encyrtidae was less significant but still worthy of men¬ tion. This family contains about 500 genera and more than 3,300 species. All species are primary or secondary parasites. Most hosts are Homoptera and con¬ sequently the family holds importance in control of many pests. Compere worked on some important genera, such as Metaphycus and Aphycus, but he did not publish improvements in the classification of the family. The Encyrtidae were too large and knowledge was too diffuse for Compere to make a significant improve¬ ment in their taxonomy. Harold Compere retired as Specialist on 1 Jul 1963. He did not hold an academic appointment and consequently he was not entitled to the privileges accorded emeriti. A petition was made in his behalf and he was accorded emeritus status. A petition was made to award him an honorary Ph.D., but this was denied because the University of California did not engage in this practice at the time. Once, Compere had been appointed to the faculty. However, this placed him under a less desirable retirement system. He successfully sought to have the appointment eliminated and his status as a Specialist reinstated. As a Specialist, Compere could not supervise graduate students, but he did influence some of them who passed through Riverside. Paul DeBach was first attracted to biological control after seeing a photograph in the Los Angeles Times of Harold Compere on a camel in Eritrea. During the 1950s and early 1960s Compere worked with the Italian Giulio Zinna. Unfortunately, after returning to Italy, Zinna committed suicide and prematurely terminated a potentially pro¬ ductive partnership. Compere also collaborated with the noted South African chalcidoid taxonomist David P. Annecke (1928-1981). Compere had a strong interest in morphology and held a great admiration for Gordon Floyd Ferris (1893-1958). During the latter years of life Harold Compere became obsessed with principles of morphology as they related to chalcidoid classification (see his last publications below). He felt that classifications should be phylogenetic in the sense that they reflect evolutionary modification of ana¬ tomical features. The higher classification of the Chalcidoidea has always been confused, in part due to the large number of species and bewildering array of anatomical modifications. Compere thought that his retirement pastime would be the clarification of taxonomic relationship through detailed analysis of ana¬ tomical structure. Compere undertook a detailed morphological analysis of all Hymenoptera with a goal of demonstrating homology and tracing modification through descent. His approach was sound in that he would take representatives of many families of chalcidoids and study them part-by-part in a comparative manner. His notion of a perfected classification embraced the concept of shared derived characters. He was tormented by the fact that he could not find a common groundplan for many important structural features in the groups which he studied. He agonized over these problems for nearly fifteen years following retirement. He would draft manu¬ scripts considering each anatomical feature in some detail. Unfortunately the gaps between taxa he studied were tremendous and he could not account for modifi¬ cations or trends of change given his limited selection of data points. In the end he quit and all that remains of his studies are boxes of slides with his dissections, 1994 GORDH: HAROLD COMPERE HISTORY 203 photomicrographs and reams of unpublished text. Final abandonment of mor¬ phological studies came around 1975. Simply put, the problem exceeded his ability. Harold Compere died 3 Feb 1978 at his home at 1900 Bonnie Brae, above Tequesquite Arroyo, at Riverside, California. He was cremated and his ashes interred at Rosedale Cemetery in Los Angeles, a few city blocks from the house in which he was born. Acknowledgment I thank Ted Fisher, David Headrick and Sergei Trjapitzin for reading the manu¬ script and commenting upon it. Literature Cited Anonymous. 1935. Jordi Compere (1858-1928). Arx. Esc. Sup. Agr. Barcelona (n.s.), 1: 293-299. Boyce, A. M. 1969. History of the Citrus Research Center and Agricultural Experiment Station. Proc. 1st Intern. Citrus Symp., pp 49-55. Doutt, R. 1958. Vice, virtue and the Vedalia. Bull. Entomol. Soc. Am., 4.1: 119-123. Essig, E. O. 1931. A history of entomology. The MacMillan Company, New York. Howard, L. O. 1930. A history of applied entomology. Smithsonian Miscellaneous Coll. 84, 564 pp. Richards, L. & J. Morse. 1992. A survey of black scale, Saissetia oleae [Horn.: Coccidae] parasitoids [Hym.: Chalcidoidea] in southern California. Entomophaga, 37: 373-390. Rosen, D. & P. DeBach. 1978. Diaspididae. pp. 78-128. In Clausen, C. P. (ed.). Introduced parasites and predators of arthropod pests and weeds: a world review. USDA Agric. Handbook 480. Rosen, D. & P. D. DeBach. 1979. Species of Aphytis of the world (Hymenoptera: Aphelinidae). Dr. W. Junk BV Publishers, The Hague. Russell, L. M. 1981. Edson J. Hambleton. Proc. Entomol. Soc. Wash., 83: 564-569. Wade, J. S. 1955. Carl Heinrich. Proc. Entomol. Soc. Wash., 57: 249-255. Bibliography of Harold Compere 1. 1916. Notes on the tomato psylla. Mon. Bull. Calif. State Com. Hort. 5: 189-191. 2. 1916. (with H. S. Smith) Observations on the Lestophanus, a dipterous parasite of the cottony- cushion scale. Mon. Bull. Calif. Com. Hort. 5: 383-389. 3. 1920. (with H. S. Smith) The life history and successful introduction into California of the black scale parasit e, Aphycus lounsburyi How. Mon. Bull. Calif. State Dept. Agr. (8): 311-320. 4. 1921. Seasonal history of the black scale and relation to biological control. Calif. Citrog. 6 (6): 197. April. 5. 1922. The black scale problem. Calif. Cultiv. 59 (2): 29-30. 6. 1924. A preliminary report on the parasitic enemies of the citricola scale, Coccus pseudomag- nolarum (Kuwana) with descriptions of new chalcidoid parasites. Bull. So. Calif. Acad. Sci. 23: 113-123. 7. 1925. New chalcidoid (hymenopterous) parasites and hyperparasites of the black scale, Saissetia oleae Bernard. Univ. Calif. Publ. Entomol. 3: 295-326. 8. 1925. A new genus and species of Aphelinidae (Hymenoptera) from China. Trans. A m. Entomol. Soc. 51 (871): 129-134. 9. 1926. Descriptions of new coccid-inhabiting chalcidoid parasites (Hymenoptera). Univ. Calif. Publ. Entomol. 4: 1-31. 10. 1926. New coccid-inhabiting parasites (Encyrtidae, Hymenoptera) from Japan and California. Univ. Calif. Publ. Entomol. 4: 33-50. 11. 1926. Descriptions of new coccid-inhabiting chalcidoid parasites (Hymenoptera). Univ. Calif. Publ. Entomol. 4: 51-61. 12. 1926. (with H. S. Smith) The establishment in California of Coccophagus modestus Silv. (Aphe¬ linidae, Hymenoptera) with notes on its life history. Univ. Calif. Publ. Entomol. 4: 51-61. 204 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) 13. 1927. (with H. Smith) Notes on the life history of two oriental chalcidoid parasites of Chrys- omphalus. Univ. Calif. Publ. Entomol. 4: 63-73. 14. 1928. New coccid-inhabiting chalcidoid parasites from Africa and California. Univ. Calif. Publ. Entomol. 4: 209-230. 15. 1928. (with H. S. Smith) A preliminary report on the insect parasites of the black scale Saissetia oleae (Bernard). Univ. Calif. Publ. Entomol. 4: 231-334. 16. 1928. Successful importation of five new natural enemies of citrophilus mealybug from Australia. Calif. Citrog. 13 (9): 318. 17. 1928. Establishment in state of newly introduced mealybug parasites. Calif. Citrog. 14c (1): 5. November. 18. 1929. (with H. S. Smith) New insect enemies of the citrophilus mealybug from Australia. Mon. Bull. Calif. State Dept. Agric. 18: 214-218. 19. 1929. Description of a new species of Coccophagus recently introduced into California. Univ. Calif. Publ. Entomol. 5: 1-3. 20. 1930. An account of a trip to Eritrea to obtain insect enemies of the black scale, Saissetia oleae (Bern.). Calif. Citrog. 15: 533, 562-568. 21. 1931. Revision of the species of Coccophagus—a. genus of hymenopterous coccid-inhabiting parasites. Proc. U.S. Nat. Mus. 78: 1-132. 22. 1931. A revision of the genus Diversinervus Silvestri, encyrtid parasites of coccids (Hymenoptera). Univ. Calif. Publ. Entomol. 5: 233-245. 23. 1931. A discussion of the parasites of Saissetia oleae (Bern.) collected in Eritrea. Univ. Calif. Publ. Entomol. 5: 247-255. 24. 1931. The African species of Baeoanusia, an encyrtid genus of hyperparasites (Hymenoptera). Univ. Calif. Publ. Entomol. 5: 257-264. 25. 1931. New encyrtid (hymenopterous) parasites of Pseudococcus species from Eritrea. Univ. Calif. Publ. Entomol. 5: 265-274. 26. 1931. (with H. S. Smith) An imported parasite attacks the yellow scale. Calif. Citrog. 16 (7): 328. 27. 1931. (with H. S. Smith) Notes on Ophelosia crawfordi. J. Econ. Entomol. 24: 1109-1110. 28. 1932. (with H. S. Smith) The control of the citrophilus mealybug, Pseudococcus gahani by Australian parasites. Hilgardia 6: 585-618. 29. 1933. The parasites of Pseudococcus comstocki Kuw. Canad. Entomol. 65: 243-247. 30. 1933. Tracking down red scale parasites in the mysterious Orient. Citrus Leaves 13 (5): 1-3, 12. 31. 1933. Proposed distribution of the black scale parasite Coccophagus trifasciatus Compere. Univ. Calif. Coll. Agr. News Letter No. 7, Div. Benef. Insect Invest. Citrus Exp. Sta. 32. 1933. (with S. E. Flanders) Anar ho pus Sydneyensis Timb., an encyrtid parasite of Pseudococcus longispinus (Targ.) recently introduced into California from Australia. J. Econ. Entomol. 28: 966-973. 33. 1935. Exploratory search for natural enemies of the red scale. (Extracts from an unpublished manuscript arranged by the editors of the California Citrograph). Calif. Citrog. 20 (12): 371, 383-388. 34. 1935. Red scale parasite search in South America described by Compere. Citrus Leaves 15 (10): 1, 2, 14, 18. 35. 1936. A new genus and species of Encyrtidae parasitic in the pineapple mealybug, Pseudococcus brevipes (Ckll.). Proc. Haw. Entomol. Soc. 9: 171-174. 36. 1936. Notes on the classification of the Apholinidae, with descriptions of new species. Univ. Calif. Publ. Entomol. 6: 277-322, 19 figs. 37. 1936. A new species of Habrolepis parasitic in Chrysomphalus aurantii Mask. Bull. Entomol. Res. 27: 493-496. 38. 1937. Coccid-inhabiting parasites from Africa, with descriptions of new Encyrtidae and Aphe- linidae. Bull. Entomol. Res. 28: 43-51, 3 figs. 39. 1937. The species of Aenasius, encyrtid parasites of mealybugs. Proc. Haw. Entomol. Soc. 9: 383-404, 4 figs. 40. 1937. Collecting red and black scale parasites in Africa. Calif. Citrog. 23 (2): 58, 59, 87-89. 41. 1938. Things not entomological on tour of Africa in search of parasites. Calif. Citrog. 23 (3): 122, 124. 42. 1938. Description of a new species of Leptomastix parasitic i n Phenacoccus hirsutus Green. Bull. Soc. Fouad Entomol, pp. 36-38 “Se’ance du It.” 1994 GORDH: HAROLD COMPERE HISTORY 205 43. 1939. A report o n some miscellaneous African Encyrtidae i n the British Museum. Bull. Entomol. Res. 29: 315-337. 44. 1939. A second report on some miscellaneous African Encyrtidae in the British Museum. Bull. Entomol. Res. 30: 1-26. 45. 1939. Mealybugs and their insect enemies in South America. Univ. Calif. Publ. Entomol. 7: 57-74. 46. 1939. The insect enemies of the black scale, Saissetia oleae (Bern.), in South America. Univ. Calif. Publ. Entomol. 7: 75-90. 47. 1940. The African species of Metaphycus Mercet. Bull. Entomol. Res. 31: 7-33. 48. 1940. A new species of Metaphycus (Hymenoptera, Encyrtidae) from Australia parasitic in Eriococcus coriaceus Maskell. Trans. Roy. Soc. So. Austral. 65: 46-47. 49. 1940. Parasites of the black scale, Saissetia oleae, in Africa. Hilgardia 13: 387-425. 50. 1941. (with S. E. Flanders & H. S. Smith) Use air transport from China for introduction of parasites. Calif. Citrog. 26: 291, 300-301. 51. 1943. A new species of Metaphycus parasitic on psyllids. Pan-Pac. Entomol. 19: 71-73. 52. 1943. A new and economically important species of Anagyrus from Africa. Bull. Entomol. Res. 34: 129-130. 53. 1947. A new species of Encyrtidae parasitic in Coccus hesperidum, L. Bull. Entomol. Res. 38: 281-283. 54. 1947. A new genus and species, Eurymyiocnema aphelinoides (Hymenoptera, Aphelinidae), and a history of the genera Euryischia Riley and Myiocnema Ashmead. Bull. Entomol. Res 38: 381-388. 55. 1947. A report on a collection of Encyrtidae with descriptions of new genera and species. Univ. Calif. Publ. Entomol. 8: 1-24. 56. 1953. An appraisal of Silvestri’s work in the orient for the University of California, some misidentifications corrected and two forms of Casca described as new species. Bol. Lab. Zool. Gen. Agr. “Filippo Silvestri” (Portici) 33: 35-46. 57. 1955. A systematic study of the genus Aphytis Howard (Hymenoptera, Aphelinidae) with de¬ scriptions of new species. Univ. Calif. Publ. Entomol. 10: 271-320. 58. 1955. (with Giulio Zinna) Tre nuovi genrei e cinque nuove specie di Encyrtidae. Stabilimento Tipografico Guglielmo Genovese (Napoli). Bol. Lab. Entomol. Agr. ‘Filippo Silvestri’ (Portici) 14:94-116. 59. 1957. Descriptions of species of Metaphycus recently introduced into California and some corrections. Bol. Lab. Entomol. Agr. “Filippo Silvestri” (Portici) 15: 221-230. 60. 1960. (with B. R. Subba Rao & R. B. Kaur) Two species of Encyrtidae parasitic in the pink mealybug of sugarcane in India—(Hymenoptera). Proc. Nat. Inst. Sci. India 26 B (1): 45-50. 61. 1960. (with D. P. Annecke) A reappraisal of Aphycus Mayr, Metaphycus Mercet, and allied genera (Hymenopt.: Encyrtidae). J. Entomol. Soc. So. Afr. 23: 375-389. 62. 1961. (with D. P. Annecke) Descriptions of parasitic Hymenoptera and comments (Hymenopt.: Aphelinidae, Encyrtidae, Eulophidae). J. Entomol. Soc. So. Afr. 24: 17-71. 63. 1961. The red scale and its insect enemies. Hilgardia 31: 173-278. 64. 1962. The reality of stemites in the mesothorax of Hymenoptera. Proc. Entomol. Soc. Wash. 64: 224-228. 65. 1969. Changing trends and objectives in biological control. Proc. 1st Intern. Citrus Symp. 2: 755-764. 66. 1969. The role of systematics in biological control: a backward look. Israel J. Entomol. 4: 5-10. 67. 1970. (with D. Rosen) The prescutum in Hymenoptera. Proc. Roy. Entomol. Soc. London (A) 45 (7-9): 91-97. PAN-PACIFIC ENTOMOLOGIST 70(3): 206-211, (1994) ESTABLISHMENT OF UROPHORA SIRUNASEVA (HERING) (DIPTERA: TEPHRITIDAE) FOR BIOLOGICAL CONTROL OF YELLOW STARTHISTLE IN THE WESTERN UNITED STATES C. E. Turner, 1 R. Sobhian, 2 D. B. Joley, 3 E. M. Coombs, 4 and G. L. Piper 5 United States Department of Agriculture, Agricultural Research Service, Western Regional Research Center, Albany, California 94710 2 United States Department of Agriculture, Agricultural Research Service, European Biological Control Laboratory, BP 4168-Agropolis, 34092 Montpellier, Cedex 5, France 3 California Department of Food & Agriculture, Biological Control Program, Sacramento, California 95814 4 Oregon Department of Agriculture, Weed Control, Salem, Oregon 97310 5 Department of Entomology, Washington State University, Pullman, Washington 99164 Abstract. — Urophora sirunaseva (Hering) (Diptera: Tephritidae) is a capitulum-galling natural enemy of yellow starthistle, Centaurea solstitialis (Asteraceae). The fly was first introduced into the United States for biological control of yellow starthistle in the mid-1980s. As of 1992, field establishment of U. sirunaseva is known from California (five sites), Oregon (six sites), and Washington (one site). Field sample data for four populations of U. sirunaseva from California and Oregon in 1992 yielded a range of galled host capitula from 22.1 to 44.0%, and a range of mean galls per galled capitulum from 1.8 to 2.1. Key Words. — Insecta, gall, biological control, weed, Urophora, Centaurea Yellow starthistle {Centaurea solstitialis L., Asteraceae) is an Eurasian annual that is a highly invasive, naturalized weed of grasslands and other environments in the western United States, especially in California (~ 3,200,000 ha infested), Idaho (~ 81,000 ha infested), Oregon (~ 400,000 ha infested), and Washington (~ 54,000 ha infested) (Maddox & Mayfield 1985, Maddox et al. 1985, Roche & Roche 1988, Callihan et al. 1989, Turner et al. in press). The weed displaces native and other more desirable plants, the spiny capitula deter grazing by livestock and are a nuisance to people working or recreating on infested lands, and it is poisonous to horses (Cordy 1978). Urophora sirunaseva (Hering) (Diptera: Tephritidae) is a natural enemy of yel¬ low starthistle from Greece eastwards (White & Clement 1987). Urophora siru¬ naseva females posit fusiform eggs in closed capitula of yellow starthistle (Sobhian 1993, Turner in press). In a laboratory cage study conducted in Greece, Sobhian (1993) observed up to 270 oviposited eggs per female, and a mean of 167 ovi¬ posited eggs per female. Turner (in press) recorded an average of 136 oviposited eggs per female in a laboratory cage study in California. Lignified, unilocular galls are formed around the developing larvae within capitula. The fly is bivoltine, and overwinters as mature larvae in galls on host capitula. Groppe et al. (1990) and Clement & Sobhian (1991) conducted host specificity tests in field plots in northern 1994 TURNER ET AL.: TEPHRITID CONTROL OF THISTLE 207 Greece, and Turner (in press) carried out host specificity tests in a quarantine glasshouse. These tests as well as field host records (White & Korneyev 1989) indicate a very high level of host specificity and safety for U. sirunaseva as a biological control agent for yellow starthistle. Field Releases and Establishment The first field releases of U. sirunaseva in North America occurred in 1984, when flies imported from Greece were released in California, and flies imported from Turkey were released in Idaho (Turner et al. in press). In 1985, additional releases of flies from Greece were made in California, Idaho, Oregon, and Wash¬ ington. After the 1985 releases, U. sirunaseva releases ceased until the taxonomic confusion between U. sirunaseva and the closely related and very similar U. jaculata Rondani was clarified (Turner et al. in press) by White & Clement (1987) and White & Korneyev (1989). The fly was thought not to have established until populations were discovered in 1989 at each of its only release sites near Loomis, California (1984 and 1985 releases) and Phoenix, Oregon (1985 release). Field releases commenced in 1989 using material imported from Greece, or flies col¬ lected from the initial California and Oregon populations. As of 1992, U. sirunaseva was known to be established in California, Oregon, and Washington. The source of all established populations is the Thessaloniki area of northern Greece. Urophora sirunaseva larvae in galled yellow starthistle capitula were shipped from Greece, and releases were made with the adults that emerged in the USDA-ARS quarantine facility in Albany, California. A more detailed description of established populations through the 1992 field season follows. California. — Urophora sirunaseva is established at five sites in California: Loomis (Placer Co.), Hornbrook (Siskiyou Co.), Ukiah (Mendocino Co.), Rancho Cordova (Sacramento Co.), and Mankas Corner (Napa Co.). Flies imported from Greece were released at Loomis in 1984 in two separate releases of 142 adults (78 females, 64 males) and 42 adults (21 females, 21 males), and in a 1985 release of 60 adults (30 females, 30 males). It is not certain which of these releases resulted in estab¬ lishment. The Hornbrook population was founded by flies (« 150 females, 200 males) released in 1990 from adults collected at Phoenix, Oregon. Flies (111 females, 165 males) collected from the Phoenix, Oregon population were also released in 1990 at Mankas Corner; in addition, flies (116 females, 192 males, but unquantified mortality due to high ambient temperature at the time of release) from Greece were released in 1991 at Mankas Comer. It is not certain which of these releases resulted in establishment at Mankas Corner. Urophora sirunaseva imported from Greece were released as adults (204 females, 168 males) at the Ukiah site in 1990. The Rancho Cordova population was founded by adult flies (225 presumably mated females) released in 1991 that were collected at Loomis. Oregon. — Urophora sirunaseva is established at six sites in Oregon: Phoenix, Brownsboro, and Black Butte (Jackson Co.); Myrtle Creek and Riddle (Douglas Co.); and East Grants Pass (Josephine Co.). Flies imported from Greece were released (100 females, 100 males) in 1985 at Phoenix. All other flies released in Oregon originated from this population. Flies were released in 1989 at Myrtle Creek (75 females, 50 males) and at Riddle (75 females, 50 males). Flies (250 adults) were released at Riddle again in 1990 as well as at Brownsboro (100 adults). 208 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) In 1991 , U. sirunaseva was released at Black Butte (100 adults) and at East Grants Pass (100 adults). Where the gender ratio is not specified, it was estimated to be ~ 1 : 1 . Washington. — Urophora sirunaseva is established at Colfax (Whitman Co.). This population originated from 400 adults (200 females, 200 males) collected from Phoenix, Oregon and released in 1990. Further field releases of U. sirunaseva collected from domestically established populations are planned in the United States. Our redistribution experience thus far has shown that establishment can readily result from releases of either first or second generation adults, and from releases of relatively small numbers of flies (< 100 females). Materials and Methods A quantitative sampling study was undertaken to assess the status of established field populations of U. sirunaseva in California and Oregon. Sampling began in 1989 at the Loomis, California and Phoenix, Oregon populations; and in 1992 at the Hombrook, California and Ukiah, California populations. Except for the Ukiah population, sampling was carried out along eight major lines (N, NE, E, SE, S, SW, W, NW) established from a central stake at each release site. Ten samples of whole plants or branches of large plants were taken at random intervals generated by a hand calculator along each line. This sampling method concentrates the sample points in the central area around the release site within a population. The Ukiah population is an elongated patch of yellow star- thistle distributed along a steep, narrow hillside. Sampling here was carried out along two lines parallel to the length of the hillside, with samples taken at 40 random points along each line. All sampled capitula were dissected in the labo¬ ratory, and counts of U. sirunaseva galls were made. All samples were taken in late summer/early fall, thus galls from both generations were counted. Results and Discussion Thus far, there are similar values for percentage galled capitula and number of galls per galled capitulum among three of the populations (Table 1). The percentage of galled capitula at the Ukiah population is approximately twice that of the other populations. The Ukiah yellow starthistle population is somewhat isolated from other host populations in the surrounding area by urban development, whereas the other infested C. solstitialis populations that were sampled are essentially continuous with surrounding areas occupied by the weed. Thus the flies may be more readily spreading throughout the surrounding area at Loomis, Hombrook, and Phoenix, which would dilute fly density and reduce the percentage of infested capitula at any one sample area. Flies have been detected in Medford, Oregon, a distance of « 10 km from the Phoenix release site, and in Newcastle, California, a distance of ~ 10 km from the Loomis release site. Urophora sirunaseva was found in Yreka, California in 1993, a distance of « 21 km from Hombrook within three years of its release there. This degree of dispersal is encouraging from a biological control viewpoint. All four populations sampled had a mean of approximately two galls per galled 1994 TURNER ET AL.: TEPHRITID CONTROL OF THISTLE 209 Table 1. Infestation of yellow starthistle capitulaby Urophora sirunaseva, 1989-1992. 1989 1990 1991 1992 Loomis, CA a % Galled capitula 1.1 11.7 21.5 22.1 (No. capitula sampled) (417) (382) (1716) (483) Mean ± SEM galls per galled capitulum 1.0 ± 0.0 1.6 ± 0.1 2.3 ± 0.1 2.1 ± 0.1 (No. galled capitula sampled) (5) (45) (370) (107) Maximum no. galls per galled capitulum 1 4 10 7 Phoenix, OR b % Galled capitula 2.2 18.1 19.6 22.9 (No. capitula sampled) (311) (253) (265) (353) Mean ± SEM galls per galled capitulum 1.4 ± 0.2 1.7 ± 0.1 1.8 ± 0.1 1.8 ± 0.1 (No. galled capitula sampled) (7) (46) (52) (81) Maximum no. galls per galled capitulum Hombrook, CA c 3 5 6 7 % Galled capitula 23.7 (No. capitula sampled) (270) Mean ± SEM galls per galled capitulum 2.1 ± 0.1 (No. galled capitula sampled) (64) Maximum no. galls per galled capitulum Ukiah, CA c % Galled capitula (No. capitula sampled) 6 44.0 (724) Mean ± SEM galls per galled capitulum 1.9 ± 0.1 (No. galled capitula sampled) (319) Maximum no. galls per galled capitulum 7 a Field released 1984 and 1985. b Field released in 1985. c Field released in 1990. capitulum in 1992 despite galled capitula of < 25% at three out of four locations (Table 1). Figure 1 shows the frequency distribution of galls per galled head for these populations in 1992. More than half of the galled capitula at each site had two or more galls. The distribution of galls per galled capitulum from field pop¬ ulations is quite similar to that obtained during host specificity testing of the fly in quarantine glasshouse studies (Turner in press), and may indicate a tendency towards aggregation, i.e., capitula infested by multiple larvae. This could be im¬ portant from a control standpoint, as due to the modest size of a single gall, infestation by multiple larvae appears to be necessary to destroy a meaningful percentage of seeds (Turner, unpublished data). The maximum number of galls per capitulum observed in field populations of the fly is nine in northern Greece (Sobhian, unpublished data); in a separate sample of 215 capitula for a seed destruction study in 1992, we recorded 12 galls in a capitulum at Loomis. We do not know with certainty whether multiple galls in a capitulum originate from one or more females. However, dissected capitula can reveal multiple eggs grouped together indicating that they originate from a single female in at least some instances. Over time we would expect an increase both in percentage galled capitula and in galls per galled capitulum, which is evident 210 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) 05 3 'a. 03 O ■o 0) la CD 5 H— o 1 2 3 4 5 6 7 J5 1 "cl 03 O TJ 0 To C5 M— o vO 0 s - 1 2 3 4 5 6 7 No. Galls per Galled Capitulum No. Galls per Galled Capitulum No. Galls per Galled Capitulum No. Galls per Galled Capitulum Figure 1. Number of Urophora sirunaseva galls per galled yellow starthistle capitulum in 1992 at (A) Loomis, California, (B) Phoenix, Oregon, (C) Hombrook, California, (D) Ukiah, California. at the Loomis and Phoenix populations (Table 1). Both types of increase should be necessary to significantly reduce seed production by yellow starthistle. Urophora sirunaseva is presently one of five capitulum-attacking insects from Greece established in the western United States for biological control of yellow starthistle (Turner et al. in press). The other established species are the weevils Bangasternus orientalis (Capiomont), Eustenopus villosus (Boheman), and Larinus curtus Hochhut, and the tephritid fly Chaetorellia australis Hering (Turner et al. in press). The entire course of capitulum development (from the very early closed bud stage to flowering) in yellow starthistle is covered by the timing of oviposition among these five insect species (Turner et al. in press), which provides some basis for optimism in terms of the combined biological control potential of these insects. 1994 TURNER ET AL.: TEPHRITID CONTROL OF THISTLE 211 Acknowledgment We thank K. L. Chan for technical assistance throughout the course of the field work. D. M. Maddox and A. Mayfield made the 1984 and 1985 field releases in California, and facilitated the releases in other states during those years; M. A. Garcia assisted with the Ukiah release; B. Villegas helped collect the flies and released them at the Rancho Cordova site; T. Allen first detected the establishment of the Mankas Comer population; L. Knutson, J. P. McCaffrey, N. J. Mills, and M. Pitcairn provided critical reviews of the manuscript. Literature Cited Callihan, R. H., F. E. Northam, J. B. Johnson, E. L. Michalson & T. S. Prather. 1989. Yellow starthistle: biology and management in pasture and rangeland. Univ. Idaho Coop. Ext. Curr. Inf. Ser., No. 634. Clement, S. L. & R. Sobhian. 1991. Host-use patterns of capitulum-feeding insects of yellow star- thistle: results from a garden plot in Greece. Environ. Entomol., 20: 724-730. Cordy, D. R. 1978. Centaurea species and equine nigropallidal encephalomalacia. pp. 327-336. In Keeler, R. F., K. R. Van Kampen & L. F. James (eds.). Effects of poisonous plants on livestock. Academic Press, New York. Groppe, K., R. Sobhian & J. Kashefi. 1990. A field experiment to determine host specificity of LarinuscurtusHochhut(Co\., Curculionidae) and Urophorasirunaseva Hg. (Dipt., Tephritidae), candidates for biological control of Centaurea solstitialis L. (Asteraceae), and Larinus minutus Gyllenhal, a candidate for biological control of C. maculosa Lam. and C. diffusa Lam. J. Appl. Entomol., 110: 300-306. Maddox, D. M. & A. Mayfield. 1985. Yellow starthistle infestations are on the increase. Calif. Agric., 39 (11 and 12): 10-12. Maddox, D. M., A. Mayfield & N. H. Poritz. 1985. Distribution of yellow starthistle ( Centaurea solstitialis ) and Russian knapweed ( Centaurea repens ). Weed Sci., 33: 315-327. Roche, C. T. & B. F. Roche, Jr. 1988. Distribution and amount of four knapweed ( Centaurea L.) species in eastern Washington. Northw. Sci., 62: 242-253. Sobhian, R. 1993. Life history and host specificity of Urophora sirunaseva (Hering) (Dipt., Tephrit¬ idae), an agent for biological control of yellow starthistle, with remarks on the host plant. J. Appl. Entomol., 116: 381-390. Turner, C. E. (in press). Host specificity and oviposition of Urophora sirunaseva (Hering), a natural enemy of yellow starthistle. Proc. Entomol. Soc. Wash. Turner, C. E., J. B. Johnson & J. P. McCaffrey, (in press). Yellow starthistle, Centaurea solstitialis L. (Asteraceae). In Nechols, J. R., L. A. Andres, J. W. Beardsley, R. D. Goeden & C. G. Jackson (eds.). Biological control in the U.S. Western Region: accomplishments and benefits of Regional Project W-84 (1964-1989). Univ. of California, DANR, Berkeley. White, I. M. & S. L. Clement. 1987. Systematic notes on Urophora (Diptera, Tephritidae) species associated with Centaurea solstitialis (Asteraceae, Cardueae) and other Palaearctic weeds ad- ventive in North America. Proc. Entomol. Soc. Wash., 89: 571-580. White, I. M. & V. A. Korneyev. 1989. A revision of the western Palaearctic species of Urophora Robineau-Desvoidy (Diptera: Tephritidae). Syst. Entomol., 14: 327-374. PAN-PACIFIC ENTOMOLOGIST 70(3): 212-221, (1994) NEUROPTEROIDEA FROM MOUNT ST. HELENS AND MOUNT RAINIER: DISPERSAL AND IMMIGRATION IN VOLCANIC LANDSCAPES Patrick M. Sugg, 1 Lita Greve, 2 and John S. Edwards 1 Zoology Department, University of Washington, Seattle, Washington 98195; 2 Museum of Zoology, University of Bergen, Museplass 3, N-5007, Bergen, Norway Abstract.— Neuroptera (= Planipennia) and Raphidioptera were collected from barren, unveg¬ etated habitats at Mount St. Helens following the 1980 eruption and from summer snowfields in the alpine zone on Mount Rainier. A total of 291 specimens were taken in pitfall or flight traps at Mount St. Helens or hand collected from snowfields on Mount Rainier. The data represent individuals engaged in long distance dispersal flight as opposed to local, within habitat movement. Phenological patterns of dispersal are detailed for species and evidence is presented for female sex bias of dispersers f or several species. Taxa in four families are represented: 24 specimens of Coniopterygidae ( Coniopteryx sp., Conwentzia californica Meinander, Semidalis sp.); 219 Hemerobiidae ( Hemerobius bistrigatus Currie, H. humulinus Linnaeus, H. kokeeanus Currie, H. neadelphus Gurney, H. pacificus Banks, H. simulans Walker, H. stigma Stephens, Micromus borealis Klimaszewski & Kevan, M. vari- olosus Hagen, Wesmaelius involutus (Carpenter), W. longifrons (Walker), W. nervosus (Fabricius), W. pretiosus (Banks)); 44 Chrysopidae ( Chrysopa coloradensis Banks, C. nigricornis Burmeister, C. oculata Say, Chrysoperla carnea Stephens, Eremochrysa punctinervis (McLachlan), Meleoma dolicharthra (Navas), M. emuncta (Fitch), Nothochrysa californica Banks); 4 specimens Raphi- diidae (Agulla adnixa (Hagen)). Key Words. — Insecta, Neuroptera, Raphidioptera, Mount St. Helens, Mount Rainier, dispersal, colonization, phenology The eruption of Mount St. Helens in May 1980 devastated at least 600 km 2 . In an area immediately north of the volcano, now called the Pumice Plain, the biota was completely removed. That area, comprising more than 50 km 2 of bare mineral surface, provided sites on which the immigration of arthropods could be monitored in the complete absence of local populations. We report here the diversity and aspects of the phenology of Neuropteroidea, mainly Neuroptera (= Planipennia) but including some Raphidioptera, collected on or in the vicinity of Mount St. Helens and an adjacent Cascade volcano, Mount Rainier. The collections from Mount St. Helens were part of surveys documenting patterns of survival and recovery of arthropod populations in the area of the volcano following the 1980 eruption. We focus on material taken in the first several years following the eruption from sampling sites in the Pumice Plain which lay at least 3 km from potential source habitats and at least 10 km from relatively undisturbed source areas. This material, therefore, reflects long distance dispersal rather than local movement of individuals. A few records are from sites where some residual vegetation survived. We include data from collections made from alpine snowfields on Mount Rainier, which also indicate long distance dispersal. The Neuroptera as a whole were a minor component of the samples taken in arrays of pitfall and flight traps at Mount St. Helens, and they were not among the first colonists (predatory and scavenging beetles, primarily carabids, and spi- 1994 SUGG ET AL.: NEUROPTERA FROM MOUNT ST. HELENS 213 ders). Neuroptera nonetheless play a significant role as predators, particularly of homopterans like aphids, and the data presented here imply a steady source of immigrants to incipient populations as the devastated area becomes vegetated. Little has been published on the Neuropteroidea of the Pacific Northwest. The only faunal survey listing species is from long term ecological work on the H. J. Andrews Experimental Forest in the Cascade Mountains of Oregon (Parsons et al. 1991). We add here a number of taxa to the list for Cascade Neuroptera, including a number of new records for the area. Our list is also unique in that the captures represent individuals all engaged in a biologically critical activity, namely dispersal. Study Area The most severe impact of the eruption occurred north of Mount St. Helens, between the volcano and Spirit Lake, an area now called the Pumice Plain (Figs. 1 and 2a), and some distance down the Toutle River to the west. In this area virtually all surfaces were buried by landslide deposits, covered by pyroclastic surges, with emplacement temperatures of several hundred degrees centigrade, or scoured clean (Christiansen & Peterson 1981). Between 1981 and 1983, three study sites were established in the Pumice Plain with 6 more added by 1985 (Figs. 1 and 2A). The post-eruption surfaces presented a mosaic of pyroclastic flow deposits (ranging from pumice boulders to fine volcanic ash), rock outcrops and landslide debris. Immediately following the eruption there was no emergent vege¬ tation in the area but during this study, a number of plant species colonized the area. However, plants became only locally abundant in a small number of isolated patches (Wood & del Moral 1988). Sites were also established on the southern slopes of the volcano where survival of vegetation was extensive. Site elevations ranged from 1000-1200 m in the Pumice Plain to 1300-1500 m for sites on the southern slopes of Mount St. Helens. Southwest winds predominate throughout the annual April-October sampling period. Brief sequences of days with easterly winds, alternating with westerlies, become more frequent and prolonged in September and October. Westerly surface winds reach the sampling sites via the agricultural Puget lowland, forested Toutle River valleys, and volcano-impacted mudflow and blowdown areas. Summer air temperatures rarely exceed 25° C. Maximum surface temperatures at the sampling sites are generally in the range 40-50° C. Summer rainfall can vary greatly; most precipitation occurs October through April and summers are generally dry. In 1983 rainfall was abundant while exceptional drought conditions prevailed in 1984. Mount Rainier is located approximately 80 km to the north of Mount St. Helens (Figs. 1 and 2B). Samples were taken from snowfields ranging in elevation from about 2200 m to the volcano summit (4392 m). Materials and Methods Sampling Method. — At Mount St. Helens, arthropods were sampled using pitfall traps and flight traps made of 30 cm x 60 cm sheets of plexiglass suspended over a bucket containing a 50% solution of ethylene glycol based anti-freeze. Some hand collecting was also done. Pitfall traps were plastic cups (Lilylite 9 oz. tumblers) which set snugly in 7.6 214 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) Figure 1. Map of the devastated area of Mount St. Helens showing location of sampling sites in the Pumice Plain and on the volcano. Inset shows location of Mount St. Helens and Mount Rainier in Washington State. cm diameter plastic (PVC) pipe sections set in the ground. The PVC sleeve allowed swift change of cups without disturbing the surrounding ground surface. Pitfalls were partially filled with a 2:1 solution of antifreeze and water which served as a killing agent and preservative, and were protected from rain and disturbance by a square piece of plywood supported approximately 2 cm over the trap. From 5 to 26 pitfalls were set at each site. Pitfalls were generally collected at two week intervals. Sampling periods at sites ranged from 5 weeks to 30 weeks of continuous sampling. Some specimens were collected from overwinter samples where pitfalls were left in place and collected the following spring. Since snowpack tended to press the pitfall cover down, effectively closing the pitfall, these samples probably represent individuals caught in the fall, after mid-October but before significant snowfall. At Mount Rainier collections were made from 2 x 100 m transects on snowfield 1994 SUGG ET AL.: NEUROPTERA FROM MOUNT ST. HELENS 215 £Sm % 60 tmi ■y, .V? : 7 X /?r.~‘•'Wit ‘vx-N.d-. "yfcjf'Jr ‘ tx&rM llllfil Figure 2. A.) View of the Pumice Plain of Mount St. Helens in 1984. B.) View of snowfield (elevation 2500 m) on Mount Rainier. 216 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) sites at elevations ranging from 2200 to over 4000 m. Hand collections were also made at various locations including the summit crater. Collections were made from mid-May to mid-October with most in early to mid-July at elevations between 2400 and 3000 m. Specimens were stored in alcohol until identified to species by one of us (LG). Confirmation on identification of several taxa was made by several others (noted in Acknowledgment). Voucher specimens are stored at the Thomas Burke Me¬ morial Museum, University of Washington, Seattle, Washington. A phenogram was assembled from original collection data, mainly from pitfall traps, taken over several years, each year with a different set of sampling dates. Sampling periods varied but were generally from 10-14 days. Collection dates for samples were standardized by dividing the season from May to October into 10 day intervals, assigning each sample to the 10 day interval in which the midpoint of the sampling period fell. Results Diversity. —A total of 250 specimens from 23 species were collected in traps at Mount St. Helens from 1981 to 1986. Another 41 specimens from 10 species were collected from snowfields on Mount Rainier from 197 5 to 19 76. This presents a total of 291 specimens representing 25 species from 12 genera in four families (Table 1). Phenology.— Consistent with previous findings of adult activity for most Neu- roptera, adults of most taxa were active from spring to fall (Fig. 3). Peak activity, as indicated by captures, varied among taxa. The coniopterygid Conwentzia cal¬ if or nica Meinander was present from spring through fall but the majority of captures occurred from mid-summer to fall (Fig. 4A). For most hemerobiids the capture rate increased through the season with a peak in late summer-early fall. An exception was Wesmaelius involutus (Carpenter). It was abundant in early summer but absent in our fall samples (Fig. 4D). Several species of Hemerobius were active from spring to fall with a general increase in numbers of dispersers through the season (Fig. 4E-I). Taken as a whole, there were 3 peaks in the number of Hemerobius dispersers through the season, with an increase in number for each successive peak (Fig. 41). The chrysopids as a group are similar to the Hemerobius spp. in having the greatest numbers of dispersers in late summer-early fall (Figs. 3 and 4). Chrysoperla carnea Stephens, the only chrysopid caught in any number, had a major dispersal period from September-October. Sex Ratios.—A Chi-square goodness of fit test (Zar 1974) on the 6 taxa with 10 or more total individuals in which males and females were identifiable revealed 3 cases where females were significantly more common than males, the coniop¬ terygid Conwentzia californica and the hemerobiids Micromus variolosus (Hagen) and Hemerobius stigma Stephens (Table 1). Among species of Hemerobius, if H. stigma is subtracted from the rest, there is still a female bias (53 males: 63 females), but it is not significant. There is bias in species of Wesmaelius taken as a whole (total 8 males: 15 females), but it is not significant. In contrast, the chrysopids (22 males: 20 females) show no indication of a shift from a 1:1 sex ratio. All specimens of the snakefly Agulla adnixa (Hagen) were female but the small number found (4) precludes analyses. 