75C.C HARVARD UNIVERSITY Library of the Museum of Comparative Zoology » V t SPECIAL PUBLICATIONS THE MUSEUM TEXAS TECH UNIVERSITY MUS. COMP. ZOOL.. UBRARY SEP 2 1 197B Revision of North American Trichodes (Coleoptera: Cleridae) David E. Foster No. 11 September 1976 TEXAS TECH UNIVERSITY Cecil Mackey, President Glenn E. Barnett, Executive Vice President Regents. — ^Judson F. Williams (Chairman), J. Fred Bucy, Jr., Bill E. Collins, Clint Form- by, John J. Hinchey, A. J. Kemp, Jr., Robert L. Pfluger, Charles G. Scruggs, and Don R. Workman. Academic Publications Policy Committee. — ^J. Knox Jones, Jr. (Chairman), Dilford C. Carter (Executive Director and Managing Editor), C. Leonard Ainsworth, Harold E. Dregne, Charles S. Hardwick, Richard W. Hemingway, Ray C. Janeway, S. M. Kennedy, Thomas A. Langford, George F. Meenaghan, Marion C. Michael, Grover E. Murray, Robert L. Packard, James V. Reese, Charles W. Sargent, and Henry A. Wright. The Museum Special Publications No. 11 86 pp., 17 figs. 17 September 1976 $2.00 Special Publications of The Museum are numbered separately and published on an irregular basis under the auspices of the Dean of the Graduate School and Director of Academic Pub¬ lications, and in cooperation with the International Center for Arid and Semi-Arid Land Studies. Copies may be obtained on an exchange basis from, or purchased through, the Ex¬ change Librarian, Texas Tech University, Lubbock. Texas 79409. Texas Tech Press. Lubbock. Texas 1976 SPECIAL PUBLICATIONS THE MUSEUM TEXAS TECH UNIVERSITY Revision of North American Trichodes (Herbst) (Coleoptera: Cleridae) David E. Foster No. 11 September 1976 TEXAS TECH UNIVERSITY Cecil Mackey, President Glenn E. Barnett, Executive Vice President Regents. — Judson F. Williams (Chairman), J. Fred Bucy, Jr., Bill E. Collins, Clint Form- by, John J. Hinchey, A. J. Kemp, Jr., Robert L. Pfiuger, Charles G. Scruggs, and Don R. Workman. Academic Publications Policy Committee. — ^J. Knox Jones, Jr. (Chairman), Dilford C. Carter (Executive Director and Managing Editor), C. Leonard Ainsworth, Harold E. Dregne, Charles S. Hardwick, Richard W. Hemingway, Ray C. Janeway, S. M. Kennedy, Thomas A. Langford, George F. Meenaghan, Marion C. Michael, Grover E. Murray, Robert L. Packard, James V. Reese, Charles W. Sargent, and Henry A. Wright. The Museum Special Publications No. 11 86 pp., 17 figs. 17 September 1976 $2.00 Special Publications of The Museum are numbered separately and published on an irregular basis under the auspices of the Dean of the Graduate School and Director of Academic Pub¬ lications, and in cooperation with the International Center for Arid and Semi-Arid Land Studies. Copies may be obtained on an exchange basis from, or purchased through, the Ex¬ change Librarian, Texas Tech University, Lubbock, Texas 79409. Texas Tech Press, Lubbock, Texas 1976 Revision of North American Trichodes (Herbst) (Coleoptera: Cleridae) David E. Foster The holarctic genus Trichodes contains more than 85 species, most of which occur in the Mediterranean Basin region. About 20 species occur in eastern and northern Europe and Asia, 10 are recorded from central and southern Africa, and 13 are from North America (Corporaal, 1950). Included in the genus are some of the most showy and commonly collected clerids. In spite of its popularity with collectors, and consequently the large amount of museum material available for study, the genus has never been revised. Old World species have been studied following geographic or political boundaries (Escherich, 1893; Champenois, 1900; Zimmerman, 1971). Studies of the North American fauna have been made by Le Conte (1849), Horn (1876), and Wolcott (1910a, 1944). Biological knowledge of North American Trichodes is limited to a study of the life history of T. ornatus (Linsley and MacSwain, 1943) and fragmentary infor¬ mation on T. horni (Krombein, 1967) and T. simulator (Stephen et al., 1969). As might be expected in a group with poor biological coverage, immature stages have received little attention. Only larvae of T. ornatus have been described and figured. During recent years, numerous specimens representing all species of North American Trichodes have been collected and are available for study. Con¬ sequently, a comprehensive study of the classification of the North American fauna now is possible. This study emphasizes the biosystematic approach, which is applied in the delineation of species, clarification of intraspecific variation, and establishment of relationships among North American taxa. Biological in¬ formation and larval taxonomy are used in collaboration with adult features to achieve the objectives. Eleven species of Trichodes are recognized. Two subspecies of T. apivorus, three of T. peninsularis, and five of T. ornatus, including one new subspecies, are treated. Larval taxonomy and comparative biologies are presented following the adult taxonomy. Historical Resume The name Trichodes first appeared in the literature when Herbst (1792) used it in place of Clerus Fabricius (1775). In his Systema Elevtheratorum, Fabricius (1801) split Clerus and used the junior synonym Trichodes as a generic name for Old World species, T. alvearius, T. ammios, T. apiarus, T. bifaciatus, T. crabron- iformis, T. cyanius, T. octopunctatus, T. sipylus, and T. tricolor. Latreille (1804) incorrectly used Clerus for species Fabricius placed under Trichodes. 3 4 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Clerus of Latreille is isotypic with Trichodes Herbst but not with Clerus of Fabri- cius. A number of nineteenth-century workers, including Curtis (1824), followed Latreille’s usage of Clerus. The validity of the name Trichodes generally became recognized during the nineteenth century and remained unchallenged until Wolcott (1944) erroneously concluded that Trichodes eventually would have to be replaced with Clerus Fab- ricius due to the isotypic nature of the names and the principle of priority. A review of the situation as stated above dispels Wolcott’s contention and reaffirms the validity of the name Trichodes. Wolcott (1944) also erred in this review of type designations for the genus. He stated that Curtis (1824) and Hope (1840) both designated Attelabus apiarus L. as the type of Clerus Latreille. In reality, Curtis designated Clerus alvearius Fabr. as the type of Clerus Latreille, and Hope designated Clerus ammios Fabr. as the type of Pachyscelis Hope. Pachyscelis was synonymized with Trichodes by Lacordaire (1857). Clerus alvearius stands as the generotype of Trichodes and is recognized correctly as such by Wolcott (1947) and Corporaal (1950). The first comprehensive study of North American Trichodes (Le Conte, 1849) treated three species; the next (Horn, 1876) included six species. Wolcott’s re¬ vision of the North American fauna (1910a) treated 10 species. Wolcott (1944) updated his earlier work to include a new variety of T. ornatus and two species described during the interim. A total of 12 species was recognized by the cata¬ loguers, Wolcott (1947) and Corporaal (1950). One additional species, T. oregonensis, has been described (Barr, 1952). Barr also has presented keys to the Trichodes species of Lower California (1950) and the Pacific Northwest (1961). Methods and Materials Taxonomy The classification presented in this revision is based on a study of adult speci¬ mens representing the 1 1 North American species recognized herein. It is sup¬ ported by data from larvae belonging to eight species and by available biological information for all species. Type specimens are referred to as they are labeled in collections or are designated in the literature. The order of specific and subspecific accounts in text is determined by the order in which they appear in the keys, the only exception being Trichodes simu¬ lator in the section on larval taxonomy. Collection data. — Locality data for adult specimens on which this study was based were presented in Foster (1973). Only localities for specimens examined since that date are listed herein, but the total number of specimens examined, including those in Foster (1973), is given in parentheses. Lending institutions (abbreviations taken from Arnett and Samuelson, 1969, whenever possible) are indicated for each entry and are as follow: AMNH — American Museum of Natural History, New York City (Lee Herman, Jr.) ANSP — Academy of Natural Sciences, Philadelphia (M. G. Emsley) CASC — California Academy of Sciences, San Francisco (Hugh B. Leech) FOSTER— REVISION OF NORTH AMERICA TRICHODES 5 CISC — California Insect Survey, University of California, Berkeley (J. A. Chemsak) FMNH — Field Museum of Natural History, Chicago (H. S. Dybas) MCZC- — Museum of Comparative Zoology, Harvard University, Cambridge (J. F. Lawrence) NDSU — North Dakota State University, Fargo (R. L. Post) NMNH — National Museum of Natural History, Washington, D.C. (P. J. Spangler) TTCC — Texas Tech University, Lubbock (C. W. O’Brien) UICM — University of Idaho, Moscow (W. F. Barr) Preparation and examination of material. — The external anatomy of adults and larvae was studied using an American Optical stereoscopic microscope at 10 to 80 X magnification. All measurements were made with a calibrated micrometer eyepiece and are given in millimeters. Male genitalia were studied by relaxing specimens in a warm water bath using a Bransonic ultrasonic cleaner. Genitalia were removed with a pair of micro¬ shears by making a small incision through the pleural regions of the fourth, fifth, and sixth abdominal segments and extracting the structures with a pair of fine forceps. The genitalia then were placed in a heated solution of 10 per cent KOH for 10 to 20 minutes until soft tissues were dissolved. Cleared genitalia were bathed in distilled water and stored in 70 per cent ethyl alcohol. Genitalia were prepared by removing the median lobe and mounting both it and the tegmen on a glass slide. They were examined using a compound micro¬ scope and transmitted light. Larvae were studied by removing the right mandible from some specimens. The structures were cleared in 10 per cent KOH and mounted on glass slides. Illustrations. — Drawings of antennae, male genitalia, and larval structures were made from unmounted material viewed through a Wild stereoscopic micro¬ scope fitted with a cameralucida. Later mounted material was viewed through a compound microscope with transmitted light to add details, such as setal patterns. Explanation of terms. — Throughout this study, the word sternite is used with reference only to visible abdominal sterna; that is, the term “first abdominal sternite” refers to the true third abdominal sternite. Biology Field studies. — Objectives during field studies were to collect adult and larval specimens, particularly from areas of specific taxonomic interest, associate larvae with host organisms, and study the biology of species occurring in the western United States. Collections and biological observations were made during several extensive field trips to areas in the western United States between 1969 and 1972. Larvae were collected and associated with hosts through the use of artificial nesting sites and through the excavation of naturally occurring host nests in areas of high Trichodes density. Artificial nests of three types were placed in the field before seasonal activity commenced. Block traps, constructed by cutting six-inch sections of 2 X 2 inch or 2 X 4 inch lumber and drilling four-inch deep holes of 1/4 inch or 3/8 inch diameter in one end, were tied to wooden fence posts, fallen trees, or standing trees and shrubs. Bundle and individual stick traps were constructed by cutting 6 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY dried stems of elderberry, Samhucus spp., into appropriate lengths and drilling a single hole in one end. Six-inch bundles of 10 to 12 sticks were set out in the manner described for block traps. Sticks 12 to 15 inches long were sharpened at the end opposite the hole and driven individually into the soil at about a 60-de¬ gree angle. Trap-nests have not been used previously to study predaceous beetles, but have been used extensively to study hymenopteran biology. For a detailed account of nest-trapping, see Krombein (1967). Whenever possible, trap-nests were allowed to remain in the field until the fol¬ lowing spring, a practice that increased rearing success. Nests were returned to the laboratory and their contents examined. Some immature specimens of both Tric/iodes and their hosts were removed, treated in KAAD solution (Peterson, 1959), and preserved in 70 per cent ethyl alcohol. Cast skins from trap-nests and laboratory rearings were preserved in 70 per cent ethyl alcohol. Traps then were resealed; remaining larvae were allowed to pupate and emerge as adults. Lxihoratory studies. — An experiment was conducted to study the effect of larval and pupal exposure to various combinations of temperature and humidity on elytral coloration of ensuing adults. Field collected, third instar specimens of Tricliodes ornalus honnevillensis from Elba-Basin Pass, Cassia County, Idaho, and Trichodes ornatus hartwegianus from near Touchet, Walla Walla County, Washington, were used. Ninety-eight per cent of all adult speci¬ mens from the former locality have red base colored elytra, whereas the latter population has individuals only with yellow base coloration. Each larva was allowed to burrow into a cork and construct its pupal chamber. Larvae were divided into groups containing two honnevillensis and eight hartwegianus larvae each. One group was held at room temperature and relative humidity as well as under each of the following combinations until adults had emerged: 4°C — 50% RH, 4°C— 75%RH, 20°C— 50%RH, 20°C— 75%RH, 36°C— 50%RH, 36°C— 75%RH. Systematics Adult Taxonomy Trichodes Herbst, 1792:154; Fabricius, 1801:283; Le Conte, 1849;17; Lacordaire, 1857:459; Horn, 1876:231; Wolcott, 1910a:366; Leng, 1920:150; Balduf, 1935:107; Wolcott, 1944:54; Barr, 1961:109; Papp, 1960:85; Arnett, 1960:599; Barr, 1962:121; Ekis and Gupta, 1971:60. Clerus, Latreille, 1804:150; Curtis, 1824:table 44; Lacordaire, 1857:459; Crotch, 1870:42; Wolcott, 1944:54. Pachyscel is Hope, 1840:139; Lacordaire, 1857:459. Description. — Male. Small to large sized, slender to robust; clothed with short and long, moderately dense to very dense, recumbent to erect hairs. Head finely and sparsely or coarsely and densely punctate; eyes finely faceted, inner margin deeply, triangularly notched, notched area of eyes bearing a tuft of stiff hairs; antenna eleven segmented, first segment elongate and feebly arcuate, second segment short and rounded, third through eighth segments subcylindrical with each segment shorter and broader than preceding one, segments nine through FOSTER— REVISION OF NORTH AMERICA TRICHODES 7 eleven forming a broad, compact, triangular club; maxilla with galea elongate, at least one-half as long as distance between eyes across front, palpus with termi¬ nal segment produced as a narrow, elongate triangle; labial palpus with terminal segment broad and triangular. Pronotum subquadrate; disc convex with a shal¬ low, transverse impression in front of middle and a deep constriction in front of hind margin. Scutellum transverse, disc flat or slightly convex. Elytra more than twice as long as humeral width; sides subparallel or expanded at posterior one- fourth; umbones prominently elevated and subbasal tumescence faintly indicated. Legs densely, irregularly roughened and variously punctured; coxa scabrous; metafemur sometimes enlarged; tibia never carinate, paired tibial spurs subequal. Abdomen irregularly roughened and sparsely, finely punctate; fifth sternite with hind margin deeply emarginate; sixth sternite longer than broad. Female. Similar to male except: metafemur never enlarged; abdomen with hind margin of fifth sternite truncate or broadly, shallowly emarginate, sixth sternite transverse. Type species. — Clerus alvearius Fabr. was designated as the type species of Trichodes by Curtis ( 1 824). Discussion. — Wolcott (1947) placed Trichodes between the exclusively New World genera Aulicus and Enoclerus. Although Aulicus is considered its nearest relative in North America, Trichodes also is related closely to the Old World genus Opilo. Trichodes is separated from the related North American Clerinae by a number of characters. The antennal club of Trichodes is broadly triangular, somewhat dorsoventrally flattened, and comprised of three compact segments. Enoclerus has a compact, three segmented club that is elongate, more cylindrical than flattened, and more subrectangular than triangular. The antennal club of Aulicus is composed of three loosely joined segments, is more or less quadran¬ gular, and never triangular. Another unique characteristic of Trichodes is the elongate and conspicuous maxillary galeae, which are at least one-half as long as the distance between the eyes in front. Other North American Clerinae have short and often inconspicuous galeae. A third reliable character distinguishing Trichodes is the shape of the apical segment of the maxillary palpus. Trichodes is the only North American genus in the subfamily in which the apical segment is subequal in length to the remaining segments, and the only one in which that segment takes the form of a narrow, elongate triangle. Enoclerus has a short, subcylindrical apical maxillary segment; in Aulicus, the segment is expanded greatly and triangular in shape. The generic description presented here is based on the North American fauna. It has been compared with several Old World species and appears to be repre¬ sentative of the entire genus. Relationship Among North American Trichodes A review of published work on North American Trichodes reveals that species groups never have been recognized formally. However, both Horn and Wolcott constructed their keys in such a way as