1994 SUGG ET AL.: NEUROPTERA FROM MOUNT ST. HELENS 217 Table 1. List of species and numbers of individuals from families of Neuropteroidea from Mount St. Helens (MSH) and Mount Rainier (MR). Numbers of males and females shown if known (males: females) and individuals with abdomens missing shown in parentheses. Significant deviations (P < 0.05) from 1:1 in sex ratio noted with asterisks. Total for Neuroptera: 4 families, 12 genera, 24 species (110 males, 172 females, 9 specimens); Raphidioptera: 1 family, 1 genus, 1 species. Numbers (males: females) Family Species MSH MR Total Coniopterygidae Coniopteryx sp. -:1 — -:1 Conwentzia califomica Meinander 3:18 — 3:18* Semidalis sp. -:1 — -:1 Hemerobiidae Micromus borealis Klimaszewski & Kevan 1:1 — 1:1 M. variolosus Hagen 4:14 -:2 4:16* Wesmaelius involutus (Carpenter) 6:8 — 6:8 W. longifrons (Walker) 2:5 — 2:5 W. nervosus (Fabricius) -:1 — -:1 W. pretiosus (Banks) -:1 -:1 -:2 Wesmaelius sp. -:1 0) — -:1 (1) Hemerobius bistrigatus Currie 6:3 1:- 7:3 H. humulinus Linnaeus 1:- — 1:- H. kokaneeanus Currie 5:2 _ 5:2 H. neadelphus Gurney 26:13 a (2) 3:-x 29:13 a (2) H. pacificus Banks 9:-x l:-x 10:x H. simulans Walker 1:- — 1:- H. stigma Stephens 14:26 4:8 18:34* Hemerobius sp., females 15 37 8 45 Chrysopidae Meleoma dolicharthra (Navas) 2:1 1:2 3:2 M. emuncta (Fitch) -:1 — -:1 Chrvsona coloradensis Banks 1:- _ 1:- C. nigricornis Burmeister 1:1 — 1:1 C. oculata Say -:1 — -:1 Chrysoperla carnea Stephens 14:13 (1) 1:1 15:14 (1) Eremochrysa punctinervis (McLachlan) -:1 — -:1 Nothochrysa californica Banks — 2:1 (1) 2:1 (1) Raphidiidae Agulla adnixa (Hagen) — (4) 4 a 13 females are tentatively placed here. Females cannot be determined with certainty. Criterion for assignment of females here was presence of males and absence of male H. pacificus in the same sample. b Most females are probably H. neadelphus or H. pacificus. Discussion Diversity and Dispersal. — Previous sampling of Neuropteroidea in the Cascade Mountains of Oregon recorded 17 species of Neuroptera and 5 species of Ra¬ phidioptera (Parsons et al. 1991). Our sampling adds 11 taxa to the list of Cascade Neuroptera, including several new records for the region or for Washington State. More importantly, the majority of these records indicate the prevalence of long distance dispersal. Some records (20) come from sites on the southern slopes of Mount St. Helens (Fig. 1, PC and BCa) where there was some local survival of vegetation and the possibility that these records represent local movement cannot be discounted. We include them because in nearly every case the time of capture coincides with capture of individuals in the non-vegetated habitats of the Pumice Plain of Mount St. Helens or the snowfields of Mount Rainier. For M. variolosus they add significantly to the total number of records (7 of 20 for that species). 218 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) n May June July Aug Sept Oct Overwinter Conloptervgldae Coniopteryx sp. Conwentzia calif omica Meinander Semldalis sp. Hemeroblldae Micromus borealis KUmaszewski flc McE. Kevan M. ixirlolosus (Hagen) Wesmaelius Involutus (Carpenter) W. longtfrons (Walker) W. nervosus (Fabriclus) W. pretiosus (Banks) Wesmaelius sp. Hemerobius bistrigatus Currie H. humulinus Linnaeus H. kokaneeanus Currie H. neadelphus Gurney H. paciflcus Banks H. simultms Walker H. stigma Stephens Hemerobius sp.. females Chrvsopldae Meleoma dolicharthru (Navas) M. emuncta (Fitch) Chrysopa coloradensis Banks C. nigricomis Burmelster C. occulata Say Chrysopcda cornea Stephens Eremochrysa punctineruis (McLacl Nothochrysa calif omica Banks RhaDhldlldae Agultn adnixn (Hagen) 4 * * * Figure 3. Phenological pattern showing periods in which neuropteran species were caught by traps at Mount St. Helens (solid bars) or hand collected at Mount Rainier (asterisks). Overwinter samples represent individuals caught in traps from late October on, and not collected until the following spring. Long distance dispersal is not effected solely by the active flight of individuals; wind currents play a major role in the distribution of dispersers (Johnson 1969). Our data on Neuropteroidea do not allow for a distinction between active flight and passive carriage on the wind, but the high incidence of ballooning spiders among the arrivals at the sampling sites (Crawford & Edwards 1986) suggests that prevailing winds from the southwest may also play a part in the distribution of Neuroptera. Female Dispersal Bias.—We suggest that the difference in numbers for those species with significant female bias has biological significance and reflects a ten¬ dency for females rather than males to undergo long distance dispersal. Another possibility is that the significant female bias is a reflection of the primary sex ratio for these species, although divergence from a 1:1 sex ratio is rare and should occur only under special circumstances (Hamilton 1967). We doubt the other alternate explanation, that the sex ratio is a reflection of behavior, with females more likely to be caught in traps than males. For H. stigma the same bias is seen from collections on snowfields of Mount Rainier (4 males: 8 females) as is found for individuals trapped at Mount St. Helens (14 males: 26 females), and the snow- fields should retain individuals regardless of sex. Number of Individuals Caught 1994 SUGG ET AL.: NEUROPTERA FROM MOUNT ST. HELENS 219 1 0 I Conwenczia calilomica A 5 4 I Hemerobius bistrigatus E Micromus vanolosus c ■ Hemerobms pacificus ImalesI 4 G Figure 4. Phenology of select Neuroptera taxa found at Mount St. Helens. Results show number of pitfall or flight trap captures occurring in 10 day time periods from 1981-1985. The results for Hemerobius neadelphus and H. pacificus are shown for males only because females cannot be reliably identified. Vegetation Association. — Vegetation data are sparse for many neuropteran taxa. In many cases the published records for a species are mainly from crop plants but the ecological breadth of most species is undoubtedly wider. For example, Meinander (1972) recorded C. californica from citrus trees and cherries but the Mount St. Helens records indicate association with a broader spectrum of vege¬ tation. We do not know from where our samples originated but the vegetation of habitats near our sampling sites is dominated by conifers with stream drainages 220 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) including hardwoods such as alder, willows and maple. Openings along rivers, roads and clearcuts provided habitats dominated by shrubs and herbs. Hemerobiid larvae were common in pitfall traps near herbaceous vegetation (mainly fireweed, Epilobium angustifolium (L.)) of clearcut areas in the devastated area of Mount St. Helens but none were reared to adult so the species are not known. Miscellaneous Notes on Select Taxa.—O ur data provides new information on several of the taxa collected including extensions of the recorded seasonal activity or range. Conwentzia californica. Meinander (1972) records adults from March through August; our data indicates dispersal activity into October. Semidalis sp. Females of most Semidalis species cannot currently be identified to species (Meinander 1972). Ours may represent the first record of the genus from Washington State. Micromus borealis Klimaszewski & Kevan. Ours represent the first records from the lower 48 states. Micromus variolosus Hagen. M. variolosus was the most common neuropteran found at sites on the southern slopes of Mount St. Helens. Our samples had a significant female bias (Table 1). Wesmaelius involutus (Carpenter). W. involutus contrasts with all other taxa for which we have reasonable numbers in our samples in having a peak dispersal activity period in the early summer (Fig. 4D). Wesmaelius nervosus (Fabricius). This is the first record known to us from the western U.S.A. Wesmaelius pretiosus (Banks). This is apparently the first record from Wash¬ ington State. Hemerobius spp. Numbers for individual species suggest periods of dispersal which may correlate with brood cycles. If species totals are combined, there are three clear peaks in dispersal, late spring, mid-summer, and late summer-early fall (Fig. 41). This suggests three brood cycles annually in this area. Hemerobius bistrigatus Currie. Before this study, adults were known from March to September (Kevan & Klimaszewski 1987). Most of our records are from Sep¬ tember and October, but a few individuals were caught in early June or July. Hemerobius neadelphus Gurney. Mitchell (1962) considered three generations per year to be possible even at high elevations. The data presented here (Fig. 4F) also suggest three generations. Acknowledgment We thank P. A. Adams, U. Aspock, J. Klimaszewski, and M. Meinander for their assistance in identifying our samples. We also acknowledge the late D. K. McE. Kevan for assistance and advice. C. A. Tauber gave helpful comments and suggestions on an early draft. P. C. Banko, C. B. Becker, R. I. Gara, D. Mann and M. Peterson assisted with collecting and processing samples. U.S. Forest Service personnel from Randle and St. Helens Districts, Gifford Pinchot National Forest and those of Mount Rainier National Park were helpful in the course of the field work. This work was supported by NSF Grants DEB 80-21460, DEB 81-0742 and BSR 84-07213; LG was provided support by University of Bergen. 1994 SUGG ET AL.: NEUROPTERA FROM MOUNT ST. HELENS 221 Literature Cited Christiansen, R. L. & D. W. Peterson. 1981. Chronology of the 1980 eruptive activity, pp. 17-30. InThe 1980 eruption of Mount St. Helens, Washington. Lipman, P. W. & D. R. Mullineaux (eds.). Geological Survey Professional Paper 1250. U.S. Government Printing Office, Wash¬ ington, D.C., USA. Crawford, R. L. & J. S. Edwards. 1986. Ballooning spiders as a component of arthropod fallout on snowfields of Mount Rainier, Washington, U.S.A. Arctic and Alpine Res., 18: 429-437. Hamilton, W. D. 1967. Extraordinary sex ratios. Science, 156: 477-488. Johnson, C. J. 1969. Migration and dispersal of insects by flight. Methuen, London. Kevan, D. K. McE. & J. Klimaszewski. 1987. The Hemerobiidae of Canada and Alaska. Genus Hemerobius L. G. Ital. Entomol., 3: 305-369. Meinander, M. 1972. A revision of the family Coniopterygidae (Planipennia). Acta Zool. Fenn., 136: 1-357. Mitchell, R. G. 1962. Balsalm woolly aphid predators native to Oregon and Washington. Oregon Agric. Exp. Stat. Tech. Bull., 62: 1-63. Parsons, G. L., G. Cassis, A. R. Moldenke, J. D. Lattin, N. H. Anderson, J. C. Miller, P. Hammond, & T. D. Schowalter. 1991. Invertebrates of the H. J. Andrews Experimental Forest, Western Cascade Range, Oregon. V: An annotated list of insects and other arthropods. Gen. Tech. Rep. PNW-GTR-290. U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, Portland, Oregon. Wood, D. M. & R. del Moral. 1988. Colonizing plants on the Pumice Plains, Mount St. Helens, Washington. Amer. J. Bot., 75: 1228-1237. Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall Inc., Englewood Cliff's, New Jersey. PAN-PACIFIC ENTOMOLOGIST 70(3): 222-229, (1994) LIVESTOCK DUNG AS A FOOD RESOURCE AND THERMAL REFUGE FOR RANGELAND GRASSHOPPERS (ORTHOPTERA: ACRIDIDAE) Kevin M. O’Neill Entomology Research Laboratory, Montana State University, Bozeman, Montana 59717 Abstract.— Fourteen species of grasshoppers from three subfamilies of Acrididae were observed feeding on dry cattle and horse dung at two rangeland sites in southwestern Montana. While feeding within dung cavities during the middle of the day, they attained equilibrium body temperatures well below the critical thermal maxima typically observed for grasshoppers. The implications of these observations for studies of grasshopper diet and nutrient cycling on range- land are discussed. Key Words.— Insecta, thermoregulation, manure, decomposition, rangeland, grasshopper Grasshoppers play a role in ecosystems beyond that of primary consumers of living plants. They reduce forage available to other animals by clipping, but not consuming standing vegetation (Hewitt & Onsager 1983). They are also scavengers upon insect cadavers (Lockwood 1988, O’Neill et al. in press) and dead plant matter (personal observation). In anecdotal accounts, their omnivory extends to an amazing variety of materials during outbreaks, including clothing, curtains, upholstery, rake handles, rubber, tree bark, and human flesh (Shotwell 1958, Gangwere 1961). This paper presents observations of feeding by grasshoppers on cattle and horse dung, and discusses the potential implications for grasshopper diet studies and for rangeland nutrient cycling. In addition, to determine whether the grasshoppers could use dung cavities as thermal refuges during hotter times of day, I measured environmental temperatures and grasshopper operative body temperatures outside and inside of dung cavities. Materials and Methods This study was conducted from 15 Jul through 2 Sep 1992, primarily at two sites. One area was 1 km N of Logan, Montana (latitude 45°45'N, longitude 111°35'W), and had native vegetation characterized by the grasses Stipa comat a Trinius & Ruprecht and Bouteloua gracilis (Humboldt, Bonplan, & Kunth) La- gasca y Segura ex. Steudel. The site is lightly grazed by horses, although none were present during observations. The other area (“red bam site”) was 11.5 km SSW of Three Forks, Montana (and 18 km SE of the Logan site). Observations were also made at a third site (“dead cow pasture”) 8 km S of Three Forks. The native vegetation at both Three Forks sites is the same as at the Logan site, but both areas have been plowed and reseeded with crested wheatgrass, Agropyron cristatum (L.) Gaertner and alfalfa, Medicago sativa L. Both sites are grazed by cattle, but cattle were not present when observations were being made. The identity of grasshoppers feeding on dung was determined at undisturbed dung masses and at those where I had broken open the hardened and dried surface crust to expose the darker and somewhat more friable material within. The latter 1994 O’NEILL: GRASSHOPPER THERMAL REFUGES 223 method allowed me to increase the sample size of feeders over a shorter period of time. Feeding grasshoppers were distinguished from those simply resting on or within the dung. To determine grasshopper community composition, two to four hundred 180° arc sweep samples were taken on several days in each area. Samples were returned to the lab and frozen until the frequency distribution of species, developmental stages, and sexes could be recorded. Temperatures of soil and dung surfaces (T s ) were measured to the nearest 0.1° C with copper/constantan thermocouples and a Cole-Parmer thermocouple ther¬ mometer. T s was measured with the tip of the probe shaded from direct solar radiation. Operative body temperatures (T E ) (Tracy 1982) were measured by inserting the tip of a thermocouple (wire diameter = 0.25 mm) posteroventrally into the enter of the thorax of dead Melanoplus sanguinipes (Fabr.), which were then dried before being used in the field. The T E of the grasshoppers was then determined in two locations: on fully insolated soil surfaces outside of the dung cavities and in the shaded confines of cavities in which grasshoppers had been observed perched within the previous 2 min. Comparisons of T s or T E from different locations were made using Wilcoxon signed-ranks tests. Results At the red barn site, 15 species of Acrididae were either collected in sweep samples or observed in the habitat (Table 1). Among the species present in a 200 sweep sample (n = 488 grasshoppers) taken on 29 Jul, the following species predominated: Aulocara elliotti (Thomas) (51.2% of sample), Ageneotettix deorum (Scudder) (17.2%), M. sanguinipes (11.6%), M. infantilis (Scudder) (7.2%), and Phoetaliotes nebrascensis (Thomas) (5.3%). At the Logan site, 17 species were present (Table 1), with the following species in the greatest abundance in a 200 sweep sample (n = 245 grasshoppers) taken on 27 Jul: Psoloessa delicatula (Scud¬ der) (all 1st and 2nd instar nymphs, 55.1%), A. deorum (15.9%), M. infantilis (10.6%), M. sanguinipes (4.9%), and Amphitornus coloradus (Thomas) (3.2%). It is not known when the dung at the two sites was deposited. However, both the cattle and horse dung were gray with a somewhat weathered and bleached appearance on the outside, and dark brown, dry, and friable with little moisture present inside. When I broke open cattle dung at the red bam site, grasshoppers typically began arriving and feeding on the newly exposed inner material within 10 minutes. At undisturbed dung (i.e., that not tread upon by large vertebrates), they usually fed within cavities in the dung that were presumably created by the grasshoppers themselves. Many of the dung piles were hollowed out due to grasshopper feeding and some eventually collapsed, so that only dried fragments of the surface crust remained. Unlike cattle dung, horse dung is deposited in piles of individual egg- shaped pieces. The result of feeding by grasshoppers on these pieces was quite distinctive. Like the cattle dung, the horse dung was often hollowed out and even burrowed through and the surface on which feeding occurred was relatively smooth due to even clipping by the grasshoppers’ mandibles. Dung on which extensive feeding had occurred also had large deposits of grasshopper feces. These were particularly extensive beneath hollowed out cattle dung, where a large mat of grasshopper feces up to one cm deep sometimes accumulated. Evidence of grasshopper feeding on dung was extensive at all three sites. Of 50 224 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) Table 1. Grasshopper species present in the communities of the two research sites and observed feeding on cattle (red bam site at Three Forks) or horse (Logan) manure. Red bam site Logan site Feeding category 0 Species Present Feeding Present Feeding Gomphocerinae Acrolophitus hirtipes (Say) — — + — F Aeropedellus clavatus (Thomas) — — + — G Ageneotettix deorum (Scudder) + + + + G Amphitornus coloradus (Thomas) + + + + G Aulocara elliotti Thomas + + + + G Phlibostroma quadrimaculatum (Thomas) — — + + G Psoloessa delicatula Scudder (nymphs) + — + — G Oedopodinae Arphia pseudonietana (Thomas) + — + + G Camnula pellucida Scudder + + — - G Dissosteira Carolina Saussure — — — — MH Encoptolophus costalis Scudder + + + — G Hadrotettix trifasciatus (Say) — — + — MH Hesperotettix viridus (Scudder) — — + — F Metator pardalinus (Saussure) + + + + G Spharagemon equale (Say) + + + — MG Trachyrachis kiowa Thomas + + + + G Xanthippus corallipes Haldeman (nymphs) — — — — G Melanoplinae Melanoplus bivittatus (Say) + — — — MF Melanoplus infantilis Scudder + + + + G Melanoplus packardii Scudder + — + + MH/F Melanoplus sanguinipes (Fabr.) + + + + MH Phoetaliotes nebrascensis (Thomas) + + — — G Total number of species 17 11 18 10 a Feeding preference, where F = forbivorous, MF = mixed forbivorous, MH = mixed herbivorous, MG = mixed gramnivorous, and G = gramnivorous (classification from Mulkem et al. 1969, with the exception of A. elliotti, C. pellucida, M.pardalinus, and X. corallipes where designation was based on Capinera & Sechrist, 1982). undisturbed dung piles surveyed at the red bam site on 29 Jul, 34 showed evidence of grasshopper feeding and 19 had feeding grasshoppers present. On the same day, 30 of 50 undisturbed dung piles surveyed at the nearby dead cow pasture showed evidence of grasshopper feeding and 26 had feeding grasshoppers present. At the Logan site on 27 Jul, 40 of 50 undisturbed piles of horse dung showed evidence of grasshopper feeding and 2 had feeding grasshoppers present. The 50 piles had an average of 3.2 feeding sites (SE = 0.4). Extensive feeding was also observed on 24 individual pieces of horse dung, broken open and placed along a transect on 27 Jul. When I returned to the site on 29 Jul, 20 had been fed on (presumably by grasshoppers), with three having at least 10% missing. By 2 Sep, no others had been fed upon, but I estimated that 11 had at least 25% of the original mass missing and three had at least 50% missing. Fourteen of the 22 species present at the two sites were observed feeding on dung (Table 1). At the red bam site, the species most commonly observed feeding 1994 O’NEILL: GRASSHOPPER THERMAL REFUGES 225 on cattle dung in 6 days of observation were A. elliotti (48.9% of 288 observations), A. deorum (24.3%), M. sanguinipes (14.9%; included adults and 4th and 5th instar nymphs), Spharagemon equate (Say) (4.5%), and P. nebrascensis (2.8%). At Logan, those most commonly observed feeding on horse dung in 4 days of observation were A. deorum (32.6% of 43 observations), A. elliotti (18.6%), M. infantilis (14.0%), and Phlibostroma quadrimaculatum (Thomas) (11.6%). At this site, I also ob¬ served two female A. elliotti and one female Metator pardalinus (Saussure) feeding on dung while in copula. Adults of both sexes of all of these species were observed feeding. The probability that a species was observed feeding on dung was related to its abundance in the community. The number of individuals of a species taken in a sweep sample at the red barn site on 29 Jul was significantly correlated with the number of each species observed feeding on dung at this site on 28 and 29 Jul (r = 0.97, P < 0.0001, n = 14). Non-feeding grasshoppers were also observed sitting within shaded cavities, atop dung, and in shade next to dung, particularly during hotter periods of sunny days. At Logan on 6 Aug, the surface temperatures of the dung or soil within dung cavities during the afternoon were 19.7° C lower on average than the bare soil surface temperature outside of the cavity (Fig. 1; Wilcoxon test, P < 0.001). At the red bam site on 28 Jul, the temperatures within dung cavities were 16.6° C lower on average than the soil surface temperatures outside of the cavity (Wilcoxon test, P < 0.001). Similarly, at the same site on 7 Aug, the temperatures within dung cavities were 18.9° C lower on average than the soil surface temperatures outside of the cavity (Wilcoxon test, P < 0.001). Thus, while T s outside of the dung cavities ranged from 50 to 62° C in the 60 observations, inside of the cavities they were >40° C in only 10% of the reading and >50° C in < 2%. The only two cavities in which T s was >45° C were the only two in which grasshoppers were not present before the reading was taken. The lower temperatures and solar radiation loads within the dung cavities correlated with substantially lower operative body temperatures (Fig. 1). The mean operative body temperature of grasshopper models (= dried grasshoppers) placed within shaded dung cavities was 17.5° C lower on average than that of models placed in a standard posture on fully insolated soil surfaces nearby (Wilcoxon test, P < 0.001). The equilibrium T E of the 20 grasshoppers inside dung cavities varied from 33.5 to 40.0° C, while those in full sun ranged from 47.5 to 63.6° C. Discussion There have been a number of surveys of dung insect communities, but most have been confined to the early successional stages of decomposition in the month following dung deposition (Duffield 1937; Mohr 1943; Sanders & Dobson 1966; Poorbaugh et al. 1968; Valiela 1969, 1974;Blume 1970, 1972; Wingo et al. 1974; Merritt & Anderson 1977; Schoenly 1983; Hanski & Cambefort 1991). The major coleopteran and dipteran scavengers of dung exploit it soon after it is deposited, when it still has a high moisture content (e.g., Valiela 1974, Wingo et al. 1974, Schoenly 1983). Like termites (Johnson & Whitford 1975) and tenebrionid beetles (Matthews 1976), the grasshoppers that I observed fed on older, drier material. The condition of the droppings corresponded to that described by Mohr (1943) for the period after the major coleopteran and dipteran dung feeders have com¬ pleted development. 226 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) LOGAN RED BARN Figure 1. Mean (± 2 SE) soil surface temperatures (T s ) and operative body temperatures (T E ) within dung cavities (DC) and on bare (fully insolated) soil (BS) for different dates and sites (n = 20 for all means). I know of no previous extensive observations on the association of grasshoppers with livestock dung. Lavigne & Pfadt (1964) observed consumption of dried dung by nine species of grasshoppers in Wyoming and Colorado, but provided no other details. Their list included 3 species observed as dung feeders at my sites: A. coloradus, A. deorum, and A. elliotti. Gangwere (1961) also notes that dung has been reported as grasshopper food, but does not specify the species or the con¬ ditions under which scavenging occurs. The propensity for grasshoppers to con¬ sume manure was also recognized earlier in this century. A poisoned bait, known as Criddle Mixture and consisting of horse manure laced with insecticide (e.g., arsenic), was used with some effectiveness to control grasshopper outbreaks in Manitoba (Criddle 1920). Rentz (1970) reports observations of a species of Ma- crobaenetes (Orthoptera: Gryllacrididae) feeding on kangaroo-rat and lizard feces. 1994 O’NEILL: GRASSHOPPER THERMAL REFUGES 227 Dung feeding was observed in all 3 subfamilies of Acrididae present at the research sites (Table 1). With the exception of Melanoplus packardii Scudder and M. sanguinipes, all of the species of grasshoppers that were observed feeding on dung are classified as either gramnivorous or mixed gramnivorous by Mulkem et al. (1969). Five of the species that I did not observe to feed on dung are not classified as gramnivorous. However, they were either rare at my sites or they occupied different microhabitats. For example, Dissosteira Carolina (L.) was abun¬ dant in weedy roadside vegetation, but uncommon where my observations took place. Furthermore, three of these species (i.e., Hesperotettix viridis (Scudder), Hadrotettix trifasciatus (Say), and Xanthippus corralipes Haldeman) were ob¬ served feeding on dried dung by Lavigne & Pfadt (1964). Therefore, the apparent association of preference for grasses with feeding on dung may be an artifact of the low population density of non-gramnivores at my sites. The correlation be¬ tween abundance of a species and its frequency in the sample of feeders at the red bam site indicates that more exhaustive sampling at the two sites may have lengthened the list of observed feeders. Studies of possible spatial and temporal variation in the propensity for grass¬ hoppers to feed on dung will be needed to determine how common dung feeding is in grasshopper communities. Interestingly, because of frequent rain during the summer of 1992, the mid- to late-summer vegetation was relatively lush compared with other years. Thus, the grasshoppers fed on dung even though relatively lush vegetation was abundant. Grasshoppers reared on material with a high moisture content are known to prefer dry food when given the choice (Chapman 1990), so desiccated dung may be an attractive alternative food during times of high water availability. An opposite trend may occur in dry years. In 1988, at a nearby site, grasshoppers were observed feeding on fresh cattle dung at an extremely dry and heavily grazed site (J. Holmes, personal communication). The existence of grasshopper feeding on livestock dung has several possible implications, the significance of which depends on how widespread it is. First, field studies of diet mixing may have to take into account not only the host plants, but other sources of nutrition such as livestock feces and insect cadavers (Lock- wood 1988, O’Neill et al. 1993). Many of the species that I have observed feeding on grasshopper cadavers at this site (O’Neill et al. 1993 and unpublished data) were also among the most common dung feeders: A. elliotti, A. deorum, M. infantilis, M. sanguinipes, and S. equale. Second, grasshopper diet studies based on crop content analysis (e.g., Mulkem et al. 1969, Gangwere 1961) may not always reflect host plant choice by grass¬ hoppers if some of the material in the gut was derived from dung feeding. This problem is reduced in studies at sites at which domestic grazers were absent (e.g., Joem 1985). However, it is possible that grasshoppers also feed on the excrement of non-domestic herbivores (e.g., ground squirrels, antelope). For example, during the same field season, I observed grasshoppers feeding on grasshopper feces (O’Neill, unpublished data). Third, because desiccation and hardening inhibits microbial decomposition of dung (Merritt & Anderson 1977) grasshoppers may play a valuable role in degradation of older dung piles. By converting large masses of livestock dung into smaller grasshopper feces, both physically- and microbially- mediated rates of decomposition and nutrient cycling could be enhanced. The observations also suggest that grasshoppers used the cavities they created 228 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) in dung while feeding to maintain non-lethal or even optimum body temperatures during the hottest periods of the day. Within the shade of cavities, they experienced lower environmental temperatures and lower solar radiation loads. As a result, the operative body temperatures recorded for grasshoppers in dung cavities were well below 1) temperatures they would experience on fully insolated soil surfaces and 2) the known critical thermal maxima for acridids (Chappell & Whitman 1990). The mean T E (36.7° C, SE = 0.51) measured for the grasshoppers in dung cavities was in the range of pref erred temperatures observed for many grasshoppers (Chappell & Whitman 1990). Although dung cavities may be useful as thermal refuges, they are not critical in areas where sufficient standing vegetation is avail¬ able (as at my sites). Grasshoppers typically use perches on vegetation as a means of lowering mid-day body temperatures via convective heat loss (Chappell & Whitman 1990). However, the thermal properties of the dung cavities did allow grasshoppers to continue feeding on dung even during the hottest periods of the day and may be important in areas were grazing livestock have removed most of the tall vegetation. Furthermore, visually hunting predators such as sparrows (Joem 1988), may also be less likely to find grasshoppers thermoregulating in dung cavities than those perched on plant stems. Acknowledgment I thank Ruth O’Neill for assistance with the research, Maria Marta Cigliano, Anthony Joem, William Kemp, Jerome Onsager, and Bret Olson for reviewing the manuscript, Jerome Onsager for information on the use of horse dung in poisoned baits, and Jeff Holmes for sharing his unpublished observations. This work was supported by the Montana Agricultural Experiment Station. Contri¬ bution #J-2822 from the Montana Agricultural Experiment Station. Literature Cited Blume, R. R. 1970. Insects associated with bovine droppings in Kerr and Bexar counties, Texas. J. Econ. Entomol., 63: 1023-1024. Blume, R. R. 1972. Additional insects associated with bovine droppings in Kerr and Bexar counties, Texas. J. Econ. Entomol., 65: 621. Capinera, J. L. & T. S. Sechrist. 1982. Grasshoppers (Acrididae) of Colorado: identification, biology, and management. Colo. St. Univ. Univ. Expt. Sta. Bull., 584S. Chapman, R. F. 1990. Food selection, pp. 39-72. In Chapman, R. F. & A. Joem (eds.). Biology of grasshoppers. John Wiley and Sons, New York. Chappell, M. A. & D. W. Whitman. 1990. Grasshopper thermoregulation, pp. 143-172. In Chapman, R. F. & A. Joem (eds.). Biology of grasshoppers. John Wiley and Sons, New York. Criddle, N. 1920. Locust control in the prairie provinces. Dominion of Canada, Department of Agriculture, Entomological Branch, Circular 13. Duffield, J. E. 1937. Notes on some animal communities ofNorwegian Lapland: an account of the dung and carrion communities and of those insects found in human dwellings. J. Anim. Ecol., 6: 160-168. Gangwere, S. K. 1961. A monograph on food selection in Orthoptera. Trans. Amer. Entomol. Soc., 87: 67-230. Hanski, I. & Y. Cambefort. 1991. Dung beetle ecology. Princeton University Press, Princeton, New Jersey. Hewitt, G. B. & J. A. Onsager. 1983. Control of grasshoppers on rangeland in the United States: a perspective. J. Range Manag., 36: 202-207. Joem, A. 1985. Grasshopper dietary (Orthoptera: Acrididae) from a Nebraska Sand Hills prairie. Trans. Nebr. Acad. Sci., 13: 21-32. Joem, A. 1988. Foraging behavior and switching by the grasshopper sparrow Ammondramus sa- 1994 O’NEILL: GRASSHOPPER THERMAL REFUGES 229 vannarum searching for multiple prey in a heterogeneous environment. Amer. Midi. Nat., 119: 225-234. Johnson, K. A. & W. G. Whitford. 1975. Foraging ecology and relative importance of subterranean termites in chihuahuan desert ecosystems. Environ. Entomol., 4: 66-70. Lavigne, R. J. & R. E. Pfadt. 1964. The role of rangeland grasshoppers as scavengers. J. Kans. Entomol. Soc., 37: 1-4. Lockwood, J. A. 1988. Cannibalism in rangeland grasshoppers (Orthoptera: Acrididae): attraction to cadavers. J. Kans. Entomol. Soc., 61: 379-387. Matthews, E. G. 1976. Insect ecology. University of Queensland Press, St. Lucia. Merritt, R. W. & J. R. Anderson. 1977. The effects of different pasture and rangeland ecosystems on the annual dynamics of insects in cattle droppings. Hilgardia, 45: 31-71. Mohr, C. O. 1943. Cattle droppings as ecological units. Ecol. Monogr., 13: 273-298. Mulkem, G. B., K. P. Pruess, H. Knutson, A. F. Hagen, J. B. Campbell & J. D. Lambley. 1969. Food habits and preferences of grassland grasshoppers of the north central Great Plains. N. Dakota Agric. Expt. Sta. Bull., 481. O’Neill, K. M., S. A. Woods, D. Streett & R. P. O’Neill. 1993. Aggressive interactions and feeding success of scavenging rangeland grasshoppers (Orthoptera: Acrididae). Environ. Entomol., 22: 751-758. Poorbaugh, J. H., J. R. Anderson & J. F. Burger. 1968. The insects inhabitants of undisturbed cattle droppings in northern California. Calif. Vector Views, 15: 17-36. Rentz, D. C. 1970. An observation of the feeding behavior of a sand-treader cricket (Orthoptera: Gryllacrididae; Raphidophorinae). Ent. News, 81: 289-291. Sanders, D. P. & R. C. Dobson. 1966. The insect complex associated with bovine manure in Indiana. Ann. Entomol. Soc. Amer., 59: 955-959. Schoenly, K. 1983. Arthropods associated with bovine and equine dung i n a n ungrazed chihuahuan desert ecosystem. Ann. Entomol. Soc. Amer., 76: 790-796. Shotwell, R. L. 1958. The grasshopper, your sharcropper. Univ. Missouri Agric. Expt. Stn. Bull., 714. Tracy, C. R. 1982. Biophysical modeling in reptilian physiology and ecology. In Gans, C. & F. H. Pough (eds.). Biology of the reptilia, Volume 12, Physiology C, Physiological Ecology. Academic Press, New York. Valiela, I. 1969. The arthropod fauna of bovine dung in central New York and sources on its natural history. J. N.Y. Entomol. Soc., 77: 210-220. Valiela, I. 1974. Composition, food webs and population limitation in dung arthropod communities during invasion and succession. Amer. Midi. Nat., 92: 370-385. Wingo, C. W., G. D. Thomas, G. N. Clark & C. E. Morgan. 1974. Succession and abundance of insects in pasture manure: relationship to face fly survival. Ann. Entomol. Soc. Amer., 76: 386— 390. PAN-PACIFIC ENTOMOLOGIST 70(3): 230-239, (1994) THE SPATIAL DISTRIBUTION OF ENDEMIC AND INTRODUCED FLOWER-BREEDING SPECIES OF DROSOPHILA (DIPTERA: DROSOPHILIDAE) DURING THEIR EARLY HISTORY OF ENCOUNTER ON THE ISLAND OF HAWAII William T. Starmer 1 and Jane M. Bowles 2 department of Biology, Syracuse University, Syracuse, New York 13244; department of Plant Sciences, University of Western Ontario, London, Ontario N6A 5B7 Abstract. —The spatial distribution of two flower-breeding drosophilids (an endemic, Scaptomyza caliginosa Hardy and an exotic, Drosophila floricola Sturtevant) is reported for morning and evening censuses of four sites at one locality on the island of Hawaii. The increase in the relative frequency of the introduced species over the last 10 years appears to be due to the increase in the number of adult D. floricola and not a reduction in number of endemic adult S. caliginosa. The analysis indicates that the two species are sympatric and adults occupy the same individual blossoms of morning glory. Afternoon and evening aggregation behavior of adult S. caliginosa may explain some variation in the joint distribution of the two species. Although there is some evidence for the adults using different blossoms, there is considerable overlap of adults of the two species in the same blossoms and it is concluded that the potential for larval-larval com¬ petition between the two species is high. Key Words. — Insecta, Diptera, Scaptomyza {Exalloscaptomyza) caliginosa, Drosophila ( Phlori- dosa ) floricola, morning glory, flower breeders, Hawaiian drosophilids, spatial distribution, co¬ existence For 12 years we have been monitoring the population densities of two drosoph- ilid species that breed in blossoms of the morning glory, Ipomoea acuminata (Vahl) Roemer & Schultes, on the island of Hawaii in the Hawaiian Archipelago. One species, Scaptomyza {Exalloscaptomyza) caliginosa Hardy, is endemic to Hawaii and uses only morning glory blossoms as a larval and adult feeding sub¬ strate (Hardy 1965, 1966; Ibara 1976). The other species, Drosophila {Phloridosa) floricola Sturtevant, was introduced to Hawaii about 15-20 years ago and also uses flowers, including morning glories, as breeding sites. Montague & Kaneshiro (1982) reported the percentage of adults to be 96% S. caliginosa and 4% D. floricola in morning glory flowers growing in Kipuka Puaulu (Bird Park) Hawaii Volcanoes National Park on Hawaii Island during September and November 1980. Since then we have several estimates of the numbers of both species in the same and neighboring sites. These counts reveal a striking change in the relative numbers of the two species and prompted us to determine the distribution of the two species in the same area. This paper reports counts of both species made in Bird Park and neighboring areas since 1980 and analyzes the spatial distribution of the adults occupying individual flowers during October 1991. This analysis provides a basis for studying the competitive interactions of two species in early stages of sympatry (Montague & Kaneshiro 1982). The species evolved their flower breeding habit independently and have different life-history characteristics. Montague & Kaneshiro (1982) em- 1994 STARMER & BOWLES: FLOWER-BREEDING DROSOPHILA 231 phasized differences in reproductive potential and in the larval stage as important facets of the ultimate fate of the two species in this habitat. The introduced species has a greater reproductive potential (i.e., 13.5 ovarioles per female for D. floricola and 1.05 ovarioles per female for S. caliginosa). The endemic female, however, either deposits a single large egg, that hatches soon after being laid, or a precocious larva in a flower. The introduced female lays several eggs that must mature before hatching. These differences might be important if adults and larvae of the two species share the same resources to an extent. There are several levels of resource sharing that can be considered. These include 1) sharing the same sites where flowers grow, 2) sharing the same individual blossoms of the morning glory, and 3) using the resources available in a shared blossom in the same way. The first two considerations are the concerns of this report. We focus on adult distributions because this could separate the two species spatially and thus preclude larval competition. Material and Methods The spatial distribution of adult S. caliginosa and D. floricola was determined by counting the number of each species in individual morning glory blossoms over a two day period (30-31 Oct 1991). Sites in Kipuka Puaulu (Bird Park) and alongside the road between Bird Park and Kipuka Ki were censused in the morning (8:00-10:00 h = AM) and afternoon (15:00-17:00 h = PM). Adult flies present in 100 blossoms were counted at each site and each blossom was mapped by systematically walking through the area and plotting their Cartesian coordinates. The number of adults in most of the blossoms in a contiguous area were counted. In some cases when flies were inadvertently frightened away, the blossom was skipped. Flies were counted either directly in the blossom or after being aspirated into a glass vial using a magnifying glass (2 x). All flies that were aspirated were released after counting. Some adults were trapped by placing a plastic bag over the blossom and returning it to the laboratory. In these cases the site was being censused for the second and last time. Two sites (A & B) within Bird Park and two sites (A & B) along the road between Bird Park and Kipuka Ki were censused. The area surveyed at each site varied as a function of flower density because the number of blossoms mapped was held constant at 100 blossoms. The approximate area mapped for each site was 227, 177, 60 and 62 m 2 for Bird Park-A, Bird Park-B, Road-A and Road-B respectively. The Bird Park sites were located along the main trail in open fields and were separated by intervening forest with no morning glories present. Site A was north of the entrance to the park and site B was east of the entrance. The road sites were between Bird Park and Kipuka Ki. Although morning glory vines were continuous along the road in this area, the two sites were on opposite sides of the road at a distance of 1.1 km (site A) and 1 km (site B) from the gate at Bird Park. Two sites (Road-A, Bird Park-A) were censused on the first morning (30 Oct 1991) and again on the afternoon of the next day (31 Oct 1991). Two sites (Road- B, Bird Park-B) censused on the first afternoon were censused again on the fol¬ lowing morning. This procedure was followed because disturbance in the morning is more likely to affect the afternoon distribution, while an afternoon census is less likely to affect the census on the following morning when most flies move to and occupy newly opened blossoms. 232 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) Table 1. Percentage of adults of the endemic species Scaptomyza caliginosa relative to the intro¬ duced species Drosophila floricola present in Bird Park and neighboring areas. Date of census Bird Park Road Collector 1976 100.0 Ibara (1976) Aug 1978 100.0 J. R. Montague & W. T. Starmer 3 Sep-Nov 1980 96.3 Montague & Kaneshiro (1982) 3-8 Dec 1987 97.9 W. T. Starmer, D. Droney, & J. Bowles 3 2 Jun 1989 ~50 K. Kaneshiro 3 Jan 1990 ~50 M. Kambysellis 3 Site A Site B Site A Site B 16 Aug 1990 57.4 60.5 7.5 3.3 M. Tlusty 3 27 Aug 1990 27.5 46.3 49.4 D. Droney 3 30-31 Oct 1991 83.9 35.7 87.0 52.7 This study a Personal communications. The joint occurrence of the two species was analyzed with correlations for the number of adults of each species in a flower and by calculating the G-statistic on 2-way tables of presence or absence of the two species. In addition 2-way tables with three levels (absent, 1 adult and > 1 adult per flower) for each species were analyzed for each census. A plot of the autocorrelation coefficients as a function of distance is termed an “I correlogram” when the correlations calculated are product-moment correla¬ tions (Sokal & Oden 1978a, b). I correlograms of the number of adults per flower were calculated for S. caliginosa versus S. caliginosa, D.floricolaversusD. floricola and for S. caliginosa versus D. floricola. The correlations were determined for each census locality (Bird Park-A, Bird Park-B, Road-A, Road-B) at each time of day (AM, PM). A step size of 2 m was chosen and spatial correlations (i.e., product-moment correlations for number of adults in a flower) for all pairs of flowers at distances of 0-2, 2-4, 4-6 . . . 