to suggest relationships. Horn (1876) recognized T. illustris as distinctive because of its truncate elytral apices, grouped 8 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY T. nutalli, T. ornatus, and T. bisignatus on the basis of fine elytral and sparse thoracic punctations, and associated T. apivorus with T. bihalteatus due to their coarse, deep elytral and dense thoracic punctations. In his first revision, Wolcott (1910a) associated T. oresterus and T. peninsular- is with T. illustris. He reaffirmed the distinctiveness of what herein is considered to be the ornatus group and aligned T. nexus with it. However, he failed to relate T. apivorus to T. bibalteatus and considered T. simulator as a separate entity. In his later paper, Wolcott (1944) associated T. basalis with T. oresterus and treated T. bicinctus as a separate entity. The present study recognizes three species groups. T. oresterus, T. peninsular- is, and T. simulator are assigned to the peninsularis group. The ornatus group contains the species placed together by earlier authors, with the addition of T. oregonensis. The bibalteatus group includes T. bicinctus in addition to T. bibal¬ teatus and T. apivorus. Members of the peninsularis group are not related closely to one another, but do comprise a distinct entity well separated from the remainder of our fauna. The shape of the elytra and the nature of the punctations; the shape, punctations, and vestiture of the pronotum; and shared biological features distinguish the peninsularis group. The bibalteatus group contains two closely related species, T. bibalteatus and T. apivorus. T. bicinctus is the most distinctive member of the group on the basis of its more robust form, elongate galea, and distinctive antennal club. Its affinities are with members of the bibalteatus group, and it is treated most logical¬ ly as a member of that group. The ornatus group is the most coherent and homogeneous sector of the North American fauna. Its members are associated by their small size and robust form, inconspicuous nature of their prothoracic and elytral punctations, and distinctive larvae. Within the group, T. ornatus and T. nexus show a close relationship, as do T. oregonensis, T. birnaculatus, and T. nutalli. Another distinctive feature of these species groups is their distributions. The peninsularis group occurs in arid and semiarid regions of the southwestern United States and adjacent Mexico. Its members have relatively broad distributional ranges and are principally allopatric. Members of the bibalteatus group occur in the eastern United States and from the southern Great Plains to southwestern Texas. T. apivorus is distributed along the Atlantic and eastern Gulf coasts. The remaining species have somewhat restricted distributions and display sympatry. The ornatus group is distributed widely, occurring in much of North America except southern Mexico and the southeastern states. Its distribution is more northerly than other groups. Four of the five species occur in the northern half of the United States, and two extend into Canada. Only one species, T. nexus, has a restricted southwestern distribution. Included in the group are species with the broadest and most restricted distributions in the North American fauna. FOSTER— REVISION OE NORTH AMERICA TRICHODES 9 Relationship to Old World Trichodes There is little doubt that Trichodes evolved somewhere on the Eurasian land mass and subsequently reached North America. However, the nature of its radia¬ tion poses some interesting questions not covered in this study. Elements of the Old World fauna closely resemble all three species groups delineated herein. It is unclear whether these represent extensions of various Old World elements that arrived during successive invasions, or if they are the result of parallel evolution. Relationships among the varied elements of Trichodes can be analyzed logical¬ ly through the application of the phylogenetic principles set forth by Hennig (1966). Until that segment of the study is complete, I have elected to refrain from designating subgenera and to refer only to species groups. Anatomical Characters Markings and coloration. — Horn (1876) and, to a lesser extent, Wolcott (1910, 1944) relied on coloration and color patterns for separating species. In some species, these conspicuous features are constant and serve well in species differ¬ entiation. In others, they are highly variable and only serve to supplement more reliable anatomical features. Color features useful as primary characters include coloration of abdominal sternites, spots on the elytral umbones, shape of the dark elytral bands, and the degree to which these bands attain the lateral margins or are expanded along them. All species in the fauna have bicolored elytra except T. peninsularis, which is usually tricolored. The base elytral color ranges from red to yellow, is highly variable in some species, and is of little value as a taxonomic character. Post- mortum changes frequently result in an alteration of coloration, compounding its unreliability. Superimposed upon the base color are two or three variously modified, dark colored bands. These may appear transverse, as in T. apivorus, T. bibalteatus, T. nutalli, and T. simulator, or one or more of them may be expanded or reduced. The most extreme examples of expansion are seen in T. bimaculatus and T. oregonensis, where all that remains of the base color is a pair of medial macula- tions. The elytral bands of T. ornatus show both expansion and reduction, ranging from a condition very similar to that of the two aforenamed species to an extreme reduction in which the subbasal band is absent and the postmedial and apical bands are reduced greatly. The head, thorax, abdomen, and appendages of most species are blue, purple, or verdigris, sometimes with variously expanded testaceous or red markings. Only in some populations of T. peninsularis do these body regions and structures appear entirely reddish. Antenna. — The length to width ratio of the antennal club, and the shape of its apical segment, provide characters useful in differentiating species of peninsular¬ is and bibalteatus groups. Antennal shape is relatively consistent within the ornatus group, providing a useful character for its recognition. 10 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Galea. — The length of the galea falls into one of three categories, one-half as long, about as long, or distinctly longer than the distance between the eyes across the front. This serves as a useful character for distinguishing several species in the peninsular is and hihalteatus groups. Punctations. — Punctations of the prothorax and elytra are useful at the group level. Members of the ornatus group have the pronotum and elytra sparsely, finely, and often inconspicuously punctate. The other groups have the pronotum and elytra coarsely punctate. Vestiture. — The occurrence and nature of hairs on the pronotum and abdomen are important at both the species and group level. At the group level, the dense¬ ness and relative coarseness of the hairs on the pronotum provide useful charac¬ ters. At the specific level, the presence of dense patches of hair on the abdominal sternites serves as a primary character for the separation of T. peninsularis. Dense and recumbent hairs on the mesosternum are important in T. peninsularis and T. oresterus. Elytral apices. — Elytral apices of North American Trichodes may be either rounded or variously truncated. The ornatus and bibalteatus groups have rounded apices, whereas in the peninsularis group the apices are truncate, with the nature of the truncation serving as a primary character for separation of species. Previous workers have suggested incorrectly that the elytral apices exhibit sexual dimorphism in T. peninsularis (Horn, 1876) and T. simulator (Wolcott, 1910a). These statements apparently can be attributed to infraspecific variation and a lack of adequate study material. Aedeagus. — The shape of the lateral lobes affords positive identification of most species. The tip of the median lobe is distinctive in T. apivorus. The shape of the tegmen in dorsal or lateral view is often distinctive, but must be used with the understanding that the tube is quite flexible and its shape is dependent some¬ what upon the position of the median lobe within it. Several other features, in¬ cluding the shape of the median struts, tegminal strut, median orifice, and median foramen are quite variable and of no value for species identification. Terminology used herein is based on the work of Sharp and Muir (1912). Sexually dimorphic characters. — The shape of the hind margins of the fifth and sixth abdominal sternites and sixth abdominal tergite provide reliable dimorphic characters in all species. Males of T. peninsularis also are recognized by a swollen hind femur. T. oresterus can be sexed on the basis of abdominal coloration: females have the venter entirely reddish, whereas in males it is darkly pigmented, at least toward the apex. Before beginning the systematic treatment, it is important to discuss briefly the concept of species and subspecies employed therein. A “biological” species concept, which holds that species are groups of actually or potentially interbreed¬ ing populations reproductively isolated from other such groups (Mayr, 1963), is employed. North American Trichodes exhibits no evidence of hybridization among sympatric species; the “biological” species concept thus is applied easily. Subspecies are viewed as relatively homogeneous, anatomically and geo¬ graphically distinctive infraspecific groups of interbreeding populations that FOSTER— REVISION OF NORTH AMERICA TRICHODES 1 1 collectively represent a partially isolated, recently evolved, or separately evolving lineage with its own evolutionary tendencies. Three North American species, T. apivorus, T. peninsularis, and T. ornatus, have distinctive populations that fit this concept and can be considered subspecies in an evolutionary sense rather than solely as geographic variants. Key to Species of Adult Trichodes 1. Elytra coarsely, deeply reticulately punctate, umbones pale . 