28-30 m were calculated. These corre¬ lations were based on an average sample size of 555 pairs of flowers for any one distance. At this level, correlations between -0.12 and +0.12 would be considered nonsignificant at a = 0.01. Results The numbers of D. floricola were low until 1988 when they increased (Table 1). Our recent records (Table 2) indicate that the number of S. caliginosa per blossom has remained about the same as that observed in 1980 (0.77, 2.56 and 3.12 adults per blossom for covered, sunlit and shrub blossoms respectively, table 1 of Montague & Kaneshiro 1982) and on 16 Aug 1990 (2.77 adults per blossom). Only the Bird Park-B site had lower numbers in the present survey. Although morning glory blossoms are considered to remain open for only one day, some blossoms will stay open for two days at lower temperatures. We recorded the number of 1st and 2nd day (i.e., new and old) blossoms during each census. The average ratio was 3 new: 1 old for all sites. An analysis of variance on the average number of adults in new versus old blossoms for each census (site, time, species) did not reveal a significant effect of flower age on the average number of 1994 STARMER & BOWLES: FLOWER-BREEDING DROSOPHILA 233 Table 2. Number of adults per blossom, correlation between number of adults of the two species and the percentage of Scaptomyza caliginosa for each census. Site S. caliginosa D. floricola Correlation 2 Average adults/ flower Standard deviation Average adults/ flower Standard deviation Percentage Empty blossoms S. caliginosa Included Excluded Bird Park A (AM) 2.19 4.35 0.34 0.77 86.51 0.104 0.044 (PM) 1.93 4.04 0.45 0.94 81.09 -0.101 -0.243* Bird Park B (AM) 0.28 0.51 0.48 0.94 36.84 -0.093 -0.574*** (PM) 0.37 0.65 0.69 1.17 33.65 -0.047 -0.356** Road A (AM) 1.67 2.03 0.47 0.76 78.04 -0.023 -0.109 (PM) 4.06 13.48 0.39 0.98 91.24 -0.023 -0.098 Road B (AM) 1.01 1.26 0.54 0.93 65.16 -0.074 -0.253* (PM) 0.67 0.87 0.97 2.34 40.85 -0.045 -0.256* a The correlations were estimated from the number of adults of each species present in each blossom. The included column includes blossoms that had no flies present, whereas the excluded column does not include empty blossoms. *, **, *** Represent significance at 0.05, 0.01 and 0.001, respectively. adults per blossom (F = 0.91; df = 6, 1; P > 0.10) or significant interactions between age of flower with the other factors (site, time or species). We considered all flowers in each census to be equivalent in the following analyses. The Bird Park sites show different frequencies of D. floricola at each site but morning versus afternoon percentage for each site remained constant. The Road site percentages differed from site to site and for the two sampling periods. Most variance to mean ratios are greater than one, indicating that the flies have aggre¬ gated rather than random or evenly dispersed distributions. This is especially apparent for S. caliginosa at Road site A in the evening where six flowers had greater than 26 flies per flower and accounted for 7 5% of the adults in the 100 flowers sampled (Table 3). The correlations for number of adults of each species in a flower are significantly negative (if empty blossoms are excluded) for 5 of the 8 censuses (Table 2). This indicates a general trend for the adults of the two species to occupy different flowers. If the average number of flies per blossom for each species is used as observations (Table 2) to estimate site to site correlations in the density of each species the estimate is also negative but non-significant (r = -0.58, df = 6, P > 0.10). In six of the eight analyses in Table 4 the two species are independently dis¬ tributed. However, during the morning at both Road sites the distribution of one species is dependent on the distribution of the other. In these cases the observed number of flowers where 1) both species are present and 2) both are absent (i.e., empty) are lower than expected. A similar analysis where three categories (absent, 1 adult and > 1 adult per blossom) for each species were considered revealed the same pattern. That is, the only significant G-statistics were for the two Road sites in the morning and these showed lower than expected numbers for flowers con¬ taining both species and flowers that were empty. The I correlograms for S. caliginosa versus S. caliginosa, D. floricola versus D. floricola and for S. caliginosa versus D. floricola are shown in Figs. 1, 2 and 3 respectively. The correlograms show that there is little spatial structure in the populations of each species and between the two species. Another spatial auto- 234 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) Table 3. Number of adult flies per blossom of Scaptomyza caliginosa (Sc) and Drosophila floricola (Df) at the four sites. Bird Park Road A B A B Adults per blossom AM PM AM PM AM PM AM PM Sc Df Sc Df Sc Df Sc Df Sc Df Sc Df Sc Df Sc Df 0 33 76 47 75 75 71 71 61 27 67 49 79 42 67 53 68 1 29 19 21 13 22 18 22 23 35 21 23 13 34 18 32 14 2 15 2 16 6 3 6 6 9 21 10 13 4 12 11 11 7 3 9 2 2 5 3 1 3 6 2 4 9 3 3 2 4 3 2 1 3 1 1 2 2 1 4 5 3 1 2 1 1 4 1 2 1 1 6 4 2 2 1 7 3 1 2 8 1 1 1 9 1 2 1 10 1 11 12 1 ' 1 14 1 15 1 16 17 1 21 1 22 1 25 1 26 1 27 1 28 1 1 32 1 52 1 77 1 89 1 Total flies counted 219 34 193 45 28 48 37 69 167 47 406 39 101 54 67 97 correlation analysis was conducted with the empty flowers at each site removed from the data set. This analysis resulted in different estimates of the correlations but did not reveal significant spatial structure for either species or the two species considered together. Discussion Variation in the number of adults of each species present at each site can be explained in several ways. The two sites in Bird Park (Table 2) were similar in many respects except that the B-site was contiguous with an area that had been 1994 STARMER & BOWLES: FLOWER-BREEDING DROSOPHILA 235 Table 4. Two-way tables for the distribution of Scaptomyza caliginosa (Sc) and Drosophila floricola (Df) in 100 blossoms for each census. The G statistic has 1 degree of freedom. Morning Evening Sc Sc Df + - G Df + - G Bird Park A + 13 11 2.28 + 11 14 1.08 — 54 22 — 42 33 Bird Park B + 6 23 0.42 + 8 31 2.30 — 19 52 — 21 40 Road A + 19 14 5.73* + 8 13 1.78 — 54 13 — 43 36 Road B + 14 19 4.88* + 12 20 1.72 — 44 23 — 35 33 * Significant at a < 0.05. defoliated with herbicide (applied after the 27 Aug 1990 census). Some of the variance for the number and relative proportions of the adults of the two species at the road sites may be due to the vertical distribution of the flowers at the two sites. The flowers along the side of the road at the A-site were below two meters and accessible, while the vines at the B-site exceeded 2 to 3 meters making the highest flowers in the vicinity inaccessible. Because adults of S. caliginosa tend to accumulate in higher blossoms in the afternoon and evening, it was not possible to count those flowers with large numbers of flies higher in the canopy. Thus the relative increase in D. floricola adults during the afternoon at the B-site (Table 2) is probably due to the movement of S. caliginosa adults into the higher blossoms. This may also be an important consideration in interpreting the counts made beside the road on 16 Aug 1990 (Table 1) since those blossoms were collected in the evening after the movement of S. caliginosa into focal blossoms. Most of the variation in percentage of D. floricola is due to changes in numbers of S. caliginosa. This could be due to movement of S. caliginosa as mentioned for the Road-site B as well as between site differences in the number of S. caliginosa present. This can be seen in the mean to variance ratios across sites (observations from Table 2) for S. caliginosa (1.52:1.56) as compared to (0.54:0.04) for D. floricola. Although the average number of S. caliginosa per blossom has not changed in the Bird Park area over years, the variation from site to site is con¬ siderable. Although the correlograms (Figs. 1, 2 and 3) show little spatial structure, there does appear to be a trend in the S. caliginosa population for positive correlations at the closer distances and negative correlations at the further distances (Fig. 1). This is possibly due to the aggregation behavior of S. caliginosa. This behavior would result in patches of blossoms with higher than average numbers of adults in the flowers close to the focal flowers as S. caliginosa adults aggregate in afternoon and evening and would have the same effect as they move to newly opened blossoms in the morning. That is, many newly opened blossoms close to the focal aggregate blossoms would have higher numbers of adults due to their proximity to that blossom. Fisher (1991) showed that when both species are kept together in laboratory 236 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) Meters - Road A - Road B - Bird Park A - Bird Park B Figure 1. Correlogram for the number of S. caliginosa versus S. caliginosa adults per blossom as a function of distance (meters) between blossoms. The four lines represent the four sites. The upper and lower figures are for morning and evening censuses, respectively. 1994 STARMER & BOWLES: FLOWER-BREEDING DROSOPHILA 237 0.3 AM I -0.4-1-r- 0 10 - 1 - 20 Meters “»- 1 30 40 I - Road A - Road B - Bird Park A - Bird Park B Figure 2. Correlogram for the number of D. Jloricola versus D. jloricola adults per blossom as a function of distance (meters) between blossoms. The four lines represent the four sites. The upper and lower figures are for morning and evening censuses, respectively. 238 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) ■-- Road A - Road B - Bird Park A - Bird Park B Meters Figure 3. Correlogram for the number of S. caliginosa versus D. JJoricola adults per blossom as a function of distance (meters) between blossoms. The four lines represent the four sites. The upper and lower figures are for morning and evening censuses, respectively. 1994 STARMER & BOWLES: FLOWER-BREEDING DROSOPHILA 239 population cages at densities similar to that in the held and provided with fresh morning glory blossoms, adults of both species are reared from the same individual blossoms. This result in conjunction with the spatial structure of the two species we have documented in this report makes it likely that the opportunity for larval competition exists under held conditions. However, other factors such as the use of blossoms of other plant species as a refuge by the introduced species (Montague & Kaneshiro 1982) could inhuence coexistence of the two species (Shorrocks 1990). We are also aware of the possibility that because most of our observations are conhned to one season over several years, we are missing important seasonal variation in the distribution of the two species. It is important to establish the status of these species over time and space during this relatively early stage of encounter in this unique habitat. Acknowledgment We thank Andre Lachance, Ken Kaneshiro, Mike Kambysellis, David Droney and Michael Tlusty for assistance in collecting hower breeding drosophilids from Bird Park. Discussions with Larry Wolf and Gwen Wehbe were helpful in pre¬ paring the manuscript. We also thank Charles Stone and Danielle Stone for their help in facilitating our research in Hawaii Volcanoes National Park. Permission to work in Kipuka Puaulu was given by the National Park Service and the work was funded by NSF grants to WTS. Literature Cited Fisher, G. M. 1991. Interactions between two flower-breeding drosophilids: Exalloscaptomyza cali- ginosa and Drosophila floricola. M.S. Thesis, Syracuse University. Hardy, D. E. 1965. Diptera: Drosophilidae. pp. 604-606. In Insects of Hawaii. Vol. 12. University of Hawaii Press, Honolulu. Hardy, D. E. 1966. Description and notes on Hawaiian Drosophilidae. Univ. of Texas Bull., 6615: 243-314. Ibara, W. 1976. The ecology of endemic Hawaiian Scaptomyza ( Exalloscaptomyza ), (Diptera: Dro¬ sophilidae) in relation to the morning glory, Ipomoea spp. M.S. Thesis, University of Hawaii- Manoa. Montague, J. R. & K. Y. Kaneshiro. 1982. Flower-breeding species of Hawaiian drosophilids in an early stage of sympatry. Pacific Insects, 24: 209-213. Shorrocks, B. 1990. Coexistence in a patchy environment, pp. 91-106. In Shorrocks, B. & I. R. Swingland (eds.). Living in a patchy environment. Oxford University Press, New York. Sokal, R. R. &N. L. Oden. 1978a. Spatial autocorrelation in biology 1. Methodology. Biol. J. Linn. Soc., 10: 199-228. Sokal, R. R. & N. L. Oden. 1978b. Spatial autocorrelation in biology 2. Some biological implications and four applications of evolutionary and ecological interest. Biol. J. Linn. Soc., 10: 229-249. PAN-PACIFIC ENTOMOLOGIST 70(3): 240-252, (1994) EUROPEAN ELM SCALE (HOMOPTERA: ERIOCOCCIDAE) ABUNDANCE AND PARASITISM IN NORTHERN CALIFORNIA Steve H. Dreistadt 1 and Kenneth S. Hagen Division of Biological Control, University of California, Berkeley, California 94720 Abstract. — European elm scale, Gossyparia spuria (Modeer) (= Eriococcus spurius) (Homoptera: Eriococcidae), infested elms ( Ulmus spp.) at all 12 sites that we sampled in northern California during 1987-1989. Scales were more abundant on English elm ( Ulmus procera Salisbury) than Siberian elm ( Ulmus pumila L.). Female scale density peaked at 301 (SD = 76) degree-days above 11° C accumulated from 1 March, mid-April through June depending on location and weather. Scale density and defoliation by elm leaf beetle, Xanthogaleruca (= Pyrrhalta ) luteola (Muller) (Coleoptera: Chrysomelidae) were apparently associated. We reared one to three species of parasitoids from scales at each of five locations. We found no parasitoids at seven other sites. Only one species, Coccophagus insidiator (Dalman) (Hymenoptera: Aphelinidae), has previously been reported as established on European elm scale in California. We recovered Trichomasthus coeruleus Mercet (Hymenoptera: Encyrtidae) at two locations; this species at the time of intro¬ duction was apparently misidentified as Trichomasthus cyanifrons (Dalman). Both Trichomas¬ thus species may have been introduced, but we recovered only T. coeruleus. A Microterys sp. (Encyrtidae) of unknown origin occurred at two locations. Limited parasitoid distribution and high scale populations at some sites provide an excellent opportunity for further biological control efforts against European elm scale. Key Words.— Insecta, Eriococcus spurius, Coccophagus insidiator, Trichomasthus coeruleus, Xanthogaleruca luteola, biological control, Gossyparia spuria European elm scale, Gossyparia spuria (Modeer), is of Palearctic origin and was first discovered in the western U.S. on the Stanford University campus in Palo Alto, California in 1893 (Herbert 1924). Scales mature on bark in the spring, developing into tiny white “cocoons” containing males or purple to dark brown females partially enclosed by a white, waxy fringe. Crawlers emerge from females and settle on leaves or bark where they feed through the summer before moving to overwinter on bark. High populations of this univoltine pest produce copious honeydew and branch dieback. Classical biological control of European elm scale was conducted in California from 1939 through the mid-1950s. One introduced species, Coccophagus insi¬ diator (Dalman), was recovered and reportedly controlled scales at the one site where it was established (Flanders 1952). Finding no published data on European elm scale and parasitoid populations in California since the 1950s, we investigated the current status of the scale and its biological control agents in northern Cali¬ fornia. Materials and Methods Scale Abundance.— We sampled European elm scales at 5 northern California sites during 1987, 8 locations in 1988, and 7 sites during 1989 (Figs. 1 and 2), a 1 Statewide IPM Project, University of California, Davis, California 95616-8620. 1994 DREISTADT & HAGEN: EUROPEAN ELM SCALE 241 total of 12 different locations were used (Fig. 5). We examined 30 cm long branch terminals and recorded the presence or absence of apparently viable female scales about every 2 weeks during the spring and early summer from an average of 3 to 4 elms ( Ulmus spp.) at each site. We sampled 40 terminals per tree (5 each from the inner and outer half of the canopy in each cardinal direction) from the lower one third of the canopy of each tree, except in Cloverdale and Hopland where we collected 24 terminals per tree (3 each inner and outer half in each cardinal direction). We determined the proportion of infested terminals (x; SD) for 1987 by pooling samples within each inner and outer quadrant (8 per tree as above) and by pooling samples by tree during 1988 and 1989. Unless otherwise stated, we report scale densities for each site and species by pooling samples on the date when the maximum proportion of terminals were infested. We used degree-days as a standard among sites to compare when the maximum proportion of terminals were infested by female scales. Because no studies on development rate or threshold temperature have been reported for G. spuria, we used the same model developed for elm leaf beetle, Xanthogaleruca luteola (Mul¬ ler). We accumulated degree-days beginning 1 Mar each year using the single sine wave method and daily maximum and minimum temperatures obtained from recorders near our study sites (Dreistadt & Dahlsten 1990). We employed a lower threshold temperature of 11° C, the same as for elm leaf beetle and intermediate to the threshold determined in California for two other scales: San Jose scale, Quadraspidiotus perniciosus (Comstock) with a threshold of 10.6° C (Jorgensen et al. 1981) and California red scale, Aonidiella citri (Maskell), with a threshold of 11.5° C (Yu & Luck 1988). We compared means with Utests using STAT-SAK (Dallal 1986). Elm Leaf Beetle Relationship. — We investigated the relationship between de¬ foliation by elm leaf beetle and the density of female scales. We sampled elm leaf beetle damage at 1 to 3 week intervals from spring until beetles largely disappeared in the fall using the same terminals that were inspected for scales. For each sample, we rated leaf area consumed from 0 to 10, where 10 is 100% or total defoliation (Dreistadt & Dahlsten 1989). We performed least squares linear regressions using PROC REG (SAS Institute Inc. 1988) to correlate beetle defoliation with scale density. We pooled samples by date for each tree and compared the maximum proportion of scale-infested terminals versus maximum beetle defoliation. We regressed maximum defoliation against maximum scale density during the same year and maximum defoliation during the current year against maximum female scale density the subsequent year. We conducted separate regressions for each elm species for all sites and years pooled. Parasitism. — During 1988 and 1989, we clipped and returned to the laboratory the terminals sampled as above. We also randomly collected scales in Albany (Alameda Co.) from five Chinese elm ( Ulmus parvifolia Jacquin) on 3 dates in May and June, 1990. We clipped a 3-6 cm long section of twig containing one or more female scales from each infested terminal and placed it in a 22 by 93 mm cotton-stoppered shell vial. We held vials at 21-27° C 16:8 (L:D) during 1989 and 1990, and at about 24° C and uncontrolled artificial and diffuse natural light during 1988. Any European fruit lecanium, Parthenolecanium corni (Bouche) or black scale, Saissetia oleae (Olivier) present were scraped off before holding sam¬ ples for emergence. We recorded the number of scales in each sample and the THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) 242 emergence of any crawlers or parasitoids. To detect any unemerged parasitoids, we dissected scales without emergence after 3 or more months. We report apparent parasitism at each site for all sample dates pooled during the two month period when female univoltine scales were found to be most abundant. We determined apparent parasitism by dividing the number of para¬ sitized scales by the sum of parasitized and viable scales, then multiplying by 100. Parasitized scales were those from which parasitoids emerged or from which they were dissected. Viable scales were those from which crawlers emerged in the laboratory. Some samples contained scales that were too immature when collected to produce crawlers or from which all crawlers or parasitoids had emerged before they were collected; we excluded these from parasitism estimates. For twig samples producing crawlers and containing more than one unparasitized scale, we did not determine whether one or more than one of the unparasitized scale had produced crawlers. For these samples, we estimated the number of viable scales (V): V = pVs x N where N is the number of unparasitized scales in each sample with crawlers and pVs is the proportion of viable scales in all samples pooled from that date and location that contained only one unparasitized scale. Parasitoid introduction data through 1950 (Table 2) are from Flanders (1952); those data from 1952-1956 are from Biological Control of Elm Scale, unpublished University of California colonization reports from Riverside and the Gill Tract in Albany authored by R. L. Doutt, S. E. Flanders, and T. W. Fisher from 15 Jul 1952 through 15 Jul 1956 and summarizing work conducted in part by K.S.H. We identified Trichomasthus coeruleus Mercet using the published keys and descriptions of Mercet (1921, 1923), Nikol’skaya (1952), Graham (1958, 1969a), Alam (1957), and Jensen & Sharkov (1989). Publications used to identify the other parasitoids were: C. insidiator (Compere 1931), Microterys sp. males (Trjapitzn & Gordh 1978), Microterys sp. females (Tryapitsyn & Gordh 1978), Tetrastichus sp. (Burks 1943), and Pachyneuron sp. (Graham 1969b). We sent specimens of the former three genera to the Taxonomic Services Unit (T5U), USDA-ARS, Beltsville, Maryland for confirmation of identification. We deposited voucher specimens from our recoveries in the Bohart Museum, University of California, Davis. Results and Discussion Scale Abundance. —European elm scales occurred at all 12 of our northern California study sites. Scales were more abundant on English elm, Ulmus procera Salisbury, than Siberian elm, Ulmus pumila L. in Adin (t = 27.7; n = 240; P < 0.001) and Susanville (t = 22.2; n = 320; P < 0.001), where we sampled both host species during 1987 (Figs. 1 and 2). Significantly more English elm terminals were infested than Siberian elm terminals when samples from all sites on all dates were pooled during 1987 (t = 46.9; n = 2,625; P < 0.001), 1988 ( t = 54.4; n = 2,340; P < 0.001), and 1989 {t = 50.2; n = 5,938; P < 0.001), but climate and natural enemy differences among sites may partly account for this (see below). Significantly more American elm, Ulmus americana L., terminals were scale in¬ fested than Siberian elms in Princeton during 1988 (t = 226.0; n = 120; P < 0.001) and 1989 (t = 26.2; n = 160; P < 0.001), but too few American elms were 1994 DREISTADT & HAGEN: EUROPEAN ELM SCALE 243 Month/Day Figure 1. Mean (+ SD) proportion of Siberian elm branch terminals (30 cm long) infested with apparently viable female European elm scales on the sample date of maximum infestation. T) 0 ■+j CO 0 _co u Cl E k_ 0 o c o • — 11 o Q_ O 1.0 0.8 0.6 0.4 + I—American elms—I I-English elms-1 1 0.0 oo 00 CD >\ c o -O 0 . 2 - < CD oo CD >N C o _Q oo CD c o •+-» 0 o c * MM L_ CL oo oo CD c o -M 0 o c CD 00 c o -M 0 o c 00 CD ~a < 00 00 CD >\ c o XJ 1^ oo CD O O • — o 1 oo oo CD o o • — o CD 00 CD JD ft KB > 0 b o oo oo CD 0 C • — _Q _Q O DC 00 0 ft KM > c o 0 13 (/) CO (N ro CM CM M - CM CM CM 1^ CM CM CD m CM \ \ \ \ \ \ \ \ CO LD m LO m lO LO LO Month/Day Figure 2. Mean (+ SD) proportion of American elm and English elm branch terminals (30 cm long) infested with apparently viable European elm scales on the sample date of maximum infestation. 244 Vol. 70(3) THE PAN-PACIFIC ENTOMOLOGIST sampled to generalize about their susceptibility to scales relative to the other elm species. The maximum proportion of infested terminals occurred about 10 June (SD = 16 days) in northeastern California (Adin, Fall River Mills and Susanville), which was significantly later ( t = 5.3; n = 25; P < 0.001) than all other sites pooled, where infestations peaked about 5 May (SD = 16 days). Northeastern sites are intermountain valleys with colder winters and a shorter growing season than other study locations. Because of the varying climate among our sites, which span about 500 km and range from about 10 to >1200 m above sea level, we used degree- day accumulations to compare peak scale occurrence. The maximum proportion of infested terminals in northeastern California, other sites, and all sites pooled occurred at 301 (SD = 76), 305 (SD = 96), and 303 (SD = 89) degree-days, respectively; differences were not significant (P > 0.05). Degree-days accumula¬ tions can assist in timing scale population monitoring, parasitoid collections and releases, and insecticide applications. However, laboratory development rate and threshold temperature data should be developed for G. spuria to validate our estimates (see Materials and Methods). Elm Leaf Beetle Relationship. — We observed and photographed scale nymphs and elm leaf beetle larvae feeding on the same leaves. Because beetles can pre¬ maturely defoliate elms before scale nymphs would normally migrate in fall from leaves to bark, we hypothesized that high defoliation would reduce scale densities the subsequent season because many scale nymphs would die on leaves that are killed or drop because of beetle feeding. We observed the opposite effect; maxi¬ mum beetle defoliation during the current year was positively correlated with maximum female scale density during the next year on English elm (F = 15.7; n = 6; P < 0.05) (Fig. 3B). American elms that experienced little beetle feeding also had lower scale densities, but results may have been an artifact of small sample size. The apparent association between beetle feeding and scale density may be because trees are similarly predisposed to attack by both pests; for example, English elms are more susceptible to both elm leaf beetle (Dreistadt & Dahlsten 1989) and European elm scale than are Siberian elms (Fig. 3). Stress associated with beetle damage may increase elm susceptibility to scales, or vice versa. There was no association between current season beetle defoliation of Siberian elm and female scale density the subsequent year (Fig. 3B). Siberian elm foliage growth is indeterminate and anytime from spring through fall trees will drop individual leaves that are partially damaged by beetle feeding, leaves that may also contain scale nymphs. English and American elm growth is determinate. Trees retain damaged leaves, dropping most foliage over a short period after damage becomes severe then sometimes growing a second flush of foliage; reten¬ tion of damaged leaves may allow scale nymphs to move from foliage to bark before leaves drop. The maximum proportion of female scale-infested terminals was correlated with maximum beetle defoliation during the same season on English elms (F = 5.3; n = 25-P< 0.05) and Siberian elms {F = 4.1; n = 40; P < 0.05); however, R 2 values were very low (Fig. 3A) and it is unlikely that female scale density on twigs is affected by beetle feeding during the same season because scales mature and produce crawlers in the spring and maximum beetle defoliation occurs later in summer. The distribution, abundance, and behavior of scale nymphs on bark Proportion of Terminals Infested by Scales (y) 1994 DREISTADT & HAGEN: EUROPEAN ELM SCALE 245 0 2 4 6 8 10 Foliage Damage by Beetles (e) Figure 3. Foliage damage by elm leaf beetle (e), rated 0 to 10 where 10 = 100%, and A) proportion of terminals infested with female scales during the same season or B) proportion of terminals infested with scales during the next season on English and Siberian elms in northern California, 1987-1989. Significant (P < 0.05) regression equations are shown for English elm (y') and Siberian elm (y*). 246 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) Table 1. Apparent parasitism of European elm scale in northern California, 1988-1990. 3 Percent of total parasitism by species Location Year Sp n Percent parasitism Coc- . cophagus insidiator Trich- omasthus coeru- leus Micro- terys sp. Other b Mum¬ my 0 X SD Adin 1988 E 87 15.9 35.7 90 0 0 0 10 Adin 1988 S 108 59.7 39.1 85 0 0 0 15 Adin 1989 S 202 9.9 24.5 42 0 0 0 58 Albany 1988 A & E 245 0 0 0 0 0 0 0 Albany 1989 A & E 123 0 0 0 0 0 0 0 Albany 1990 C 345 1.1 6.3 0 100 0 0 0 Fall River 1988 S 287 1.5 10.5 100 0 0 0 0 Fall River 1989 S 204 2.9 14.1 100 0 0 0 0 Stockton 1988 s 168 13.4 33.8 0 67 0 33 0 Stockton 1989 s 851 4.6 16.3 0 81 14 0 5 Susan ville 1988 E 35 11.1 32.3 100 0 0 0 0 Susan ville 1988 s 148 0 0 0 0 0 0 0 Susanville 1989 E 72 18.1 34 43 0 14 0 43 Susanville 1989 S 828 12.6 27.2 44 0 24 2 30 a Host species (Sp) are American elm (A) English elm (E), Siberian elm (S), or Chinese elm (C); n is number of viable scales + parasitized scales sampled. b Other species are apparently secondaries: Tetrastichus sp. (in Stockton); Pachyneuron sp. (Susan- ville). c Parasitoids dissected from mummified scales were not identified. and leaves has not been reported and must be investigated in order to better understand the relationship between elm leaf beetle feeding and scale density. Coccophagus insidiator. — We reared at least three species of primary parasitoids and two apparently secondary species from European elm scale; of these only C. insidiator has previously been recovered in California. Coccophagus insidiator was the only scale parasitoid in Adin (Modoc Co.) and Fall River Mills (Shasta Co.) and it was the predominant species in Susanville (Table 1, Figs. 4 and 5). In comparison with its univoltine host, C. insidiator reportedly has more than one annual generation on female scales and parasitizes overwintering male scales prior to females becoming susceptible (Griswold 1927, Flanders 1952). Nearly all male scales and any parasitoids they may have contained emerged prior to sampling. Generational parasitism (van Driesche 1983) is likely higher than indicated by the apparent parasitism of female scales on any one date (Fig. 4) or from all dates pooled (Table 1). Coccophagus insidiator was introduced into the southern two-thirds of Cali¬ fornia through the mid-1950s (Table 2). It was previously reported as established only in Redlands (San Bernardino Co.) in southern California (Flanders 1952); which is about 900 km south of our recoveries. Although we collected many scales in Albany and Stockton (Table 1) and held an average of 370 scales for parasitoid emergence from each of 7 other locations in the Central Valley or coastal valleys of northern California (Fig. 5), we did not detect C. insidiator at those locations even though they encompass original release sites (Table 2) and are 300 km or more closer to Redlands than our recovery sites. Our identification of C. insidiator was confirmed by M. E. Schauff, TSU, USDA-ARS. Scales Parasitized Tips Infested or Viable Scales DREISTADT & HAGEN: EUROPEAN ELM SCALE 247 1994 0.4 - 0.3 - 0.2 - 0.1 - 0 Tips Infested Viable Scales Mummy Pachyneuron Microterys □ Coccophagus B m 5/26 6 / 16 6/30 Month/Day 7/28 Figure 4. A) Mean (+ SD) proportion of terminals (tips) infested and scales from which crawlers emerged (viable scales) and B) proportion of European elm scales parasitized by three species in Susanville, California, on each sample date in 1989. Trichomasthus coeruleus. — We recovered T. coeruleus in Stockton (San Joaquin Co.), where it was the most abundant parasitoid, and in Albany, where it was the only species parasitizing European elm scale (Table 1). Prior to its introduction in the Central Valley and the San Francisco Bay Area in 1952 and 1954, this 248 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) Adin ■ Fall River Mills Susanville •Chico • Princeton Hopland .•Marysville *Cloverdale Robbins Calistoga Albany Stockton 100 km ESTABLISHED PARASITOIDS: • None ■ Coccophagus ▲ Trichomasthus O Microterys \ Pachyneuron /Xletrastichus Figure 5. Northern California study sites where three species of European elm scale primary parasitoids ( Coccophagus insidiator, Trichomasthus coeruleus, Microterys sp.) and two apparently secondary parasitoids ( Pachyneuron sp., Tetrastichus sp.) were found or not detected, 1988-1990. species was keyed using Mercet (1921) and identified as Trichomasthus cyanifrons (Dalman). That key was in error (Mercet 1923). Based on more recent keys and descriptions that distinguish between T. cyanifrons and T. coeruleus, we believe that the Trichomasthus introduced from France into Albany, Berkeley, Madera, 1994 DREISTADT & HAGEN: EUROPEAN ELM SCALE 249 Table 2. Parasitoid introductions reported against European elm scale in California. 3 Parasitoid species Release year Parasitoid source Number released Release city (county) C. insidiator 1939 Italy 50 Los Gatos (Santa Clara) 1949 France & Germany 175 Redlands (San Bernardino) 80 San Jose (Santa Clara) 65 San Anselmo (Marin) 1950 France & Germany 61 San Anselmo 78 Berkeley (Alameda) 1952 France 312 Sacramento (Sacramento) 27 Berkeley 1953 Redlands, Calif. 60 Claremont (Los Angeles) 1954 France “several Albany (Alameda), Berkeley, hun- Madera (Madera) dred” 1955 Redlands, Calif. 400 San Bernardino (San Bernardino) 500 Claremont 1956 310 Claremont T. cyanifrons b 1949 France & Germany 27 Redlands 13 Balboa (Los Angeles) 16 Pasadena 1952 France 39 Sacramento 1954 31 Albany, Berkeley, Madera Metaphycus sp. 1954 France 26 Albany, Berkeley, Madera 3 Source: Flanders 1952 (1939-1950 data); Biological Control of Elm Scale, unpublished University of California colonization reports from Riverside and the Gill Tract in Albany (1952-1956). b Probably T. coeruleus, see text. and Sacramento during 1952 and 1954 (Table 2) was T. coeruleus, not T. cyani- frons as previously reported. The Trichomasthus introduced in southern California in 1949 from France and Germany may have been T. cyanifrons as reported by Flanders (1952), but we did not recover that species. Ours is the first reported recovery of a Trichomasthus species from G. spuria in North America. Specimens we sent to TSU were identified as Trichomasthus sp. Microterys sp.—European elm scales in Stockton and Susanville were parasit¬ ized by a Microterys sp. Two peaks of Microterys sp. emergence (early June and late July) were observed in Susanville in 1989 (Fig. 4B), indicating that Microterys sp. has more than one annual generation and inflicts greater mortality on its univoltine host than is indicated by apparent parasitism on any one sample date (Fig. 4B) or all dates pooled (Table 1). An undescribed Microterys sp. was imported into quarantine in California in 1949, but only males were produced and no releases were made according to Flanders (1952) and the recollection of K.S.H., who worked on that project. We believe that Clausen (1978) was wrong in stating that “a very few” Microterys sp. were released from quarantine and introduced. Clausen (1956) makes no mention of this species even though it discusses releases through 1953 and was published closer to the actual event than Clausen (1978). Morphological distinctions between Microterys in Susanville and Stockton indicate they may be different species. At least 8 species of Microterys have been reported in California, but their taxonomy is uncertain and this genus of parasitoids is poorly known (Rosen 1976, Gordh 250 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) 1979). We do not know the origin of our Microterys sp. and specimens sent to the TSU were returned as genus and species undetermined. Biological Control. — Our methods were not designed to assess parasitoid im¬ pact; we lacked sufficient time and there were no previous data on scale abundance or the species and distribution of any parasitoids. Differences in climate, elm species, and natural enemy distribution and possible effects from elm leaf beetle defoliation confound efforts to evaluate biological control. We also recovered three scale feeding Coccinellidae on our study elms: Hyper- aspis quadrioculata (Motschulsky) in Adin, Fall River Mills and Susanville, Chi- locorus bipustulatus (L.) in Stockton, and Rhyzobius forestieri (Mulsant) in Albany. Predators may also influence scale densities, but our methods were not designed to detect or sample predators or assess their impact. There is evidence that parasitoids can provide biological control of European elm scale. We excluded Argentine ants, Iridomyrmex humilis (Mayr) from Chinese elm branches in Albany in 1990 and 1991 and parasitism by T. coeruleus ap¬ parently increased and scale densities declined (Dreistadt et al. 1992). Flanders (1952) reported that C. insidiator parasitized up to 85% of mature European elm scales in Redlands and significantly reduced host populations there, but he pro¬ vided no methods or other data. Flanders (1952) states it is a “well-known fact that elm scale in Europe is under excellent natural control.” We found no published data on European elm scale density and parasitism in Europe; however, Burger et al. (1985) report that in 1984 a “heavy outbreak of the uncommon coccid Gossyparia spuria (Modeer) occurred at ’S-Gravenhage” in the Netherlands and that this may have been a secondary pest problem. Conversely, Casado (1985) states that European elm scale is, “another weakening factor to add to the very severe problems that the elm stands [around Madrid, Spain] already have.” Because of the copious and annoying honeydew it excretes, European elm scale infested elms have been treated with insecticide by at least one city (Chico) where we sampled scales but found no parasitoids. There are reports of European elm scale damage or control soon after its introduction in the West (Doten 1908, Herbert 1924) and more recently (Cranshaw et al. 1989). Conclusions. — Although one to three parasitoid species occurred at each of five of our study sites, no parasitoids were detected at seven locations. Substantial rates of parasitism and the apparently limited distribution of parasitoids in Cal¬ ifornia provides opportunity to introduce and study natural enemies before and after introduction at sites where they are not present. Ant exclusion may improve biological control at locations where parasitoids are established. European elm scale is an excellent candidate for further biological control research. Acknowledgment Sampling was conducted during travel for elm leaf beetle research funded by grants to Donald L. Dahlsten, Division of Biological Control, University of Cal¬ ifornia, Berkeley from the University of California Elvenia J. Slossen Endowment Fund for Ornamental Horticulture and the Statewide Integrated Pest Management Project. Susan M. Tait, Mabel Fong, David L. Rowney, and William A. Copper with the Division of Biological Control collected or processed some samples or helped to manage data. M. E. Schauff and M. Lacey-Theisen, Taxonomic Services 1994 DREISTADT & HAGEN: EUROPEAN ELM SCALE 251 Unit, Agricultural Research Service, United States Department of Agriculture, provided parasitoid identifications. Donald L. Dahlsten and anonymous reviewers critiqued our manuscript. Whitney S. Cranshaw, Colorado State University, pro¬ vided helpful suggestions. Literature Cited Alam, S. Mashhood. 1957. On the taxonomy of some British encyrtid parasites (Hymenoptera) of scale insects (Coccidoidea). Trans. Royal Ent. Soc. London, 109: 446. Burger, H. C., A. van Frankenhuyzen, L. J. W. de Goffau & S. A. Ulenberg. 1985. Bijzondere aantastingen door insekten in 1984. Entomologische Berichten, 45: 157-165. Burks, D. B. 1943. The North American parasitic wasps of the genus Tetrastichus— a contribution to biological control of insect pests. Proc. U.S. Nat. Muse., 93: 505-608. Casado, J. R. 1985. Gossyparia ulmi Geoffroy (Homoptera: Eriococcidae) una causa mas de debi- litamiento de los olmos: estudio morfologico y bionomico. Bol. Serv. Plagas., 11: 45-58. Clausen, C. P. 1956. Biological control of insect pests in the continental United States. U.S. Dept. Agric. Tech. Bull., 1139. Clausen, C. P. 1978. Introduced parasites and predators of arthropod pests and weeds. U.S. Dept. Agric. Handb., 480. Compere, H. 1931. A revision of the species of Coccophagus a genus of hymenopterous coccid- inhabiting parasites. Proc. U.S. Nat. Muse., 78: 1-132. Cranshaw, W. S., R. J. Zimmerman & D. Patrick. 1989. European elm scale insecticide evaluations. Insecticide & Acaricide Tests, 14: 341. Dallal, G. E. 1986. STAT-SAK, Version 2.14. Malden, Mass. Doten, S. B. 1908. The European elm scale ( Gossyparia spuria, Modeer). Univ. Nevada Agric. Exper. Stat. Bull., No. 65. Dreistadt, S. H. & D. L. Dahlsten. 1989. Density-damage relationship and presence-absence sampling of elm leaf beetle (Coleoptera: Chrysomelidae) in northern California. Environ. Entomol., 18: 849-853. Dreistadt, S. H. & D. L. Dahlsten. 1990. Relationships of temperature to elm leaf beetle (Coleoptera: Chrysomelidae) density and damage in the field. J. Econ. Entomol., 83: 837-841. Dreistadt, S. H., M. P. Parrella & M. L. Flint. 1992. Integrating chemical and biological control of ornamental pests, pp. 26-41. In Ward, C. R. (ed.). Proceedings of the 1992 First Southwest Ornamental Pest Management Workshop. New Mexico State University, Albuquerque. Flanders, S. E. 1952. A parasite of the European elm scale established in California. J. Econ. Entomol., 45: 1078-1079. Gordh, G. 1979. Encyrtidae. In Krombein, K. V., P. D. Hurd, Jr., D. R. Smith & B. D. Burks (eds.). Catalog of Hymenoptera in America north of Mexico. Vol. 1. Smithsonian Institution Press, Washington, D.C. Graham, M. W. R. de V. 1958. Notes on some genera and species of Encyrtidae (Hym.,Chalcidoidea), with special reference to Dalman’s types. Ent. Tidskr., 79: 172. Graham, M. W. R. de V. 1969a. Synonymic and descriptive notes on European Encyrtidae (Hym., Chalcidoidea). Bull. Entomol. de Pologne, 39: 267. Graham, M. W. R. de V. 1969b. The Pteromalidae of northwestern Europe (Hymenoptera: Chal¬ cidoidea). Bull. Brit. Muse. Nat. Hist. Entomol., Supplement 16. Griswold, G. H. 1927. The development of Coccophagus gossypariae Gahan, a parasite of the European elm scale. Ann. Entomol. Soc. Amer., 20: 553-555. Herbert, F. B. 1924. The European elm scale in the West. U.S. Dept. Agric. Bull., 1223. Jensen, P. B. & A. V. Sharkov. 1989. Revision of the genus Trichomasthus (Hymenoptera: Encyrtidae) in Europe and Soviet Asia. Ent. Scand., 20: 23-54. Jorgensen, C. D., R. E. Rice, S. C. Hoyt & P. H. Westigard. 1981. Phenology of the San Jose scale (Homoptera: Diaspididae). Can. Entomol., 113: 149-159. Mercet, R. C. 1921. Hymenopteros: Fam. Encirtidos. Fauna Iberica. Museo Nacional de Ciencias Naturales, Madrid. 732 pp. Mercet, R. G. 1923. Adiciones a la fauna espanola de Encirtidos. Bol. R. Soc. Esp. Hist. Nat., 23: 49-56, 174-179. Nikol’skaya, M. N. 1952. The chalcid fauna of the U.S.S.R. (Chalcidoidea). Academy of Sciences 252 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) of the U.S.S.R. Translated from Russian for the National Science Foundation, Washington, D.C., by the Israel Program for Scientific Translations, Jerusalem, 1963. Rosen, D. 1976. The species of Microterys (Hymenoptera: Encyrtidae): an annotated world list. Ann. Entomol. Soc. Amer., 69: 479-485. SAS Institute Inc. 1991. SAS/STAT user’s guide, Release 6.03. Cary, North Carolina. Trjapitzn(Tryapitsyn), V. A. & G. Gordh. 