2 Elytra indistinctly punctate, umbones nearly always dark (ornatus group) . 7 2. Elytral apices truncate or emarginate; pronotum clothed with moderately dense, fine hairs (peninsularis group) . 3 Elytral apices rounded; pronotum clothed with very dense, coarse hairs (hihalteatus group) . 5 3. Antennal club at least one and one-half times longer than broad; elytra variously colored, dark bands never transverse . 4 Antennal club less than one and one-half times longer than broad; elytra orange with three dark, transverse bands, including dark apices; North Dakota, southern Wyoming to northeastern Arizona (Fig. 3) . simulator 4. Abdominal sternites one through four bearing dense patches of white hairs on either side; elytra usually tricolored, yellow with three broad brown bands bordered with pur¬ ple, purple borders sometimes greatly expanded; southwestern New Mexico, west into southern California, and south into Sonora and Baja California (Fig. 3) . . . peninsularis Abdominal sternites without dense patches of hairs; elytra bicolored, red with blue or greenish bands; from southwestern Arizona, across southern New Mexico to the Trans- Pecos region of Texas (Fig. 3) . oresterus 5. Elytra orange or red with three dark, complete or incomplete transverse bands or macu- lations including the dark apical tips . 6 Elytra orange with two dark, complete transverse bands, apices never dark; southern Great Plains from Kansas to the Mexican border and west to New Mexico (Eig. 5) . . hihalteatus 6. Galea much shorter than distance between eyes across front; antennal club almost twice as long as broad (Fig. 4); pronotum with dark red hairs; Atlantic and Gulf states to Mississippi (Fig. 5) . apivorus Galea longer than distance between eyes across front; antennal club only slightly longer than broad (Fig. 4); pronotum clothed with orange hairs; southwestern Texas (Fig. 5) . . . hicinctus 1. Variously colored; elytra bearing more than one pair of maculations, anterior angles yellow or red . 8 Metallic blue or green; elytra with a single pair of maculations, anterior angles dark . . . 10 8. Elytra with umbone spots produced to front margin, dark subbasal band never attaining lateral margin . 9 Elytra with umbone spots usually completely surrounded by yellow or red, if umbone spots extend to medial suture, then dark subbasal band attains lateral margin; United States, Canada, and northern Mexico west of the one-hundredth meridian (Fig. 10) .... . ornatus 9. Elytra marked with alternating incomplete testaceous and complete dark bands, middle and posterior testaceous bands narrow; lower Baja California (Fig. 11) . nexus Elytra marked with alternating complete red and incomplete purple transverse bands of about equal width; southern Canada and northern United States east of the Continen¬ tal Divide (Fig. 13) . nutalli 10. Elytra about two and three-fourths times longer than humeral width; elytral maculations attaining lateral margins or separated from them by a distance much less than their 12 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY diameter; coast ranges of northern and southern California (Fig. 11) . himaculatus Elytra about two and one-fourth times longer than humeral width; elytral maculations separated from lateral margins by a distance subequal to their diameter; coastal ranges of southern Oregon and extreme northern California (Fig. 11) . oregonensis Peninsiilaris Group Description. — Medium to large sized with body moderately slender; antenna variable (Fig. 1); prothorax coarsely punctate and clothed with moderately dense, fine hairs; elytra with sides subparallel, apices truncate or emarginate, surface coarsely and reticulately punctate. Species included. — T. simulator, T. peninsularis, and T. oresterus. Trichodes simulator Horn (Figs. 1, 2, 3) Trichodes simulator Horn, 1880:149; Cockerell, 1898:155; Schenkling, 1903:plate 2, fig. 4; Wolcott, 1910«;370; Wickham and Wolcott, 1912:59; Clemens, 1916:397; Wolcott, 1944:58; Thompson and Simmonds, 1965:30; Stephen et al., 1969:127. Trichodes simulator v2lt. flavescens CockexeW, 1898:155; Wolcott, 1910a:370. Trichodes apivorus. Snow, 1907; 176; Tanner, 1934:45 (misidentifications). Trichodes hihalteatus, Wickham and Wolcott, 1912:59 (misidentification). Description. — Male. Medium to large sized, moderately robust; clothed with short to long, recumbent to erect fulvous hairs; head, thorax, scutellum, venter, and legs atropurpureus, margins of abdominal sternites, tibial apex, and tarsus sometimes reddish orange; antenna with at least under side of funicle testaceous, club piceous; elytra reddish orange with atropurpureus subbasal, medial, and apical transverse bands, umbones pale. Head densely punctate with many puncta- tions confluent; front broadly impressed above clypeus and between eyes; antenna with club nearly as broad as long in side view (Fig. 1); clypeus testaceous and glabrous; labrum dark brown, finely, sparsely punctate, front margin shal¬ lowly emarginate; galea long, subequal in length to distance between eyes across front. Pronotum very coarsely, densely punctate with few punctations confluent, moderately densely clothed with long, suberect and erect fine hairs. Scutellum densely punctate and roughened; disc medially depressed to the subtruncate hind margin. Elytra about two and three-fourths times longer than humeral width, coarsely, reticulately punctate, umbones coarsely punctate and roughened; sides subparallel, apices emarginate with outer and sutural angles acute to acuminate. Mesosternum densely rugosopunctate; moderately densely clothed with suberect, long hairs. Metafemur not swollen disproportionately. Abdomen with sternites one through four coarsely punctate and roughened in front of hind margin, punc¬ tations widely scattered or evanescent toward base; fifth stemite with hind margin deeply, arcuately emarginate; sixth sternite elongate, disc convex with baso- medial surface glabrous, lateral margin feebly arcuate, hind margin truncate; eighth tergum narrower and longer than sixth sternite, disc broadly, deeply im¬ pressed with the impression clothed with a distinct row of dense, suberect hairs along either side, hind angles bearing conspicuous tufts of long, stiff hairs, hind margin arcuately rounded and broadly deflexed. Aedeagus distinctly arcuate; FOSTER— REVISION OF NORTH AMERICA TRICHODES 13 Fig. 1.— Male genitalia and antennal clubs: a, Trichodes oresterus, dorsal view; b, Tri- chodes oresterus, lateral view; c, Trichodes simulator, lateral view; d, Trichodes peninsularis horni, lateral view; e, Trichodes oresterus, lateral view. lateral lobes gradually tapered to podoform apices in lateral view, subparallel in dorsal view; median lobe laterally compressed, apex ovoid (Fig. 2). Length: 8.7-16.4. Female. Similar to male except: abdomen with hind margin of fifth sternite broadly, shallowly emarginate; sixth sternite transverse, disc feebly convex; eighth tergum impressed to fimbriate hind margin; hind margin not deflexed. Length: 10.3 to 1 6.7. 