1978. A review of the Nearctic Encyrtidae (Hymenoptera, Chalcidoidea). Communication 1. Entomol. Review, 57: 257. Translated from Entomologi- cheskoye Obozreniye. Tryapitsyn (Tijapitzin), V. A. & G. Gordh. 1978. A review of the Nearctic Encyrtidae (Hymenoptera, Chalcidoidea). Communication 2. Entomol. Review, 57: 437. Translated from Entomologi- cheskoye Obozreniye. van Driesche, R. G. 1983. Meaning of “percent parasitism” in studies of insect parasitoids. Environ. Entomol., 12: 1611-1622. Yu, D. S. & R. F. Luck. 1988. Temperature-dependent size and development of California red scale (Homoptera: Diaspididae) and its effect on host availability for the ectoparasitoid Aphytis melinus DeBach (Hymenoptera: Aphelinidae). Environ. Entomol., 17: 154-161. PAN-PACIFIC ENTOMOLOGIST 70(3): 253-258, (1994) HOST-SPECIFIC DEMOGRAPHIC STUDIES OF WILD BACTROCERA TAU (WALKER) (DIPTERA: TEPHRITIDAE) Pingjun Yang, 1 James R. Carey, 13 and Robert V. Dowell 2 department of Entomology, University of California, Davis, California 95616 2 California Department of Food and Agriculture, 1220 N Street, Sacramento, California 95814 Abstract. — Developmental time and survival rates of preadult stages and adult survival and fecundity of Bactrocera tau (Walker) were examined when reared on six common hosts at 25° C. The durations of the egg and pupal stages were independent of host. The larval developmental time ranged from 4.5 days, for those reared on cucumber, to 7.3 days, for those reared on eggplant. The survival rate of larvae ranged from 48%, when reared on bitter melon, to 77%, when reared on cucumber. Net reproductive rates (R 0 ) were similar when the files were reared on bitter melon, papaya, and cucumber, and they were twice that of flies reared on eggplant. The generation time ranged from 59.1 days, for flies reared on pumpkin, to 40.7 days, for those reared on eggplant. The intrinsic rate of increase ranged from 0.087, for the flies reared on water¬ melon, to 0.123, for those reared on cucumber. The data indicate that B. tau is capable of reaching high population densities quickly on a number of hosts, and that this fly poses a significant threat to agriculture in California. Key Words.— Insecta, demography, reproductive parameters, life history traits, Bactrocera tau Demographic studies of fruit flies (Diptera: Tephritidae) are important for de¬ veloping of effective control programs, efficient mass rearing of sterile flies, and the interpretation of trap data (Carey 1993, Vargas & Carey 1990). Several eco¬ nomically important species including Ceratitis capitata (Weidemann) (Carey 1984, Vargas et al. 1984, Vargas & Carey 1990); Bactrocera cucurbitae (Coquillett) (Vargas et al. 1984, Carey et al. 1985, Vargas & Carey 1990); Bactrocera dorsalis (Hendel) (Vargas et al. 1984, Foote & Carey 1987, Vargas & Carey 1990); Bac¬ trocera Malaysian A and B (Chua 1991a, b) and Bactrocera latifrons (Hendel) (Vargas & Nishida 1985) have been studied. Because larval host is an important factor determining fruit fly geographic distribution and abundance, many studies have investigated the effect of host plants on fruit fly demographic characters (Carey 1984, Krainacker et al. 1987, Celedonio-Hurtado et al. 1988, Yang et al. 1990). Bactrocera tau (Walker), the pumpkin fly, is an economically important fruit fly distributed in South and Southeast Asia and some Pacific islands (Hardy 1973). It has a host range similar to that of B. cucurbitae, but is considered a more destructive species in southern China (Chao & Ming 1986). This study investigated B. tau host-specific, pre-adult survival and development; determined the age- specific fecundity of its adults, when reared on six different hosts; and analyzed the effect of host on its population growth. 3 To whom reprint requests should be sent. 254 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) Table 1. Mean larval and pupal development times and survivorship for Bactrocera tau reared on seven hosts at 25° C. Host Larva Pupa Jc (± SD) a % survival X (± SD) % survival Bitter melon 5.2 (1.1) 48 9.2 (0.4) 94 Cucumber 4.5 (0.7) 77 9.6 (0.5) 99 Eggplant 7.3 (1.2) 56 9.4 (0.6) 91 Papaya 5.3 (0.8) 68 8.8 (0.5) 95 Pumpkin 6.2 (0.9) 59 9.2 (0.8) 92 Watermelon 5.3 (0.9) 75 9.3 (0.7) 96 In days. Materials and Methods Bactrocera tau were reared from infested bitter melon {Momordica charantia L.) growing near the campus of Zhongshan (Sun Yat-sen) University, Guangzhou, China. The colony was reared in a room maintained at 25° ± 0.5° C, 50-75% RH, and 12:12 L:D. Pumpkin [Cucurbita moschata (Duchesne) Duchesne ex Poiret] was used as the larval host. After three generations, the offspring were used in the following experiments. All tests were replicated three times. Duration and mortality of larval and pupal stages were evaluated as follows. A pumpkin slice was exposed to gravid females for one hour, after which the eggs were removed with a knife and placed on moist black cloth in a Petri dish. Fifty newly hatched larvae each were seeded on a piece of host, which was held in a 1 liter glass jar with a layer of sand; the hosts were: bitter melon [Momordica charantia L.], cucumber [ Cucumis sativus L.], eggplant [Solanum melonena L.], papaya [Carica papaya L.], pumpkin [ Cucurbita moschata (Duchesne) Duchesne ex Poiret], or watermelon [Citrullus lanatus (Thunberg) Matsumato & Nakai]. Larvae were checked daily and food was added as needed. As pupation occurred, Table 2. Reproductive parameters (eggs per female), mean age of reproduction, and expectation of life (days) of adult B. tau reared on six different hosts. Host Gross rate 2 Net rate b Eggs per day Mean age reproduction 0 Expectation of life d Bitter melon 1289 819 7.5 81.3 109 Cucumber 911 538 7.5 59.5 72 Eggplant 886 372 7.2 58.3 52 Papaya 1064 563 6.6 77.2 85 Pumpkin 1512 640 5.8 129.2 103 Watermelon 1700 554 7.4 105 75 a Gross fecundity rate is expressed as the number of eggs per female that lives to the last possible day of life. b Net fecundity rate is expressed as the number of eggs per female, considering adult survival. c Mean age of reproduction is the age, in days, at which an average female has laid half of the total number of eggs. d Expectation of life is the average age of death expressed in days; calculated using data from males and females. 1994 YANG ET AL.: BACTROCERA DEMOGRAPHY 255 40 80 120 160 200 240 40 80 120 160 200 240 AGE (days) Figure 1. Number of eggs per female/day for B. tau reared on six hosts. the sand was sifted daily to remove pupae. Fifty pupae were placed in a Petri dish in a 1 liter glass jar and adult emergence was recorded daily. Fifty pairs of newly emerged adults were placed in a cubical cage (25 cm per side) provisioned with water and a 3:1 volumetric mixture of commercial sugar and enzymatic yeast hydrolysate. A thin slice of pumpkin was placed in the cage daily to determine egg production. Mortality was recorded daily until all adults had died. All tests were conducted in a room maintained at 25° ± 0.5° C, 50- 75% RH, and 12:12 L:D. The demographic parameters were calculated and sum¬ marized according to Carey (1993). Results Duration and survival of B. tau eggs were independent of host and averaged approximately 1.5 days and 87% respectively. Larval developmental time ranged from 4.5 days, for flies reared on cucumber, to 7.3 days, on eggplant. Larval survival rates ranged from 48%, on bitter melon, to 77% on cucumber. There are no significant differences among pupal developmental times or survival rates among the different hosts (P > 0.05; Analysis of Variance) (Table 1). Expectation of life of adult females ranged from 52 days, for flies reared on eggplant, to 109 days, for those reared on bitter melon. Gross fecundity rate ranged from 886 eggs/female, for flies reared on eggplant, to 1700 eggs/female, for those reared on watermelon (Table 2). The reproduction patterns of females reared on the six hosts were similar (Fig. 1). Age at first reproduction was approximately 16 days, and peak egg laying occurred shortly after this. Females reared on bittermelon, cucumber, eggplant, and papaya had egg production that peaked early in life (40-80 days) and declined 256 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) Table 3. Life table parameters of Bactrocera tau reared on six common hosts as computed from laboratory birth and death rates. Host Parameter 1 b d r X Ro T DT Bitter melon 0.159 0.063 0.096 1.100 184 54.4 7.2 Cucumber 0.125 0.004 0.123 1.104 180 42.3 5.6 Eggplant 0.155 0.043 0.112 1.118 95 40.7 6.2 Papaya 0.128 0.027 0.099 1.104 183 52.3 7.0 Pumpkin 0.132 0.032 0.092 1.097 169 59.1 7.5 Watermelon 0.120 0.033 0.087 1.091 138 56.8 8.0 a b = intrinsic birth rate; d = intrinsic death rate; r = intrinsic rate of increase (daily); X = finite rate of increase (daily); R 0 = net reproductive rate; T = mean generation time (days); DT = population doubling time (days). thereafter. Flies reared on pumpkin and watermelon layed eggs over a longer time span and lacked a clear decline in egg production. Average daily egg production ranged from 5.8 to 7.5 days (Table 2). There was a small number of highly fecund and longer-lived individuals in every cohort, especially among those reared on pumpkin and watermelon (Fig. 1). Demographic parameters of B. tau reared on six different hosts were calculated from the development and survival information for preadults and the life history information for adults (Table 3). Flies reared on eggplant had the lowest net reproductive rate. Flies reared on bitter melon, papaya, and cucumber had similar net fecundity rates that were twice that of flies reared on eggplant. Generation times ranged from 40.7 to 59.1 days. Intrinsic rates of increase ranged from 0.087, for flies reared on watermelon, to 0.123, for those reared on cucumber. The stable age structure of B. tau populations reared on different hosts are shown in Table 4. The fraction of flies in each stage was similar among the different hosts except for eggplant. The greater proportion of the population in the larval stage, and smaller proportion in the adult stages, reflect reduced larval survival and increased larval development times in eggplant (Table 1). It takes more B. latifrons larvae in eggplant to give rise to each adult compared to the other host plants. Discussion Our results are similar to those obtained for B. tau reared at 25° C on cucumber (Zhou et al. in press) for larval (4.5 versus 6.1 d) and pupal (9.6 versus 10.0 d) developmental times. Rates of gross and net fecundity (979 versus 911 eggs, and 665 versus 538 eggs respectively) were greater in Zhou et al., but our daily egg Table 4. Stable age distribution (percent) for B. tau reared on six different hosts. Stage Bitter melon Cucumber Eggplant Papaya Pumpkin Watermelon Egg 22.1 20.0 21.9 17.9 18.7 17.2 Larva 38.6 37.0 51.1 38.4 42.5 39.9 Pupa 23.7 30.0 18.0 26.2 22.5 26.3 Adult 15.6 13.0 9.0 17.5 16.3 16.6 1994 YANG ET AL.: BACTROCERA DEMOGRAPHY 257 deposition rate was greater (7.5 versus 6.0 eggs per day). The intrinsic rate of increase (0.11 versus 0.12) and finite rates of increase (1.11 versus 1.10) were almost identical. Similar rates of population increase (represented by the finite rate of increase) for B. tau reared on six different hosts indicate that these hosts had similar over¬ all effects on B. tau populations. Although flies reared on eggplant have the lowest fecundity and the shortest life expectancy, they have the highest finite rate of increase because their short generation time offsets lower egg production. Similar life history traits have been found in C. capitata (Carey 1984, Krainacker et al. 1987). B. cucurbitae (Carey et al. 1985), and Anastrepha spp. (Celedonio-Hurtado et al. 1988). Finite rates of increase of B. tau in this study are similar to those of B cucurbitae in Hawaii and southern China, where these two species of flies were reared on the same hosts (Carey et al. 1985). For example, the finite rate of increase for B. cucurbitae was 1.12 to 1.05 when reared on cucumber in Hawaii and southern China, respectively, but that for B. tau was 1.10. Our data agree with the results of other comparisons of B. tau and B. cucurbitae (Yang 1992). Bactrocera cucur¬ bitae is considered an important and potentially dangerous exotic pest to agri¬ culture in California. Our data indicate that B. tau is as dangerous a pest as B. cucurbitae, based upon their similar host ranges and population growth capacities. Bactrocera tau ranges further north into China than B. cucurbitae (Yang 1992) and, thus, poses a greater threat in California, especially in those areas that are marginally available to B. cucurbitae, such as the San Francisco Bay area. Acknowledgment We thank Pu Zhelong in Zhongshan University, Guangzhou, China for his suggestions and support. Literature Cited Carey, J. R. 1984. Host-specific demographic studies of the Mediterranean fruit fly, Ceratitis capitata. Ecol. Entom., 9: 261-270. Carey, J. R. 1993. Applied demography for biologists with special emphasis on insects. Oxford Univ. Press, New York. Carey, J. R., E. J. Harris & D. O. Mclnnis. 1985. Demography of a native strain of melon fly Dacus cucurbitae, from Hawaii. Entomol. Exp. Appl., 38: 195-199. Celedonio-Hurtado, H., P. Liedo, M. Aluja & J. Guillen. 1988. Demography of Anastrepha ludens, A. obhqua and A serpentina (Diptera: Tephritidae) in Mexico. Fla. Entomol., 71: 111-120. Chao, Y. S. &. Y. Ming. 1986. The investigation on fruit flies (Trypetidae-Diptera) injurious to fruits and vegetables in South China. Tech. Bull. Inst. Plant Quar. Press, Beijing, (in Chinese, with English abstract). Chua, T. H. 1991a. Comparison of demographic parameters in wild Bactrocera sp. (Malaysian A) (Diptera: Tephritidae) from different hosts. J. Pit Prot. Tropics, 8: 161-166. Chua, T. H. 1991b. Demographic parameters of wild Bactrocera sp. (Malaysian B) (Diptera: Te¬ phritidae). J. Pit. Prot. Tropics, 8: 139-144. Foote, D. & J. R. Carey. 1987. Comparative demography of a laboratory and a wild strain of the Oriental fruit fly, Dacus dorsalis. Entomol. Exp. Appl., 44: 263-268. Hardy, D. E. 1973. The fruit flies (Tephritidae-Diptera) of Thailand and bordering countries. Pac. Ins. Mono., 31. Krainacker, D. A., J. R. Carey & R. I. Vargas. 1987. Effect of larval host on life history traits of the Mediterranean fruit fly, Ceratitis capitata. Oecologia (Berlin), 73: 583-590. Vargas, R. I., D. Miyashita & T. Nishida. 1984. Life history and demographic parameters of three laboratory-reared tephritids (Diptera: Tephritidae). Ann. Entomol. Soc. Am., 77: 651-656. 258 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(3) Vargas, R. I. & T. Nishida. 198 5. Life history and demographic parameters of Dacus latifrons (Hendel) (Diptera: Tephritidae). J. Econ. Entomol., 78: 1242-1244. Vargas, R. I. & J. R. Carey. 1990. Comparative survival and demographic statistics for wild Oriental fruit fly, Mediterranean fruit fly, and melon fly (Diptera: Tephritidae) on papaya. J. Econ. Entomol., 83: 1344-1349. Yang, P., 1992. Demographic investigations on selected dacine fruit flies in southern China and Hawaii. Ph.D. Dissertation, University of California, Davis. Yang, P., C. Zhou, H. Chen & J. R. Carey. 1990. Demographic analysis of melon fly, Dacus cucurbitae rearing on five common hosts in China. Ecological Science, No. 2. (in Chinese, with English abstract). Y ang, P. & C. Zhou. 1988. Demography of a native strain of pumpkin fly, Dacus tau from Guangzhou, pp. 61-63. In The Proceeding of International Workshop on Statistical Ecology and its Appli¬ cation in Fisheries and the Second National Conference on Mathematical Ecology and its Application. July, 1988, Wuxi, China. Zhou, C., K. Wu, H. Chen, P. Yang & R. V. Dowell, (in press). Effect of temperature on the population growth of Bactrocera tau (Walker) (Diptera: Tephritidae). J. Appl. Entomol. PAN-PACIFIC ENTOMOLOGIST Information for Contributors See volume 66(1): 1-8, January 1990, for detailed general format information and the issues thereafter for examples; see below for discussion of this journal’s specific formats for taxonomic manuscripts and locality data for specimens. Manuscripts must be in English, but foreign lan¬ guage summaries are permitted. Manuscripts not meeting the format guidelines may be returned. Please maintain a copy of the article on a word- processor because revisions are usually necessary before acceptance, pending review and copy-editing. Format. — Type manuscripts in a legible serif font IN DOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 X 11 in, non¬ erasable, high quality paper. THREE (3) COPIES of each manuscript must be submitted, EACH INCLUDING REDUCTIONS OF ANY FIG¬ URES TO THE 8.5 X 11 IN PAGE. 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Volume 70 THE PAN-PACIFIC ENTOMOLOGIST July 1994 Number 3 Contents ICHINOSE, K.—Limited multiple-mating in males and single-mating in females of the ant species, Paratrechinaflavipes (Fr. Smith) (Hymenoptera: Formicidae).. 183 GORDH, G.—A biographical account of Harold Compere (1896-1978), biological control foreign explorer....... 18 8 TURNER, C. E„ R. SOBHIAN, D. B. JOLEY, E. M. COOMBS & G. L. PIPER—Establishment of Urophora sirunaseva (Hering) (Diptera: Tephritidae) for biological control of yellow starthistle in the western United States.... 206 SUGG, P. M., L. GREVE & J. S. EDWARDS—Neuropteroidea from Mount St. Helens and Mount Rainier: dispersal and immigration in volcanic landscapes__ 212 O’NEILL, K. M.—Livestock dung as a food resource and thermal refuge for rangeland grass¬ hoppers (Orthoptera: Acrididae).... 222 STARMER, W. T. & J. M. BOWLES—The spatial distribution of endemic and introduced flower-breeding species of Drosophila (Diptera: Drosophilidae) during their early his¬ tory of encounter on the island of Hawaii...... 230 DREISTADT, S. H. & K. S. HAGEN—European elm scale (Homoptera: Eriococcidae) abun¬ dance and parasitism in northern California_____ 240 YANG, P., J. R. CAREY & R. V. DOWELL—Host-specific demographic studies of wild Bactrocera tau (Walker) (Diptera: Tephritidae).. 253 The PAN-PACIFIC ENTOMOLOGIST Volume 70 October 1994 Number 4 Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES (ISSN 0031-0603) The Pan-Pacific Entomologist EDITORIAL BOARD J. T. Sorensen, Editor R. V. Dowell, Associate Editor R. E. Somerby, Book Review Editor Paul H. Arnaud, Jr., Treasurer R. M. Bohart J. T. Doyen J. E. Hafernik, Jr. J. A. Powell Published quarterly in January, April, July, and October with Society Proceed¬ ings usually appearing in the October issue. All communications regarding non-re¬ ceipt of numbers, requests for sample copies, and financial communications should be addressed to: Paul H. Arnaud, Jr., Treasurer, Pacific Coast Entomologi¬ cal Society, Dept, of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Application for membership in the Society and changes of address should be ad¬ dressed to: Stanley E. Vaughn, Membership Committee chair, Pacific Coast Ento¬ mological Society, Dept, of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Manuscripts, proofs, and all correspondence concerning editorial matters (but not aspects of publication charges or costs) should be sent to: Dr. John T. Sorensen, Editor, Pan-Pacific Entomologist, Insect Taxonomy Laboratory, California Dept, of Food & Agriculture, 1220 N Street, Sacramento, CA 95814. See the back cover for Information-to-Contributors, and volume 66(1): 1-8, January 1990, for more detailed information. Information on format for taxonomic manuscripts can be found in volume 69(2): 194-198. Refer inquiries for publication charges and costs to the Treasurer. The annual dues, paid in advance, are $25.00 for regular members of the Society, $26.00 for family memberships, $12.50 for student members, or $40.00 for institu¬ tional subscriptions or sponsoring members. Members of the Society receive The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes are available; see 67(1): 80 for current prices. Make checks payable to the Pacific Coast Entomological Society. Pacific Coast Entomological Society OFFICERS FOR 1994 Kirby W. Brown, President Paul H. Arnaud, Jr., Treasurer Julieta F. Parinas, Assist. Treasurer Curtis Y. Takahashi, President-Elect Vincent F. Lee, Managing Secretary Keve Ribardo, Recording Secretary THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly by the Pacific Coast Entomological Society, c/o California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Second-class postage is paid at San Francisco, CA and additional mailing offices. Postmaster: Send address changes to the Pacific Coast Entomological Society, c/o California Acade¬ my of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. This issue mailed 31 October 1994 The Pan-Pacific Entomologist (ISSN 0031-0603) PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A. © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PAN-PACIFIC ENTOMOLOGIST 70(4): 259-266, (1994) GENETIC DIVERSITY IN OVERWINTERED AND NON-OVERWINTERED IPS PINI (SAY) (COLEOPTERA: SCOLYTIDAE) IN IDAHO Sandra J. Gast 1 and Molly W. Stock 2 ‘USDA Forest Service, 3 Timber Cooperative Forestry and Pest Management, Coeur d’Alene, Idaho 83814 department of Forest Resources, University of Idaho, Moscow, Idaho 83843 Abstract.— The pine engraver, Ips pint (Say), has two generations per year in ponderosa pine (Pinus ponderosa Lawson) in Idaho, one that develops in pine slash in the spring and one that develops in live trees in the summer. The summer generation overwinters to produce the spring generation the following year. Non-overwintered and overwintered beetles were sampled from two sites. Average heterozygosity was significantly higher in overwintered beetles in both groups. While the proportion of homozygous individuals did not differ significantly between overwintered and non-overwintered beetles from either site, the proportion of heterozygous individuals was significantly greater after overwintering in beetles from one of the two sites. The increase in genetic diversity after overwintering is consistent with observations that heterozygosity is favored by severe environmental conditions. Key Words.— Insecta, Scolytidae, genetics, heterozygosity, stress The pine engraver, Ips pini (Say), is a widely distributed bark beetle infesting several species of pine in coniferous forests throughout North America. In Idaho, ponderosa pine ( Pinus ponderosa Lawson) is most commonly infested. Typically, two generations of I. pini are produced each year in Idaho, one in spring and one in summer (Fig. 1). The FI progeny of overwintered adults emerge in late spring and infest either fresh slash or live, standing trees that are typically immature, in dense stands, and moisture stressed. Like other bark beetle species, the pine engraver can mass attack live trees and kill them by feeding on the phloem, which disrupts the trees’ food supply, and introducing pathogenic fungi. They may also infest tops of more mature trees, including those previously attacked by the moun¬ tain pine beetle, Dendroctonus ponderosae Hopkins, or the western pine beetle, D. brevicomis LeConte. The second generation (F2, or summer generation) ma¬ tures in late summer and overwinters as adults in the litter beneath trees killed in the summer, or under the bark of these trees, most often at their base (Livingston 1979). Overwintered adults emerge in April and May and fly to infest fresh slash (recently felled trees and branches) or the tops of trees broken off by wind or snow. At that time, live trees are generally not attacked. Ips pini exhibits the high levels of heterozygosity (average frequency of hetero¬ zygous individuals per locus) that characterize bark beetles in general. Average heterozygosity for 17 bark beetles —10 Dendroctonus species (Bentz & Stock 1986) and seven Ips species (Cane et al. 1990)—was 16.7 percent. Ips pini was 15.9 percent heterozygous (Cane et al. 1990). The advantage of heterozygosity is frequently expressed as higher survival of 3 1201 Ironwood Drive. 260 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) F2 generation adults overwinter under the bark or in the duff and litter Adults j- F2 Generation { Pu P ae *- Larvae i- Eggs i- Green Trees or New Slash FI generation adults emerge from slash to attack standing green trees or new slash FI Generation < 1 Adults t I Pupae I Larvae t- ■ Eggs i- Fresh Slash, Snowbreakage, or Windthrow F2 overwintered adults leave duff and litter to attack fresh slash, snowbreakage, or windthrow Overwintering adults Bark, Duff, and Litter Overwintering adults "l-r DEC MARCH APRIL MAY JUNE Figure 1. Ips pini life history in Idaho (modified from Livingston 1979). T T -1-1-r JULY AUGUST SEPT OCT NOV heterozygous individuals under severe or stressful environmental conditions, even when these conditions are transient or episodic (e.g., Bryant 1974, 1976; Milkman 1978; Smith et al. 1975; Parsons 1971, 1987). Samollow & Soule (1983), for example, discovered a striking case of superior heterozygote survivorship among toads during the winter. Development of the mountain pine beetle in stressful environmental conditions—dry or thin phloem in the laboratory or in high-density infestations in the field—has been associated with increased levels of genetic diversity, measured as average heterozygosity (Stock & Amman 1983, Amman and Stock in press). The high levels of inherent genetic diversity in I. pini populations, combined with differences in the severity of environmental conditions affecting a population over the course of a year, make the pine engraver an ideal subject for further studies of the relationship between heterozygosity and environmental stress. To initiate studies of this type, we measured heterozygosity in non-overwintered and overwintered F2 I. pini from two sites in northern Idaho. Materials and Methods Collections. — F2 beetles were obtained in August 1985 from standing ponderosa pine trees at Moscow Mountain (Latah Co., Idaho) and Greer (Clearwater Co., Idaho, approximately 80 km from Moscow Mtn.). Fifteen to 20 logs, approxi¬ mately 1.5 m long and averaging 20 cm in diameter, were collected from each site. Half of the logs from the Greer and Moscow sites were brought indoors in the fall, and beetles were collected as they emerged and flew to an illuminated white sheet. These beetles represented the F2 generation before overwintering. The other half of the logs from Greer were kept over the winter in the outdoor cage containing a 10 cm layer of soil and litter, and emerging beetles (overwintered F2s) were 1994 GAST & STOCK: GENETIC DIVERSITY IN IPS 261 Table 1. Enzyme characteristics, allele frequencies, heterozygosity (h), and average heterozygosity (H) in samples of Ips pini taken from two sites. F2 = summer generation (non-overwintered) and OW = overwintered F2s. Asterisks mark loci where observed genotype frequencies differed from expected Hardy-Weinberg values. Locus Anodal or cath¬ odal 11 Mono¬ meric or dimeric b RM C Moscow Mountain Greer F2 ow F2 ow AAT A D n ( 144 ) ( 150 ) ( 89 ) ( 128 ) .45 P ( l ) .05 .08 .04 .07 .38 P ( 2 ) .89 .84 .92 .88 .31 P ( 3 ) .06 .07 .04 .05 .24 P ( 4 ) — — — — h (. 202 ) (. 280 ) (. 150 ) (. 218 ) ADH C D n ( 150 ) ( 150 ) ( 145 ) ( 148 ) ** .27 Pd ) .07 .05 .07 .06 .22 P ( 2 ) .91 .83 .92 .82 .17 P ( 3 ) .01 .12 .01 .11 .12 P ( 4 ) — — — .01 h (. 167 ) (. 294 ) (. 149 ) (. 312 ) AK A M n ( 78 ) ( 149 ) ( 113 ) ( 150 ) .76 P(D — — — .01 .69 P ( 2 ) .24 .29 .22 .24 .61 P ( 3 ) .74 .70 .76 .72 .51 P ( 4 ) .02 .01 .01 .02 .40 P ( 5 ) — — .01 .02 h (. 394 ) (. 426 ) (. 374 ) (- 423 ) CAT A D n ( 147 ) ( 108 ) ( 155 ) ( 150 ) .26 P ( l ) .02 .02 .01 .04 .21 P ( 2 ) .96 .91 .93 .91 .16 P ( 3 ) .02 .07 .05 .06 h (. 078 ) (. 167 ) (. 132 ) (. 167 ) G6PDH A M n 150 149 148 146 .20 P ( l ) 1.0 1.0 1.0 1.0 h ( 0 ) ( 0 ) ( 0 ) ( 0 ) GAPDH C D n ( 150 ) ( 150 ) ( 155 ) ( 148 ) ** ** .27 Pd ) .06 .05 .06 .05 .22 p(2) .93 .84 .94 .78 .17 P ( 3 ) .01 .11 — .16 .12 P ( 4 ) — — — .01 h (. 131 ) (. 280 ) (. 113 ) (. 363 ) GUS A D n ( 149 ) ( 111 ) ( 132 ) ( 150 ) .38 Pd ) .01 .03 — .04 .32 P ( 2 ) .99 .92 .99 .93 .26 P ( 3 ) — .05 — .03 h (. 020 ) (. 150 ) (. 020 ) (. 133 ) IDH1 A D n ( 150 ) ( 150 ) ( 158 ) ( 150 ) .42 P(l) — — — — .36 p(2) .89 .87 .89 .84 .28 P ( 3 ) .08 .09 .11 .11 262 THE PAN-PACIFIC ENTOMOLOGIST Yol. 70(4) Table 1. Continued. Locus IDH2 LAP LDH MDH1 MDH2 ME MPI PEP1 Anodal Mono- Moscow Mountain Greer or cath- meric or_ odal‘ dimeric b RM C F2 ow F2 ow .22 P ( 4 ) .02 .04 .01 .05 h (. 201 ) (. 233 ) (. 196 ) (. 280 ) c M n ( 150 ) ( 150 ) ( 155 ) ( 150 ) .16 P ( l ) 1.0 1.0 1.0 1.0 h (0) ( 0 ) ( 0 ) ( 0 ) A M n ( 148 ) ( 150 ) ( 158 ) ( 150 ) .34 P ( l ) .02 .05 — .03 .28 P ( 2 ) .98 .95 1.0 .97 h (. 039 ) (. 095 ) ( 0 ) (. 058 ) C D n ( 150 ) ( 150 ) ( 154 ) ( 148 ) ** ** .27 P(D .06 .05 .06 .05 .22 P ( 2 ) .93 .84 .93 .78 .17 P ( 3 ) .01 .11 .01 .16 .12 P ( 4 ) — — — .01 h (. 131 ) (. 280 ) (. 131 ) (. 363 ) A D n ( 150 ) ( 150 ) ( 156 ) ( 150 ) .42 Pd ) .01 — — — .34 p ( 2 ) .01 .01 — .01 .26 P ( 3 ) .96 .97 .96 .96 .18 P ( 4 ) — — .02 .02 .10 P ( 5 ) .02 .01 .01 — h (. 078 ) (. 059 ) (. 078 ) (. 078 ) C D n ( 150 ) ( 150 ) ( 158 ) ( 150 ) .33 P(l) — — — — .25 P ( 2 ) 1.0 1.0 .99 .98 .17 P ( 3 ) — — .01 .01 h ( 0 ) ( 0 ) (. 020 ) (. 039 ) A D n ( 150 ) ( 150 ) ( 158 ) ( 150 ) ** .31 P(l) .01 .03 — .03 .26 p ( 2 ) .98 .96 .99 .97 .21 P ( 3 ) — .01 .01 — h (. 039 ) (. 077 ) (. 020 ) (. 058 ) A M n ( 150 ) ( 149 ) ( 77 ) ( 150 ) .71 P(D .93 .89 .94 .93 .66 P ( 2 ) .07 .09 .05 .05 .56 P ( 3 ) - .02 .01 .02 h (. 130 ) (. 200 ) (. 114 ) (. 132 ) A D n ( 150 ) ( 150 ) ( 158 ) ( 150 ) .44 P ( l ) — — .01 — .40 P ( 2 ) .99 1.0 .99 .99 .36 P ( 3 ) .01 — — — h (. 020 ) ( 0 ) (. 020 ) (. 020 ) A D n ( 148 ) ( 146 ) ( 149 ) ( 150 ) ** * .27 P ( l ) .01 .01 .02 .02 PEP2 1994 GAST & STOCK: GENETIC DIVERSITY IN IPS 263 Table 1. Continued. Locus Anodal or cath¬ odal 8 Mono¬ meric or dimeric 1 ’ RM C Moscow Mountain Greer F2 ow F2 ow .21 P(2) .47 .45 .54 .50 .15 P(3) .15 .52 .43 .45 .09 p(4) .35 .02 .02 .02 .03 P(5) .01 — — — h (.634) (.527) (.523) (.547) PGI A D n (150) (150) (158) (150) .37 P(l) .01 — .01 .01 .32 P(2) .74 .77 .68 .74 .27 P(3) .24 .22 .31 .24 .24 P(4) .01 .01 — .01 h (.394) (.358) (.441) (.395) SOD A M n (150) (150) (156) (150) .27 Pd) 1.0 1.0 1.0 1.0 h (0) (0) (0) (0) H (%) 14.0 18.3 13.1 18.9 a Direction of migration during electrophoresis. b Molecular structure. c Relative mobility. collected in spring 1986. The other half of the logs from Moscow Mtn. were left on site over the winter and brought indoors in the spring for collection of beetles as they emerged and flew to an illuminated white sheet. Voucher specimens of male and female beetles from each location were placed in the William F. Ban- Entomological Museum, University of Idaho. Genetic Analysis. — The genetic makeup of populations was estimated using data obtained by horizontal starch gel electrophoresis. Approximately 150 overwin¬ tered and non-overwintered beetles from each site were analyzed. Techniques and stains used followed those of Higby & Stock (1982) and Bentz & Stock (1986). Analyses of data were performed using BIOSYS-1 (Swofford & Selander 1981) and SAS (SAS Institute 1988). Initially, genotype frequencies from male and female beetles in each group were compared using a contingency chi-square test. Where no significant differences occurred, data on males and females were pooled for further analysis. Observed genotype frequencies were compared to values derived from random-mating (Hardy-Weinberg) expectations using a chi-square test. Levels of heterozygosity were compared using two different approaches. In the first, Nei’s (1975) average heterozygosity (H) was calculated and compared with two-tailed 7-tests on transformed data to identify differences between non-over¬ wintered and overwintered beetles from the two sites. In the second approach, the proportion of heterozygotes, taken by direct count, was compared using a procedure for categorical data modeling. Results Nineteen loci from 16 enzyme systems were assayed. Of these, three loci (G6PDH, IDH2, and SOD) were monomorphic in all samples and 16 loci (AAT, ADH, 264 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Table 2. Percent of heterozygous individuals at 16 polymorphic loci in non-overwintered (F2) and overwintered (OW) Ips pini from two sites. Locus Moscow Mountain Greer F2 ow F2 OW AAT 16.7 19.8 10.1 23.4 ADH 14.7 13.3 15.2 23.0 AK 41.0 47.0 43.4 33.3 CAT 8.3 12.2 9.7 13.3 GAPDH 12.7 12.7 4.5 25.7 GUS 1.8 7.4 1.5 11.3 IDH1 16.0 21.3 17.7 22.0 LAP 1.3 6.7 0 4.0 LDH 13.3 12.7 13.0 24.3 MDH1 6.0 4.0 5.8 7.3 MDH2 0 1.3 1.3 2.0 ME 0.7 0.6 0 0 MPI 12.7 16.8 9.1 14.7 PEP1 0 0 1.3 1.3 PEP2 52.7 37.7 48.3 46.7 PGI 46.0 42.7 47.5 37.3 Average % heterozygotes per locus 15.2 16.0 14.3 18.1 AK, CAT, GAPDH, GUS, IDH1, LAP, LDH, MDH1, MDH2, ME, MPI, PEP1, PEP2, and PGI) were polymorphic in one or more samples. No significant dif¬ ferences occurred between genotype frequencies of male and female beetles from different locations within overwintered and non-overwintered groups. Therefore, the males and females were pooled for comparison of the genetic makeup of overwintered and non-overwintered beetles from the two sites (Table 1). Deviations from expected genotype frequencies occurred not at all in non- overwintered F2s from Greer and only once (PEP2) in non-overwintered F2s from Moscow Mtn.. Deviations from Hardy-Weinberg expectations were observed in overwintered F2s from Moscow Mtn. (at four loci: ADH, GAPDH, LDH, and PEP2) and in overwintered F2s from Greer (three loci: GAPDH, LDH, and ME). At both sites, average heterozygosity (H) was significantly higher in the overwin¬ tered beetles than in non-overwintered beetles: 18.3 vs. 14.0% (P < 0.05) in the Moscow Mtn. beetles and 18.9 vs. 13.1% (P < 0.01) in Greer beetles. The proportion of homozygotes taken by direct count (Table 2) did not differ between overwintered and non-overwintered beetles from either site. However, the proportion of heterozygotes taken by direct count was significantly higher (P < 0.01) in overwintered than in non-overwintered beetles from Greer (18.1 vs. 14.3%). Although there was also a slightly larger proportion of heterozygotes in overwintered beetles from Moscow Mtn. (16.0 vs. 15.2%), this difference was not significant. Discussion The classical view of genetic diversity in nature assumes that the fittest form of virtually every locus is the homozygous form. However, electrophoretic studies have revealed a large amount of genetic diversity within natural populations (e.g., 1994 GAST & STOCK: GENETIC DIVERSITY IN IPS 265 Selander 1976, Ferguson 1980), and have shown that heterozygous individuals are more adaptable than their more homozygous counterparts. In a number of plant and animal species, relatively heterozygous individuals display superior growth and survival (e.g., Garton et al. 1984, Mitton & Grant 1980, Mitton & Koehn 1975, Soule 1980), and under laboratory conditions, heterozygous pop¬ ulations are often able to maintain larger population sizes or biomass than less heterozygous populations (Beardmore 1983). The precise biochemical basis of heterozygote superiority is unknown, but it has been suggested that the presence of multiple forms of a gene product in heterozygous individuals confers relatively greater flexibility and latitude of a biochemical process. These advantageous effects can, in most cases, be attributed to heterozygosity per se, not to the effects of specific gene combinations (Mitton & Grant 1984). Heterozygosity appears to broaden the range of physiological tolerance and function relative to homozygosity (Mitton & Grant 1984, Smith et al. 1975). Similarly, a genetically diverse population is considered more adaptable to changing environmental conditions and more likely to survive over long time periods. A central difference between non-overwintered and overwintered populations of Ips pini is the severe environmental conditions to which the latter group is exposed. Our primary question concerned the effect of overwintering on the level of heterozygosity in the F2 generation of Ips pini. Overwintering significantly increased levels of average heterozygosity in beetles from both sites that were studied, and the proportion of heterozygous individuals was significantly greater after overwintering at one of the two sites. Thus, this study lends some support to the hypothesis that severe environmental conditions tend to select for a more heterozygous population. Acknowledgment We thank Malcolm M. Fumiss and Ronald W. Stark for their advice and encouragement during this study, Barbara Wilton for assistance with the labo¬ ratory work, and Morgan Stage, Zoran Antonijevic, and Calib Baldwin for help with the computer analyses. Critical reviews of the manuscript were provided by Jeffry Mitton, University of Colorado; Gene D. Amman, U.S. Forest Service, Ogden, Utah; the late Gerald N. Lanier, State University of New York, Syracuse; Daniel R. Miller, Simon Fraser University, British Columbia; and Malcolm Fur- niss, University of Idaho. Literature Cited Amman, G. D. & M. W. Stock, (in press). The effect of phloem thickness on heterozygosity in laboratory-reared mountain pine beetles (Dendroctonus ponderosae Hopkins, COLEOPTERA: SCOLYTIDAE). USDA Forest Service, Intermountain Research Station Research Paper. Beardmore, J. A. 1983. Extinction, survival, and genetic variation, pp. 125-151. In Schonewald- Cox, C. M., S. M. Chambers, B. MacBryde & W. L. Thomas (eds.). 1983. Genetics and conservation. The Benjamin/Cummings Publ. Co., Inc., Menlo Park, California. Bentz, B. J. & M. W. Stock. 1986. Phenetic and phylogenetic relationships among ten species of Dendroctonus bark beetles (Coleoptera: Scolytidae). Ann. Entomol. Soc. Amer., 79: 527-534. Bryant, E. H. 1974. On the adaptive significance of enzyme polymorphisms in relation to environ¬ mental variability. Am. Nat., 108: 1-19. Bryant, E. H. 1976. A comment on the role of environmental variation in maintaining polymorphisms in natural populations. Evolution, 30: 188-189. 266 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Cane, J. H., M. W. Stock, D. L. Wood & S. J. Gast. 1990. Phylogenetic relationships of Ips bark beetles (Coleoptera: Scolytidae): electrophoretic and morphometric analyses of Hopping’s Group IX. Biochem. Syst. Ecol, 18: 359-368. Ferguson, A. 1980. Biochemical systematics and evolution. John Wiley and Sons, New York. Garton, D. W., R. K. Koehn & T. M. Scott. 1984. Multiple-locus heterozygosity and the physiological energetics of growth in the coot clam, Mulinia lateralis, from a natural population. Genetics, 108: 445—455. Higby, P. K. & M. W. Stock. 1982. Genetic relationships between two sibling species of bark beetle (Coleoptera: Scolytidae), Jeffrey pine beetle and mountain pine beetle, in northern California. Ann. Entomol. Soc. Amer., 75: 668-674. Livingston, R. L. 1979. The pine engraver, Ips pini (Say), in Idaho: life history, habits and man¬ agement recommendations. Idaho Dept. Lands, Coeur d’Alene, Idaho. Rept. 79-3. Milkman, R. 1978. Selection differentials and selection coefficients. Genetics, 88: 391^103. Mitton, J. B. & M. C. Grant. 1980. Observations on the ecology and evolution of quaking aspen, Populus tremuloides, in the Colorado Front Range. Amer. J. Bot., 67: 202-209. Mitton, J. B. & M. C. Grant. 1984. Associations among protein heterozygosity, growth rate, and developmental homeostasis. A nn . Rev. Ecol. Syst., 15: 479-499. Mitton, J. B. & R. K. Koehn. 1975. Genetic organization and adaptive response of allozymes to ecological variables in Fundulus heteroclitus. Genetics, 79: 97-111. Nei, M. 1975. Genetic variability in natural populations, pp. 127-174. In Molecular population genetics and evolution. North Holland Research Monographs, Frontiers of Biology 40, Amer¬ ican Elsevier Publ. Co., Inc., New York. Parsons, P. A. 1971. Extreme-environment heterosis and genetic loads. Heredity, 26: 479-483. Parsons, P. A. 1987. Evolutionary rates under environmental stress. Evol. Biol., 21: 311-347. Samollow, P. B. & M. E. Soule. 1983. A case of stress related heterozygote superiority in nature. Evolution, 37: 646-649. SAS Institute. 1988. SAS/STAT user’s guide. SAS Institute Inc., Gary, North Carolina. Selander, R. K. 1976. Genic variation in natural populations, pp. 21-45. In F. J. Ayala (ed.). 1976. Molecular Evolution, Sinauer Associates, Sunderland, Massachusetts. Smith, M. H., C. T. Garten & P. E. Ramsay. 1975. Genetic heterozygosity and population dynamics in small mammals, pp. 85-102. In Markert, C. L. (ed.). Isozymes, genetics, and evolution. Academic Press, New York. Soule, M. E. 1980. Threshold for survival: maintaining fitness and evolutionary potential, pp. 151- 169. In Soule, M. E. & B. A. Wilcox (eds.). Conservation biology: an evolutionary-ecological perspective. Sinauer Associates, Sunderland. Stock, M. W. 1984. Genetic variation among mountain pine beetle sub-populations along an endemic to epidemic gradient across the north slope of the Uinta Mountains in Utah. Research Report submitted to USDA Forest Service, Intermountain For. and Range Expt. Sta., Ogden, Utah. Stock, M. W. & G. D. Amman. 1983. Host effects on the genetic structure of mountain pine beetle, Dendroctonus ponderosa, populations, pp. 83-95. In Safranyik, L. (ed.). The role of the host in the population dynamics of forest insects. Banff, Alberta, Canada. Swofford, D. L. & R. B. Selander. 1981. BIOSYS-1, a computer program for the analysis of allelic variation in genetics. User’s manual. Dept, of Genetics and Development, University of Illinois, Urbana. PAN-PACIFIC ENTOMOLOGIST 70(4): 267-268, (1994) A KEY TO THE XYLEBORUS OF CALIFORNIA, WITH FAUNAL COMMENTS (COLEOPTERA: SCOLYTIDAE) K. R. Hobson 1 and D. E. Bright 2 Department of Entomological Sciences, University of California, Berkeley, California 94720; 2 CLBRR, Canada Agriculture, Ottawa, Ontario K1A 0C6 Canada Abstract.— A key to the Xyleborus species of California is presented. The collection and estab¬ lishment of two very rare species, Xyleborus californicus and X. xylographus is reported from the central Sierra Nevada. Key words.