14 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Fig. 2. — Male genitalia: a, Trichodes simulator, dorsal view; b, Trichodes sioinlator, lateral view; c, Trichodes peninstdaris horni, dorsal view; d, Trichodes peninsidaris horni, lateral view. Type locality. — “Arizona”; of T. simulator var. flavescens, Santa Fe, New Mexico. Type material. — Lectotype, male, no. 3521, ANSP. Cockerell apparently did not designate a type specimen of T. simulator var. flavescens. Distribution. — Colorado Plateau region of eastern Utah, western Colorado, northwestern New Mexico, and northeastern Arizona, and extreme southwestern North Dakota (Fig. 3). Discussion. — Elytral coloration and markings of T. simulator most closely resemble those of T. apivorus and T. bibalteatus. This species can be separated easily from either of these allopatric species on the basis of the elytral apices. FOSTER— REVISION OF NORTH AMERICA TRICHODES 15 Fig. 3. — Distribution of the peninsularis group: Trichodes simulator, horizontal lines; Trichodes peninsularis peninsularis, shaded; Trichodes peninsularis basal is, diagonal lines; Trichodes peninsularis horni, squares; Trichodes oresterus, stippled. which are emarginate and pointed in T. simulator and rounded in the other two species. Additionally, T. simulator has the pronotum clothed in moderately dense, long, silky hairs that never obscure its surface. T. apivorus and T. bibalteatus have the pronotum covered with very dense, coarse hairs that at least partially obscure its surface. 16 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY T. simulator occurs over a rather limited range and exhibits a remarkable con¬ sistence in appearance except for some variation in elytral coloration. Ninety-one per cent of all specimens studied have bright reddish orange elytra. The re¬ mainder, comprised primarily of material from Sandoval and Santa Fe counties, New Mexico, but also including a few specimens from scattered localities throughout the range, have testaceous elytra. It was apparently this variant to which Cockerell applied the name flavescens. Specimens examined (S2). — North Dakota: Slope Co., Burning Coal Vein, 17 August 1972 (Tim L. McCabe), 1 d, 1 9 (NDSU). For additional records, see Foster (1973). Trichodes peninsularis Horn (Figs. 1, 2, 3) Trichodes peninsularis Horn, 1 894:382; Wolcott, 1910«:369, 1944:55; Barr, 1950:507. Description. — Male. Small to large sized, slender to moderately robust; clothed with short to long, recumbent to erect whitish hairs; head, thorax, scutellum, venter, and legs atropurpureus, verdigris, or testaceous; antennal funicle atro- purpureus to testaceous, club fuscus to testaceous; elytra atropurpureus with basal, medial, and subapical yellow bands, atropurpureus areas sometimes bear¬ ing small to large brown spots, umbones pale. Head densely punctate with about one-half of the punctations confluent; front not impressed above clypeus or between eyes; antenna with club more than one and one-half times as broad as long in side view (Fig. 1); clypeus testaceous and glabrous; labrum testaceous, surface finely, sparsely punctate, front margin broadly, shallowly emarginate; galea short, slightly longer than one-half the distance between eyes across front. Pronotum very coarsely, densely punctate with few punctations confluent, moderately densely clothed with long, suberect and erect fine hairs. Scutellum densely punctate and roughened; disc convex; hind margin subtruncate or rounded. Elytra about two and three-fourths times longer than humeral width, coarsely, reticulately punctate with umbones finely, sparsely punctate, sides subparallel, apices truncate or sinuatotruncate with sutural tooth well developed. Mesosternum densely, moderately coarsely punctate, densely clothed with re¬ cumbent, long hairs. Metafemur swollen disproportionately. Abdomen with sternites one through four finely, sparsely punctate, punctations most numerous near hind margin of each stemite, evanescent toward base; first four sternites bearing dense patches of appressed white hairs at sides; fifth stemite with hind margin deeply, arcuately emarginate; sixth stemite elongate, disc convex with mesobasal surface glabrous, lateral margin feebly arcuate, hind margin rounded; eighth tergum narrower and longer than sixth stemite, disc shallowly, longitudi¬ nally impressed, impression clothed with a distinct row of dense, suberect hairs on either side, hind angles bearing conspicuous tufts of long hairs, hind margin semicircularly rounded and deflexed at middle. Aedeagus feebly arcuate with lateral lobes gradually tapered to bluntly rounded apices in lateral view, sub¬ parallel in dorsal view; median lobe laterally compressed, apex ovoid (Fig. 2). Female. Similar to male except; metafemur not disproportionately swollen. Abdomen with hind margin of fifth stemite truncate or shallowly emarginate; FOSTER— REVISION OF NORTH AMERICA TRICHODES 17 sixth sternite transverse, dise slightly convex; eighth tergum with disc feebly convex, clothed with scattered long, suberect hairs, hind margin truncate and not deflexed. Discussion. — When Horn described T. peninsularis, he suggested that it might be conspecific with T. illustris ( = T. horni). However because his limited series of T. illustris showed little variation, he chose to consider T. peninsularis a dis¬ tinct species. The large number of specimens now available indicates that T. horni and T. peninsularis, as well as T. basalis, are conspecific. They nevertheless represent taxonomically and geographically distinctive entities worthy of subspecific designation. This distinctiveness is evidenced further by temporal and habitat differences. T. p. basalis occurs in the very arid lower Colorado River Valley. Adults are present from late June through mid-September. T. p. horni occurs primarily in transitional zones between the Sonoran and Chihuahuan deserts. Adults appear following the first fall rains in late August and September. T. p. peninsularis occurs in the foothills of southwestern Sonora and in the cape region of lower Baja California. Adults start to appear by late July. Populations of all three subspecies show evidence of hybridization. A narrow band of primary intergradation occurs between T. p. horni and T. p. basalis in Yuma County, Arizona. Some specimens of T. p. horni from the Huachuca Mountains, Cochise County, Arizona, and Nogales, Sonora, bear a strong resem¬ blance to T. p. peninsularis in overall coloration. However, the presence of brown elytral marking as well as size and temporal occurrence conclusively align the populations with T. p. horni. Further evidence of the conspecific nature of the three taxa can be found in the similarity of body form and shape, punctations, and pubescence. Elytral coloration, although quite distinctive, conforms to a basic pattern shared by all three subspecies. Additionally, the aedeagus, which has proven a useful specific level character in this study as well as in studies of Palearctic Trichodes {E.sc\\&x'\ch, 1893; Zimmerman, 1971), is indistinguishable among the three subspecies of T. peninsularis. Key to Subspecies ^^/Trichodes peninsularis Horn 1. Elytra with basal yellow band attaining scutellum, apices usually tipped with brown; femora usually testaceous . 2 Elytra with basal yellow band separated from scutellum by a purple border, apices purple; femora purple; lower Baja California and southwestern Sonora (Fig. 3) . . peninsularis 2. Small to medium sized; elytra with yellow bands expanded, covering about one-half of total elytral surface, brown areas reduced or absent; abdomen testaceous except for dark medial spots on sternites one through five of some specimens; extreme southwestern Arizona and southeastern California (Fig. 3) . basalis Medium to large sized; elytra with yellow bands narrow, covering about one-fourth of total elytral surface, brown areas large; abdomen usually dark, sometimes with marginal areas testaceous; extreme southwestern New Mexico, southern Arizona, and northern Sonora (Fig. 3) . horni 18 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Trichodes peninsularis peninsularis Horn Trichocic’s peninsularis Horn, 1 894:382; Wolcott, 1910fl;369, 1944:55; Barr, 1950:507. Description. — Small sized, slender; clothed with whitish hairs; head, thorax, scutellum, and legs purpurescent, scutellum bounded by a narrow purpurescent band, elytra with basal, medial, and subapical yellow bands narrow, purpurescent areas large, sometimes bearing very small brown spots; abdomen purpurescent with apex testaceous in some specimens. Length: male, 7.8-10.7; female, 7.8. Type locality. — El Chinche, Baja California = Sierra El Chinche. Type material. — Type, female, no. 37, Department of Entomology, CASC. Distribution. — Lower Baja California and southeastern Sonora (Fig. 3). Discussion. — T. p. peninsularis is separated from most specimens of T. p. hasalis and T. p. horni on the basis of its dark coloration and lack of extensive brown areas on the elytra. A few specimens of T. p. horni from the Huachuca Mountains of Cochise County, Arizona, and Nogales, Sonora, resemble T. p. peninsularis but are larger and usually have more brown coloration. Specimens examined (5). — Baja California Sur: Las Parras, 1 October 1923 (W. M. Mann), 1 d (TTCC). Trichodes peninsularis basalis Van Dyke, new status Trichodes hasalis Wan Dyke, 1943:41; Wolcott, 1944:56. Description. — Small to medium sized, slender; clothed with whitish hairs; head, thorax, scutellum, and legs verdigris or testaceous; elytra with basal, medial, and subapical yellow bands covering about one-half of surface, purpures¬ cent areas