—Xyleborus, Scolytidae, California, ambrosia beetle During investigations into the insect associates of Dendroctonus valens LeConte, a few specimens of two very rare species of Xyleborus were collected. Insects were sampled during the summer months of 1986-1990 at the University of California’s Blodgett Forest Research station near Georgetown, California in El Dorado Coun¬ ty. The research forest is between 1200 and 1400 m in a mixed conifer association dominated by ponderosa pine [Pinus ponderosa Lawson], sugar pine [J°. lamber- tiana Douglas], incense cedar [Calocedrus decurrens (Torrey) Florin], Douglas-fir [Pseudotsuga menziesii (Mirbel) Franco], white fir [Abies concolor (Gordon & Glendinning) Lindley], California black oak [Quercus helloggii Newberry] and several other broad-leaved tree species. The specimens were collected in Lindgren flight traps (Lindgren 1983) baited with turpentine, a distillate of ponderosa pine resin or blank controls. One specimen of Xyleborus californicus Wood was collected by KRH on 6 May 1990 in a trap baited with myrcene. This species was previously known only from a series of six specimens from Stanford University (collected 1944), one specimen from Yolo Co., California (collected 1949), and one specimen from Marion Co., Oregon. The Stanford University specimens were known to DEB for many years, but were considered to be intercepted or introduced but not established. It was, therefore, omitted from Bright & Stark (1973). Wood (1975) described and named the species but remarked that it was probably already named and was undoubtedly introduced into California, probably from either South America or Southeast Asia. Because of the lack of taxonomic information about the species of Xyleborus and the huge number of names available, Wood was unable to obtain a name for the species. The collection by KRH confirms the establishment of this species in California, if indeed it is introduced, or indicates that the species may be an extremely rare, endemic native species. The host of this species is unknown. Two specimens of Xyleborus xylographus Say were collected by KRH in a trap baited with turpentine on 28-31 May 1986 and in a trap baited with a mixture of a and 0-pinene on 6 May 1990. This species is known from Minnesota, Ontario, 1 Current address: Department of Forest Resources, Utah State University, and Forest Science Lab, USFS Intermountain Experiment Station, Logan, Utah 84322. 268 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) and Quebec south to Texas and Florida. One specimen from China Flat, California is noted by Wood (1982) with the notation “introduction or labeling error?” That specimen was also known to DEB in the early 1960s but was considered an accidental occurrence in California and, therefore, omitted from Bright & Stark (1973). The recent collection of two specimens by KRH confirms the occurrence of this species in California. This species is recorded in the east from Quercus spp. and we assume that this is the host in California. Because both species mentioned above were omitted in Bright & Stark (1973), a new key is presented below to aid in the recognition of the species. California Species of the Tribe Xyleborini 1. Scutellum conical, not filling scutellar notch, adjacent sides of scutellar notch pubescent; lower margin of declivity, beginning about inter¬ space 7 bearing a series of pointed tubercules, the 2 at the apex of interspace 2 large, more prominent ... Xyleborinus saxeseni (Ratzeburg) Scutellum flat, filling scutellar notch, pubescence in scutellar notch absent; lower margin of declivity smooth . 2 2( 1 b). Pronotum wider than long, coarsely asperate in front; body stout, about 2.2-2.4 x longer than wide . Xyleborus dispar (Fabricius) Pronotum longer than wide, finely asperate in front; body elongate, 2.8-3.0 x longer than wide . 3 3 (2b). Declivity steep, somewhat flattened, its surface dull, minutely reticu¬ late, granules on interspace 1 and 3 minute . . Xyleborus xylographus Say Declivity broadly sloping, convex, its surface shining, granules on in¬ terspaces 1 and 3 minute or coarse . 4 4(3b). Interstrial pubescence on elytra abundant, randomly placed, long; in¬ terior of strial punctures on declivity reticulate; granules on declivital interspaces 1 and 3 minute; length 2.0-2.2 mm. . Xyleborus calif or nicus Wood Interstrial pubescence on elytra sparse, arranged in uniseriate even row; interior of strial punctures on declivity shining; granules on declivital interspaces 1 and 3 distinct; length 2.2-2.7 mm . . Xyleborus intrusus Blandford Material Examined.—Xyleborus californicus : CALIFORNIA. EL DORADO Co.: Blodgett Forest, 6 May 1990, K. R. Hobson, flight trap with myrcene, 1 female. Xyleborus xylographus: CALIFORNIA. EL DORADO Co.: Blodgett Forest, 6 May 1990, K. R. Hobson, flight trap with a and /3-pinene, 1 female; same loc. 28-31 May 1986, K. R. Hobson, flight trap with turpentine, 1 female. Literature Cited Bright, D. E. &R. W. Stark. 1973. The bark and ambrosia beetles of California (Coleoptera: Scolytidae and Platypodidae). California Insect Survey Bull., 16. Lindgren, B. S. 1983. A multiple funnel trap for scolytid beetles (Coleoptera). Canad. Entomol., 115: 299-302. Wood, S. L. 1975. New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part II. Great Basin Nat., 35: 391—401. Wood, S. L. 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Nat., Memoirs, 6. PAN-PACIFIC ENTOMOLOGIST 70(4): 269-275, (1994) TEMPERATURE STUDIES ON A CHINESE STRAIN OF BACTROCERA CUCURBITAE (COQUILLETT) (DIPTERA: TEPHRITIDAE) P. Yang, 15 C. Zhou, 1 G. Liang, 2 Robert V. Dowell, 3 and James R. Carey 4 6 Research Institute of Entomology, Zhongshan University, Guangzhou, People’s Republic of China 2 Guangzhou Animal and Quarantine Service, People’s Republic of China California Department of Food and Agriculture, Sacramento, California 95814 4 University of California, Davis, California 95616 Abstract. — We examined preadult and adult survival, development and fecundity of Bactrocera cucurbitae (Coquillett) from China at six constant temperatures between 19° and 36° C. Devel¬ opment of immature stages and ovaries was inversely related to temperature. Preadult mortality was greatest at 36° C. Average female longevity was inversely related to temperature but male longevity was not. The intrinsic rate of increase was greatest at 25° C. No eggs or larvae survived exposure to constant temperatures of 2° to 3° C for longer than seven days. Key Words.— Insecta, Bactrocera cucurbitae, commodity treatment, demographic parameters Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae) is a cucurbit pest found in Kenya, Mauritius, Sri Lanka, India, China, Malaya, Indonesia, and the Phil¬ ippines (White & Elson-Harris 1992). In the last century, it has expanded its range to a number of Pacific islands including Hawaii. It has been detected in California on two occasions. Each of these California infestations, along with those on several Pacific islands, have been eradicated (Dowell & Gill 1989; Shiga 1989; RVD, unpublished data). Bactrocera cucurbitae is one of five species of economically important fruit flies found in mainland China, and one of two species attacking cucurbits; the other is Bactrocera tau (Walker). Recent efforts by the Chinese to expand agricultural exports have increased the importance of developing information about the bi¬ onomics of B. cucurbitae in China, because the fly is quarantined by a number of countries including the United States (Yang et al. in press). Temperature is an important environmental factor influencing fruit fly popu¬ lation dynamics. Data on the effects of temperature on fruit fly development and survival are critical: to developing models that predict fly phenology, to estimate the age structure of field populations, to the timing of control activities, and to developing quarantine compliance protocols (Smith 1977, Carey 1993). We stud¬ ied the effect of temperature on the development, survival and reproduction of the immature and adult stages of a Chinese strain of B. cucurbitae. We also evaluated the survival of B. cucurbitae eggs and larvae when they were subjected to cold treatments similar to those used to meet USD A quarantine regulations for Ceratitis capitata (Wiedemann) (Fiskaali 1991). 5 Current address: 2729 Kapiolani Blvd. #203, Honolulu, Hawaii 96823. 6 To whom correspondence should be sent. 270 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Table 1. Average developmental time and survival of immature B. cucurbitae reared at six constant temperatures. Temperature (° C) Stage 19= 22° 25° 28° 30° 36° Developmental time ab Egg 3.00 2.00 2.00 1.50 1.00 1.00 (1.1) (0.5) (0.7) (0.4) (0.1) (0.1) Larva 7.31 6.42 5.56 3.41 3.26 4.42 (1.4) (1.7) (2.5) (0.9) (0.9) (1.5) Pupa 14.80 14.30 10.20 9.00 7.90 6.50 (0.5) (0.5) (1.5) (0.6) (0.2) (0.8) Percent mortality Egg 5 4 4 5 5 12 Larva 15 16 19 9 14 7 Pupa 3 1 5 8 9 22 Total 23 21 28 22 28 41 a Days, mean ± (SD). b Three replicates, n = 30 eggs and 50 larvae or pupae per replicate. Materials and Methods Effect of Temperature on Growth and Survival. —Bactrocera cucurbitae were collected from the Parcel Islands, in the South China Sea, and maintained in a colony for several generations at the Guangzhou Animal and Plant Quarantine Service prior to use. Tests were conducted at 12:12 L:D cycle and 80% to 90% RH. Immature stages and adults were held at test temperatures (± 0.5° C) of 19°, 22°, 25°, 28°, 30°, or 36° C. All trials were replicated three times. Duration and survival of immature stages were determined as follows. Thirty newly-laid eggs were placed on a piece of wet black cloth in a petri dish and checked for hatch every eight hours. Fifty neonate larvae were placed on pieces Table 2. Adult longevity and reproduction of B. cucurbitae held at six constant temperatures. Temperature (° C) 19° 22° 25° 28° 30° 36° Longevity a ' bc Female 103.0 100.8 97.8 72.2 69.0 31.7 (45.7) (62.1) (45.3) (51.9) (49.5) (21.5) Male 95.9 107.8 111.4 80.9 71.1 29.7 (50.8) (66.2) 50.8) (51.5) (48.9) (16.0) Reproduction Preoviposition period 33.0 19.0 16.0 12.0 11.0 9.0 Gross fecundity bd 317.9 644.5 509.8 452.0 468.2 434.7 Net fecundity bd 171.8 317.6 249.7 201.9 191.2 86.2 Eggs/day/female 1.7 3.2 2.6 2.9 2.8 2.7 a Average number of days. b Three replicates with 50 pairs of flies per replicate. 0 Mean ± (SD). d Eggs per female. FECUNDITY 1994 YANG ET AL.: TEMPERATURE EFFECTS ON BACTROCERA 271 Figure 1. Daily gross egg production for B. cucurbitae female reared at six constant temperatures (19°, 22°, 25°, 28°, 30°, 36° C), given at left top of each graph. of cucumber ( Cucumis sativus L.) held in glass bottles (10 cm dia x 10 cm high) containing a layer of moist sand. The larvae were checked daily and food was added as needed. The sand was sifted daily to recover pupae. Fifty newly formed pupae were held in petri dishes and checked daily for emergence. Adult life history traits were determined by placing 50 pairs of newly emerged adults in cages 25 cm on a side. Water was provided and fresh orange juice was used as adult food. A small piece of cucumber was placed daily in each cage for egg collection. Mortality was recorded daily until the last female died. Life history data were analyzed using the methods of Carey (1993). Effect of Cold Temperatures on Egg and Larval Survival.— The survival of B. cucurbitae larvae held in four host plants at low temperatures was determined by placing newly molted second instar larvae in bottles (10 cm dia x 10 cm high) with cut pieces of each plant (Table 4). The bottles were then held at 2° C for one to five days to simulate quarantine treatment conditions. Each day one-fifth of the bottles were removed and the number of living and dead larvae was deter¬ mined. All subsequent tests were run using cucumber as larvae feeding on it took the longest time to reach 100% mortality. Another series of tests was run as above holding third instars at 2° C, and eggs, first and third instars at 3° C for eight to ten days. Results Preadult Development and Survival.— Developmental times for B. cucurbitae eggs and larvae were inversely related to temperature from 19° to 30° C (eggs: r 272 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Table 3. Demographic parameters for B. cucurbitae reared at six constant temperatures. Temperature (° C) Parameter 19° 22° 25° 28° 30° 36° Intrinsic rate of increase a 0.044 0.065 0.069 0.064 0.066 0.053 Mean generation time b 95.7 65.2 65.2 65.4 62.8 51.8 Doubling time b 16.9 10.7 10.1 10.9 4.7 5.7 a Per day. b Days. = -0.92, F = 20.49, P = 0.01, n = 5; larvae: r = -0.98, F = 71.23, P = 0.004, n = 4). The duration of the egg stage did not change at 36° C, but that of the larval stage increased. The duration of the pupal stage and total preadult devel¬ opment time were inversely related to temperature at all test temperatures (pupae: r — -0.95, F = 38.85, P = 0.003, n = 5; preadult: r = —0.95, F = 35, P = 0.004, n = 5) (Table 1). Egg mortality was relatively uniform between 19° and 30° C, but it increased Table 4. Mortality of second instar B. cucurbitae larvae reared at 2° C in four hosts. Host/days exposed Number alive Number dead Total Percent mortality Sponge Gourd 3 1 13 40 53 75.4 2 3 8 11 72.7 3 1 15 16 93.8 4 0 21 21 100.0 5 0 15 15 100.0 Balsam Pear b 1 9 39 48 81.3 2 7 35 42 83.3 3 1 88 89 98.9 4 0 54 54 100.0 5 0 26 26 100.0 Cucumber c 1 14 7 21 33.3 2 8 58 66 87.9 3 4 80 84 95.2 4 1 19 20 95.0 5 0 15 15 100.0 Wax Gourd d 1 49 4 53 7.6 2 5 18 23 78.3 3 0 27 27 100.0 4 0 43 43 100.0 5 0 21 21 100.0 a Luffa aegyptiaca Miller. b Momordica charantia L. c Cucumis sativus L. d Benincasa hisida (Thunberg). 1994 YANG ET AL.: TEMPERATURE EFFECTS ON BACTROCERA 273 Table 5. Mortality of immature stages of B. cucurbitae in cucumber held at cold temperatures. Days Eggs* Percent mortality 1st instar* 2nd instar* 3rd instar b 1 21.7 14.0 12.4 13.3 2 14.6 43.8 11.1 64.7 3 40.2 41.0 100.0 100.0 4 81.8 35.2 40.3 100.0 5 94.7 79.1 94.4 100.0 6 97.5 90.1 96.9 86.7 7 100.0 100.0 100.0 100.0 8 100.0 100.0 100.0 100.0 9 100.0 10 100.0 a Test run at 2° C, n = 15 to 20 larvae per temperature per day. b Test run at 3° C, n = 15 to 20 larvae per temperature per day. 2.4 fold at 36° C. Larval mortality was lowest at 28° and 36° C and varied little among the other test temperatures. Pupal mortality increased 4.4 fold between 25° and 36° C. Preadult mortality was greatest at 36° C (Table 1). Adult Survival and Reproduction.— Survival of B. cucurbitae females was in¬ versely related to temperature (r = —0.96, F = 48.41, P = 0.002, n = 5), but that of the males increased with temperature between 19° and 25° C and decreased with increasing temperature thereafter. The preovipositional period was inversely related to temperature (r = —0.87. F = 12.65, P = 0.02, n = 5). Gross and net fecundity, and eggs per female per day were greatest at 22° C (Table 2) and were not related to temperature ( r — 0.02, r = 0.64, r = 0.40 respectively, P > 0.05). Daily egg production fluctuated widely, with no clear trend regardless of rearing temperature. Females continued to lay eggs for at least 140 days at temperatures at or below of 30° C and for up to 180 days at 22° to 25° C (Fig. 1). The intrinsic rate of population increase was greatest at 25° C, but there was little difference among the values between 22° and 30° C. Mean generation time was shortest at 36° C and there was little difference among the values between 22° and 30° C. Population doubling time was shortest at 30° C, with nearly identical times between 22° and 28° C (Table 3). Effects of Cold Temperatures on Survival.— No second instar B. cucurbitae survived beyond four days when held at a constant 2° C in any of the test plants (Table 4). No third instars survived beyond six days when held at a constant 2° C and no eggs, first or second instars survived beyond six days when held at 3° C (Table 5). Increasing the temperature 1° C, from 2° to 3° C, increased the time needed to kill all second instars from four to six days (Tables 4 and 5). Discussion Our preadult developmental times and survivorships of B. cucurbitae fall within the range of those from previous studies of wild flies in culture six or fewer generations (Miyatake 1993). Egg and pupal development are mainly dependent upon temperature and larval development upon temperature and host (Tables 1 and 6). 274 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Table 6. Stage specific duration and survival of wild B. cucurbitae from previous studies. Stage" •c Duration b Host Reference Eggs 20 2.0 (73) Bhatia & Mahto 1970 25 1.1 (77) Bhatia & Mahto 1970 25 1.0 (74) Carey et al. 1985 Larvae 20 6.7 (85) pumpkin Bhatia & Mahto 1970 25 3.7 (85) pumpkin Bhatia & Mahto 1970 27.5 3.4 (83) pumpkin Bhatia & Mahto 1970 25 4.1 (88) cucumber Carey et al. 1985 25 7.4 (38) eggplant Carey et al. 1985 24 9.8 (90) papaya Vargas & Carey 1990 25 9.0 (na) media Miyatake 1993 Pupae 20 15.1 (92) pumpkin Bhatia & Mahto 1970 25 7.8 (91) pumpkin Bhatia & Mahto 1970 28 6.8 (93) pumpkin Bhatia & Mahto 1970 25 13.0 (89) cucumber Carey et al. 1985 24 9.8 (61) papaya Vargas & Carey 1990 25 11.0 (na) media Miyatake 1993 Preoviposition 25 16.0 cucumber Carey et al. 1985 23.8 14.8 tomato Keck 1951 26.7 13.5 tomato Keck 1951 25 14.0 unknown Back Pemberton 1918 a Considered wild if in colony six or fewer generations (Miyatake 1993), na = not available. b Percent survival in parentheses. The adult reproductive parameters, however, differ considerably among the studies. The gross fecundity of wild B. cucurbitae from China is approximately half that of wild B. cucurbitae from Hawaii. Wild Chinese B. cucurbitae lay one- half to one-third the eggs per day and have population doubling times 1.5 times greater than those from Hawaii. The greater variation in adult responses suggests that this is the stage in which local environmental factors have their greatest influence and, thus, the stage in which the fly adapts to them (Tables 2, 3 and 7). In culture, the response time to selection for a characteristic of adult flies was faster than that for larvae (Miyatake 1993). Although not definitive, our results suggest that cold treatments may be effective as a disinfestation treatment for produce harboring B. cucurbitae eggs and larvae. Table 7. Adult demographic parameters for wild B. cucurbitae from previous studies. Parameter Value" Reference Gross fecundity 1293 eggs Carey et al. 1985 Net fecundity 709 eggs Carey et al. 1985 Doubling time 6.9 days Carey et al. 1985 12.0 days Vargas & Carey 1990 Eggs/female/day 7.2 eggs Carey et al. 1985 4.7 eggs (21.1° C) Keck 1951 8.9 eggs (23.9° C) Keck 1951 8.2 eggs (29.4° C) Keck 1951 a Considered wild if in colony for six or fewer generations (Miyatake 1993). 1994 YANG ET AL.: TEMPERATURE EFFECTS ON BACTROCERA 275 Further, large scale tests will be required before cold treatments of B. cucurbitae hosts can be certified for use as a quarantine treatment from countries having the pest. Literature Cited Back, E. A. & C. E. Pemberton. 1918. The melon fly. USDA Bull., 643. Bhatia, S. K. & Y. Mahto. 1970. Influence of temperature on the speed of development of melon- fly, Dacus cucurbitae Coquillett (Diptera: Tephritidae). Indian J. Agric. Sci., 40: 821-828. Carey, J. R. 1993. Applied demography for biologists with special emphasis on insects. Oxford Univ. Press., New York. Carey, J. R., E. J. Harris & D. O. Mclnnis. 1985. Demography of a native strain of the melon fly, Dacus cucurbitae, from Hawaii. Ent. Exp. Appl., 38: 195-199. Dowell, R. Y. & R. Gill. 1989. Exotic invertebrates and their effects on California. Pan-Pacif. Entomol., 65: 132-145. Fiskaali, D. A. 1991. Commodity treatment manual, Vol. I. Treatments. Calif. Dept. Food & Agric., Sacramento, California. Keck, C. B. 1951. Effect of temperature on development and activity of the melon fly. J. Econ. Entomol., 44: 1001-1002. Miyatake, T. 1993. Difference in the larval and pupal periods between mass-reared and wild strains of the melon fly, Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae). Appl. Entomol. Zool., 28: 577-581. Nakamori, H. 1987. Variation of reproductive characters in wild and mass-reared melon flies, Dacus cucurbitae Coquillett (Diptera: Tephritidae). Jpn. J. Appl. Entomol. Zool., 31: 309-314. Shiga, M. 1989. Current programme in Japan. Chap. 9.5.2. In Robinson, A. S„ & G. Hooper (eds.). Fruit flies, their biology, natural enemies and control. Elsevier, New York. Smith, E. S. C. 1977. Studies on the biology and commodity control of the banana fruit fly Dacus musae (Tryon), in Papau New Guinea. Papau New Guinea Agric. J., 28: 47-56. Vargas, R. I. & J. R. Carey. 1990. Comparative survival and demographic statistics for wild oriental fruit fly, Mediterranean fruit fly, and melon fly (Diptera: Tephritidae) on papaya. J. Econ. Entomol., 83: 1344-1349. White, I. M. & M. M. Elson-Harris. 1992. Fruit flies of economic significance: their identification and bionomics. C.A.B. Int., London. Yang, P., J. R. Carey & R. V. Dowel. 1994. Tephritid fruit flies in China: historical perspective and current status. Pan-Pacif. Entomol., 70: 159-167. PAN-PACIFIC ENTOMOLOGIST 70(4): 276-282, (1994) REPRODUCTIVE BEHAVIOR OF COPITARSIA CONSUETA (WALKER) (LEPIDOPTERA: NOCTUIDAE): MATING FREQUENCY, EFFECT OF AGE ON MATING, AND INFLUENCE OF DELAYED MATING ON FECUNDITY AND EGG FERTILITY Julio C. Rojas 12 and Juan Cibrian-Tovar Laboratorio de Ecologia Quimica, Centro de Entomologia y Acarologia, Colegio de Postgraduados, Chapingo, Edo de Mexico, C.P. 56230, Mexico Abstract. — This study determined: mating frequency, effect of age on mating, and effect of delayed mating on fecundity and egg fertility of Copitarsia consueta (Walker). Females copulated (mean ± SD) 2.5 ± 1.2 times (range 1-6), but only once per night. Mating frequency increased with moth age. Males mated an average of 2.4 ± 1.5 times during their lives (range 1-7), although only once per night. Copitarsia consueta start mating two days after emergence, although only 33% of the pairs mated this early. Mating frequency increased with moth age, reaching a peak 7 days after eclosion. Females mated when 2 days old laid significantly more eggs than those in which mating was delayed till 4, 6, and 8 days after eclosion. Fertility was 70.2% in females mating when 2 days old. It decreased to 50.9, 6.5, and 0.4% when mating was delayed until 4, 6, and 8 days after eclosion, respectively. Key Words.— Insecta, Copitarsia consueta, mating, fecundity, fertility The moth, Copitarsia consueta (Walker), is a polyphagous pest found in Mexico and Central and South America (Angulo & Weigert 1975, Gutierrez & MacGregor 1983) where it is a important pest of cultivated plants. In some regions of Mexico it is a key pest of cabbage, but it also attacks many other plants (Gutierrez & MacGregor 1983, Guevara & Cervantes 1991). The quality and quantity of cab¬ bage are greatly affected by C. consueta as only one larvae is sufficient to destroy the plant (Monge et al. 1984). Information on biology of C. consueta is scarce (Artigas & Angulo 1973), and its reproductive behavior is unknown. This study determined: mating frequency, effect of age on mating, and effect of delayed mating on fecundity and egg fertility of C. consueta. Materials and Methods Insects.— Moths used in this study were reared on an artificial diet (Cibrian- Tovar & Sugimoto 1992) at 25 ± 2° C and 65 ± 5% RH. Male and female pupae were held separately in growth chambers under 14:10 (L:D) photoperiod. The night after emergence, the insects were placed in glass cages (20 x 20 cm) with 10% sugar solution dispensed from shell vials plugged with cotton, and maintained at the conditions described above. Female Mating Frequency. — Newly emerged females and an equal, or greater, number of males were confined in a glass cage (20 x 20 cm) and provided a solution of 10% sugar in water. After dying, 100 females were dissected and the 1 Laboratorio de Ecologia Quimica, Centro de Investigaciones Ecologicas del Sureste, Carretera Antiguo Aeropuerto Km 2.5, Tapachula, C.P. 30700. Mexico. 2 To whom correspondence should be addressed. 1994 ROJAS & CIBRIAN-TOVAR: COPITARSIA REPRODUCTION 277 number of spermatophores in the bursa copulatrix counted to determine the number of times the female had mated. To determine the effect of age on mating frequency, groups of females (n = 15) were placed with 3-5 d old males in plastic containers (4x4x7 cm). Females were held for 2 to 7 d before being removed from the cage, killed and dissected to determine the number of spermatophores in the bursa copulatrix. The maximum number of matings per night was determined by holding a female (2-3 d old) in a plastic container with 2 to 3 virgin males for one night. The female was killed, dissected, and the number of spermatophores counted. The experiment was repeated 20 times. Male Mating Frequency. —Aid old virgin male and a 2-3 d old female were placed in a plastic container. Every 2 d the female was replaced with another 2- 3 d old until the male died. The experiment was repeated 15 times. The females were dissected and the number of spermatophores present was counted. The number of times a male could copulate per night was determined by placing a 2-3 d old male with 2 or 3 virgin females. Females were killed the next day, dissected and the number of spermatophores counted. The experiment was re¬ peated 20 times. Effect of Age on Mating. — Virgin males and females were placed individually in plastic containers. The experiments were conducted using insects of the fol¬ lowing age groups: 1) male and female of equal age, ranging from 1 to 7 d old; 2) 1 d old female with a 3-5 d old male; and 3) 1 d old male with a 3-5 d old female. After each scotophase 15 females were killed, dissected, and the number of sper¬ matophores counted. Effect of Delayed Mating on Fecundity and Egg Fertility. — Five treatments were run: 1) 2 d old females with 3-5 d old males; 2) 4 d old females with 3-5 d old males; 3) 6 d old females with 3-5 d old males; 4) 8 d old females with 3-5 d old males; and 5) unmated females. When the male died it was replaced with another 3-5 d old male. All treatments were repeated 14 times. For each group, preovi- position period (days), oviposition period (days), fecundity (eggs/female), and fertility (%) were recorded. Dead females were dissected and the number of sper¬ matophores determined. Statistical Analysis.— The data were analyzed by ANOVA and the Student- Neumann-Kuels test. The preoviposition, oviposition, and fecundity data were square-root transformed and fertility data were square root arcsine transformed before analysis. The percentage of mating were analyzed using the X 2 test. Un¬ transformed data are presented. Results and Discussion Female Mating Frequency.— Females had an average of (mean ± SD) 2.5 ± 1.2 spermatophores (range 1-6). Twenty-three females had 1 spermatophore, 28 had 2 spermatophores, 25 females had 3 spermatophores, 17 females had 4 sper¬ matophores, 5 females had 5 spermatophores, and only 2 females received 6 spermatophores. Similar results were observed in Pseudoplusia includens (Walker), with an av¬ erage of 2.2 ± 0.2 spermatophores per female (ranging 0-5) (Mason & Johnson 1987). By comparison, females of Chilo partellus (Swinhoe) copulated only once (Unnithan & Paye 1991). There are two mating systems in Lepidoptera, those 278 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) that copulate only once, and those that have multiple matings. The benefits of multiple matings differ among species. Watanabe (1988) showed that multiple matings increased the fecundity of Papilio xuthus L., but fertility was not increased in Orgyia pseudotsugata (McDonaough) despite females copulating several times (Swaby et al. 1987). The most probable benefits of multiple matings are to remedy an inadequate initial copula, to enhance genetic diversity, and to facilitate a paternal nutritional investment (Byers 1978). Traditionally, the role of multiple matings in Lepidoptera has been discussed in terms of sexual selection (Thornhill & Alcock 1983). The C. consueta females only mated once per night, although they copulated repeatedly during their lives. Spodoptera littoralis (Boisduval) females (84%) mat¬ ed only once per night, but the rest (16%) mated twice a night (Kehat & Gordon 1975). Older females mated more frequently (Fig. 1). Landolt & Curtis (1991) had similar results with field and laboratory populations of Amyelois transitella (Walk¬ er). We do not know if the mating frequency of C. consueta in the wild is equal or different from that observed in the laboratory. It could argue that the mating frequency in the field is less because under natural conditions the insects are distributed over a great area. Euxoa ochrogaster (Guenee), Euxoa declarata (Walk¬ er) and Euxoa messoria (Harris) had similar mating frequencies in field and laboratory populations (Byers 1978). Long-term laboratory strains of Heliothis virescens (F.) and Pectinophora gossypiella (Saunders) have a higher mating fre¬ quency than recently colonized strains (Proshold & Bartell 1972, LaChance et al. 1975). This study used recently colonized insects (F 1 ), therefore, it is likely that 1994 ROJAS & CIBRIAN-TOVAR: COPITARSIA REPRODUCTION 279 Figure 2. Mating frequency in C. consueta males. mating frequency observed in the laboratory will be similar to that under natural conditions, as suggested for Euxoa spp. (Byers 1978). Male Mating Frequency. — Males copulated an average of 2.4 ± 1.5 times during their life span (range 1-7) (Fig. 2). Similar trends were observed in other noctuid moths: S. littoralis males mated an average of 5-6 times (Kehat & Gordon 1975), Spodoptera frugiperda (J. E. Smith) males mated an average of 6.7 times (range 0-15) (Simmons & Marti 1992), and C. partellus males mated a mean of 4.6 ± 0.4 times (Unnithan & Paye 1991). The males of C. consueta mated only once per night. Similar results were reported for S. littoralis (Kehat & Gordon 1975) and Spodoptera exempta (Walker) (Khasimuddin 1978), whereas, in C. partellus, 90% mated once and 6% mated twice (Unnithan & Paye 1991). Effect of Age on Mating. — Copitarsia consueta start mating two days after emer¬ gence, although only 33% of the pairs mated this day (Fig. 3). Mating frequency increased with the age reaching a peak 7 d after eclosion in this experiment. The differences in the percentage of mating pairs for 4, 5, 6 and 7 d are not significant (X 2 = 1.18; df = 3 , P > 0.05). The age of the female is important if mating is to be successful, as the pairs that had 1 d old females failed to mate. The pairs with 3-4 d old females mated 12.4% of the time. These results mirror the calling behavior of the females. No female called the day following emergence. Females called for the first time during the second or third scotophase after eclosion (JCR, unpublished data). Moths can be placed in two groups according to age at which mating, those that mate soon after eclosion, such as C. partellus (Unnithan & Paye 1991) and Spodoptera ornithogalli (Guenee) (Shorey et al. 1968); and those that need time 280 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Figure 3. Effect of age on the mating capacity of C. consueta. to reach sexual maturity like C. consueta. This group includes S. exempta (Khasi- muddin 1978) and Corcyra cephalonica (Stainton) (Etman et al. 1988). That only a few 1 d old males copulated with mature females could be due to: 1) that they are not sexually mature and thus do not respond to sex pheromone; as occurs in S. exempta (Khasimuddin 1978) and 2) that males are rejected because they do not provide the proper stimulus for mating, as has been suggested for Anticarsia gemmatalis (Hubner) males (Leppla et al. 1987). We do not know which mechanism occurs in C. consueta ; this will be investigated in the future. Effect of Delayed Mating and Fecundity and Egg Fertility. — The percentages of females mating after 2, 4, 6, and 8 d did not differ significantly (X 2 = 1.6; df = 3; P > 0.05). These results differ from those reported for Homona magnanima Diakonoff and P. gossypiella, in which the percentage of mated females decreased with an increase in the number of days elapsing before pairing (Kiritani & Kanoh 1984, Lingren et al. 1988). The preoviposition period increased significantly with delayed mating (F = 7.3; df = 3, 52; P < 0.05) (Table 1). Similar results were reported in C. partellus (Unnithan & Paye 1991). The ovipositional period differed significantly among treatments (F = 3.6; df = 3, 52; P < 0.05). The same phenomenon was reported for H. magnanima, P. gossypiella, and C. partellus (Kiritani & Kanoh 1984, Lingren et al. 1988, Un¬ nithan & Paye 1991). Females mating at 2 d old laid significantly more eggs than those in which mating was delayed till 4, 6, and 8 d (F = 17.7; df = 3, 52; P < 0.05). The effect of delayed mating on eggs fertility was significant (F = 36.7; df = 3, 52; P < 0.05). Fertility was 70.2% in females mating at 2 d old, and it decreased with age. The 1994 ROJAS & CIBRIAN-TOVAR: COPITARSIA REPRODUCTION 281 Table 1. Influence of delayed mating on reproductive biology of C. consueta. Age at mating (days) %of mating Preoviposition period (days) (X ± SE) Oviposition period (days) (X ± SE) Fecundity (eggs/female) (X ± SE) Fertility (%) (X ± SE) 2 86.6 5.0 ± 0.5 a 12.4 ± 1.3“ 1638.8 ± 199.7 a 70.2 ± 8.6 a 4 83.3 5.4 ± 0.4 a 7.0 ± 1.6 b 976.8 ± 248.l b 50.9 ± 11.6 b 6 78.5 6.3 ± 0.4 a 8.4 ± 1.5 b 283.5 ± 84.9 C 6.5 ± 6.4 C 8 78.5 10.6 ± 0.8 b 7.0 ± 1.3 b 220.0 ± 67.8 d 0.4 ± 0.2 d Unmated females — (14.2 ± 3.3) a (2.3 ± 0.7) a (131.5 ± 56.9) a — Means within columns followed by the same letters are not significantly different (Student-Newman- Keuls test, P < 0.05). a Data are not included in analysis. effect of delaying mating on fecundity and fertility in Lepidoptera is variable. Barrer (1976) found in Ephestia cautella (Walker) that when mating was delayed, the fecundity and fertility were reduced. Similar results were reported in S. lit- toralis, H. magnanima, P. gossypiella, and C. partellus (Ellis & Steele 1982, Kiritani & Kanoh 1984, Lingren et al. 1988, Unnithan & Paye 1991). However, in Erias insulana (Boisduval) (Kehat & Gordon 1977) and the Israeli strain of S. littoralis (Kehat & Gordon 1975) delays in mating do not have an effect on fecundity and fertility. Reduced fecundity of C. consueta females that experience a delay in mating could be due to the resorbing of eggs as has been suggested for E. cautella (Barrer 1976). The use of sex pheromones to delay mating, and hence provide control of C. consueta, requires a delay of 8 d between adult emergence and mating to prevent the laying of viable eggs. Delays of less than 8 d would lead to the production of progeny. Delaying of mating has been proposed as the principal mechanism lead¬ ing to successful mating disruption against P. gossypiella (Lingren et al. 1988). Acknowledgment JCR thanks CONACyT for a graduate fellowship (register No 62158). This research was supported by a CONACyT grant (0691-N9111). Literature Cited Angulo, O. A. & G. Th. Weigert. 1975. Estados inmaduros de Lepidopteros noctuidos de importancia economica en Chile y claves para su determination (Lepidoptera: Noctuidae). Soc. Biol. Con¬ cepcion, publication especial 2. Artigas, J. N. & A. O. Angulo. 1973. Copitarsia consueta (Walker), biologia e importancia economica en el cultivo de raps (Lepidoptera: Noctuidae). Bol. Soc. Biol. Concepcion, 46: 199-216. Barrer, P. M. 1976. The influence of delayed mating on the reproduction of Ephestia cautella (Walker) (Lepidoptera: Noctuidae). J. Stored Prod. Res., 12: 165-169. Byers, J. R. 1978. Biosystematics of the genus Euxoa (Lepidoptera: Noctuidae). X. Incidence and level of multiple mating in natural and laboratory populations. Can. Ent., 110: 193-200. Cibrian-Tovar, J. & A. Sugimoto. 1992. Elaboration de una dieta artificial para la cria de Copitarsia consueta (Walker) (Lepidoptera: Noctuidae). pp. 416. In Memorias del XXVII Congreso Na¬ tional de Entomologia. Sociedad Mexicana de Entomologia. Ellis, P. E. & G. Steele. 1982. Effects of delayed mating on fecundity of females of Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae). Bull. Entomol. Res., 72: 295-302. Etman, A. A. M., F. M. A. El-Sayed, N. M. Eesa & L. E. Moursy. 1988. Laboratory studies on the 282 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) development, survival, mating behavior and reproductive capacity of the rice moth, Corcyra cephalonica (Stainton). J. Appl. Ent., 106: 232-240. Guevara, A. R. & J. F. Cervantes. 1991. Insectos plaga de hortalizas en la zona chinampera de Xochimilco, D. F. pp. 528-529. In Memorias del XXVI Congreso Nacional de Entomologia. Sociedad Mexicana de Entomologia. Gutierrez, O. & R. MacGregor. 1983. Guia de insectos nocivos para la agricultura en Mexico. Alhambra Mexicana, Mexico, D.F. Kehat, M. & D. Gordon. 1975. Mating, longevity, fertility and fecundity of the cotton leaf-worm, Spodoptera littoralis (Boisd.) (Lepidoptera: Noctuidae). Phytoparasitica, 3: 87-102. Kehat, M. & D. Gordon. 1977. Mating ability, longevity and fecundity of the spiny bollworm, Earias insulana (Lepidoptera: Noctuidae). Entomol. Exp. Appl., 22: 267-273. Khasimuddin, S. 1978. Courtship and mating behavior of the African armyworm, Spodoptera ex- empta (Walker) (Lepidoptera: Noctuidae). Bull. Entomol. Res., 68: 195-202. Kiritani, K. & M. Kanoh. 1984. Influence of delayed mating on the reproduction of the oriental tea tortrix, Homona magnanima Diakonoff (Lepidoptera: Tortricidae), with reference to phero¬ mone-based control. Prot. Ecol., 6: 137-144. LaChance, L. E., R. D. Richard & F. I. Proshold. 1975. Radiation response in the pink bollworm: a comparative study of sperm bundle production, sperm transfer, and oviposition response elicited by native and laboratory-reared males. Environ. Entomol., 4: 321-324. Landolt, P. J. & C. E. Curtis. 1991. Mating frequency of female navel orangeworm moths (Lepi¬ doptera: Pyralidae) and patterns of oviposition with and without mating. J. Kansas Entomol. Soc, 64: 414-420. Leppla, N. C., R. H. Guy, R. R. Heath & B. Dueben. 1987. Laboratory studies of the courtship of the velvetbean caterpillar moth, Anticarsia gemmatalis (Hubner) (Lepidoptera: Noctuidae). Ann. Entomol. Soc. Am., 80: 278-283. Lingren, P. D., W. B. Warner & T. J. Henneberry. 1988. Influence of delayed mating, on egg production, egg viability, mating, and longevity of female pink bollworm (Lepidoptera: Gele- chiidae). Environ. Entomol., 17: 86-89. Mason, L. J. & S. J. Johnson. 1987. Observations on the mating behavior of Pseudoplusia includens (Lepidoptera: Noctuidae). Fla. Entomol., 70: 411-413. Monge, V. L., J. G. Vera, S. I. Gil & J. L. Carrillo. 1984. Efecto de las practicas culturales sobre las poblaciones de insectos y dano causado al cultivo del repollo ( Brassica oleraceae var. capitata). Agrociencia, 57: 109-126. Proshold, F. I. & J. A. Bartell. 1972. Difference in radiosensitivity of two colonies of tobacco budworm, Heliothis virescens (Lepidoptera: Noctuidae). Can. Ent., 104: 995-1002. Shorey, H. H., S. U. McFarland & L. K. Gaston. 1968. Sex pheromone of noctuid moths. XIII. Changes in pheromone quantity, as related to reproductive age and mating history, in females of seven species of Noctuidae (Lepidoptera). Ann. Entomol. Soc. Am., 61: 372-376. Simmons, A. M. & O. G. Marti Jr. 1992. Mating by the fall armyworm (Lepidoptera: Noctuidae): frequency, duration, and effect of temperature. Environ. Entomol., 21: 371-375. Swaby, J. A., G. E. Daterman & L. L. Sower. 1987. Mating behavior of douglas-fir tussock moth, Orgyia pseudotsugata (Lepidoptera: Lymantriidae), with special reference to effects of female age. Ann. Entomol. Soc. Am., 80: 47-50. Thornhill, R. & J. Alcock. 1983. The evolution of insects mating systems. Harvard University Press, Cambridge. Unnithan, G. C. & S. O. Paye. 1991. Mating, longevity, fecundity, and egg fertility of Chilo partellus (Lepidoptera: Pyralidae): effects of delayed or successive matings and their relevance to pher¬ omonal methods. Environ. Entomol., 20: 150-155. Watanabe, M. 1988. Multiple mating increase the fecundity of the yellow swallowtail butterfly, Papilio xuthus L., in summer generations. J. Insect. Behav., 1: 17-29. PAN-PACIFIC ENTOMOLOGIST 70(4): 283-300, (1994) TAXONOMIC REVIEW OF CALLIOPSIS SUBGENUS HYPOMACROTERA (HYMENOPTERA: ANDRENIDAE), WITH SPECIAL EMPHASIS ON THE DISTRIBUTIONS AND HOST PLANT ASSOCIATIONS Bryan N. Danforth Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853 Abstract. — Two names, previously used to designate subspecies of Calliopsis ( Hypomacrotera ) callops, are herein used to refer to distinct species: C. (H.) callops (Cockerell & Porter) and C. ( H .) persimilis (Cockerell). These two species are described and the data on floral host association and distributions are listed and illustrated. These two species, plus Calliopsis ( Hypomacrotera) subalpinus Cockerell, comprise the monophyletic subgenus Hypomacrotera. Floral association and distribution data from over 850 specimens are analyzed. Calliopsis callops and C. persimilis are oligolectic on a group of closely related genera in the family Solanaceae; the former on Chamaesaracha and Quincula and the latter on Physalis. Calliopsis subalpinus is clearly oli¬ golectic on mallows in the genus Sphaeralcea. Calliopsis persimilis and C. callops are parapatric with a narrow region of overlap in the San Simon Valley, near the Continental Divide in southern Arizona. Calliopsis subalpinus ranges widely across the southwestern deserts from southern California to southwestern Texas and southward to northern Mexico. Key Words. — Insecta, Apoidea, taxonomy, floral associations This paper establishes that the previously recognized subspecies of Calliopsis (. Hypomacrotera ) callops (Cockerell & Porter) are, in fact, two easily distinguish¬ able species. Rozen (1970) anticipated these taxonomic changes in a study on the nesting biology of C. callops. In addition to this minor taxonomic point, an account of the distributions and floral associations of the three valid species in the subgenus Hypomacrotera is given. Quantitative investigations into floral specialization are rare (except see Heit- haus [1979]), in part because in many groups of oligolectic, or pollen specialist, bees there are not enough specimens with associated floral data collected over large areas to provide sufficient data for such an analysis. Because the species within Hypomacrotera have been collected in the southwestern United States and northern Mexico during the last century by many different collectors, large num¬ bers of specimens with associated floral data are available in museum collections. As a result, Hypomacrotera makes an excellent case study in bee floral special¬ ization. Hypomacrotera was first named by Cockerell & Porter (1899) to include H. callops (the type species) and H. subalpinus (previously placed in Calliopsis ). For the purposes of this study, I accept the view that Hypomacrotera is a monophyletic group, as indicated by Ruz (1991). The monophyly of Hypomacrotera was sup¬ ported by her characters 52 (propodeal triangle smooth) and reversal to the ple- siomorphic state in character 71 (tarsomeres 2-4 of male hind leg expanded; they are not expanded in Hypomacrotera ). The presence of darkened areas at the apices of the forewings in males (and females in two of the three species) is another common character, but one not unique to Hypomacrotera within Calliopsis (Ruz 1991:232). 284 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) In the taxonomic descriptions given below, I have used the surface sculpturing terms explained in Harris (1979). Morphological terms follow Michener (1944) except the sternum and tergum of the first metasomal segment (homologous to the second abdominal segment) are called tergum 1 (abbreviated Tl) and sternum 1 (abbreviated SI), respectively. The following metasomal sclerites are numbered sequentially thereafter. Measurements are expressed as mean ± standard error of the mean. Depository Abbreviations.— The locations of specimens used in this study are indicated with the following abbreviations: American Museum of Natural History (AMNH), Snow Entomological Museum, University of Kansas (KU), Los Angeles County Museum of Natural History (LACM), University of California, Riverside (UCR), California Academy of Sciences (CAS), National Museum of Natural History, Smithsonian Institution (NMNH), Central Texas Melittological Institute (CTMI), and Universidad Nacional Autonoma de Mexico (UNAM). In Material Examined the locality data are listed hierarchically, and the numbers of females and males are indicated in brackets as follows: [number females, number males]. Calliopsis ( Hypomacrotera ) persimilis (Cockerell) Hypomacrotera callops persimilis Cockerell (1899:8) [male, female]; Calliopsis (Hypomacrotera) persimilis, Danforth (1990) [biology]. Types. — Cotypes, male; data: ARIZONA. MARICOPA Co.: Phoenix, 7 Oct [no year], Tribulus grandiflora, 1 male; deposited: California Academy of Sciences, San Francisco. Cotypes, females; data: same as male except collected 9 Oct on flowers of Physalis, unknown number; deposited: unknown. Although the male of the cotype series is clearly the basis for Cockerell’s account of this species, to the best of my knowledge he did not formally designate a holotype and I here designate this male the lectotype. Description.— Female.— Head: (1) width 1.88-2.00 mm (x = 1.91 ± 0.02; n = 10); (2) 1.35-1.48 (x = 1.41 ± 0.01; n = 10) x broader than long, as measured from vertex to lower margin of clypeus; (3) clypeus distinctly punctate with weak imbrication; (4) frons mostly shiny with scattered punctations, more imbricate above antennal sockets; (5) vertex shiny and nearly impunctate; (6) gena shiny and nearly impunctate; (7) head coloration dark brown to black, no maculation; (8) head lightly clothed in erect white setae, most dense and longest setae on gena, posterior surface of head and vertex; (9) inner margins of eyes diverging slightly below; eyes brown; (10) lateral ocelli separated from median ocellus by 1 ocellar diameter; (11) facial foveae weakly impressed, concave surface slightly dull; (12) scape equal in length to flagellar segments 1-6; flagellum lighter brown ventrally and apically. Mouth- parts: (13) labrum with proximal impunctate concave area separated from distal punctate area by ridge; (14) mandible dark brown basally becoming light brown apically; simple; (15) glossa short, two- thirds length of prementum; (16) paraglossae broad and blade-like; (17) labial palpus 4-segmented with segments 2-4 equal in length to segment 1; (18) galeal comb present; (19) maxillary palpus 6-segmented, with first segment longest. Mesosoma: (20) pronotum brown, imbricate-punctate with fine pilosity on dorsal and lateral surfaces; pronotal lobe with elongate, finely branched white setae; (21) mesoscutum shiny with widely scattered punctures dorsally, becoming more closely-spaced lat¬ erally; elongate, erect setae 0.20 mm long over most of the surface; notauli lacking; parapsidal lines weak; (22) mesoscutellum shiny at center becoming punctate around edges, erect setae as on meso¬ scutum; metanotum imbricate-punctate; (23) mesopleuron distinctly imbricate with weak punctations; erect, white setae; (24) metapleuron weakly imbricate, with fine pilosity, no long, erect setae; (25) propodeum imbricate laterally with short, fine pilosity; patch of erect setae of varying lengths on either side of entirely glabrous propodeal triangle; (26) intertegular distance 1.28-1.40 mm (5c = 1.34 ± 0.02; n = 10); (27) forewing length 3.80-4.20 mm (5c = 3.99 ± 0.03; n = 10); wings clear with brown wing veins and weak dark spot at apex of forewing; (28) legs brown except for white spot at base of foretibia; 1994 DANFORTH: REVIEW OF CALLIOPSIS (. HYPOMACROTERA) 285 Figure 1. Calliopsis persimilis. Male: (a) sixth stemite, (b) seventh stemite, (c) fifth stemite, (d) eighth stemite, (e) genital capsule, (f) seventh tergite. Female: (g) midleg. mesobasitarsus slender (length 2.5-2.9 x width; Fig. lg); (29) basitibial plate distinctly kidney-shaped, with setae lining concave surface; (30) scopal hairs simple, erect; (31) hindtibial spurs serrate, inner longer than outer; (33) tarsal claws bifid. Metasoma: (34) terga dark brown; (35) T1 shiny, impunctate; T2-T4 minutely imbricate-punctate with small, posteriorly directed recumbent, brown setae; (36) weakly developed fovea on lateral edge of T2; (37) lateral angles of T3, T4 and all of T5 with elongate, 286 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) erect, finely-branched setae, T5 distinctly punctate; (38) pygidial plate rounded apically, surface convex and clothed in setae; (39) sterna similar in color and sculpturing to T2-T4; (40) S2 to S5 with graduli; (41) S6 like other Calliopsis, with medial paired laminar lobes on proximal margin between paired apodemal arms; apex simple with fringe of short setae. Male.— Head: (42) width 1.50-1.80 mm (3c = 1.66 ± 0.03; n = 10); (43) 1.35-1.43 (3c = 1.39 ± 0.01; n = 10) x broader than long; (44) clypeus granulate and distinctly punctate; (45) frons coarsely granulate with weak punctures; (46) vertex granulate to imbricate; (47) gena imbricate; (48) head black to dark brown with creamy white maculation entirely covering clypeus, subantennal plates, median supraclypeal patch and lower paraocular areas extending upward along inner margin of eyes to just above level of antennal sockets; (49) head clothed in erect, white setae; (50) inner margins of eyes converging below; eyes brown; (51) lateral ocelli separated from median ocellus by 1 ocellar diameter; (52) facial fovea slender, weakly impressed; (53) scape equal in length to flagellar segments 1-4; flagellum yellow below, with dark band above. Mouthparts: (54) labrum concolorous with frons; with central depressed glabrous area; (55) mandible yellow basally, becoming reddish apically; simple and acutely pointed; (56-60) mouthparts as in female except paraglossae slender and acutely pointed. Mesosoma: (61) pronotum imbricate dorsally and laterally with fine, short setae; pronotal lobe with erect, finely branched setae; (62) mesoscutum shiny to weakly imbricate with widely scattered, weak punctation; (63) mesoscutellum shiny, becoming distinctly punctate around lateral and posterior edges; metanotum imbricate with distinct punctation; (64) mesopleuron imbricate-punctate with erect white setae; (65) metapleuron imbricate with fine, short setae; (66) propodeum as in female; propodeal triangle glabrous with fine striations on dorsal surface; (67) intertegular distance 0.70-1.12 mm (3c = 1.04 ± 0.01; n = 25); (68) forewing length 3.80-4.35 mm (3c = 4.09 ± 0.03; n = 25); wings as in female except with more slender stigma and more distinct dark coloration to wing tip; (69) legs with yellow maculation on anterior surface of forefemur and all of foretibia and foretarsus, at apex of femur, and tibia and tarsus on mid and hind legs, except for dark, longitudinal spots on outer surfaces of mid and hind tibiae; (70) basitibial plate slender but distinct, with shiny concave surface; (71) mid and hindtibial spurs slender and weakly serrate; (72) tarsal claws weakly bifid. Metasoma: (73) terga brown; (74) T1-T4 imbricate-punctate with simple recumbent setae; T5-T6 with more elongate, erect, finely-branched setae; (75) fovea on lateral comers of T2 barely visible; (76) T7 with slender pygidial plate (width = 0.12-0.13 mm) well-defined by salient rim (Fig. If); surface colliculate; surrounding cuticle of T7 imbricate-punctate; (77) S1-S4 similar in color and sculpturing to terga; (78) S5 with elongate medial process (Fig. lc); process length equal to width of stemite along midline; (79) S6 with distal, paired, vertically oriented, quadrate processes fringed with a comb of setae apically (Fig. la); (80) S7 with distal, slender, vertically oriented lamellate lobes (Fig. lb); (81) S8 notched apically (Fig. Id); (82) genital capsule as in Fig. le. Diagnosis. — This species is very similar to C. callops but differs in the following respects. Male: apex of S8 notched medially (Fig. Id); apical prong of S5 more acutely pointed and longer (Fig. lc); apical paired projections of S6 broad and quadrate in lateral view, with a comb of apical setae (Fig. la); small foramen in the genital capsule (Fig. le); male T7, pygidial plate slender and deeply concave (Fig. If); male clypeus with less dense covering of hairs; yellow of clypeus typically extends above the fronto-clypeal suture along midline of face, commonly reaching level of antennal sockets. Female: pygidial plate more obtuse, surface slightly more concave; meso-basitarsus slender (length 2.5-2.9 x width; Fig. lg); faint black spot at apex of forewing; yellow spot absent or weakly developed on protibia and absent on mesotibia; eyes brown in pinned specimens. Material Examined.— USA. ARI Z ONA. COCHISE Co.: Apache, 21.7 km SW, 27 Aug 1969, J. G. & B. L. Rozen, AMNH [0, 2]; same loc., 14 Aug 1969, J. G. & K. C. Rozen, AMNH [0, 2]; same loc., 20 Aug 1971, Rozen & Favreau, AMNH [2, 9]; same loc., 14 Aug 1974, Rozens, AMNH [0, 7]; same loc., 23 Aug 1971, Rozen & Favreau, AMNH [0, 1]; Apache, 23.3 km SW, 4 Aug 1961, J. G. Rozen, Kallstroemia grandiflora Torrey ex. Gray, AMNH [9, 7]; Douglas, 1.6 km E, 16 Aug 1974, Rozens & Favreau, AMNH [0, 1]; Douglas, 26.7 km N, 24 Aug 1987, J. H. Cane, KU [6, 0]; Douglas, 30 km NE, 16 Aug 1971, Rozen & Favreau, AMNH [7, 3]; Portal, 15 km NE, 18 Aug 1992, B. N. Danforth, Physalis wrightii Gray [11, 20]. GRAHAM Co.: Safford, 29 Jul 1954, G. D. Butler, cotton, 1994 DANFORTH: REVIEW OF CALLIOPSIS (. HYPOMACROTERA ) 287 LACM [0, 1]. PIMA Co.: Sahuarita, 13 Jul 1956, R. H. Beamer, Eurphorbia sp., KU [0, 1]; Sahuarita, 13 Aug 1946, L. P. Wehme, LACM [0, 2]; Silver Bell Bajada, J. L. Neff, LACM [1,31]; Tucson, San Xavier, 24 Jul 1916, share w/ Clark etc., AMNH [2, 2]. YUMA Co.: Yuma, 14 Oct 1936, Lauderdale, NMNH [2, 0], CALIFORNIA. IMPERIAL Co.: Jun 1912, J. C. Bridwell, KU [3, 3]; Jun 1912, J. C. Bridwell, NMNH [38, 39]; May 1911, J. C. Bridwell, NMNH [13, 4]; Calexico, 14 Sep 1959, C. R. Wagner, LACM [1, 0]; Calexico, 1.6 km E, 28 Jun 1953, R. R. Snelling, Melilotus alba Medicus, LACM [0, 1]; Calexico, 1.6 km E, 28 Jun 1953, R. R. Snelling, Sida hederacea (Douglas) Torrey, LACM [1, 1]; Experimental Farm, Jun 1912, J. C. Bridwell, Physalis, NMNH [0, 1]. RIVERSIDE Co.. Indio, Keosegan Ranch, 17 Jul 1970, M. E. Irwin, cotton, UCR [1,0]. NEW MEXICO. HIDALGO Co.: Animas, 1 km N, 7 Aug 1988, B. N. Danforth, Physalis wrightii, KU [50, 50]. MEXICO. SINALOA: Culiacan, 27.8 km S, 30 Sep 1976, George & Snelling, LACM [0, 5]; Los Mochis, 16 km N, 152 m, 30 Sep 1976, George & Snelling, LACM [2, 3], SONORA: Guaymas, 13 km N, 1 Oct 1976, George & Snelling, LACM [1, 1]; Hermosillo, 85 km ENE, El Gavilan, 13 Aug 1991, Rozen & Pember, Kallstroemia grandiflora, AMNH [0, 1]; Los Alamos, 440 m, 3 Apr 1991, R. Ayala, [1, 0]. BAJA CALIFORNIA NORTE: Mexicali, 12 km SW, 19 Jul 1953, R. R. Snelling, Sida hederacea, LACM [0, 1], BAJA CALIFORNIA SUR: Loreto, 48 km S, 425 m, 7 Sep 1977, R. R. Snelling, LACM [0, 1], Calliopsis ( Hypomacrotera ) callops (Cockerell & Porter) Hypomacrotera callops Cockerell & Porter (1899: 419) [male, female]; Cockerell (1937:3) [holotype designation]; Rozen (1970) [biology]; Hurd & Linsley (1972) [parasite]. Types. -Holotype, male; data: NEW MEXICO. SAN MIGUEL Co.: Las Vegas, 1 Aug [no year], T. D. A. Cockerell, Chamaesaracha coronopus\ deposited: Amer¬ ican Museum of Natural History, New York. Cockerell & Porter (1899) designated an unknown number of males and females as the original type series. Cockerell (1937:3) later designated one male from this series as the holotype Description.—Female.— Head: (1) width 1.7-2.12 mm (5c = 1.95 ± 0.04; n = 10); (2) 1.39-1.47 (5c = 1.43 ± 0.01; n = 10) times broader than long; (3-19) as in C. persimilis except (7) head coloration dark brown to black with minute, creamy white spots on paraocular area immediately above man¬ dibular acetabulum in some specimens; (9) eyes blue. Mesosoma: (20-33) as in C. persimilis except (20) lateral surfaces of pronotum distinctly shiny, glabrous, not imbricate; (26) intertegular distance 1.16-1.50 mm (5c = 1.30 ± 0.03; n = 10); (27) forewing length 3.40-4.20 mm (5c = 3.86 ± 0.08; n = 10); wings without dark spot at apex; (28) legs brown with white spots at base of both foretibia and mesotibia; mesobasitarsus broad (length 1.75-2.4 times width; Fig. 2g). Metasoma: (34-41) as in C. persimilis except (38) pygidial plate slightly more acute. Male. —Head: (42) width 1.4-2.1 mm (x = 1.71 ± 0.04; n = 10); (43) 1.26-1.38 (* = 1.33 ± 0.01; n = 10) times broader than long; (44) clypeus granulate-punctate; (45) frons shiny, glabrous except for widely-scattered punctation; (46) vertex shiny with scattered punctation; (47) gena weakly imbricate, shiny; (48) head black to dark brown with white maculation as in C. persimilis except less extensive, barely reaching above antennal sockets and often evanescent or absent on subantennal plates and subantennal area; (49) head setae as in C. persimilis but setae on clypeus much more dense, mostly obscuring surface of clypeus; (50-60) as in C. persimilis. Mesosoma: (61-72) as in C. persimilis except (61) lateral surface of pronotum more shiny; (67) intertegular distance 0.75-1.10 mm (5c = 0.99 ± 0.02; n = 25); (68) forewing length 3.20-4.65 mm (5c = 3.98 ± 0.06; n = 25); (69) yellow on legs as in C. persimilis but dark spots on outer surface of mid and hind tibiae larger. Metasoma: (73-82) as in C. persimilis except (76) T7 with broad (width >0.16 mm), weakly-defined, blunt pygidial plate (Fig. 2f); surface mostly shiny, weakly rugulose; (78) median process on S5 shorter (length two-thirds width of stemite along midline; Fig. 2c); apex laterally flattened, blade-like with dense apical setae (Fig. 2c); (79) apical prongs of S6 more slender in lateral view but with similar comb of setae (Fig. 2a); (80) S7 with distal lamellate lobes broad and horizontal (Fig. 2b); (81) S8 tapering to acute apex, not notched apically (Fig. 2d); (82) genital capsule as in Fig. 2e, foramen large. Diagnosis. — This species is very similar to C. persimilis but is slightly larger 288 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Figure 2. Calliopsis callops. Male: (a) sixth stemite, (b) seventh stemite, (c) fifth stemite, (d) eighth stemite, (e) genital capsule, (f) seventh tergite. Female: (g) midleg. 1994 DANFORTH: REVIEW OF CALLIOPSIS {HYPOMA CROT ERA) 289 and differs in the following structural features: Male: apex of S8 rounded, coming to a point medially (Fig. 2d); apical prong of S5 less acutely pointed and shorter (Fig. 2c); apical paired projections of S6 acutely pointed in lateral view (Fig. 2a); large foramen in the genital capsule (Fig. 2e); male T7, pygidial plate broader and less deeply concave (Fig. 2f); male clypeus with more dense covering of hairs, which overlap the labrum; yellow of clypeus rarely extending above the fronto- clypeal suture along midline of face. Female: pygidial plate more acute; meso- basitarsus broad (length 1.75-2.4 times width; Fig. 2g); black spot at apex of wing lacking; well-developed yellow or white spot at base of pro- and mesotibiae; eyes blue in pinned specimens. Material Examined.— USA. ARIZONA. COCHISE Co.: Douglas, 22 Aug 1968, Rozen & Favreau, AMNH [11, 7]; Douglas, 1 Sep 1968, Rozen & Favreau, AMNH [0, 1]; Douglas, 14 Aug 1969, J. G. 6 K. C. Rozen, AMNH [0, 1]; Douglas, 31 Aug 1968, Rozen & Favreau, AMNH [1, 0]; Douglas, 17 Aug 1970, AMNH [0, 1]; Douglas, 3 May 1969, Rozen & Favreau, AMNH [2, 2]; Douglas, 1.6 km E, 17 Aug 1971, Rozen & Favreau, AMNH [4, 0]; same loc., 21 Aug 1968, Rozen & Favreau, AMNH [2, 1]; same loc., 20 Aug 1968, Rozen & Favreau, AMNH [1, 0]; same loc., 31 Aug 1971, Rozen & Favreau, AMNH [2, 0]; same loc., 18 Aug 1971, Rozen & Favreau, AMNH [1, 0]; same loc., 29 Aug 1971, Rozen & Favreau, AMNH [4, 0]; same loc., 21 Aug 1974, J. G. & B. L. Rozen, AMNH [1, 1]; same loc., 29 Aug 1971, Rozen & Favreau, AMNH [0, 1]; same loc., 19 Aug 1968, J. G. Rozen, AMNH [2, 1]; same loc., 24 Aug 1970, J. G. Rozen, AMNH [0, 1]; Douglas, 28.3 km E, 4 Aug 1958, P. A. Opler, LACM [0, 1]; Portal, 24 Aug 1971, Rozen & Favreau, AMNH [1, 0]. COLORADO. BAVA Co.: Regnier, 1372 m, 6 Jun 1919, T. D. A. Cockerell, AMNH [0, 1]. PROWERS Co.: Lamar, 1097 m, 4 Jun 1919, T. D. A. Cockerell, AMNH [0, 1], KANSAS. BARBER Co.: Aetna, 4.2 km S, 7 Aug 1962, Kerfoot & Michener, Quincula lobata (Torrey) Rafinesque, KU [10, 1]; Medicine Lodge, 25 km W, 12 May 1962, Michener & party, Quincula lobata, KU [1, 1]. DOUGLAS Co.: Lawrence, 24 Sep 1952, R. R. Snelling, Helianthus petiolaris Nuttal, LACM [0, 1]; Lawrence, 23 Aug 1952, J. A. Mathewson, Helianthus petiolaris, LACM [1, 0]. HAMILTON Co.: 1021 m, F. H. Snow, KU [1, 1], STANTON Co.: Johnson, 16 Jun 1949, Michener & Beamer, Quincula lobata, KU [1, 1], NEW MEXICO. EDDY Co.: Artesia, 5 km S, 20 May 1969, Brothers et al., Chamaesaracha conioides (Moricand) Britton, KU [1, 0]. HIDALGO Co.: Animas, 6.7 km S, 24 Aug 1974, Rozen & Favreau, AMNH [2, 4]; Cienega Ranch, 14 May 1987, J. G. Rozen, Chamaesaracha, AMNH [1, 0]; Rodeo, 1.6 km N, 19 Aug 1971, Rozen & Favreau, AMNH [0, 1], SIERRA Co.: Hot Springs, 22 Jul 1950, R. H. Beamer, Chamaesaracha conioides, KU [4, 2]; Hot Springs, 58.3 km N, 22 Jul 1950, R. H. Beamer, Baileya multiradiata Harvey & Gray, KU [1, 1]. TEXAS. ARMSTRONG Co.: Claude, 36.7 km S, Palo Duro Canyon, 4 Jun 1979, C. D. Michener, Quincula lobata, KU [2, 0]. BREWSTER Co.: Big Bend Park, Cooper’s Store, 11 Apr 1949, Michener & Beamer, Phacelia popei Torrey & Gray, KU [2, 0]. JEFF DAVIS Co.: Fort Davis, 33.3 km N, Davis Mts., 16 Apr 1961, Rozen & Schramel, AMNH [0, 2], DIMMIT Co.: Carrizo Springs, 14 Apr 1949, Michener & Beamer, KU [0,2], HIDALGO Co.: Progresso, 12 Apr 1950, Michener et al., Quincula lobata, KU [5, 2], MA VERICK Co.: Quemado, 14 Apr 1949, Michener & Beamer, Quincula lobata, KU [13, 5]. REEVES Co.: Toyahvale, 2.5 km S, 25 Apr 1979, R. R. Snelling, LACM [0, 1]; Toyahvale, Balmorhea State Park, 16 Apr 1961, Rozen & Schramel, AMNH [0, 1]. STARR Co.: Rio Grande (City?), 12 Apr 1950, R. H. Beamer, et al., Quincula lobata (1 male), KU [2, 1]. TERRELL Co.: Dryden, 21.7 km SE, 13 Apr 1949, Michener & Beamer, Chamaesaracha conioides, KU [0, 1]. VAL VERDE Co.: Langtry, 23.8 km NW, 549 m, 22 Apr 1973, R. R. Snelling, Chamaesaracha sordida (Dunal) Gray, LACM [0, 1]. MEXICO. CHI¬ HUAHUA: Camargo, 26 km N, 27 Aug 1991, J. G. Rozen, Quincula lobata, AMNH [0, 1]; same loc., 27 Aug 1991, J. G. Rozen, Euphorbia, AMNH [1, 0]; same loc., 27 Aug 1991, J. L. Neff, Quincula lobata, CTMI [2, 0]; Ceballos, 49 km NE, 15 Mar 1992, D. Yanega, KU [0, 1]; Chihuahua, 38 km S, 27 Aug 1991, R. L. Minckley, Dyssodia, KU [2, 2]; Jimenez, 18 km NW, 26 Aug 1991, J. G. Rozen, Dyssodia sp., AMNH [0, 1]; same loc., 26 Aug 1991, J. G. Rozen, AMNH [1, 1]; Jimenez, 5 km E, 21 Aug 1991, J. L. Neff, Chamaesaracha conioides, CTMI [2, 0]; Ojinaga, 31 km W, 28 Aug 1991, R. L. Minckley, KU [0, 2). COAHUILA: San Rafael, 1210 m, 24 Mar 1992, R. Brooks, Chamaesaracha crenata Rydberg, KU [2, 7]; San Rafael, 1170 m, 24 Mar 1992, J. L. Neff, Chamaesaracha coronopus (Dunal) Gray, CTMI [1, 0]. DURANGO: La Loma, 1249 m, 20 Aug 1947, C. D. Michener, Physalis, AMNH [9, 0]; Reserva Biosfera Mapimi, 23 Aug 1991, J. G. Rozen, Chamaesaracha, AMNH [5, 2]; 290 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) same loc., 22 Aug 1991, J. G. Rozen, Chamaesaracha crenata, AMNH [1,4]; same loc., 23 Aug 1991, J. G. Rozen, Dyssodia aurea (Gray) A. Nelson, AMNH [0, 1]; same loc., 21 Aug 1991, R. L. Minckley, Xylothamia triantha [sic?], KU [0, 1]; same loc., 21 Aug 1991, R. L. Minckley, Euphorbia, KU [0, 1]; Estacion Biologica, Mapimi, 28 Aug 1991, J. L. Neff, Malvella leprosa (Ortega) Krapovickas, CTMI [0, 1]; same loc., 23 Aug 1991, R. Ayala, UNAM [1, 2], Calliopsis {Hypomacrotera) subalpinus Cockerell Calliopsis subalpinus Cockerell (1894:235) [male]. Calliopsis semirufus Cockerell (1896:219) [female.]. Hypomacrotera andradensis Cockerell (1937:3) [male, female]. Hypomacrotera subalpinus, Rozen (1970) [biology]. Types.—Calliopsis subalpinus, Holotype, male; data: NEW MEXICO. DONA ANA Co.: Las Cruces, 1893, T. D. A. Cockerell; deposited: Academy of Natural Sciences, Philadelphia. Paratypes: same data as holotype, 1 male, 1 female; de¬ posited: National Museum of Natural History, Smithsonian Institution, Wash¬ ington, DC. Diagnosis.— Forewing length, male: 5.0-6.4 mm; female: 5.5-6.5 mm. This species is considerably larger than C. callops and C. persimilis (roughly 1.5 to 2.0 times the size in forewing length) and differs from those two species in the following structural and coloration characters: Male: maculation on legs restricted to anterior surface of foretibia and tarsus, and a small spot at base of mesotibia; head more quadrate with yellow maculation on clypeus broken up by dark area on disk of clypeus, below fronto-clypeal suture, and yellow maculations along inner orbit of eyes slender; male T7 with elongate, well developed, concave pygidial plate (Fig. 3f); male genitalia and apical sclerites as in Figs. 3a-e; Female: T1-T4 and prox¬ imal % of T5 reddish dorsally, becoming chocolate brown laterally, S1-S6 and distal l h of T5 deep chocolate brown; face with yellow maculation on subantennal sclerites, subantennal area immediately above fronto-clypeal suture, on lateral portions of clypeus and small spots of yellow on inner orbits of eyes just above fronto-clypeal suture; stigma elongate and slender, barely distinguishable from prestigma; marginal cell elongate and slender (length 5.0 times greatest width; length roughly 4.0 times width in C. callops and persimilis). Synonyms. —Calliopsis semirufus, Holotype, female; data: NEW MEXICO. DONA ANA Co.: Las Cruces, 25 Aug 1895, Sphaeralcea angustifolia\ deposited: National Museum of Natural History, Smithsonian Institution, Washington, DC. Hypomacrotera andradensis, Holotype, female; data: CALIFORNIA. IMPE¬ RIAL Co.: Andrade, near Yuma, 19 Apr 1937, Sphaeralcea-, deposited: American Museum of Natural History, New York. Discussion. —No description of this species is given here because previous de¬ scriptions are adequate. Although Hurd (1979) recognized andradensis as a sub¬ species of subalpinus, I see no reason for doing so. Calliopsis {H.) andradensis was originally distinguished from C. subalpinus based on color pattern differences: in male specimens from west of the Arizona-Califomia border the yellow mac¬ ulation on the clypeus is restricted to the lateral portions, immediately beneath the eyes, but eastern specimens show yellow across the entire clypeus, and more yellow coloration overall. The width of the male pygidial plate is a correlated character. Western specimens in general show broader pygidial plates (> 0.22 mm) but eastern specimens have more slender pygidial plates (> 0.22 mm). This 1994 DANFORTH: REVIEW OF CALLIOPSIS ( HYPOMACROTERA ) 291 Figure 3. Calliopsis subalpinus. Male: (a) sixth stemite, (b) seventh stemite, (c) fifth stemite, (d) eighth stemite, (e) genital capsule, (f) seventh tergite. character, like clypeal coloration shows a gradual transition from east to west. Neither character can be used unambiguously to separate eastern and western specimens. Therefore, I consider this insufficient basis for distinguishing two species, or even two subspecies. 292 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Material Examined. -USA. ARIZONA. COCHISE Co.: Apache, 23.3 km SW, 7 May 1989, J. G. Rozen, AMNH [0, 1]; Apache, 3.3 km E, 17 May 1987, J. G. Rozen, AMNH [1, 0]; Bisbee, 20 km W on Hwy 92, 14 Aug 1991, B. N. Danforth, Lepidium, KU [2, 0]; same loc., 14 Aug 1991, B. N. Danforth, Sphaeralcea, KU [17, 9]; Douglas, 23 Aug 1968, Rozen & Favreau, AMNH [1, 0]; Douglas, 3 May 1969, Rozen & Favreau, AMNH [0, 5]; Douglas, 21 Aug 1968, Rozen & Favreau, AMNH [1, 2] ; Douglas, 22 Aug 1968, Rozen & Favreau, AMNH [1,0]; Douglas, 31 Aug 1968, Rozen & Favreau, AMNH [1, 0]; Douglas, 1.6 km E, 21 Aug 1968, Rozen & Favreau, AMNH [0, 1]; same loc., 19 Aug 1968, Rozen & Favreau, AMNH [0, 1] same loc., 16 Aug 1962, M. Statham, Sphaeralcea, AMNH [2, 1]; Douglas, 20 km NW, 30 Aug 1989, Rozen et al., AMNH [0, 1]; Portal, 16.7 km NE, 24 Aug 1966, Rozens, AMNH [2, 0]; Portal, 14 km NNE, 31 Aug 1989, B. N. Danforth, Sphaeralcea, KU [0, 1]; Portal, 13.3 km NE, 14 Aug 1990, Rozen & Krieger, AMNH [0, 1]; same loc., 31 Aug 1990, J. G. & B. L. Rozen, AMNH [0, 1]; same loc., 23 Aug 1989, Rozen & Foster, AMNH [0, 1]; Rodeo vicinity, 11 Jun 1987, B. N. Danforth, Solarium, KU [0, 1]; San Simon, 3.3-10 km S, 3 Sep 1977, J. G. Rozen, AMNH [5, 6]; San Simon, 8.3 km S, 12 May 1987, J. G. Rozen, AMNH [4, 0]; San Simon, 10 km S, 16 May 1987, J. G. Rozen, AMNH [3, 0]; same loc., 10 May 1987, J. G. Rozen, Sphaeralcea, AMNH [1, 0]. LA PAZ Co.: Salome, 30 Aug 1979, E. M. Fisher, LACM [0, 1] MARICOPA Co.: Gila Bend, 26 Mar 1940, R. H. Crandall, LACM [1,0]; Gila Bend, 28.3 km S, 14 Apr 1968, E. M. Fisher, LACM [1, 1]; Sentinel, 25 Mar 1960, Gertsch & Schramel, AMNH [0, 9]; Tonopah, 8.3 km E, 24 Apr 1961, Rozen & Schramel, AMNH [0, 3], MOHAVE Co.: Kingman, 50 km W, 13 May 1980, Rozens, AMNH [0, 2], Nixon Springs, 60 km NW, 5 Aug 1969, R. R. Snelling, Sphaeralcea, LACM [0, 1], PIMA Co.: Tucson, 10 Jun 1938, R. H. Crandall, LACM [2, 1], PINAL Co.: Sacaton, (no date), T. H. Kearney, Sphaeralcea, NMNH [1, 0]. YAVAPAI Co.: Chino Valley, 6.7 km N, 31 Jul 1961, J. G. Rozen, AMNH [0, 1]; Congress, 33.3 kmNW, 29 Apr 1991, J. G. Rozen, AMNH [1,0]; Morristown, 35 km E, 24 Apr 1991, J. G. Rozen, AMNH [1, 0]. YUMA Co.: Dome Valley, 3 May 1991, J. G. Rozen, Sphaeralcea, AMNH [5, 2]; Ligurta, 20 Apr 1973, Rozen, AMNH [0, 2]; (no specific locality data), Ashmead, NMNH [0, 1], CALIFORNIA. IMPERIAL Co.: May 1911, J. C. Bridwell, Sphaer¬ alcea orcuttii Rose, NMNH [34, 2]; Apr 1911, J. C. Bridwell, NMNH [5, 7]; Andrade, 21 Jun 1953, R. R. Snelling, Sphaeralcea orcuttii, LACM [0, 2]; Calexico, 1.6 km E, 28 Jun 1953, R. R. Snelling, Sphaeralcea orcuttii, LACM [1, 0]; Calexico, 20 km E, 20 Apr 1949, R. C. Dickson, Sphaeralcea orcuttii, LACM [0, 1]; Experimental Fann, 21 May 1912, J. C. Bridwell, Sphaeralcea orcuttii, NMNH [0, 1]; Experimental Farm, Jun 1912, J. C. Bridwell, NMNH [0, 1]; Glamis, 28.7 km NW, 3 May 1958, E. L. Sleeper, LACM [2, 0]; Imperial, 29 Apr 1950, C. D. MacNeil, AMNH [0, 1]; Imperial, 16.7 km W, 26 Apr 1951, C. D. MacNeil, AMNH [2, 0]; Imperial, 8.3 km NW, 27 Apr 1951, C. D. MacNeil, AMNH [2, 7], INYO Co.: Eureka Valley Dunes, 4 May 1977, J. C. Hall, Sphaeralcea, LACM [0, 1]. RIVERSIDE Co.: Blythe, 30 km W, 8 Apr 1979, E. M. Fisher, Sphaeralcea, LACM [2, 0]; same loc., 8 Apr 1979, E. M. Fisher, Baileya, LACM [0, 11]; same loc., 17 Apr 1973, Rozens, AMNH [1, 1]; same loc., 17 Apr 1973, Rozens, Sphaeralcea, AMNH [2, 17]; same loc., 17 Apr 1973, Rozens, Malacothrix, AMNH [0, 1]; Blythe, 30-33 km W, 29 Mar 1958, Menke & Stange, LACM [1, 0]; Desert Center, 45 km E, 25 Apr 1961, Rozen & Schramel, AMNH [2, 1]; Joshua Tree National Monument, 14 Jun 1965, Sleeper & Jenkins, LACM [1, 0]. SAN BERNARDINO Co.: Adelanto, 6.7 km NW, 884 m, 18 Sep 1978, R. R. Snelling, Sphaeralcea ambigua Gray, LACM [7, 2], NEW MEXICO. BERNALILLO Co.: Albuquerque, 1524 m, 28 May 1944, W. O. Griesel, Oryzopsis, LACM [0, 1]. DONA ANA Co.: Mesilla, 1 Jul 1923, Cockerell, Sphaeralcea angustifolia (Cavanilles) G. Don, NMNH [1, 0]. HIDALGO Co.: Animas, 1 km N, 1 Aug 1988, B. N. Danforth, Sphaeralcea, KU [7, 3] ; Animas, 33.3 km S, 12 Sep 1977, Rozens, AMNH [1, 0]; same loc., 14 Sep 1977, B. L. Rozen, AMNH [1, 0]; same loc., 13 Sep 1977, Rozens, AMNH [2, 2]; Animas, 35 km S, 25 Aug 1975, Rozens, AMNH [3, 2]; same loc., 21 Aug 1975, Rozens, AMNH [0, 1]; same loc., 18 Aug 1975, Rozens, AMNH [1, 0]; Animas, 38.3 km S, 28 Aug 1975, Rozen & McGinley, AMNH [1, 0]; Animas, 41.6 km S, 30 Aug 1975, Rozens, AMNH [1, 0]; Animas, 6.7 km S, 24 Aug 1974, Rozen & Favreau, AMNH [1, 1]; Cienega Ranch, 16 Aug 1974, Rozen & Favreau, AMNH [2, 0]; Cotton City, 6.7 km NW, 22 Aug 1983, Rozen & Favreau, AMNH [2, 1]; Rodeo, 24 Jun 1987, B. N. Danforth, Sphaeralcea, KU [1, 0]; Rodeo, 20 km N, 15 Aug 1976, J. G. Rozen, AMNH [0, 1]; Rodeo, 21.7 km N, 19 May 1987, J. G. Rozen, AMNH [1,0]; Rodeo, 2.5 km N, 12 Aug 1991, B. N. Danforth, Sphaeralcea, KU [1, 0]; Rodeo, 7.5 km N, 21 Sep 1962, J. G. Rozen et al., AMNH [1, 0], LINCOLN Co.: Carrizozo, S on Rte 54, 15 May 1987, B. N. Danforth, Sphaeralcea sp., KU [8, 0]. VALENCIA Co.: Pueblo Laguna, 23 Jun 1959, Snelling & Snelling, Sphaeralcea ambigua, LACM [1, 1]. NEVADA. WASHOE Co.: Patrick, 16 May 1964, A. Gillogly, LACM [1, 2], TEXAS. BREWSTER Co.: Big Bend National Park, Rio Grande Village, 18 Apr 1970, L. B. & C. W. O’Brien, LACM [0, 3]. CULBERSON Co.: 1994 DANFORTH: REVIEW OF CALLIOPSIS (. HYPOMACROTERA) 293 Van Horn, 12.7 km S, 27 Apr 1979, R. R. Snelling, LACM [1, 2], HUDSPETH Co.: Dell City, 3.3 km N, 31 Jul 1950, R. F. Smith, AMNH [4, 0], PECOS Co.: Imperial, 17 Apr 1961, Rozen & Schramel, AMNH [1, 0]. REEVES Co.: Balmorhea, 16 Apr 1961, Rozen & Schramel, AMNH [2, 3]; Pecos, 53.3 km ESE, 17 Apr 1961, Rozen & Schramel, AMNH [1, 0]; Toyahvale, 2.5 km S, 25 Apr 1979, R. R. Snelling, LACM [4, 3]. TERRELL Co.: Dryden, 17.8 km S, 670 m, 22 Apr 1973, R. R. Snelling, LACM [9, 0]. UVALDE Co.: Uvalde, Nueces River, 11 Jul 1941, J. J. duBois, LACM [2, 0]. WARD Co.: Monahans, 10 km S, 17 Apr 1961, Rozen & Schramel, AMNH [0, 1]. ZAPATA Co.: San Ygnacio, 15 Apr 1952, Michener et al., Lindheimera texana, LACM [0, 1]. MEXICO. CHI- HUAUA: Salaices, 1584 m, 20 Aug 1947, G. M. Brandt, AMNH [0, 1]; Samalayuca, 17 km S, 31 Aug 1992, B. N. Danforth, Sphaeralcea incana Torrey, KU [2, 2], COAHUILA: General Cepeda, 3 km W, 1550 m, 23 Mar 1992, D. Yanega, Sphaeralcea angustifolia, KU [0, 1]; Guadalupe, 23 Aug 1947, M. Cazier, AMNH [2, 0]; San Lorenzo, 2 km N, 1430 m, 24 Mar 1992, B. Alexander, Thelocactus bicolor (Galeotti) Britton & Rose, KU [0, 1]. DURANGO: La Loma, 1250 m, 20 Aug 1947, C. D. Michener, AMNH [9,2]; Mapimi, 12 km E, 1350 m, 25 Mar 1992, R. Brooks, Sphaeralcea angustifolia, KU [0, 1], SONORA: Pueblo el Molinote, 21 Apr 1990, B. N. Danforth, Sphaeralcea, KU [0, 2]; San Jose de Guaymas, 10 Apr 1900, L. O. Howard, NMNH [1, 0], ZACATECAS: Concepcion del Oro, 10 Aug 1981, J. L. Neff, Sphaeralcea, LACM [1, 3], Discussion Phenology. — Figure 4a shows the collection data expressed as number of spec¬ imens collected per month from February through November. Calliopsis callops and C. subalpinus show clearly bimodal patterns corresponding to the bimodal rainfall typical of the Sonoran and Chihuahuan deserts (Sellers & Hill 1974). Calliopsis persimilis, however, shows a large peak in June, typically a very dry month. This peak results from a single collection of 83 specimens made by J. C. Bridwell in Imperial County, California in 1912. The spring records of C. callops were primarily collected in the eastern part of the range, in Texas and Coahuila, but the late summer specimens were collected further west, in Arizona, Chihuahua and Durango, where late summer “monsoon” rainfalls are common. Floral Associations. —There can be little doubt that H. subalpinus females re¬ strict their pollen foraging to species of Sphaeralcea (Malvaceae) (Fig. 4b). Species visited include S. ambigua Gray, S. angustifolia, S. incana and S. orcuttii. The other two species of Hypomacrotera appear to restrict their pollen-collecting to members of solanaceous genera: 89.6% of C. callops females and 75.6% of C. persimilis females were collected on solanaceous genera including Physalis, Quin- cula and Chamaesaracha. These three genera, along with five others, belong to the “physaloid genera” (Averett 1979), a group of low-growing, desert plants with non-poricidal anthers. While the single species of Quincula, Q. lobata, has been placed in Physalis by various sources (Correll & Johnston 1970, Kearney & Peebles 1960, Martin & Hutchins 1981), Quincula continues to be recognized as a distinct genus closely related to either Chamaesaracha or Physalis (Averett 1979; M. Nee, personal communication). Calliopsis persimilis and C. callops also show slight differences in plant pref¬ erence (Fig. 4b). Although C. persimilis shows a clear preference for species of Physalis, in particular P. acutifolia (Miers) Sandwith (= P. wrightii), female C. callops are more commonly collected on Chamaesaracha (C. conioides, C. crenata, and C. sordida) and Quincula lobata than on Physalis. Field studies support these results. Danforth (1990) found C. persimilis col¬ lecting pollen exclusively from Physalis acutifolia (as P. wrightii) near Animas, New Mexico, and Rozen (1970) found C. subalpinus visiting Sphaeralcea sp. near 294 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) a. m SUBALP □ PERSIM ■ CALLOPS Figure 4. (a) Collection data expressed as percent of all specimens collected per month, (b) Host plant association records expressed as percent of female specimens with associated plant data collected on each of 12 plant genera. Plant genera arranged in order of families given in USDA National List of Scientific Plant Names. (CHAMAESARA = Chamaesaracha, KALLSTROEM = Kallstroemia .) Douglas, Arizona. Rozen’s (1970) report of C. callops (as Hypomacrotera callops callops) visiting Physalis sp. near Douglas, Arizona is in error. The bees were collecting pollen and nectar from a species of Chamaesaracha that still grows abundantly at the site (J. Rozen, personal communication). If one included all the female C. callops collected by Rozen and colleagues at the Douglas locality (n = 31) in the histogram of plant preferences (Fig. 4b), the preference of C. callops for Chamaesaracha over Physalis would be even more apparent. Calliopsis callops and C. persimilis are clearly more closely related to each other than either is to C. subalpinus, and the plant data support that hypothesis. The species pair of callops + persimilis are solanaceous specialists, whereas C. sub¬ alpinus is an unambiguous oligolege on Sphaeralcea. Oligolecty, or restricted pollen foraging, is widespread among panurgine bees. Examples include Arhyso- sage species foraging exclusively on Opuntia (Cactaceae) (Jorgensen 1909), Cal- lonychium petuniae Cure & Wittman on Petunia (Solanacea) (Cure & Wittmann 1990), and Perdita species specializing on particular genera in over 30 plant families (Danforth 1991). Within Calliopsis sensu lato (Ruz 1991), there is con¬ siderable variation in the degree of specialization and in the plant groups visited. The subgenus Calliopsis includes species that are fairly polylectic, such as C. (C.) andreniformis, which collects pollen from 12 different plant families (Shinn 1967). The remaining subgenera of Calliopsis typically show much more oligolectic hab¬ its: most C. {Perisander) species are oligolectic on Euphorbia, C. ( Calliopsima ) species typically collect composite pollen, and the Calliopsis subgenera Noma- dopsis, Macronomadopsis and Micronomadopsis are almost all oligolectic, spe¬ cializing on one or two genera within various plant families including Legumi- nosae, Liliaceae, Hydrophyllaceae, Euphorbiaceae, Rosaceae and Boraginaceae (Rozen 1958). Because of this diversity in host plant usage among closely-related Calliopsis species, it is impossible to polarize host-plant association in the subgenus Hy¬ pomacrotera. Based on outgroup comparison, one cannot say whether the host plant shift has gone from Sphaeralcea to Solanaceae, vice versa, or whether the 1994 DANFORTH: REVIEW OF CALLIOPSIS ( HYPOMACROTERA) 295 common ancestor of these three species had a completely different source of pollen. The most interesting Calliopsis subgenus in this regard is Liopoeum because it is the sister group to Hypomacrotera (Ruz 1991). However, little is known of the host plants used by this group of South American bees. Geographical Distribution. — Calliopsis subalpinus (Fig. 6) is widespread throughout the Sonoran and Chihuahuan deserts of southern California, Arizona, New Mexico, western Texas and northern Mexico. The distribution patterns of the sister species C. callops and C. persimilis show a biogeographic pattern con¬ gruent with the division of the arid southwestern U.S. and northern Mexico into lowland, western, Sonoran desert and the upland, easterly, Chihuahuan desert (Shreve 1942) (Fig. 5). The distribution of C. persimilis corresponds closely to the distribution of Sonoran desert (Shreve & Wiggins 1964). Calliopsis callops 296 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) shows a roughly Chihuahuan desert distribution in the southern portions of its range but extends northward into grassland regions of northern New Mexico, west-central Texas, Colorado, southwestern Kansas and (presumably) western Oklahoma. Although the lowland areas of southeastern Arizona to southwestern Texas, Chihuahua, Durango, and Coahuila are classified as Chihuahuan desert, in fact, these areas are a patchwork of true Chihuahuan desert and semi-desert grassland (Brown 1982, Brown & Lowe 1980). Calliopsis callops most likely in¬ habits these desert-grassland habitats in the southern parts of its range, which easily accounts for its extension into true grassland further north. The record of C. callops in northeastern Kansas (one male and one female collected near Law¬ rence, Douglas Co.), however, is almost certainly due to an error in labeling 1994 DANFORTH: REVIEW OF CALLIOPSIS (HYPOMA CROTERA) 297 Figure 7. Distributions of collecting sites in Cochise County, Arizona and Hidalgo County New Mexico showing region of range overlap for C. persimilis and C. callops. because the northern-most range of Quincula is 200 miles to the southwest of this locality. Figure 7 shows the area where the ranges of C. callops and C. persimilis overlap in southern Arizona and New Mexico. Although specimens of C. callops have, 298 THE PAN-PACIFIC ENTOMOLOGIST Yol. 70(4) in general, been collected eastward, and C. persimilis westward, there is a region of overlap roughly 25 miles (42 km) wide that runs parallel to the San Simon Valley, just west of the Arizona-New Mexico border. Within this overlap zone specimens of the two species are easily distinguishable, which supports the view that they are, in fact, good species. Plants, in particular Larrea divaricata Cava- nilles (Wells & Huntziker 1976; Yang 1961, 1970; Yang & Lowe 1968), show similar patterns of vicariance in this area. The distributions of C. callops and C. persimilis are correlated with the distri¬ butions of the plant genera that serve as their pollen sources. In order to assess this relationship, I compiled lists of the plant species that could potentially serve as pollen sources by consulting regional floras (Correll & Johnston 1970, Kearney & Peebles 1960, Martin & Hutchins 1981, Munz & Keck 1968) and then accepting synonymy decisions in the USDA National List of Scientific Plant Names (1982). The genera that primarily serve as pollen sources for C. callops (Chamaesaracha and Quincula) reach the western-most limits of their ranges in eastern Arizona. In contrast, nine of the seventeen species of Physalis occurring in the southwestern U.S. extend westward into southern California and thereby overlap all or a part of the range of C. persimilis. In other words, neither Quincula lobata nor the seven southwestern species of Chamaesaracha combined could serve as a pollen source for C. persimilis over its entire range. Acknowledgment I am grateful to the curators and collections managers who provided specimens for this study, including: Jerome G. Rozen and Eric Quinter, American Museum of Natural History; Robert W. Brooks, Snow Entomological Museum, University of Kansas; Roy R. Snelling, Los Angeles County Museum of Natural History; Saul Frommer, University of California, Riverside; Wojciech Pulawski, California Academy of Sciences; Ronald McGinley, National Museum of Natural History, Smithsonian Institution; John Neff, Central Texas Melittological Institute; and Ricardo Ayala, Universidad Nacional Autonoma de Mexico. Many specimens used in this study were provided as a result of collecting trips organized by the Programa Cooperative sobre la Apifauna Mexicana and funded by the National Science Foundation (Wallace E. LaBerge & Ronald J. McGinley, Principal In¬ vestigators, NSF BFR 90-24723). These specimens were especially important because the Mexican fauna was poorly collected previously. I thank J. G. Rozen, Jr. for originally raising the possibility that H. callops was, in fact, two species. Final preparation of this paper was made possible by an NSF Post-doctoral Fel¬ lowship (DEB-9201921). I am grateful to Jerome G. Rozen, Jr. and Charles D. Michener for comments on earlier versions of this paper, and to the two reviewers. Literature Cited Averett, J. E. 1979. Biosystematics of the physaloid genera of Solanaceae in North America, pp. 493-503. In Hawkes, J. G., R. N. Lester & A. D. Skelding (eds.). The biology and taxonomy of the Solanaceae. Linnean Society Symposium Series, No. 7. Academic Press, London. Brown, D. E. 1982. Semidesert grassland, pp. 123-131. In Brown, D. E. (ed.). Biotic communities of the American southwest—United States and Mexico. Desert Plants, Special Issue, vol. 4 (1-4). 1994 DANFORTH: REVIEW OF CALLIOPSIS ( HYPOMACROTERA ) 299 Brown, D. E. & C. H. Lowe. 1980. Biotic communities of the southwest. USDA Forest Service Technical Report RM-78, 1 p. (map). Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado. Cockerell, T. D. A. 1894. Description of new Hymenoptera. Ent. News, 5: 234-236. Cockerell, T. D. A. 1896. New North American bees. Ent. Monthly Mag., 32: 218-221. Cockerell, T. D. A. 1899. New insects from Arizona, and a new bee from New Mexico. The Entomologist, 33: 61-66. Cockerell, T. D. A. 1937. Bees collected in Arizona and California in the Spring of 1937. Am. Mus. Novitates, 948. Cockerell, T. D. A. & W. Porter. 1899. Contributions from the New Mexico Biological Station. VII. Observations on bees and descriptions of new genera and species. Ann. Mag. Nat. Hist., (7)4: 403-421. Correll, D. S. & M. C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Council, Renner, Texas. Cure, J. R. & D. Wittmann. 1990. Callonychium petuniae a new panurgine bee species (Apoidea, Andrenidae) oligolectic on Petunia (Solanaceae). Stud. Neotrop. Fauna Environ., 25: 153-156. Danforth, B. N. 1990. Provisioning behavior and the estimation of investment ratios in a solitary bee, Calliopsis ( Hypomacrotera ) persimilis (Cockerell) (Hymenoptera: Andrenidae). Behav. Ecol. Sociobiol., 27: 159-168. Danforth, B. N. 1991. Phylogeny of the bee genus Perdita (Andrenidae: Panurginae). Ph.D. Thesis, University of Kansas, Lawrence. Harris, R. A. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology, State of California, Department of Food and Agriculture, 28. Sacramento, California. Heithaus, E. R. 1979. Flower-feeding specialization in wild bee and wasp communities in seasonal neotropical habitats. Oecologia, 42: 179-194. Hurd, P. D., Jr. 1979. Superfamily Apoidea. pp. 1741-2209. In Krombein, K. V., P. D. Hurd, D. R. Smith & B. D. Burks (eds.). Catalog of Hymenoptera in America north of Mexico (Vol. 2). Smithsonian Institution Press, Washington. Hurd, P. D., Jr. & E. G. Linsley. 