reduced, sometimes bearing small brown spots; abdomen testaceous with sternites one through five sometimes darkened along midline. Length: male, 7.5-12.7; female, 7.7-13.0 Type locality. — Imperial County, California. Type material. — Holotype, male, no. 5327, Department of Entomology, CASC. Distribution. — Lower Colorado River Valley of southeastern California and extreme western Arizona (Fig. 3). Discussion. — T. p. basalis is separated from all specimens of T. p. peninsularis and from most specimens of T. p. horni. A narrow band of hybridization in Yuma County, Arizona, makes separation from T. p. horni arbitrary in some instances. Specimens from either side of the hybrid band can be identified by means of the characters presented in the key. Specimens examined (22). — See Foster (1973). Trichodes peninsularis horni Wolcott and Chapin, new status Trichodes illustris Horn, 1876:231 (nec Klug, 1842:333); Schenkling, 1906;286; Schaeffer, 1908:131; Wolcott, 1910^:369; Wickham and Wolcott, 1912:59; Wolcott and Chapin, 1918:107. Trichodes horni ^0)0011 and Chapin, 1918:107; Wolcott, 1944:56; Krombein, 1967:376. FOSTER — REVISION OF NORTH AMERICA TRICHODES 19 Description. — Medium to large sized, moderately robust; clothed with fulvous hairs; head and prothorax purpurescent; legs testaceous; elytra with basal, medial, and subapical yellow bands covering about one-fourth of surface, purpurescent areas nearly replaced by large brown spots; venter piceous with abdominal apex testaceous in some specimens. Length: male, 7.8 to 15.1 ; female, 8.2 to 16.8. Type locality. — “Arizona.” Type material. — Lectotype, female, no. 3522, ANSP. Distribution. — Southwestern New Mexico, southern Arizona, and north¬ ern Sonora (Fig. 3). Discussion. — Specimens assigned to T. p. horni are predominantly brown and are usually larger than those of other subspecies. Other characters useful in its separation are presented under each of the other subspecies. Specimens examined (686). — See Foster (1973). Trichodes oresterus Wolcott (Figs. 1, 3) Trichodes oresterus 'Wo\con, 1 9 10rt:368; Wickham and Wolcott, 1912:59; Wolcott, 1944:54. Description. — Male. Small to medium sized, slender; clothed with short to long, recumbent to erect whitish hairs; head, thorax, scutellum, and legs dark metallic blue to verdigris; antenna testaceous with club orange, dark brown, or piceous; elytra reddish with dark metallic blue or verdigris subbasal, medial, and apical bands, umbones pale; abdomen testaceous with variable dark markings. Head densely punctate with few punctations confluent; front transversely im¬ pressed above clypeus and between eyes; antenna with club twice as long as broad in lateral view (Fig. 1); clypeus testaceous to dark brown, glabrous; labrum dark brown, sometimes with front and lateral margins testaceous, surface finely, very sparsely punctate, front margin broadly, shallowly emarginate; galea short, slightly longer than one-half the distance between eyes across front. Pronotum very coarsely, densely punctate, punctations never confluent; moderately densely clothed with long, suberect and erect, fine hairs. Scutellum densely punctate and roughened, disc convex, hind margin subtruncate or rounded. Elytra about two and two-thirds times longer than humeral width, sides subparallel, apices sinuato- truncate, sutural tooth slightly developed or absent. Mesosternum densely, moder¬ ately coarsely punctate, densely clothed with long, recumbent hairs. Metafemur not swollen disproportionately. Abdomen with sternites one through five coarsely, densely punctate and roughened in front of hind margin, punctations evanescent toward base; fifth stemite with hind margin deeply, arcuately emarginate; sixth sternite elongate, disc feebly convex with basomedial surface glabrous, lateral margin arcuate, hind margin semicircularly rounded, sometimes narrowly truncate at middle; eighth tergum narrower and longer than sixth ster¬ nite, disc broadly, shallowly impressed, impression clothed with dense, suberect hairs, hind margin semicircularly rounded and deflexed at middle. Aedeagus feebly arcuate; lateral lobes strongly constricted before the bluntly rounded apices 20 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY in lateral view, subparallel in dorsal view; median lobe laterally compressed, apex ovoid (Fig. 1). Length: 6.8 to 12.0. Female. Similar to male except: abdomen entirely testaceous, fifth sternite with hind margin truncate or shallowly emarginate; sixth sternite transverse, disc feebly convex, hind margin semicircularly rounded; eighth tergum with disc slightly convex, sparsely clothed with long, suberect hairs, hind margin truncate, not deflexed. Length: 7.6 to 1 1.8. Type locality. — Alpine, Brewster County, Texas. Type material. — Type, male, NMNH. Distribution. — Southwestern Texas and northern Chihuahua to southeastern Arizona (Fig. 3). Discussion. — T. oresterus is related most closely to T. peninsularis as evi¬ denced by the nature of the punctations, pronotal and elytral shape, the pubes¬ cence of the pronotum and mesostemum, shape of the antennal club, and similari¬ ties of the aedeagus. Nevertheless, T. oresterus is a very distinctive, easily recog¬ nized species. It is the only North American species with the apical markings extended forward along the suture to the middle of the elytra and the only species having reddish elytra, the dark markings of which are adorned with whitish hairs. T. oresterus exhibits a good deal of infraspecific variation, especially in size, elytral markings, and coloration. Both sexes exhibit a wide range of sizes that cannot be correlated with distribution. It seems plausible that the nature of size variation will be clarified with adequate knowledge of the larval host range. The dark subbasal elytral band appears broad and transverse in 82 per cent of the specimens examined. In the remainder, it does not attain the suture, appearing as a pair of large spots, one on each elytron. Specimens with a reduced subbasal band usually have the median band reduced as well. A few specimens from older collections apparently have faded and the base coloration of the elytra is yellow. All living specimens observed during the study had reddish orange elytra. Specimens examined (165). — See Foster (1973). Bibalteatus Group Description. — Medium to large sized, robust; antenna variable (Fig. 4); pro¬ thorax coarsely punctate and clothed with very dense, coarse hairs; elytra with sides expanded at posterior one-fourth, apices rounded, surface coarsely and reticulately punctate. Species included. — T. apivorus, T. bibalteatus, and T. bicinctus. Trichodes bibalteatus Le Conte (Figs. 4, 5) Trichodes bibalteatus Le Conte, 1858:18; Horn, 1876:231; Wickham, 1896:163; Wolcott, 1910«:371; Wickham and Wolcott, 1912:59; Wolcott, 1944:58. Trichodes apivorus. Snow, 1907:176 (misidentification). Description. — Male. Large sized, robust; clothed with short to long, suberect and erect, reddish orange hairs; head and prothorax dull red to atropurpureus; FOSTER— REVISION OE NORTH AMERICA TRICHODES 21 Fig. 4. — Male genitalia and antennal clubs: a, Trichodes hihalteatiis, dorsal view; b, Tri- chodes hihalteatus, lateral view; c, Trichodes hihcdteatiis, lateral view; d, Trichodes apivorus apivorus, lateral view; e, Trichodes hicinctus, lateral view. scutellum, mesosternum, and legs atropurpureus; antennal funicle testaceous, club orange to piceous; elytra reddish orange or testaceous with complete and transverse, atropurpureus subbasal and medial bands, apices without a dark band, umbones pale; abdomen atropurpureus, becoming reddish orange toward apex. Head densely punctate with many punctations confluent; front feebly impressed above clypeus; antenna with club about one and one-half times as long as broad in side view (Fig. 4); clypeus testaceous and glabrous; labrum black, surface finely, sparsely punctate, front margin shallowly emarginate; galea short, slightly 22 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY longer than one-half distance between eyes across front. Pronotum moderately coarsely, densely punctate with many punctations confluent, very densely clothed with erect, coarse hairs. Scutellum densely punctate and roughened, disc flat¬ tened, hind margin truncate to deeply emarginate. Elytra about two and one- fourth times longer than humeral width, coarsely, reticulately punctate, umbones finely, sparsely punctate, sides expanded at posterior one-fourth, apices rounded. Mesosternum densely, moderately coarsely punctate; moderately densely clothed with suberect hairs. Metafemur not swollen disproportionately. Abdomen with sternites one through five sparsely punctate and roughened, some¬ times evanescent toward base; fifth sternite with hind margin deeply, arcu- ately emarginate; sixth sternite elongate, disc convex with basomedial surface glabrous, lateral margin arcuate, hind margin semicircularly rounded; eighth tergum narrower and longer than sixth sternite, disc shallowly impressed, im¬ pression clothed with dense, suberect hairs, hind angles bearing conspicuous tufts of long, stiff hairs, hind margin arcuately rounded and deflexed at middle. Aedeagus feebly arcuate; lateral lobes gradually constricted to bluntly rounded apices in lateral view, lateral lobes subparallel in dorsal view; median lobe lateral¬ ly compressed, apex ovoid (Fig. 4). Length: 8.1 to 1 7. 