1972. Parasitic bees of the genus Holcopasites Ashmead. Smithson. Contr. ZooL, 114: 1-41. Jorgensen, P. 1909. Beobachtungen iiber Blumenbesuch, Biologie, Verbreitung usw. der Bienen von Mendoza. (Hym.). Teil 1. Deutsch. Ent. Zeitschr., 1909: 53-65. Kearney, T. H. & R. H. Peebles. 1960. Arizona flora (2nd ed.). Rev. J. T. Howell & E. McClintock. University of California Press, Berkeley. Martin, W. C. & C. R. Hutchins. 1981. A flora of New Mexico (Vol. 2). J. Cramer, Vaduz, Germany. Michener, C. D. 1944. Comparative external morphology, phylogeny, and classification of the bees (Hymenoptera). Bull. Amer. Mus. Natur. Hist., 82: 151-326. Munz, P. A. & D. D. Keck. 1968. A California flora. University of California Press, Berkeley and Los Angeles. Rozen, J. G., Jr. 1958. Monographic study of the genus Nomadopsis Ashmead (Hymenoptera: Andrenidae). Univ. Calif. Publ. Entomol., 15. Rozen, J. G., Jr. 1970. Biology and immature stages of the panurgine bee genera Hypomacrotera and Psaenythia (Hymenoptera, Apoidea). Am. Mus. Novit., 2416: 1-16. Ruz, L. 1991. Classification and phylogenetic relationships of the panurgine bees: the Calliopsini and allies (Hymenoptera: Andrenidae). Univ. Kansas Sci. Bull., 54: 209-256. Sellers, W. D. & R. H. Hill. 1974. Arizona climate 1931-1972 (2nd ed.). University of Arizona Press, Tucson. Shinn, A. F. 1967. A revision of the bee genus Calliopsis and the biology and ecology of C. an- dreniformis (Hymenoptera: Andrenidae). Univ. Kansas Science Bull., 46: 753-936. Shreve, F. 1942. The desert vegetation of North America. Bot. Rev., 8: 195-247. Shreve, F. & I. L. Wiggins. 1964. Vegetation and flora of the Sonoran Desert (Vol. 1). Stanford University Press, Stanford, California. United States Department of Agriculture, Soil Conservation Service. 1982. National list of scientific plant names (Vols. 1 & 2). Washington, D.C. Wells, P. V. & J. H. Huntziker. 1976. Origin of the creosotebush ( Larrea ) deserts of southwestern North America. Ann. Missouri Bot. Garden, 63: 843-861. 300 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Yang, T. W. 1961. The recent expansion of creosotebush (Larrea divaricata) in the North American desert. Western Reserve Academy Natural History Museum Publication 1 (11 pp.). Yang, T. W. 1970. Major chromosomes races of Larrea divaricata in North America. J. Arizona Acad. Sci., 6: 41-45. Yang, T. W. & C. H. Lowe. 1968. Chromosome variation in ecotypes of Larrea divaricata in the North American desert. Madrono, 19: 161-164. PAN-PACIFIC ENTOMOLOGIST 70(4): 301-308, (1994) SYSTEMATICS AND BIOLOGY OF ACENTRELLA TURBIDA (McDUNNOUGH) (EPHEMEROPTERA: BAETIDAE) W. P. McCafferty, 1 Michael J. Wigle, 2 and R. D. Waltz 3 department of Entomology, Purdue University, West Lafayette, Indiana 47907 Abstract. — The larval stage of Acentrella turbida (McDunnough) is described for the first time. Populations from British Columbia and Oregon are the basis of the description and also represent new geographic records for the species. Acentrella Carolina (Banks), the only other described species of Acentrella lacking hindwings in North America, is shown to be a junior synonym of A. turbida. Keys are provided for distinguishing North American species of Acentrella in both the adult and larval stages. The European species A. sinaica Bogoescu is closely related to A. turbida, the two sharing nearly identical larval and adult morphology but differing mainly in the presence or absence of hindwings, respectively. Abdominal coloration of male adults of A. turbida quickly fades in alcohol-preserved specimens. Notes on emergence times and habitat of A. turbida, based mainly from studies on the Atnarko River in British Columbia, are provided. Key Words.— Insecta, Ephemeroptera, Baetidae, Acentrella turbida, North America, larval description, keys A major problem in the taxonomy of the Ephemeroptera of North America, and elsewhere, is the lack of descriptive and comparative larval data (McCafferty et al. 1990). This is particularly acute in the complex family Baetidae, where many species remain unknown in the larval stage, and where larval characteristics often are indispensable for both species diagnosis and interpreting phylogenetic rela¬ tionships (e.g., Waltz & McCafferty 1987a, b, c; McCafferty & Waltz 1990). Within Baetidae, species previously described under the name Pseudocloeon Klapalek in North America are especially poorly known as larvae. These species have been variously recombined with other genera (see McCafferty & Waltz 1990) because Pseudocloeon proved to be an artificial construct for baetine species possessing paired marginal intercalaries in the fore wings and lacking hindwings. Certain species originally described in Pseudocloeon have proven to belong to Acentrella Bengtsson. In North America this has included A. Carolina (Banks) and A. turbida (McDunnough) (Waltz & McCafferty 1987a). Acentrella has recently been comprised of five nominal species in North America, the other three being A. ampla Traver, A. insignificans (McDunnough), and A. lapponica Bengtsson (McCafferty & Waltz 1990). Of these species, only A. turbida has been thought to be unknown as larvae. Upon recently procuring an extensive series of adults and larvae of A. turbida from British Columbia, we sought to study this material and provide a first larval description. Our analysis and comparison with other Acentrella indicated that A. turbida and A. Carolina are conspecific. We report Acentrella turbida in British Columbia for the first time. The rugged coastal area of British Columbia, from where it was collected, was discussed by Wigle & Thommasen (1990). We also have studied larvae from Oregon that match 2 Box 643, Bella Coola, British Columbia V0T ICO, Canada. 3 Division of Entomology and Plant Pathology, IDNR, 402 West Washington, Indianapolis, Indiana 46204. 302 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) the associated larvae from British Columbia. Previously, A. turbida was reported from Alberta (McDunnough 1924), Utah (Edmunds 1954), and Colorado (McCafferty et al. 1993). Acentrella Carolina s. auctt. has been known from eastern and midwestem North America. In addition to nomenclatural changes, we herein give: (1) a morphological description of the larvae of A. turbida, (2) comments on possible relationships of the species, (3) diagnostic keys to the larvae and male adults of Acentrella species in North America, (4) some notes on the adult coloration in the species, and (5) notes on the habitat and biology of the species in British Columbia. The larval description will serve as a first complete description of the species, because larval morphological information previously has been restricted to incomplete data referred to A. Carolina in keys by McDunnough (1931, 1932). Materials upon which this study has been based are deposited in the Oregon State University Collection, the Purdue Entomological Research Collection, and in private collections of the authors. Acentrella turbida (McDunnough), 1924 Pseudocloeon turbidum McDunnough, 1924 Pseudocloeon Carolina Banks, 1924, NEW SYNONYM Acentrella turbida (McDunnough): Waltz and McCafferty, 1987a Acentrella Carolina (Banks): Waltz and McCafferty, 1987a, NEW SYNONYM Larval Morphology.— Length excluding caudal filaments: 4-6 mm. Head: head capsule broader than long; frons without medial process. Antennae longer than head capsule, with pedicel subequal to scape in length (each with fine setae). Labrum width approximately 2.0 x length; submarginal setae 1 + 5-6; other setae present at base. Right and left mandibles with incisors (Figs. 1 and 2) bent back¬ ward and inner surface serrate at margin, with 7-9 discernible denticles (outermost dorsally juxtaposed to second denticle); molar process of left mandible prominent (height ca. 2.0 x basal width). Maxillary palpi short and robust, subequal in length to galealaciniae. Labial palpi (Fig. 3) compact, subequal in length to apices of glossae and paraglossae; segment 2 length subequal to segment 3, with lobe weakly developed, and with 2-3 dorsal setae; innermost setal row of each paraglossa with 5-6 setae; each glossa with 5-6 setae projecting medially. Thorax: Hindwingpads absent. Legs (Fig. 4) with well-developed row of long fine setae on femora, tibiae and tarsi; femoral setae approximately 0.60-0.75 x width of femora; short, sharp setae on venter of femora and tibiae, and somewhat longer ones on venter of tarsi; claws (Fig. 5) with 8-10 denticles and lacking distal subapical setae. Abdomen: Dorsal color pattern as in Figs. 6-8. Tergal surfaces with fine setae, without scales, and with tergal marginal spines poorly developed as short, sharp, single spiculae. Paraproct surface with pores, setae, and small spines; posteromedial margins with sparse spines. Abdominal gills 1.5-2. Ox length of respective tergum, slightly asymmetrical, with posterior margins more rounded, and both margins smooth with very sparse fine setae. Caudal filaments not banded; median terminal filament consisting of one segment. Diagnosis.— See key. Species Relationships. — Acentrella turbida is most closely related to A. sinaica Bogoescu, presently known from Romania, Italy, Poland, Portugal, and Switzer- 1994 McCAFFERTY ET AL.: ACENTRELLA TURBID A 303 Figures 1-5. Acentrella turbida larva from British Columbia. Figure 1. Right mandible. Figure 2. Left mandible. Figure 3. Labium. Figure 4. Foreleg. Figure 5. Foreclaw. land (see Miiller-Liebenau 1969, Waltz & McCafferty 1987a, Studemann et al. 1992). Male adult genitalia and mouthpart, leg, and tergal structures of the larvae are symmorphic. The major difference between the two species is that A. sinaica retains vestigial hindwings. Mtiller-Liebenau (1969) and Jacob (1990) provided data for differentiating A. sinaica from other European Acentrella species. Waltz & McCafferty (1987a) previously indicated that A. turbida formed a natural grouping within Acentrella, including the Holarctic A. lapponica as well as the Palaearctic A. chatauensis (Kluge), A. fenestrata (Kazlauskas), A. sibirica (Kazlauskas), and A. sinaica. This was inferred on the basis of this grouping possessing reduced posterior spines on the abdominal terga that take the form of spiculae. It remains to be seen if the western and northwestern populations of A. turbida are disjunct from those of the east and southeast, and if any consistent differences will separate these populations. We have not been able to find consistent color differences in the larvae; however, the abdominal terga of male adults apparently differ as follows: northwestern and western males tend to have olive brown terga (although terga 3-6 are usually faded) and segment joinings of the cerci do not appear darkened; southeastern and eastern males usually have dark brown terga (also often faded in terga 3-6), and segment joinings of the cerci tend to be darkened. If, in the future, these minor differences prove to be consistent, rec¬ ognition of two geographic subspecies may be warranted. Adult Coloration.— Freshly collected male adults of A. turbida were found to 304 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Figures 6-8. Acentrella turbida larvae from British Columbia. Figure 6. Variation 1 (male). Figure 7. Variation 2 (female). Figure 8. Variation 3 (female). 1994 McCAFFERTY ET AL.: ACENTRELLA TURBID A 305 Figures 9-10. Acentrella turbida male adults from British Columbia. Figure 9. Color pattern in life. Figure 10. Color pattern after being alcohol-preserved. 306 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) be typical of the descriptions given by McDunnough (1924) and Traver (1935), having a solidly colored dorsal abdomen (Fig. 9). These workers had evidently worked with pinned material. However, we noted that the dorsal abdominal patterns of male adults quickly faded in alcohol, giving rise to an abdomen that was lighter in terga 3-6, especially terga 4 and 5 (Fig. 10) and somewhat similar to the condition found in the larvae. When we removed these latter specimens from the alcohol (some having been in alcohol for a little over a year) and allowed them to dry, much of their original coloration reappeared. We do not know if this procedure will work on specimens that have been in alcohol for prolonged periods. This is an important discovery because, to a large extent, the keys to male adult Baetis and Pseudocloeon used by Traver (1935) and others have relied heavily on these particular coloration differences to separate species. Habitat and Biology.— Larvae were collected from the Atnarko River, British Columbia, in deep riffle and rapids with rocky bottoms. Larvae were never taken in great abundance, with only a few at a time being taken with a D-frame net. Water temperature ranged from 14° C (8 Sep 1991) to 7° C (16 Oct 1990) during the period of early fall when larvae were collected. Other mayflies taken with A. turbida were Baetis tricaudatus Dodds, Rhithrogena sp., and Drunella doddsi (Needham). Serratella tibialis (McDunnough) was common until about mid Sep¬ tember. Emergence of subimagos of A. turbida took place from the first to the middle of October in large numbers from deep riffles and runs of moderate to fast flow areas of the river. The daily emergence period began at about 13:00 h and abruptly ceased at about 16:30 h. Emergence occurred during a wide variety of weather conditions, including overcast, rainy, clear, and sunny weather when wind was calm or slight. Reports of habitat and emergence of A. turbida in the east have basically been limited to Traver’s (1935) comment that larvae of A Carolina were taken abun¬ dantly in mountains of North Carolina. In addition, a table by Unzicker and Carlson (1982) indicated that A. Carolina occurred in the mountains and Piedmont of North and South Carolina and that emergence took place from April to August. Material Examined.—OREGON. COOS Co.: S Frk Coquilla Riv, 12 Jun 1982, C. W. Courtney, several male and female larvae. DOUGLAS Co.: S Umpqua Riv, 19 Jul 1980, C. W. Courtney, several male and female larvae. CANADA. BRITISH COLUMBIA: Bella Coola Watershed, Atnarko R. near Flat Rock, 1-16 Oct 1990, 8-19 Oct 1991, M. J. Wigle, several male and female larvae and adults. Keys to the North American Species of Acentrella Mature Larvae la. Hindwingpads present . 2 lb. Hindwingpads absent . A. turbida 2a (lb). Dorsum of femora, tibiae, and tarsi with dense row of long, fine setae (Morihara & McCafferty 1979: fig. 14d); western and far northern North America . A. insignificans 2b. Dorsum of tibiae and tarsi nearly bare (Morihara & McCafferty 1979: fig. 15b) or with shorter, more robust setae only (Morihara & McCafferty 1979: fig. 13c) . 3a (2b). Labial palpi with segments 1 and 2 nearly parallel sided (from dor- 3 1994 McCAFFERTY ET AL.: ACENTRELLA TURBIDA 307 soventral perspective), inner margin of segment 2 almost straight (Morihara & McCafferty 1979: fig. 15a); abdominal terga with posterior marginal spines blunt to slightly rounded, not spiculate (Morihara & McCafferty 1979: fig. 15e); claws without pair of sub- apical setae; eastern and midwestem North America. A. ampla 3b. Labial palpi with inner margins of segments 1 and 2 forming broadly rounded lobes (from dorsoventral perspective), inner margin of segment 2 decurved (Morihara & McCafferty 1979: fig. 13b); ab¬ dominal terga with very fine marginal spines; claws with paired subapical setae (Muller-Liebenau 1969: fig. 46h); far northern North America . A. lapponica Male Adults Hindwings present. 2 Hindwings absent. A. turbida Abdominal terga 2-7 translucent brown; length of distal segment of forceps 4x width; far northern North America . A. lapponica Abdominal terga 2-6 yellow-brown or tinged with smoky brown; length of distal segment of forceps ca. 3 x width; all North Amer¬ ican regions represented . 3 Turbinate portion of eyes well developed and orange; body 6-7 mm long; abdominal terga 2-6 yellow-brown; eastern and midwestem North America . A. ampla Turbinate portion of eyes relatively small and bright red; body 4-5 mm long; abdominal terga 2-6 tinged with smoky brown; western and far northern North America . A. insignificans Acknowledgment We thank Arwin Provonsha for line drawings. This paper has been assigned Purdue Experiment Station Journal No. 13729. Literature Cited Edmunds, G. F., Jr. 1954. The mayflies of Utah. Proc. Utah Acad. Sci. Arts. Lettr., 31: 64-66. Jacob, U. 1990. Ephemeroptera: zur systematik der Europaischen Baetidae auf Gattungsebene. Verh. Westd. Entomol., 1990: 271-290. McCafferty, W. P. & R. D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera) of North and Middle America. Trans. Am. Entomol. Soc., 116: 769-799. McCafferty, W. P., R. S. Durfee & B. C. Kondratieff. 1993. Colorado mayflies (Ephemeroptera): an annotated inventory. Southwest. Natural., 38: 252-274. McCafferty, W. P., B. P. Stark & A. V. Provonsha. 1990. Ephemeroptera, Plecoptera, and Odonata. pp. 43-58. In Kostarab, M. & C. Schaefer (eds.). Systematics of the North American insects and arachnids: status and needs. Va. Agri. Exper. Stat. Infor. Ser. No. 90-1, Va. Polytech. Inst. Univ., Blacksburg. McDunnough, J. 1924. New Canadian Ephemeridae with notes, II. Can. Entomol., 56: 90-122. McDunnough, J. 1931. New species of North American Ephemeroptera. Can. Entomol., 63: 82-93. McDunnough, J. 1932. New species of North American Ephemeroptera II. Can. Entomol., 64: 209- 215. Morihara, D. K. & W. P. McCafferty. 1979. The Baetis larvae of North America (Ephemeroptera: Baetidae). Trans. Am. Entomol. Soc., 105: 139-221. Muller-Liebenau, I. 1969. Revision der Europaischen Arten der Gattung Baetis Leach, 1815 (Insecta, Ephemeroptera). Gewass. Abwass., 48/49: 1-214. la. lb. 2a (la). 2b. 3a (2b). 3b. 308 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Studemann, D., P. Landolt, M. Sartori, D. Hefti & I. Tomka. 1992. Ephemeroptera, Insecta Helvetica, Yol. 9. Soc. Entomol. Suisse, Fribourg. Traver, J. R. 1935. Systematics. pp. 237-739. In Needham, J. G., J. R. Traver & Y. C. Hsu (eds.). The biology of mayflies with a systematic account of North American species. Comstock, Ithaca New York. Unzicker, J. D. & P. H. Carlson. 1982. Ephemeroptera. pp. 3.1-3.97. In Brigham, A. R., W. U. Brigham & A. Gnilka (eds.). Aquatic insects and oligochaetes of North and South Carolina. Midwest Aquat. Enterpr., Mahomet, Illinois. Waltz, R. D. & W. P. McCafferty. 1987a. Systematics of Pseudocloeon, Acentrella, Baetiella, and Liebebiella new genus (Ephemeroptera: Baetidae). J. N. Y. Entomol. Soc., 95: 553-568. Waltz, R. D. & W. P. McCafferty. 1987b. New genera of Baetidae for some Nearctic species included in Baetis Leach (Ephemeroptera). Ann. Entomol. Soc. Am., 80: 472^474. Waltz, R. D. & W. P. McCafferty. 1987c. New genera of Baetidae (Ephemeroptera) from Africa. Proc. Entomol. Soc. Wash., 89: 177-184. Wigle, M. J. & H. Y. Thommasen. 1990. Ephemeroptera of the Bella Coola and Owikeno Lake watersheds, British Columbia central coast. J. Entomol. Soc. Brit. Col., 87: 1-9. PAN-PACIFIC ENTOMOLOGIST 70(4): 309-312, (1994) A NEW SPECIES OF NALLACHIUS (NEUROPTERA: DILARIDAE) FROM COSTA RICA Norman D. Penny Department of Entomology, California Academy of Sciences, San Francisco, California 94118 Abstract.— A new species of Nallachius is described from Guanacaste Province of Costa Rica, and relationships with other species discussed. Key Words. — Insecta, Neuroptera, Dilaridae, Nallachius, Costa Rica In the course of identifying a series of 14 Nallachius from Costa Rica for the Utah State Insect Collection, four specimens of an undescribed species were en¬ countered. This species is herein described. Male genital terminology is that of Adams (1970). Original description based on four males, pinned, with genitalia mascerated in 10% KOH, stained in Chlorazol Black E, and preserved in glycerin capsules beneath the specimen. Abbreviations. — b—basal piece of MA; CuA—cubitus anterior; CuP—cubitus posterior; dl—dorsal lobe of ectoproct; ect—ectoproct; gs—gonarcus; KOH—po¬ tassium hydroxide; MA—media anterior; ma-mp—anterior-posterior medial crossvein; ml—median lobe of aedeagus; MP1 — first media posterior; mpl-mp2— first-second posterior medial crossvein; MP2—second media posterior; mu— mediuncus; r-m—radial-medial crossvein; Rl— first radial vein; Rs—radial sec¬ tor; 8S, 8T—eighth abdominal stemite and tergite; 9S, 9T—ninth abdominal stemite and tergite. Nallachius parkeri Penny, NEW SPECIES Types. -Holotype male: COSTA RICA. GUANACASTE: 3 km SE of Rio Naran¬ jo, 23-27 Jan 1992, F. D. Parker. Three paratype males from the type locality: 7-12 Feb 1992, 25 Feb-2 Mar 1992, and 10-20 Sep 1992, F. D. Parker. The holotype will be deposited with INBio (Instituto de Biodiversidad) in Santo Do¬ mingo de Heredia, Costa Rica. Two paratypes are in the Utah State University Collection, Logan, and one paratype is deposited at the California Academy of Sciences, San Francisco. Description.— Head. Frons, clypeus, and vertex dark brown. Vertex bearing 3 protuberances, each bearing numerous long setae. Antennal scape quadrangular, pale basally, dark brown apically, bearing subapical, lateral long seta; pedicel 3.0 x wider than long; 15 flagellomeres, first broader than long, others all much longer than wide, each bearing very elongate pectinate lobe, except apical 3. Thorax. Dark brown dorsally, pale yellow laterally and ventrally. Notum bearing numerous tufts of long setae. Legs. Pale yellow, with numerous long pale setae on all segments. Apex of tibiae with abundant long dark setae. Wings. Forewing length—4.1 mm: Darkly suffused transverse bands across forewing, as in Fig. 1. Costal crossveins numerous, not branched; 2 to 5 sc crossveins; 2 radial crossveins, the anterior one close to origin of first branch of Rs; 4 branches of Rs; MA two-branched; MP1 two- branched; MP2 three-branched; base of MP2 fused with CuA for short distance; 1 r-m crossvein; 2 ma-mp crossveins; two mpl-mp2 crossveins. Hindwing-length—3.0 mm: No sc crossveins anterior of pterostigmal area; 2 radial crossveins; 1 ma-mp crossvein and 1 mp2-cua crossvein (Fig. 2). Abdomen. Pale yellow. Each segment with narrow transverse ridge dorsal and ventrally bearing 310 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Figure 1. Right forewing of Nallachius parkeri NEW SPECIES. numerous long dark setae. Each ridge narrowly divided along mid-ventral and mid-dorsal lines, as well as broadly separated laterally. Male Terminalia. Ninth tergite bearing pair of blunt tooth-shaped lobes closely appressed to dorsal surface. Ectoprocts bearing medial, bipartite, dorsal lobes with smaller acute tooth at lateral base (Fig. 3). Digitiform process of ectoproct absent. Gonocoxites very narrow and elongate (extending the length of ectoproct beyond abdomen), apically upturned and acutely tapered. Mediuncus very long, narrow, apically curved ventrally and acutely tapered (Fig. 4); median lobe bifid apically. Diagnosis. — This species is a member of the N. americanus group, having fore¬ wing vein MP2 fused with CuA for a short distance. It differs from the other two species in this group by having only two r-rs crossveins, as opposed to N. loxanus Navas from Ecuador, which has five such crossveins. The only other species in this group, N. americanus (McLachlan) can be distinguished most easily by the male genitalia, which have much shorter gonocoxites, narrower, non-bifurcate dorsal lobes, a well developed digitiform process laterally on the ectoproct and a Figure 2. Right hindwing of Nallachius parkeri NEW SPECIES. 1994 PENNY: A NEW NALLACHIUS 311 Figure 3. Apex of male abdomen of Nallachius parked NEW SPECIES (ventral view). small medial point on the ninth tergite, as opposed to the flat, plate-like paired teeth of N. parkeri. Etymology.— This species is named for Frank D. Parker, a Hymenoptera sys¬ tematise who has devoted several years studying the insect fauna of northern 0.1 mm Figure 4. Apex of male abdomen of Nallachius parkeri NEW SPECIES (lateral view). 312 THE PAN-PACIFIC ENTOMOLOGIST Yol. 70(4) Costa Rica and who has collected all 14 specimens known to this author from this region. Acknowledgment The author thanks Fred Ehrmann for making the four illustrations used in this paper. A great deal of patience and keen observational skills were required to illustrate male genitalia less than 1 mm in length. Literature Cited Adams, P. A. 1970. A review of the New World Dilaridae. Postilla, 148. PAN-PACIFIC ENTOMOLOGIST 70(4): 313-317, (1994) DEPOSITIONS OF PARASITIC HYMENOPTERA (INSECTA) TYPES FROM THE UNIVERSITY OF CALIFORIA, BERKELEY 1 Robert L. Zuparko 2 and Junji Hamai Laboratory of Biological Control, University of California, Berkeley Abstract.— In 1993, 71 hymenopteran type specimens from the University of California, Berke¬ ley’s Essig Museum were deposited in four other institutions: the California Academy of Sciences (San Francisco, California), the Queensland Museum (Brisbane, Australia), the Plant Protection Research Institute (Pretoria, South Africa) and the U.S. National Museum (Washington, D.C.). The specimens are Braconidae, Ichneumonidae, Scelionidae, Eucoilidae, Perilampidae, Encyr- tidae, Aphelinidae, Eulophidae, Trichogrammatidae, Mymaridae and Bethylidae, and represent 54 holotypes, 12 allotypes and 5 syntypes. The three holotype specimens of Bactropria brasiliensis Kieffer, Xyalopria ruficornis Kieffer and Ganaspis reclusa Kieffer are reported missing. Key Words.— Insecta, Hymenoptera, Parasitica, type specimens, depositions The collection of parasitic Hymenoptera at the Laboratory of Biological Control, University of California at Berkeley, was started in 1946 by R. L. Doutt. This collection is part of U.C. Berkeley’s Essig Museum (formerly the California Insect Survey), but is housed separately at the Gill Tract in Albany, California. The Essig Museum has a standing policy of depositing type material, belonging to Hyme¬ noptera and certain other taxa, in the California Academy of Science in San Francisco, California, on indefinite loan. Since 1947, a number of type specimens have accumulated at the Gill Tract. These specimens fall into three categories: those that were specifically designated for deposition at U.C. Berkeley, those designated for deposition at other insti¬ tutions (either in their published descriptions or as notations attached to the specimens) but were never transferred for various reasons, and those with no specific instructions for deposition. In 1993, an effort was made to rectify this situation. All the type specimens in hand at the Gill Tract (54 holotypes, 12 allotypes and 5 syntypes) were transferred (type specimens then on temporary loan to other workers were retained). As per their specifications, ten specimens were deposited in the Plant Protection Research Institute (Pretoria, South Africa) (PPRI), one in the Queensland Museum (Bris¬ bane, Australia) (QM), and seven in the United States National Museum (Wash¬ ington, D.C.) (USNM). The remaining 53 types were deposited in the California Academy of Sciences (CAS). These taxa are listed below, alphabetically by specific name under each family. The receiving institutions are indicated by acronyms, and the specimens sent to the CAS have their new type number appended. Three specimens (Bactropria brasiliensis Kieffer, Xyalopria ruficornis Kieffer [both Hymenoptera: Diapriidae] and Ganaspis reclusa Kieffer [Hymenoptera: Eucoilidae]) were missing from their pins and labels before their transfers. 1 Authors’ page charges were partially offset by a grant from the C. P. Alexander Fund, PCES. 2 1050 San Pablo Avenue, Albany, California 94706. 314 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Braconidae amelanchieri, Aphidius Liu (1977). Holotype, female, pinned (missing abdomen and antennal segments). —CAS (#17035) amphorophori, Aphidius Liu (1977). Holotype, female, pinned (missing abdomen and forewings).—CAS (#17036) lineatellae, Hahrohracon Fischer (1968). Holotype, female, pinned (missing an¬ tennal segments). —CAS (#17043) liriodendrii, Aphidius Liu (1977). Holotype, female, pinned (missing abdomen).— CAS (#17037) lupini, Aphidius Liu (1977). Holotype, female, pinned (missing abdomen and antennal segments).—CAS (#17038) masonaphis, Aphidius Liu (1977). Holotype, female, pinned (tip of abdomen broken). —CAS (#17039) montereyensis, Aphidius Liu (1977). Holotype, female, pinned (missing abdomen, 3 legs and 1 antenna; head mounted on point). —CAS (#17040) proia, Alysia (Anarcha ) Wharton (1988). Holotype, female, pinned (missing an¬ tennal segments).—CAS (#17033) pteridis, Aphidius Liu (1977). Holotype, female, pinned (missing abdomen and 1 forewing).—CAS (#17041) vaccinii, Aphidius Liu (1977). Holotype, female, pinned (missing 1 forewing and antenna, 1 antenna broken). —CAS (#17042) vespertina, Alysia (Anarcha) Wharton (1988). Holotype, female, pinned.—CAS (#17034) ICHNEUMONIDAE badius, Homotropus Dasch (1964). Holotype, female, pinned. —CAS (#17032) labratus, Erromenus (Erromenus) Townes & Townes (1949). Holotype, female, pinned. —CAS (#17031) Diapriidae brasiliensis, Bactropria Kieffer (1909a). Pin & label only, missing specimen! — CAS (#17051) ruficornis, Xyalopria Kieffer (1909a). Pin & label only, missing specimen!—CAS (#17052) SCELIONIDAE erythropus, Macroteleia Cameron (1907). Holotype, male, pinned (missing ab¬ domen).-CAS (#17047) paraensis, Macroteleia Kieffer (1909a). Holotype, male, pinned (missing abdo¬ men).-CAS (#17048) rufitarsis, Pentacantha Kieffer (1906). Holotype, female, pinned (missing abdo¬ men, wings in poor condition). —CAS (#17049) rufitarsis, Trissolcus Kieffer (1906). Syntype, female, pinned (missing 3 legs and 1 antenna, head mounted on point).—CAS (#17050) Eucoilidae clarimontis, Cothonaspis (Pentarhoptra ) Kieffer (1909b). Holotype, female, pinned.-CAS (#17045) 1994 ZUPARKO & HAMAI: DEPOSITIONS OF TYPES 315 reclusa, Ganaspis Kieffer (1908). Pin and label only, missing specimen! —CAS (#17046) striatipennis, Caleucoela Kieffer (1909b). Holotype, female, pinned.—CAS (#17044) Perilampidae hesperis, Chrysomalla Darling (1986). Holotype, female, pinned. —CAS (#17059) Encyrtidae dahlsteni, Avetianella Tijapitzin (1971). Holotype, female, on slide.—CAS (#17065) inconspicuus, Anthemus Doutt (1966). Holotype, female and allotype, male, on slides.-CAS (Holotype = #17064) plethoricus, Pentalitomastix Caltagirone (1966). Holotype, female, on slide.—CAS (#17137) saipanensis, Anagyrus Doutt (1952). Syntypes, female and male, on slides (spec¬ imens labelled on slides as holotype female and allotype male, but not so designated in the published description).—CAS (female = #17062) smithi, Anagyrus Doutt (1952). Syntypes, female and male, on slides (specimens labelled on slides as holotype female and allotype male, but not so designated in the published description).—CAS (female = #17063) tanytmemus, Copidosomopsis Caltagirone (1985). Holotype, female, on slide.— CAS (#17136) Aphelinidae comperei, Coccophagoides Doutt (1966). Holotype, female, on slide. —CAS (#17060) utilis, Coccophagoides Doutt (1966). Holotype, female and allotype, male on slides.-CAS (#17061) Eulophidae glaber, Pnigalio Yoshimoto (1983). Holotype, female, pinned.—CAS (#17058) T RICHOGRAMM ATIDAE anneckei, Xiphogramma Doutt (1974a). Holotype, female and allotype, male, on slides.—PPRI californica, Paratrichogramma Doutt (1973). Holotype, female and allotype, male, on slides.-CAS (#17073) fletcheri, Oligositoides Doutt (in Doutt & Viggiani 1968). Holotype, female and allotype, male, on slides.—PPRI kusaiensis, Oligosita Doutt (1955). Holotype, female, on slide.—USNM occidentalis, Chaetogramma Doutt (1974a). Holotype, female and allotype, male, on slides.-CAS (#17072) oceanica, Oligosita Doutt (1955). Holotype, female, on slide.—USNM pacifica, Lathromeris Doutt (1955). Holotype, female, on slide.—USNM pallida, Epoligosita Doutt (in Doutt & Viggiani 1968). Holotype, female, on slide.—PPRI pretoriensis, Chaetogramma Doutt (1974a). Holotype, female, on slide.—PPRI pretoriensis, Paratrichogramma Doutt (1973). Holotype, female and allotype, male, on slides.—PPRI 316 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) salutaris, Brachygrammatella Doutt (1968). Holotype, female and allotype, male on slides. —PPRI Mymaridae annulatum, Stethynium Doutt (1947). Holotype, female, on slide.—CAS (#17071) australiensis, Dahmsia Doutt (1975). Holotype, female, on slide. —QM clarkei, Arescon Doutt (1955). Holotype, female, on slide.—USNM conferta, Anaphoidea Doutt (1949a). Holotype, female, on slide.—CAS (#17134) enocki, Dicopus Doutt (1974b). Holotype, female, on slide.—CAS (#17067) gerrisophaga, Anaphoidea Doutt (1949a). Holotype, female, on slide. —CAS (#17135) flandersi, Erythmelus Doutt (1949b). Holotype, female and allotype, male on slides.—CAS (#17069) gressitti, Nesetaerus Doutt (1955). Holotype, female, on slide.—USNM medicae, Barypolynema Annecke & Doutt (1961). Holotype, female and allotype, male on slides. —CAS (#17066) pygmaeus. Dicopus Doutt (1974b). Holotype, female and allotype, male on slides.— CAS (#17068) rete, Ptilomymar Annecke & Doutt (1961). Holotype, female, on slide. —CAS (#17070) saipanensis, Lymaenon Doutt (1955). Holotype, female and allotype, male on slide.-USNM Bethylidae bifoveatus, Dissomphalus Kieffer (1906). Holotype, male, pinned. —CAS (#17053) brasiliensis, Dissomphalus Kieffer (1909a). Holotype, male, pinned.—CAS (#17054) brasiliensis, Rhabdepyris Kieffer (1909a). Holotype, male, pinned.—CAS (#17056) paraensis, Rhabdepyris Kieffer (1909a). Holotype, male, pinned (missing 1 an¬ tennae).-CAS (#17057) rufipalpis, Dissomphalus Kieffer (1910). Holotype, male pinned.—CAS (#17055) Acknowledgment Ken Hagen and Leo Caltagirone (of the Laboratory of Biological Control, Uni¬ versity of California at Berkeley) provided immeasureable help in tracking down the appropriate literature and ensuring the completion of this project. Major improvements in the ms. were suggested by two anonymous reviewers. All the Kieffer specimens (Eucoilidae, Scelionidae, Diapriidae and Bethylidae) were orig¬ inally obtained from Pomona College by R. L. Doutt. Literature Cited Annecke, D. P. & R. L. Doutt. 1961. The genera of the Mymaridae (Hymenoptera: Chalcidoidea). Rep. S. Afr. Dept. Agr. Tech. Serv. Mem., 5. Cameron, P. 1907. Algunos Himenopteros. Ann. Estac. Agron. Cuba., 277. Caltagirone, L. E. 1966. A new Pentalitomastix from Mexico (Hymenoptera: Encyrtidae). Pan-Pacif. Entomol., 42: 145-151. Caltagirone, L. E. 1985. New Copidosomopsis (Hymenoptera: Encyrtidae) from California, with comments on the genus. Ann. Entomol. Soc. Am., 78: 705-708. 1994 ZUPARKO & HAMAI: DEPOSITIONS OF TYPES 317 Darling, D. C. 1986. Revision of the New World Chrysolampinae (Hymenoptera: Chalcidoidea). Can. Entomol., 118: 913-940. Dasch, C. E. 1964. Ichneumon-flies of America north of Mexico. 5. Subfamily Diaplazontinae. Mem. Amer. Entomol. Inst., 3. Doutt, R. L. 1947. The occurrence of the genus Stethynium in California. Pan-Pacif. Entomol., 32: 152-154. Doutt, R. L. 1949a. A synopsis of North American Anaphoidea (Hymenoptera: Mymaridae). Pan- Pacif. Entomol., 25: 155-160. Doutt, R. L. 1949b. The genus Erythmelus in California (Hymenoptera: Mymaridae). Pan-Pacif. Entomol., 25: 77-81. Doutt, R. L. 1952. Two new species of Anagyrus (Hymenoptera: Encyrtidae). Proc. Haw. Entomol. Soc., 14: 399-402. Doutt, R. L. 1955. Insects of Micronesia, Hymenoptera: Trichogrammatidae and Mymaridae. Insects of Micronesia, Bishop Museum, Honolulu, 19. Doutt, R. L. 1966. A taxonomic analysis of parasitic Hymenoptera reared from Parlatoria oleae (Colvee). Hilgardia, 37:219-231. Doutt, R. L. 1968. The genus Brachygrammatella Girault (Hymenoptera: Trichogrammatidae). Pan- Pacif. Entomol., 44: 289-294. Doutt, R. L. 1973. The genus Paratrichogramma Girault (Hymenoptera: Trichogrammatidae). Pan- Pacif. Entomol., 49: 192-196. Doutt, R. L. 1974a. Chaetogramma, a new genus of Trichogrammatidae (Hymenoptera: Chalci¬ doidea). Pan-Pacif. Entomol., 50: 238-242. Doutt, R. L. 1974b. The genus Dicopus Enock (Hymenoptera: Mymaridae). Pan-Pacif. Entomol., 50: 165-168. Doutt, R. L. 1975. Dahmsia, a new genus of Mymaridae (Hymenoptera: Chalcidoidea). Pan-Pacif. Entomol., 51: 254-256. Doutt, R. L. & G. Viggiani. 1968. The classification of the Trichogrammatidae (Hymenoptera: Chalcidoidea). Proc. Calif. Acad. Sci., 35: 477-586. Fischer, M. 1968. Ubergezuchtete raupenwespen (Hymenoptera, Braconidae). Pflanzenschutz Ber., 37: 97-140. Kieffer, J. J. 1906. Beschreibung neuer Proctotrypiden aus Nord- und Zentralamerika. Entomol. Zeit. Ber., 50: 237-290. Kieffer, J. J. 1908. Nouveaux Proctotrypides et Cynipides d’Amerique. Ann. Soc. Sci. Bruxelles, 32: 7-66. Kieffer, J. J. 1909a. Nouveaux microhymenopteres du Bresil. Ann. Soc. Entomol. France, 78: 287- 348. Kieffer, J. J. 1909b. Description de nouveaux Cynipides zoophages. Bull. Soc. Hist. Nat. Metz, 26: 57-96. Kieffer, J. J. 1910. Description de nouveaux Bethylides (Hymen.). Ann. Soc. Entomol. France, 79: 31-56. Liu, C. G. 1977. Genus Aphidius (Aphidiidae, Hymenoptera) of California. Taiwan Agric. Res. Inst. Taipei. Spec. Pub., 11. Townes, H. K. & M. C. Townes. 1949. A revision of the genera and of the American species of Tryphonini (Hymenoptera: Ichneumonidae). Ann. Entomol. Soc. Am., 42: 321-395. Trjapitzin, Y. A. 1971. A Nearctic representative of the genus Avetianella Trjapitzin, 1968 (Hy¬ menoptera, Encyrtidae). Entomol. Rev., 50: 507-508. Wharton, R. A. 1988. The braconid genus Alysia (Hymenoptera): a revision of the subgenus Anarcha. Contrib. Am. Entomol. Inst., 25. Yoshimoto, C. M. 1983. Review of North American Pnigalio Schrank (Hymenoptera: Eulophidae). Can. Entomol., 115: 971-1000. PAN-PACIFIC ENTOMOLOGIST 70(4): 318-321, (1994) A REVIEW OF OCHRERIADES (HYMENOPTERA: MEGACHILIDAE: OSMIINI) Terry L. Griswold USDA-ARS Bee Biology & Systematics Lab, Utah State University, Logan, Utah 84322-5310 Abstract. — A new Ochreriades is described from Namibia, O. rozeni NEW SPECIES. New records are presented for the only other known species, O. fasciatus Friese, restricted to the eastern Mediterranean. The disjunct distributional pattern exemplified by Ochreriades is uncommon among megachilids. The inclusion of this maculated genus in the Osmiini is confirmed and its relationship to other genera discussed. Key Words. — Insecta, Hymenoptera, Ochreriades rozeni'NEW SPECIES, Megachilidae The genus Ochreriades Mavromoustakis 1956 was erected for a single atypical species then placed in Heriades Spinola, H. fasciatus Friese 1899, which Mav¬ romoustakis recognized as having close relationship to Chelostoma Latreille. Here the generic relationships of Ochreriades are discussed, a new species is described from southwestern Africa, and new records presented for O. fasciatus (Friese). Though Ochreriades superficially resembles Anthidiini due to the light macu- lations of the body, Mavromoustakis (1956) was correct in placing it near Che¬ lostoma. Chelostoma has long been associated with Heriades, and was at one time considered a subgenus of the latter, but it is, as Peters (1978) has suggested, not closely related to Heriades and is rightly excluded from the Heriades complex (in which Peters included the genera Heriades, Protosmia, Othinosmia, Chelosto- mopsis, Noteriades, Pseudoheriades, and Archeriades). Ochreriades likewise should be excluded from this group. It differs from the Heriades complex in a number of key characters. The male has T7 exposed, transverse swellings on S2-3, S4 with dense discal pubescence, and S5 not emarginate and without modified hair. Females lack the apical hair tuft on the labrum and T6 has neither a preapical carina nor a wide hyaline flange. Shared derived characters held in common with Chelostoma include an elongate scutum, male S4 with apical hyaline flaps (some¬ times very narrow), female clypeus not overhanging labrum, and female labrum without hair fringe. It seems likely that both Chelostoma and Ochreriades were derived from a Hop litis-like ancestor. They share with such subgenera of Hoplitis as Alcidamea Cresson and Liosmia Thomson the preapical pit on male T7 and a medial swelling on male S2, both characters not found elsewhere in the Megachilidae. In the description terga are numbered T1, T2,. . . , sterna, likewise. Depositories of specimens are indicated parenthetically in data citations by names of cities (see acknowledgment). Ochreriades fasciatus (Friese) Eriades fasciatus Friese 1899. Entomol. Nachr., 21:325. Discussion.— Friese (1899) described O. fasciatus from a single male collected at Jericho, 16 Apr 1899. Alfken (1935) recorded a female from Wadi el Kelt, but 1994 GRISWOLD: A REVIEW OF OCHRERIADES 319 Figure 1. Dorsal view female Ochreriades rozeni. did not present a description of the female. Mavromoustakis (1939) recorded two males from Jerusalem, collected in early April, and later (Mavromoustakis 1956) described the female from a single specimen collected from “Mezze of Damascus” in early July. 320 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) New Records.—SYRIA. “Damas Hattite,” 2/18 May 1960, J. de Beaumont, 3 males, 1 female, (Lausanne); “Umm es Charatite,” 19 Jun 1945, K. P. Whitehom, 1 female (London). OCHRERIADES ROZEN1, NEW SPECIES (Fig. 1) Type.-Holotype female. SOUTHWEST AFRICA. 53 km SE of Omaruru, 13 Mar 1979, J. G. & B. L. Rozen. Holotype deposited in American Museum of Natural History, New York. Female. — Length, 10 mm; forewing length, 7 mm. Body black except antenna brown, mandible red subapically, apical tarsomeres yellow-red, linear transverse white maculations subapically on Tl-4, that on T1 very narrowly interrupted medially. Wings hyaline, venation dark brown. Head length equal to width; mandible narrowly tridentate; mouthparts extremely long, reaching well past base of abdomen in repose (apical part lost in holotype); occipital margin carinate; hypostomal area polished, very sparsely punctate. Pronotum with narrow raised, rounded collar medially, pronotal shoulder pronounced but not carinate; scutum and scutellum with very fine dense punctation, punctures one- half the size of those on mesopleuron; axilla produced posterolaterally to a curved point; propodeum without horizontal basal pitted zone; posterior coxa with punctures moderate-sized except absent on maculated areas and fine along posterior margins; T6 with apical subtruncate flange; scopa short, white. Male. — Unknown. Diagnosis. — Ochreriades rozeni is abundantly distinct from O. fasciatus. Prom¬ inent differences include lack of maculations on head and thorax, carinate occipital margin, noncarinate pronotum, fine punctation of thoracic dorsum, and angled axilla. The tongue of this species is remarkably long (unfortunately, its full extent is unknown because the apical portion is lost in the only known specimen). Discussion. — The presence of Ochreriades in southwestern Africa was unantic¬ ipated. Other components of the osmiine fauna from this region are either endemic to southern Africa or are widespread in the paleotropics. The only exception is Hoplitis (Anthocopa) Lepeletier, which also appears to have a disjunct distribution. It is broadly distributed across southern Africa north to southern Zaire and Ma¬ lawi, but is most numerous in the palearctic from the western Mediterranean to central Asia and southern India with its center of diversity in the Mediterranean region. It is not known from East Africa, but may be present along the Rift Valley. Material Examined. —See types. Acknowledgment Thanks to D. Cherix, Musee Zoologique, Lausanne; G. Else, British Museum of Natural History, London; and M. Fauveau and J. Rozen, American Museum of Natural History, New York, for loan of material. Travel to study specimens in European museums was funded by National Science Foundation Doctoral Dissertation Improvement Grant B5R-8313307. Greg Frehner produced the il¬ lustration. This is a contribution from Utah Agricultural Experiment Station, Utah State University, Logan, UT 84322-4810, Journal Paper No. 4401, and USDA-ARS Bee Biology and Systematics Laboratory, Utah State University, Logan, UT 84322-5310. Literature Cited Alfken, J. D. 1935. Bietrag zur kenntnis der Bienenfauna von Palastina. Veroff. Dtsch. Kol. Mus. Bremen, 1: 169-193. 1994 GRISWOLD: A REVIEW OF OCHRERIADES 321 Friese, H. 1899. Neue palaearktische Sammelbienen. Entomol. Nachr., 25: 321-346. Mavromoustakis, G. A. 1939. Some bees from Palestine. Ann. Mag. Nat. Hist., (11) 3: 225-230. Mavromoustakis, G. A. 1956. On the bees of Siria. Part I. Eos, 32: 215-229. Peters, D. S. 1978. Archeriades gen. n., eine verhaltnismassig urspriingliche Gattungder Megachilidae. Entomol. Germ., 4: 337-343. PAN-PACIFIC ENTOMOLOGIST 70(4): 322, (1994) Scientific Note NEW RECORD OF THE BEETLE, SCYMNUS FENDERI MALKIN (COLEOPTERA: COCCINELLIDAE), FROM DIURAPHIS NOXIA (MORDVILKO) (HOMOPTERA: APHIDIDAE) Scymnus fenderi Malkin is reported for the first time as a natural enemy of the Russian wheat aphid, Diuraphis noxia (Mordvilko). I seeded spring barley at the USD A—Agricultural Research Service Plant Materials Introduction Center at Central Ferry, Washington in May 1992. Weekly counts of Russian wheat aphid were made and all parasitoid mummies and predators encountered were recorded and retained. One adult female S. fenderi was recovered from the plot on 24 June 1992. Scymnus fenderi is an endemic species. No other specimens of S. fenderi have been collected to date. Scymnus fenderi differs from Scymnus frontalis Fabr. in having black elytra, whereas S. frontalis has a red spot on each elytron. Scymnus frontalis was released en mass in 1991 at a site near Central Ferry by the USD A— Animal Plant Health Inspection Service Plant Protection & Quarantine. No spec¬ imen of S. frontalis has been recovered in Washington since that release. The specimen of S. fenderi recovered was feeding on Russian wheat aphids located between the flag leaf sheath and stem of a barley tiller. Record. —WASHINGTON. GARFIELD Co.: Central Ferry, May 1992, 24 June 1992. David E. Bragg, Cooperative Extension, Washington State University, Pomeroy, Washington 99347-0190. PAN-PACIFIC ENTOMOLOGIST 70(4): 322-323, (1994) Scientific Note NOTES ON PARASITOIDS OF PLATYPTILIA CARDUIDACTYLA (RILEY) (LEPIDOPTERA: PTEROPHORIDAE) IN TRANSITION ZONE SOUTHEASTERN WASHINGTON Parasitoids and adult artichoke plume moths were reared from prepupal larvae and pupae collected in 1990 and 1991 from bull thistle, Cirsium vulgare L., growing in a pasture near Pomeroy, Washington. Larvae and pupae dissected from thistle tissue were placed on cut pieces of thistle stem in organdy screen cages of 34 cm x 46 cm with a sleeve to allow access. Fresh pieces of thistle stem were supplied as needed until all larvae in a cage pupated or parasitoids emerged from 1994 SCIENTIFIC NOTE 323 them. Adult plume moths were counted, separated by sex, and released back into the environment. Parasitoid adults were aspirated upon emergence, placed in alcohol, and retained for identification. Collections were made weekly in this manner from May through September of both years. Six species of primary par¬ asitoid were reared from the artichoke plume moth in this location over the two year period: Bracon hyslopi (Viereck) (Braconidae: Braconinae: Braconini); Cal- liephialtes notandus (Cresson) (Ichneumonidae: Ephialtinae: Pimplini); Campo- plex polychrosidis Viereck (Ichneumonidae: Porizontinae: Campoplegini); Diadeg- ma acuta (Viereck) (Ichneumonidae: Porizontinae: Porizontini); Colpognathus helvus (Cresson) (Ichneumonidae: Ichneumoninae: Alomyini); and Phaeogenes cynarae Bragg (Ichneumonidae: Ichneumoninae: Alomyini). Two species of sec¬ ondary parasitoid were reared as well: Gelus sp. (Ichneumonidae: Gelinae: Gelini); and Catolaccus aeneoviridis (Girualt) (Pteromalidae: Pteromalinae: Pteromalini). The two Alomyine Ichneumonids are apparently specific to the artichoke plume moth (Bragg, D. E. 1971. Pan-Pacific Entomol., 47: 57-62). Phaeogenes and Diadegma were reared in substantial numbers both years throughout the period of plume moth activity. Records.— WASHINGTON. GARFIELD Co.: Pataha Crk, 1 km E of Pomeroy, ex Platytilia car- duidactyla. David E. Bragg, Department of Entomology, Washington State University, P.O. Box 190, Pomeroy, Washington 99347-0190. PAN-PACIFIC ENTOMOLOGIST 70(4): 323-327, (1994) Scientific Note THE REVIVAL OF RICE-FIELD GRASSHOPPERS AS HUMAN FOOD IN SOUTH KOREA Grasshoppers have been a common food for people in many parts of the world (Bodenheimer, F. S. 1951. Insects as human food, W. Junk, The Hague). Rice- field grasshoppers (Acrididae, Oxya spp.) are eaten in most east Asian countries. In Korea, these grasshoppers are called metdugi and were a common food eaten as a side dish at meals, as a lunch box ingredient and as a drinking snack (K. S. Woo, personal communication). The use of rice-field grasshoppers declined during the 1960s and 1970s with increased insecticide use. I observed Koreans gather, pan-fry and eat metdugi during a picnic in October 1989. Samples were subsequently identified as Oxya velox (Fabr.). During four years (1989-1992) of visiting the Seoul markets, I did not see any metdugi, where they were once common (G. S. Yun, personal communication), and where silk moth pupae (Bombyx mori L.), a human food that is a by-product of the Korean silk industry, are almost always present. Some metdugi were sold in 1989 in a beer hall in the city of Suwon; a dish of about 20 was 5000 Won (US $7.57) (K. 324 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Figure 1. Korean rice-field grasshoppers (Oxya spp.) prepared for the table. S. Woo, personal communication). In 1990, a discount store for Korean Gov¬ ernment employees sold dried metdugi seasoned with soy sauce, sugar and sesame oil, in 130 g packages for 3750 Won (US $5.20). I was able to contact J.-R. Lee the president of the small food company that marketed these metdugi. He said that the metdugi were gathered in areas away from agricultural fields (and insecticides), such as in mountains and even in the outer DMZ military security zone, where agriculture is limited. The farmer-col¬ lectors sold their metdugi at local five-day markets (open one day every five days), where his company would purchase them. Some were also obtained earlier in the 1980s from artificial rearing. Rearing of metdugi proved difficult because of in¬ secticide contamination of the food and water used. The metdugi also were very sensitive to the carbon monoxide gas produced by charcoal heaters. These prob¬ lems, and increased labor costs, ended the commercial rearing attempts. Mr. Lee’s business declined because of the lack of metdugi, and in 1990 it ended. The metdugi food culture of Korea had become rare and seemed to be disappearing. Then on 8 Oct 1990, the Korean language newspaper Chungang Ilbo published an article by Huh Sang-Chun titled “The metdugi revival.” It described the rebirth of metdugi gathering and selling in Kyungsang Namdo, a province in the southern part of the country. Shortly after the article appeared, I visited the center of this revival, Chahwang Myun ( a district of Sanchung County), where I interviewed the Chahwang Myun Agricultural Cooperative Manager Park Chung-Ki, and two local farm women, Im Pun-Nam and Kim Ssang-Soon, who were active metdugi collectors. 1994 SCIENTIFIC NOTE 325 Chahwang is a small district with about 3000 people, most of whom belong to 744 farm families that cultivate 642 ha of paddy rice (1990 figures). Before in¬ secticide use intensified in the 1960s, metdugi were abundant in and around the rice fields and were collected for both home use and sale. The elevation of Chah¬ wang Myun is from 380 to 420 m, so it has cooler nights and consequently fewer problems with rice pests than areas at lower elevations. Despite this situation, the farmers could not avoid the government policy requiring at least three sprays per season. (Some Korean entomologists that I subsequently spoke with doubted that there was a government policy requiring the spraying of rice, despite the Chahwang-Myun people’s statements about such a policy.) In 1981 the rules mandating insecticide use loosened and farmers started using less, which allowed the metdugi populations to begin to increase. In 1982 some metdugi began to be collected and sold again in the local market at Sanchon. The decline in insecticide use and the desire of some Koreans to eat pesticide- free rice led to the development of organic rice farming in Chahwang Myun. This was economically viable because the yields of rice were the same in unsprayed fields as in sprayed fields, and organic rice sold (and still sells) for higher prices. In 1989, the Chahwang Agricultural Cooperative, which functions primarily to buy, mill and sell rice, began to buy dried metdugi from the farmer-collectors. In that year, more than 600 liters were purchased from more than 300 families. The farmers earned 4000 Won (US $6.06) per liter. The Cooperative sold the metdugi in bulk for 4250 Won per liter (US $6.44). The farmers probably sold another 600 liters at the five-day market and on the street. In 1990, more than 600 families (out of 744) sold 1744 liters of metdugi to the Cooperative at 5000 Won per liter (US $6.98). The Cooperative sold them for 6500 Won per liter (US $9.08). Much of the 1990 sale went to a supermarket company in Pusan, which divided the metdugi into 0.2 liter packages and sold these for 3000 Won (US $4.19). Metdugi were also sold by mail-order and to out-of-town visitors to the Cooperative. In 1990, the average collector sold 2 liters of metdugi to the Cooperative, but some collectors brought in as much as 40 liters, and one man, who had no rice field to tend, sold 160 liters to the Cooperative. Metdugi are most commonly collected by older women, usually from mid October to early November. They are collected by hand primarily from rice fields until the rice is harvested, then some are taken from other crops (such as dry beans) and from wild vegetation in the surrounding mountains. The average collection rate is about 0.25 liter per hour, while the best rate is 1.0 liter per hour. Both Mrs. Im and Mrs. Kim collect for 15 days each year on a part-time basis. Collected metdugi are steamed or boiled, then dried in the sun for one day and in a room for two more days. In 1990, Mrs. Im (age 58) collected 100 liters, with the help of her husband, and sold 40 liters to the Cooperative. She sold most of the remainder at the five-day market and gave some to relatives. She was pleased to say that her city-dwelling grandchildren get metdugi in their lunch boxes. She has been collecting and selling metdugi for five years. Mrs. Kim (age 37) has been collecting and selling metdugi for eight years. She collected 80 liters in 1990. During 1990, the income per hour for collecting metdugi for these women ranged from 1250-5000 Won (US $1.75-6.98), excluding the time spent in processing and marketing the metdugi. The average 1990 income for farm households (3.8 people) was US $16,706 (Korean Ministry of Agriculture, Forestry and Fisheries. 326 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) 1992. Statistical Yearbook of Agriculture, Forestry and Fisheries) and many fam¬ ilies in hilly areas such as Chahwang Myun earned less. The added income from metdugi collection and sale was, then, significant to these families. Mrs. Im said “metdugi helps us live.” A one liter package of metdugi was purchased from the Cooperative and the grasshoppers in a subsample of about one third of a liter (149 insects) were identified. Three species were present. Oxya velox was the most common species, comprising 84.5% of the total, then Oxya sinuosa Mistshenko with 14.8%, and a single Acrida lata Motschulsky. Oxya velox is a yellow-green grasshopper 27-37 mm in length found in Japan, Korea, China and Taiwan (Cho, B. S. 1969. Illus¬ trated encyclopedia of fauna and flora of Korea, Vol. 10, Sam Hwa Pub. Co., Seoul). Oxya sinuosa is yellow-green, 30-38 mm long and occurs throughout Korea (Lee, H. S. and C. E. Lee. 1983. Nature and Life, Taegu, South Korea, 13: 1-23). Acrida lata is a large 54-89 mm grasshopper, green or grey-brown colored, which occurs in Japan, China and Taiwan as well as Korea (Cho 1969). I did not expect the A. lata in the sample, but it is one of the grasshoppers eaten in Korea (Jang Hoon Lee, personal communication). In 1991 and 1992, large numbers of metdugi continued to be bought and sold by the Chahwang Myun Cooperative and many people came to buy directly from the farmers (Min Pyung-Hong, personal communication). In 1992, the Cooper¬ ative bought metdugi for US $9.91 per liter and sold them at a bulk rate for US $12.03 per liter. Many Koreans consider metdugi to be a health food. Indeed, metdugi (probably Oxya spp.) have high levels of iron (43 mg/100 g), vitamin B2 (5.6 mg/100 g) and protein (64.2 g/100 g) (Chai, R. S., Y. Y. Yu, Y. H. Park, K. K. Kim, Y. J. Moon & H. H. Kwon. 1962. Reports National Chemistry Laboratories, Seoul, 10: 56-64). In Chahwang, metdugi is used to prevent and cure constipation and to treat heart problems. Metdugi {O. velox ) is used as a drug in traditional Korean medicine, prescribed to treat the convulsions of children, coughs, tetanus and weakness (Kim, J. G. 1984. Illustrated natural drugs encyclopedia. Nam San Dang Pub., Seoul). The food preparations of dried metdugi vary. Sometimes they are eaten dried without seasoning. They are usually pan-fried with or without oil after the wings and legs have been removed. During or after cooking, they are flavored with sesame oil and salt, or sesame oil and sugar, or soy sauce with or without sugar. I have also seen live ones fried whole. These turn red like shrimp as they cook. Many of these preparations produce a product with good snack food essence. They are bite-sized, crispy, crunchy, and salty and/or slightly sweet. Korean prep¬ arations of rice-field grasshoppers are, to my taste, much better than the sweet sticky Japanese preparations of Oxya that are sold in tins and restaurants in Japan as imago. For older Koreans, much of the appeal of eating metdugi are the feelings of nostalgia that it brings. Korea has undergone very rapid industrialization and urbanization during the past 25 years. The metdugi revival gives at least some people a chance to taste the past. Koreans and other east Asian people, in general, use and enjoy insects more than do Americans and Europeans (Pemberton, R. W. 1988. Pan-Pacific Entomol., 64: 81-82; 1990. 66: 93-95; 66: 172-174). 1994 SCIENTIFIC NOTE 327 Acknowledgment.— I thank the following people: Im Pun-Nam, Kim Ssang- Soon and Park Chung-Ki (Chahwang Myun), Lee Jang-Hoon (USDA-ARS, Asian Parasite Laboratory, Seoul), J.-R. Lee (Seoul), Woo Kun-Suk (Seoul National Univ.), and Yun Gye-Sup (Seoul) for providing information about metdugr, Lee Jang-Hoon, for identifying the metdugi samples and acting as a translator in Seoul; Kim Jeong-Sook (Asian Parasite Laboratory) for serving as a translator in Chah¬ wang Myun and Robert D. Macke (U.S. Embassy, Seoul) for information on Korean agricultural income. James B. Johnson, University of Idaho, and Douglas W. Whitman, Illinois State University, kindly provided reviews of the manuscript. Robert W. Pemberton, Asian Parasite Laboratory, United States Department of Agriculture-Agricultural Research Service. Current address: Aquatic Weed Con¬ trol Research, USDA ARS SAA, 3205 College Ave, Ft. Lauderdale, Florida USA 33314. PAN-PACIFIC ENTOMOLOGIST 70(4): 327-328, (1994) Scientific Note XYLOCORIS GALACTINUS (FIEBER) (HEMIPTERA: ANTHOCORIDAE) NEWLY DISCOVERED IN MONTANA STORED GRAIN Herein, we note the occurrence of the predaceous bug, Xylocoris galactinus (Fieber) in stored grain in Montana, one of the top four states in small grain production and storage (Montana Agricultural Statistics Service. 1991. Helena, Montana). Xylocoris galactinus has been introduced into the New World, where it often occurs in stored grain (J. A. Slater & R. M. Baranowski. 1978. How to Know the True Bugs. Wm. C. Brown Co. Dubuque, Iowa). It is reported trans- continentally in Canada (T. J. Henry & R. C. Froeschner. 1988. Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill Publ. Co. New York), but has not been recorded from the northern great plains of the United States (Henry & Froeschner 1988). California, Idaho and Missouri are the only states recorded to harbor this species west of the Mississippi River. Based on its distribution records, this species may be better able to survive in colder northern climes than does the better known Xylocoris flavipes (Reuter). During surveys of stored grain insects, F. Dunkel found an established popu¬ lation of X. galactinus at the Montana State University Southern Agricultural Research Center near Huntley, Montana. This population represents a significant range extension of over 320 km from the closest areas previously known to harbor the species in the Alberta and Idaho grain growing regions. The population was found in a 0.25 metric ton barley spill adjacent to grain storage bins. Within this spill, the population density of X. galactinus exceeded 200 immatures and >25 adults per kg of grain. The population was sampled by 328 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) sieving the grain every two weeks through Aug and Sep 1992, and a laboratory culture of X. galactinus was derived therefrom. Other insects found in the spilled grain, and potentially available as food for X. galactinus included: Trogoderma spp. (Dermestidae); grain beetles, Cryptolestes spp. (Silvanidae); hairy fungus bee¬ tles, Typhea stercorea (L.) (Mycetophagidae); picnic beetles, Carpophilus spp. (Nitidulidae); red flour beetles, Tribolium castaneum (Herbst) (Tenebrionidae); and larger black flour beetles, Cynaeus angustus (LeConte) (Tenebrionidae). Because X. galactinus is a beneficial insect exempted from tolerance by the U.S. Environmental Protection Agency and the U.S. Food and Drug Administration (Anonymous. 1992. Federal Register, 57, No. 78, April 22, 1992), it is a possible biocontrol agent for insects destructive to stored grain. Efforts are underway to adapt X. flavipes culture techniques to X. galactinus that may provide a better control option in colder climates. Record.—Montana. YELLOWSTONE CO.: nr Huntley, Aug/Sep 1992, F. Dunkel, ex. barley spill nr storage bins. Acknowledgment.— We thank T. J. Henry, U.S. Department of Agriculture- Systematic Entomology Laboratory, Washington D.C., for confirming the iden¬ tification of X. galactinus. This paper was supported by Montana Agricultural Experiment Station (MAES) Projects 156 (M. Ivie P.I.), 157 and 161 (F. Dunkel P.I.) and is contribution J-2830 of the Montana Agricultural Experiment Station. This is a contribution to Regional Project NC-151 Delivery and Marketing of Quality Grain and Oilseeds. Florence V. Dunkel and Michael A. Ivie, Department of Entomology, Montana State University, Bozeman, Montana 59717. PAN-PACIFIC ENTOMOLOGIST 70(4): 328-330, (1994) Scientific Note DESCRIPTION OF A SLEEPING AGGREGATION OF MALE CHALICODOMA CHILOPSIS (COCKERELL) (HYMENOPTERA: MEGACHILIDAE) The occurrence of male sleeping aggregations is characteristic of many species of aculeate Hymenoptera (Linsley, E. G. 1962. Ann. Entomol. Soc. Amer., 55: 148-164). The majority of these aggregations occur on “sleeping plants,” where the bees grasp a stem with mandibles and/or legs. The bees show a preference for dead or dry, relatively rigid, moderately tall (1 to 2 m), multibanched plants. This behavior has been recorded for many species of bees and wasps, but has been poorly documented for the diverse family Megachilidae. Linsley (1962) recorded two megachilid species ( Anthidiellum notatum robertsoni Cockerell and the clep- toparasite Coelioxys deplanata Cresson) as members of larger mixed species sleep- 1994 SCIENTIFIC NOTE 329 Figure 1. A male sleeping aggregation of the megachilid bee Chalicodoma chilopsis containing thirteen bees. ing aggregations. Sleeping Anthidiellum grasp stem tips with their mandibles only, and extend wings laterally. Adjacent stems may be occupied by individuals of this species. Sleeping Coelioxys individuals grasp the periphery of stems with their mandibles and the first two pairs of legs, fold their wings back over the body, and orient head-down on the stem. This species does not aggregate with conspecifics for sleeping. Osgood (unpublished) has been cited (Stephen, W. P., G. E. Bohart & P. F. Torchio. 1969. Oregon State Univ. Press, Corvalis, page 72) for a record of aggregating male Megachile rotundata (Fabr.) that returned to a cavity under the siding of a building for several weeks. On the evening of 21 Apr 1992 at 18:00 h (MST) in Tucson, Arizona, I noticed an aggregation of megachilid bees. The bees were resting on the apical end of a dead branch on an Alepo pine (Pinus halepensis Miller) approximately 3 m above ground. At this time, the bees were actively shifting positions along the branch, but activity ceased at sundown. The bees were arranged in a single layer, 360° around the stem from the apical end to approximately 15 cm up the stem. The 330 THE PAN-PACIFIC ENTOMOLOGIST Yol. 70(4) bees were in close proximity, often in direct contact. Each individual grasped the stem with mandibles and all legs, folded the wings back over the body and oriented the head upwards (Fig. 1). There were no other megachilid bees located in the vicinity of this tree. Before sunrise the next morning, I collected the cold and immobile bees by placing a plastic bag over the branch and shaking. Upon examination, I discovered that all 87 bees were males of the same species. The bees were identified as Chalicodoma chilopsis (Cockerell) by Terry Griswold of the USDA-ARS Bee Lab in Logan, Utah. Even though the entire aggregation was collected on 22 Apr, more individuals of the same species utilized the same branch on successive nights. The site was used continuously for a total of 20 nights by 10 to 40 bees. Occa¬ sionally there were smaller aggregations of up to three individuals on a branch in close proximity to the main aggregation. Thus, megachilid’s also exhibit the monospecific male sleeping aggregations typical of anthophorids. Most likely such aggregations are not rare, but have not been recorded due to the difficulty in finding a cluster, as well as correctly iden¬ tifying megachilid bees. Acknowledgment. — I thank Terry Griswold for species identification, Stephen L. Buchmann, James Cane, and E. Gorton Linsley for manuscript reviews. Steven C. Thoenes, USDA Agricultural Research Service, Carl Hayden Bee Research Center, 2000 E. Allen Rd., Tucson, Arizona, 85719. PAN-PACIFIC ENTOMOLOGIST 70(4): 330-332, (1994) Scientific Note FAN PALM AS AN URBAN NESTING SUBSTRATE FOR XYLOCOPA CALIFORNICA ARIZONENSIS CRESSON (HYMENOPTERA: ANTHOPHORIDAE) The city of Tucson, Arizona has been expanding into the desert for years. As a result, much of the native vegetation has been removed or severely reduced. Xylocopa californica arizonensis Cresson typically nests in the dried fruiting stalks (infructescenses) of Yucca, Agave, and Dasylirion, preferring stalks that are only 1-2 years old. These plants have been virtually eliminated from the native plant communities within the Tucson area, but do exist in ornamental plantings. Fruit¬ ing stalks are removed by homeowners after they have dried, due to their “un¬ sightly” nature. This has created shortages of nesting substrates within the city, however X. c. arizonensis persists in large numbers. Thus, we began to examine whether X. c. arizonensis has begun using ornamental plants or structural timbers as nesting substrates. We discovered large numbers of X. c. arizonensis associated with the fan palm [ Washingtonia filifera (Lindley) Wendland] which is not native to the Sonoran 1994 SCIENTIFIC NOTE 331 Figure 1. A fan palm infructescense containing a Xylocopa californica arizonensis nest. Notice the entrance hole at the base of the stalk. Desert. It is, however, used extensively as an ornamental plant. There have been reports (O’Brien, L. B. & C. W. O’Brien. 1966. Pan-Pac. Entomol., 42: 27-29) that X. c. arizonensis used fan palm fronds for nesting in California, where both the bee and plant are native in palm oases. Closer examination revealed that the bees were indeed using this plant as a nesting substrate in Tucson, but they were not using the fronds; instead they were using the dried fruiting stalks (Fig. 1). This use of dried fan palm infructescenses as a nesting substrate has not been previously reported. The nest morphology was similar to that reported by O’Brien & O’Brien (1966): entrance holes were located on the underside of the stalk, near the base, with the most of the gallery extending downward away from the trunk and only a short portion extending toward the trunk. The nests are typically much longer than those reported by O’Brien & O’Brien (1966) with one measuring 35 cm long and containing 15 cells. This difference can be explained by the morphology of the two substrates: the fronds are oblong in shape and flatten out much sooner than the round fruiting stalks. Apparently, as Tucson expanded and native nesting substrates were removed, the bees found and used the fan palm fruit stalks for nest sites. The fruit stalks of native plants (Yucca, Agave, and Dasylirion) and fan palm are similar: all are cylindrical with similar diameters (except that Agave stalks are typically much bigger), have hard exterior sheaths, and a softer pithy interior. Perhaps the use of fronds for nesting substrate over fruiting stalks is more derived, and could be due to the more abundant nest sites available (i.e., more fronds than fruiting stalks per palm). Thus, in California where palms and bees have occurred together for a longer period of time, nesting in the fronds may have 332 THE PAN-PACIFIC ENTOMOLOGIST Yol. 70(4) become the dominant behavior. Unfortunately, this hypothesis does not explain why no nests in fruiting stalks have been reported from California. An alternative explanation is that use of the fronds as nest sites is simply rare and the two fronds with nests described by O’Brien & O’Brien (1966) are exceptions rather than the usual pattern. Certainly the dichotomy between nest site selection in fan palm fronds versus fruiting stalks warrants further examination, and may provide in¬ sights into nest site selection criteria used by X. c. arizonensis females. Acknowledgment.— We thank E. Gorton Lindsley, Jim Cane, and two anony¬ mous reviewers for their suggestions on this manuscript. Steven C. Thoenes, and Stephen L. Buchmann, USDA Agricultural Research Service, Carl Hayden Bee Research Center, 2000 E. Allen Rd., Tucson, Arizona 85719. PAN-PACIFIC ENTOMOLOGIST 70(4): 333-334, (1994) The Pan-Pacific Entomologist Table of Contents for Volume 70 Bowles, J. M., see Starmer, W. T. 230 Bragg, D. E.—New record of the beetle, Scymnus fenderi Malkin (Coleoptera: Coccinellidae), from Diuraphis noxia (Mordvilko) (Homop- tera: Aphididae) . 322 Bright, D. E., see Hobson, K. R. 267 Buchmann, S. L., see Thoenes, S. C. 330 Carey, J. R., see Yang, P. ... 159,253,269 Cibrian-Tovar, J., see Rojas, J. C. 276 Clark, W. H ., see Merickel, F. W. 148 Coombs, E. M., see Turner, C. E. 206 D. E. Bragg—N otes on parasitoids of Platyptilia carduidactyla (Riley) (Lepidoptera: Ptero- phoridae) in transition zone southeastern Washington. 322 Daly, H. V.—Lectotype designations and holo- types for bees of the genus Hylaeus ( Neso- prosopis) described from the Hawaiian Is¬ lands (Hymenoptera: Colletidae) .... 113 Danforth, B. N.—Taxonomic review of Calli- opsis subgenus Hypomacrotera (Hymenop¬ tera: Andrenidae), with special reference on the distribution and host plant associations . 283 Dowell, R. V., see Yang, P. ... 159, 253, 269 Dreistadt, S. H. & K. S. Hagen—E uropean elm scale (Homoptera: Eriococcidae) abundance and parasitism in northern California. . 240 Duncan, R. W.—Bionomics and life history of the gall midge Chamaediplosis nootkatensis Gagne & Duncan (Diptera: Cecidomyiidae) on yellow cypress in British Columbia .... . 103 Dunkel, F. V. & M. A. I vie —Xylocoris galactinus (Fieber) (Hemiptera: Anthocoridae) newly discovered in Montana stored grain . . 327 Edwards, J. S., see Sugg, P. M. 212 Gast, S. J. & M. W. Stock—G enetic diversity in overwintered and non-overwintered Ips pini (Say) (Coleoptera: Scolytidae) in Idaho .... . 259 Gordh, G.—A biographical account of Harold Compere (1896-1978), biological control foreign explorer . 188 Greve, L., see Sugg, P. M. 212 Griswold, T. L.—A review of Ochreriades (Hy¬ menoptera: Megachilidae: Osmiini) ... 318 Hagne, K. S., see Dreistadt, S. H. 240 Hamai, J., see Zuparko, R. L. 313 Hobson, K. R. & D. E. Bright— A key to the Xyleborus of California, with faunal com¬ ments (Coleoptera: Scolytidae) . 267 Ichinose, K. — Limited multiple-mating in males and single-mating in females of the ant spe¬ cies, Paratrechina flavipes (Fr. Smith) (Hy¬ menoptera: Formicidae). 183 Index, Pan-Pacific Entomologist, Volume 70 ... . 335 I vie, M. A., see Dunkel, F. V. 327 Joley, D. B., see Turner, C. E. 206 McCafferty, W. P., M. J. Wigle & R. D. Waltz—S ystematics and biology of Acen- trella turbida (McDunnough) (Ephemerop- tera: Baetidae). 301 Merickel, F. W. & W. H. Clark— Tetramorium caespitum (Linnaeus) and Liometopum luc- tuosm W. M. Wheeler (Hymenoptera: Formic¬ idae): new state records for Idaho and Oregon, with notes on their natural history .... 148 Moitoza, F. J.—A revision of the C. maculata species group of Conura spinola in America, north of Mexico, and a new species of the C. immaculata species group of Conura (Hy¬ menoptera: Chalcididae). 168 O’Neill, K. M.—Livestock dung as a food re¬ source and thermal refuge for rangeland grasshoppers (Orthoptera: Acrididae) . . 222 Pacific Coast Entomological Society: financial statement for 1990 and 1991 . ii Pacific Coast Entomological Society: funding an¬ nouncement for The Pan-Pacific Entomolo¬ gist, 70(1). i Pemberton, R. W.—The revival of rice-field grasshoppers as human food in Korea .... . 323 Penny, N.D.—A new species of Nallachius (Neu- roptera: Dilaridae) from Costa Rica . . 309 Piper, G. L., see Turner, C. E. 206 Polhemus, D. A.—An annotated checklist of the plant bugs of Colorado (Heteroptera: Miri- dae). 122 Rojas, J. C. & J. Cibrian-Tovar—R eproductive behavior of Copitarsia consueta (Walker) (Lepidoptera: Noctuidae): mating frequency, effect of age on mating, and influence of de¬ layed mating on fecundity and egg fertility . 276 Sobhian, R., see Turner, C. E. 206 Sorensen, J. T.—A revision of the aphid genus Essigella (Homoptera: Aphididae: Lachni- nae): its ecological associations with, and evolution on, Pinaceae hosts. 1 Starmer, W. T. & J. M. Bowles—T he spatial 334 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) distribution of endemic and introduced flow¬ er-breeding species of Drosophila (Diptera: Drosophiliadae) during their early history of encounter on the island of Hawaii ... 230 Stock, M. W., see Gast, S. J. 259 Sugg, P. M., L. Greve & J. S. Edwards— Neu- ropteroidea from Mount St. Helens and Mount Rainier: dispersal and immigration in volcanic landscapes . 212 Table of Contents, Pan-Pacific Entomologist, Volume 70 . 333 Thoenes, S. C—Description of a sleeping aggre¬ gation of male Chalicodoma chilopsis (Cock¬ erell) (Hymenoptera: Megachilidae) ... 328 Turner, C. E., R. Sobhian, D. B. Joley, E. M. Coombs & G. L. Piper— Establishment of Urophora sirunaseva (Hering) (Diptera: Te- phritidae) for biological control of yellow starthistle in the western United States .... . 206 Waltz, R. D., see McCafferty, W. P. .. 301 Wigle, M. J., see McCafferty, W. P. ... 301 Yang, P., C. Zhou, R. V. Dowell & J. R. Carey— Temperature studies on a Chinese strain of Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae) . 269 Yang, P., J. R. Carey & R. V. Dowell— Host- specific demographic studies of wild Bactro¬ cera tau (Walker) (Diptera: Tephritidae) ... . 253 Yang, P., J. R. Carey & R. V. Dowell— Te- phritid fruit flies in China: historical back¬ ground and current status . 159 Zhou, C., see Yang, P. 269 Zuparko, R. L. & J. Hamai— Depositions of par¬ asitic Hymenoptera (Insecta) types from the University of California, Berkeley ... 313 PAN-PACIFIC ENTOMOLOGIST 70 ( 4 ): 335 - 336 , ( 1994 ) Index to Volume 70 Title and Key Words, and New Taxa abundance 240 Acentrella turbida 301 Acrididae 222 age 276 Andrenidae 283 ant 183 Anthocoridae 327 Anthophoridae 330 Aonidella aurantii 188 Aphididae 322 Apoidea 283 Bactrocera 159, 253; B. cilifer 159; B. citri 159; B. cucurbitae 159, 269; B. diversa 159; B. dorsalis 159; B. latifrons 159; B. minax 159; B. occipitalis 159; B. scutellata 159; B. tau 159, 253; B. tsuneonis 159 Baetidae 301 biological control 159, 188, 206, 240 bionomics 103 black scale 188 British Columbia 103 California 267,240 California red scale 188 Calliopsis (Hypomacrotera) 283 Cecidomyiidae 103 Centaurea 206 Chalcididae 168 Chalicodoma chilopsis 328 Chamaediplosis nootkatensis 103 checklist 122 China 159 citrophilous mealybug 188 Coccinellidae 322 Coccophagus insidiator 240 coexistence 230 Coleoptera 259, 267, 322 Colletidae 113 colonization 212 Colorado 122 commodity treatment 269 Conura 168; C. dentiscapa NEW SPECIES 180; C. igneopatruelis NEW SPECIES 174; C. immaculata species group 168; C. maculata species group 168; C. pilosipartis NEW SPECIES 177 Copitarsia consueta 276 Costa Rica 309 county distributions 122 cultural control 159 damage 103 decomposition 222 demographics 269,253 Dillaridae 309 Diptera 103,159, 206, 230, 253, 269 dispersal 212 distribution 283 Diuraphis noxia 148, 322 Drosophila (Phloridosa) floricola 230 Drosophilidae 230 dung 222 Ephemeroptera 301 Eriococcidae 240 Eriococcus spurius 240 Essigella 18, 102; Essigella (Archeoessigella) NEW SUBGENUS 21; Essigella (Lambersella) NEW SUBGENUS 29; E. critchfieldi NEW SPECIES 75; E. eastopi NEW SPECIES 30; E. fusca voegtlini NEW SUBSPECIES 39; E. hillerislambersi NEW SPECIES 41 European elm scale 240 fan palm 330 fauna 267 fecundity 276 fertility 276 floral associations 283 flower breeders 230 food resources 222 foreign exploration 188 Formicidae 148, 183 Fruit flies 159 galls 103,206 gall midges 103 genetic diversity 259 genetics 259 George Compere 188 grasshoppers 222, 323 Harold Compere 188 Hawaii 230,327 Hemiptera 327 Heteroptera 122 heterozygosity 259 Hokkaido Japan 183 holotypes 113 Homoptera 240,322 host plants 283,122 host-specificity 253 human food 323 Hylaeus (Nesoprosopis) 113 336 THE PAN-PACIFIC ENTOMOLOGIST Vol. 70(4) Hymenoptera 113,148, 168, 183, 283, 313,318,328,330 Idaho 148 immigration 212 Ips pint 259 Japan 183 Korea 323 lectotype designations 113 Lepidoptera 276,322 life history 103, 253 Liometopum luctuosum 148 livestock 222 manure 222 mating 276 mating behavior 183 Megachilidae 318, 328 Miridae 122 Montana 327 morning glory 230 morphology 188 Mount Rainier 212 Mount St. Helens 212 multi pie-mating 183 Nallachius parkeri NEW SPECIES 309 natural history 148 Nesoprosopis 113 Neuroptera 212,309 Neuropteroidea 212 Noctuidae 276 Ochreriades 318, O. rozeni NEW SPECIES 320 Oregon 148 Orthoptera 222 parasitic Hymenoptera 188; types 313 parasitism 240 parasitoids 103 Paratrechina flavipes 183 pest ants 148 phenology 212 phylogenetic analyses 91,95 Pinus sp. 95 Platyptilla carduidactyla 322 Prosopis 113 Pseudococcus calceolariae 188 Pterophoridae 322 rangeland 222 Raphi diopter a 212 reproductive behavior 276 reproductive parameters 253 Saissetia oleae 188 Scaptomyza (Exalloscaptomyza) caliginosa 230 Scolytidae 259,267 Scymnus fenderi 322 sleeping aggregations 328 spatial distribution 230 species concepts 4 state records 148 stress 259 taxonomy 188 Temperature 269 Tephritidae 159, 206, 253, 269 Tetramorium caespitum 148 thermal refuges 222 thermoregulation 222 Trichomasthus coeruleus 240 University of California, Berkeley 313 Urophora sirunaseva 206 volcanic landscapes 212 Washington 322 weed 206 Xanthogaleruca luteola 240 Xyleborus 267 Xylocopa californica arizonensis 330 Xylocoris galactinus 327 yellow cypress 103 yellow starthistle 206 PAN-PACIFIC ENTOMOLOGIST Information for Contributors See volume 66(1): 1-8, January 1990, for detailed general format information and the issues thereafter for examples; see below for discussion of this journal’s specific formats for taxonomic manuscripts and locality data for specimens. Manuscripts must be in English, but foreign lan¬ guage summaries are permitted. Manuscripts not meeting the format guidelines may be returned. Please maintain a copy of the article on a word- processor because revisions are usually necessary before acceptance, pending review and copy-editing. Format. — Type manuscripts in a legible serif font IN DOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 X 11 in, non¬ erasable. high quality paper. THREE (3) COPIES of each manuscript must be submitted. EACH INCLUDING REDUCTIONS OF ANY FIG¬ URES TO THE 8.5 X 11 IN PAGE. Number pages as: title page (page 1), abstract and key words page (page 2), text pages (pages 3+), ac¬ knowledgment page, literature cited pages, footnote page, tables, figure caption page; place original figures last. List the corresponding author’s name, address including ZIP code, and phone number on the title page in the upper right corner. The title must include the taxon's des¬ ignation, where appropriate, as: (Order: Family). The ABSTRACT must not exceed 250 words; use five to seven words or concise phrases as KEY WORDS. Number FOOTNOTES sequentially and list on a separate page. Text. — Demarcate MAJOR HEADINGS as centered headings and MINOR HEADINGS as left indented paragraphs with lead phrases under¬ lined and followed by a period and two hyphens. CITATION FORMATS are: Coswell (1986). (Asher 1987a, Franks & Ebbet 1988, Dorly et al. 1989), (Burton in press) and (R. F. Tray, personal communication). For multiple papers by the same author use: (Weber 1932, 1936, 1941; Sebb 1950. 1952). For more detailed reference use: (Smith 1983: 149-153, Price 1985: fig. 7a. Nothwith 1987: table 3). Taxonomy. — Systematics manuscripts have special requirements outlined in volume 69(2): 194-198; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are applicable. These re¬ quirements include SEPARATE PARAGRAPHS FOR DIAGNOSES, TYPES AND MATERIAL EXAMINED (INCLUDING A SPECIFIC FORMAT), and a specific order for paragraphs in descriptions. List the unabbreviated taxonomic author of each species after its first mention. Data Formats. — All specimen data must be cited in the journal's locality data format. See volume 69(2), pages 196-198 for these format re¬ quirements: if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before submitting manu¬ scripts for which these formats are applicable. Literature Cited. — Format examples are: Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York. Blackman, R. L., P. A. Brown & V. F. Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometries provide some answers? pp. 233-238. In Holman. J., J. Pelikan, A. G. F. Dixon & L. Weismann (eds.). Population structure, genetics and taxonomy of aphids and Thysanoptcra. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14. 1985. SPB Academic Publishing, The Hague, The Netherlands. Ferrari. J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899. Sorensen. J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol. Illustrations. — Illustrations must be of high quality and large enough to ultimately reduce to 117 X 181 mm while maintaining label letter sizes of at least 1 mm: this reduction must also allow for space below the illustrations for the typeset figure captions. 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Authors and affiliations are listed in the last, left indented paragraph of the note with the affiliation underscored. Page Charges. — PCES members are charged $35.00 per page, for the first 20 (cumulative) pages per volume and full galley costs for pages thereafter. Nonmembers should contact the Treasurer for current nonmember page charge rates. Page charges do not include reprint costs, or charges for author changes to manuscripts after they are sent to the printer. Contributing authors will be sent a page charge fee notice with ac¬ knowledgment of initial receipt of manuscripts. Volume 70 THE PAN-PACIFIC ENTOMOLOGIST October 1994 Number 4 Contents GAST, S. J. & M. W. STOCK—Genetic diversity in overwintered and non-overwintered Ips pini (Say) (Coleoptera: Scolytidae) in Idaho -..... 259 HOBSON, K. R. & D. E. BRIGHT—A key to the Xyleborus of California, with faunal com¬ ments (Coleoptera: Scolytidae).... 267 YANG, P„ C. ZHOU, G. LIANG, R. V. DOWELL & J. R. CAREY—Temperature studies on a Chinese strain of Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae)-__ 269 ROJAS, J. C. & J. CIBRIAN-TOVAR—Reproductive behavior of Copitarsia consueta (Walk¬ er) (Lepidoptera: Noctuidae): mating frequency, effect of age on mating, and influence of delayed mating on fecundity and egg fertility... 276 DANFORTH, B. N.—Taxonomic review of Calliopsis subgenus Hypomacrotera (Hyme- noptera: Andrenidae), with special emphasis on the distributions and host plant asso¬ ciations........... 283 McCAFFERTY, W. P., M. J. WIGLE & R. D. WALTZ—Systematics and biology of Acentrella turbida (McDunnough) (Ephemeroptera: Baetidae)...... 301 PENNY, N. D.—A new species of Nallachius (Neuroptera: Dilaridae) from Costa Rica... 309 ZUPARKO, R. L. & J. HAMAI—Depositions of parasitic Hymenoptera (Insecta) types from the University of California, Berkeley___ 313 GRISWOLD, T. L.—A review of Ochreriades (Hymenoptera: Megachilidae: Osmiini) .. 318 SCIENTIFIC NOTES BRAGG, D. E.—New record of the beetle, Scymnus fenderi Malkin (Coleoptera: Coccinelli- dae), from Diuraphis noxia (Mordvilko) (Homoptera: Aphididae)... 322 BRAGG, D. E.—Notes on parasitoids of Platyptilia carduidactyla (Riley) (Lepidoptera: Pterophoridae) in transition zone southeastern Washington_ 322 PEMBERTON, R. W.—The revival of rice-field grasshoppers as human food in South Korea-— 323 DUNKEL, F. V. & M. A. IVIE —Xylocoris galactinus (Fieber) (Hemiptera: Anthocoridae) newly discovered in Montana stored grain.—.. 327 THOENES, S. C.-—Description of a sleeping aggregation of male Chalicodoma chilopsis (Cockerell) (Hymenoptera: Megachilidae).......... 328 THOENES, S. C. & S. L. BUCHMANN—Fan palm as an urban nesting substrate for Xylocopci californica arizonensis Cresson (Hymenoptera: Anthophoridae)_ 330 Table of Contents, Pan-Pacific Entomologist, Volume 70.-. 333 Index, Pan-Pacific Entomologist, Volume 70..... 335