1 . Female. Similar to male except: abdomen with hind margin of fifth sternite truncate or shallowly emarginate; sixth sternite transverse, disc feebly convex; eighth tergum with disc slightly convex, sparsely clothed with long, suberect hairs, hind margin broadly, shallowly emarginate, not deflexed at middle. Length: 9.8 to 22.0. Type locality. — Not given with original description; type specimen bears a red disk = “Texas.” Type material. — Type, female, MCZC. Distribution. — Southern Kansas south to Texas and eastern New Mexico (Fig. 5). Wickham and Wolcott’s (1912) record from Winslow, Arizona, apparently resulted from a misidentification. Discussion. — T. bibalteatus is recognized as the only species in the fauna having reddish elytra with only two dark transverse bands and reddish elytral apices. Other similarly colored species, T. bicinctus, T. apivorus, and T. simula¬ tor, have three complete or incomplete dark transverse elytral bands, with the posterior band covering the apices. T. bibalteatus exhibits relatively little variation in elytral coloration and color pattern although a few specimens are more testaceous than reddish orange. Other features are much more variable. The coloration of the head and pronotum range from dull red to black, with most specimens exhibiting a mottled coloration somewhere between the two extremes. The antennal funicle is usually testaceous but appears brown in some specimens; the club varies from testaceous to piceous. Abdominal coloration also is quite variable. Most specimens have sternites one through four dark with part of the fifth and sixth sternites reddish orange. Testa¬ ceous coloration is expanded greatly on a few specimens; the most extreme variants have the abdominal sternites reddish orange, except for a dark medial band on the first four sternites. FOSTER — REVISION OE NORTH AMERICA TRICHODES 23 Fig. 5. — Distribution of the hihaheatus group: Trichodes bihcdteatus, coarse diagonal lines; Trichodes apivorus apivorns, diamonds; Trichodes apivorus borealis, fine diagonal lines; Trichodes bicinctiis, stippled. Specimens examined (623). — Texas; Kimble Co., Junction, TTU Center (D. E. Foster), 2dd, 1 9 (TTCC). For additional records, see Foster (1973). Trichodes apivorus Germar (Figs. 4, 5, 6) Trichodes apivorus Germar, 1824;8I; Klug, 1842;332; Spinola, I844;307; Horn, 1876;23I; Chittenden, I890;154; Wickham, 1895;249; Wolcott, 1910a;375; Wickham and Wolcott, 1912;59; Boving and Champlain, 1920;635; Forbes, 1922:fig. 48; Balduf, 1935;108; Wade, 1935;! 10; Barr, 1961;109. Clems apivorus, Gibson, 191 7; 149. Trichodes trifasciatus Sturm, 1826:59, 1843:83; Wolcott, 19106;46; Corporaal, 1948:288. Trichodes nuttalli. Say, 1825:164 (misidentification); Gemminger and von Harold, 1869:1742 (misidentification); Wolcott, 1944:60. Trichodes crihripennis Spinola, 1844:307. Trichodes nuthalli, Sp'mola, 1844:307 (mididentification). 24 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Description. — Male. Medium to large sized, robust; clothed with short to long, suberect and erect, reddish brown hairs; head, thorax, scutellum, and legs piceous; antennal funicle testaceous at base, becoming dark toward apex, club black; elytra red with atropurpureus subbasal, medial, and apical transverse bands either complete or interrupted, umbones pale; abdomen piceous with lat¬ eral and hind margins red. Head densely punctate with many punctations con¬ fluent; front transversely impressed between antennae; antenna with club nearly twice as long as broad (Fig. 4); clypeus testaceous and glabrous; labrum dark brown, with lateral and front margins testaceous, surface finely, sparsely punctate, front margin shallowly emarginate; galea short, slightly longer than one-half distance between eyes across front. Pronotum moderately coarsely, densely punctate with many punctations confluent, very densely clothed with erect, coarse hairs. Scutellum densely punctate and roughened; disc flattened; hind margin truncate or shallowly emarginate. Elytra about two and one-fourth times longer than humeral width, coarsely reticulately punctate, umbones finely, sparsely punctate, sides expanded at posterior one-fourth, apices rounded. Meso- sternum densely, moderately coarsely punctate; moderately densely clothed with suberect hairs. Metafemur not swollen disproportionately. Abdomen with ster- nites one through five sparsely punctate and roughened, often evanescent toward base; fifth sternite with hind margin deeply, arcuately emarginate; sixth stemite elongate, disc convex with basomedial surface glabrous, lateral margin arcuate, hind margin arcuately rounded; eighth tergum narrower and longer than sixth sternite, disc shallowly impressed, impression clothed with dense, suberect hairs, hind angles bearing conspicuous tufts of long, stiff hairs, hind margin arcuately rounded and deflexed at middle. Aedeagus strongly arcuate with lateral lobes gradually constricted to bluntly rounded apices in lateral view, subparallel in dorsal view; median lobe laterally compressed, apex crabriform with a dorsal claw and a lateral process at middle on either side (Fig. 6). Female. Similar to male except: abdomen with hind margin of fifth stemite transverse or shallowly emarginate; sixth sternite transverse, disc feebly convex; eighth tergum with disc slightly convex, hind margin tmncate or shallowly emarginate and not deflexed. Discussion. — T. apivorus occurs throughout the Atlantic and eastern Gulf Coast states from New Hampshire to Mississippi. Within its range are two dis¬ tinct climatic regions that can be correlated with variation found in the species. The southern Atlantic and Gulf seaboards, including southern Georgia, Florida, Alabama, and Mississippi, are characterized by southern pine forests. The climate is wet and warm, with freezing temperatures occurring only rarely. Populations of T. apivorus from this region are typified by adult occurrence from April to May. Populations of T. apivorus occurring from northern Georgia to New Hamp¬ shire are influenced by colder Atlantic coastal conditions. This region, charac¬ terized by both conifer and hardwood forests, has cold winter temperatures. Adults are common from June through July. The syntypic series of T. apivorus designated by Germar (1824) is indicative of populations inhabiting the southerly climate described above. Le Conte (1849) FOSTER— REVISION OF NORTH AMERICA TRICHODES 25 Fig. 6. — Male genitalia; a, Trichodes apivorns, dorsal view; b, Trichodes apivorus, lateral view; c, Trichodes hicinctus, dorsal view; d, Trichodes hicinctus, lateral view. recognized a northern variety of the species and described it as T. apivorus var. interruptus. Anatomical distinctiveness along with the unique geographical, ecological, and temporal occurrence of the two taxa suggest that they are distinctive entities separated by an area of reduced gene flow and worthy of subspecific recognition. The conspecific nature of the taxa is evidenced by the presence of a narrow band of hybridization in northern Georgia and the low coastal plain of South Carolina. Key to Subspecies o/ Trichodes apivorus Germar Medium to large sized; elytra with subbasal and medial dark bands complete to suture; abdominal sternites black with lateral margins red; southeast Georgia to Mississippi (Fig. 5) . apivorus Small to medium sized; elytra with subbasal and medial dark bands usually interrupted before suture; abdominal sternites black, first sternite occasionally with red lateral margins; Atlantic coastal states from northern New Hampshire to Georgia (Fig. 5) . borealis 26 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Trichodes apivorus apivorus Germar Trichodes apivorus Germar, 1824:81; Le Conte, 1849:18; Wickham, 1910:401; Wolcott, 1910«:395, 1944:55. Description. — Medium to large sized, robust; elytra with subbasal and medial dark bands broad and complete to sutural margin; abdominal sternites blackish with lateral and hind margins red. Length; male, 7.3 to 13.4; female, 8.2 to 15.0. Type locality. — “North America”; of T. trifasciatus Sturm, type specimen labeled “Georgia”; of T. cribripennis Spinola, “North America.” Type material. — The syntypic series of T. apivorus is deposited at the Zoologisches Museum der Humboldt-Universitat zu Berlin, Berlin, East Ger¬ many. The type of T. trifasciatus is in the Zoologisches Staatsmuseum zu Miinchen. Distribution. — Southeastern coastal states from Georgia to Mississippi (Fig. 5). A single specimen labeled “Louisiana” and several specimens labeled “Texas” appear erroneous. Discussion. — T. a. apivorus is the only representative of the genus occurring in the southeastern United States and is identified easily on that basis. It differs from its fellow subspecies most noticeably in elytral pattern. In T. a. apivorus, the dark subbasal and medial bands are broad, transverse, and produced usually to the sutural margin; the subbasal and medial bands of T. a. borealis are reduced to paired transverse maculations that fail to attain the sutural margin. The colora¬ tion of the abdomen serves as a second, highly reliable character for separating the taxa. All specimens of T. a. apivorus examined have the margins of the ab¬ domen red. Within this character, considerable variation exists, ranging from specimens with the abdomen red except for very reduced black medial markings to specimens with the abdomen predominantly black with the lateral and hind margins narrowly bordered in red. By contrast, the abdominal coloration of T. a. borealis features black lateral and hind margins. The only variation noted during the study was a few specimens in which the lateral margins of the first abdominal segment are red. Ninety-eight per cent of all specimens examined dur¬ ing the study can be separated to subspecies by abdominal coloration, making it an ideal character to use in support of the more conspicuous but less reliable characteristics of the elytral pattern. Specimens examined (172). — See Foster (1973). Trichodes apivorus borealis Wolcott and Chapin, new status Trichodes apivorus var. interruptus Le Conte, 1849:18 (nec King, 1842:331); Wolcott, 1910fl: 376; Wickham and Wolcott, 1912:59. Trichodes apivorus, Hamilton, 1895:335; Ulke, 1902:23,48; Knull, 1951:301; Kirk, 1969:61. Trichodes apivorus var. borealis Wolcott and Chapin, 1918:108; Wolcott, 1944:60; Knull, 1951:301. Description. — Small to medium sized, robust; elytra with subbasal and medial dark bands interrupted before the sutural margin on either elytron; ab¬ dominal sternites entirely piceous, first sternite sometimes red at lateral margins. Length: male, 6.4 to 12.8; female, 7.6 to 12.6. FOSTER— REVISION OF NORTH AMERICA TRICHODES 27 Type locality. — “Saratoga”; the type specimen bears an orange disk = “New York.” Type material. — Type, female, MCZC. Distribution. — The Atlantic coastal and New England states from northern Georgia to New Hampshire (Fig. 5). Discussion. — Separation of T. a. borealis from T. a. apivorus is discussed under that subspecies. T. a. borealis might be confused with T. nutalli with which it occurs sympatrically in the northern part of its range. Although the general coloration and elytral markings are somewhat similar, the two taxa are separated easily by a number of characters. The pronotum of T. a. borealis is clothed with dense, dark red hairs that partially obscure its surface from view. Pronotal hairs of T. nutalli are not nearly so dense, are whitish, and never obscure the surface from view. Also, the elytral punctations of T. a. borealis are large, conspicuous, and reticulate, whereas those of T. nutalli are small and inconspicuous and form no discernible pattern. Wolcott and Chapin (1918) stated that Klug (1842) used the name interruptus for a variety of T. apiarius, making Le Conte’s use of the name invalid. They proposed the replacement name borealis. Specimens examined (208). — See Foster (1973). Trichodes bicinctus Green (Figs. 4, 5, 6) Trichodes hicinctns Green, 1917:367; Wolcott, 1944:56. Description. — Male. Medium sized, robust; clothed with short to long, sub¬ erect and erect, testaceous to orange hairs; head, pronotum, scutellum, venter, and legs greenish black, often with an aeneous luster, lower surface of femur, tibia, and tarsus sometimes testaceous; antenna testaceous to dark brown; elytra reddish orange with paired, blackish subbasal, medial, and apical transverse maculations, umbones pale. Head densely punctate with many punctations con¬ fluent; front broadly impressed above clypeus and between eyes; antenna with club slightly longer than broad in side view (Fig. 4); clypeus testaceous and glabrous; labrum testaceous to dark brown, surface finely, sparsely punctured, front margin truncate or shallowly emarginate; galea longer than distance between eyes across front. Pronotum moderately coarsely, densely punctate with erect, coarse hairs. Scutellum densely punctate and roughened, disc convex, hind margin truncate or shallowly emarginate. Elytra about two and one-fourth times longer than humeral width, coarsely, reticulately punctate with umbones finely, sparsely punctate, sides expanded at posterior one-fourth, apices rounded. Mesosternum densely, moderately coarsely punctate, moderately densely clothed with suberect hairs. Metafemur not swollen disproportionately. Abdomen with sternites one through five sparsely punctate and roughened, sometimes evanescent toward base; fifth sternite with hind margin broadly, deeply, arcuately emarginate; sixth sternite elongate, disc convex with basomedial surface glabrous, lateral margin arcuate, hind margin truncate or shallowly emarginate; eighth tergum narrower 28 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY and longer than sixth sternite, disc deeply impressed, impression clothed with a distinct row of dense, coarse, suberect hairs along either side, hind angles bearing conspicuous tufts of long, stiff hairs, hind margin feebly arcuate or truncate, deflexed at middle. Aedeagus strongly arcuate; lateral lobes gradually constricted to podoform apices in lateral view, subparallel in dorsal view; median lobe laterally compressed, apex ovoid (Fig. 6). Length: 6.7 to 13.4. Female. Similar to male except: abdomen with hind margin of fifth sternite truncate, sometimes notched at middle; sixth sternite transverse, disc slightly convex, hind margin arcuate to semicircularly rounded; eighth tergum with a broad, deep longitudinal medial impression extending to hind margin, impression densely clothed with fine hairs, hind margin truncate, not deflexed at middle. Length: 7.0 to 14.6. Type locality. — Green Valley, “about 20 miles south of Marfa,” Brewster County, Texas. Type material. — Cotype, male, FMNH. This specimen compares favorably with the original description and hereby is designated to stand as the lectotype of T. bicinctus. Distribution. — Trans-Pecos, Edward’s Plateau, and Rolling Plains regions of Texas (Fig. 5). Discussion. — T. bicinctus occurs sympatrically with T. bibalteatus and T. oresterus, but can be separated easily from either species on the basis of color and color pattern. T. bicinctus differs from T. bibalteatus by having its elytral apices darkly tipped. Subbasal elytral maculations of T. bicinctus never attain the suture, whereas those of T. bibalteatus always broadly attain the suture. The postmedial elytral maculations of T. bicinctus are oblique, whereas those of T. bibalteatus are transverse. Living specimens of T. bicinctus have bright orange elytra with blackish maculations; the pronotum is very densely covered with conspicuous, bright orange hairs that nearly obscure its surface from view. The elytral colora¬ tion of T. oresterus is brick red with metallic verdigris markings. The pronotum is clothed with ash grey hairs, which are neither dense nor conspicuous and never obscure its surface. In his original description of T. bicinctus. Green described coloration of legs of both males and females, alluding to sexual dimorphism in that character. The greater number of specimens now available for study reveals that leg coloration is not sexually dimorphic, but is rather one of several variable features associated with coloration. Leg coloration ranges from entirely greenish black to a condition in which the basal portion of the femur, the ventral surface of the tibia, and the tarsi are testaceous. Most specimens have legs dark with tibial apices and tarsi testaceous. Variation also exists in the elytral pattern. Wolcott (1944) figures two variants of the postmedial and apical maculations. His figure 2 is indicative of the lectotype, in which the medial and apical maculations of each elytron are joined at both the suture and lateral margin. The condition is seen most fre¬ quently in specimens from southern Brewster County, Texas. Fifty-seven per cent of the material examined from Big Bend National Park exhibited this type of elytral marking. Wolcott’s figure 3 represents an intermediate condition in FOSTER— REVISION OE NORTH AMERICA TRICHODES 29 which the apical markings are somewhat reduced and not continuous with the medial maculations. This is the most common form throughout most of the species’ range. The most extreme reduction of the apical markings is seen in specimens from southern Val Verde County, Texas, in which they are reduced greatly, do not extend along the lateral margin, and are produced only slightly along the suture. Seventy-six per cent of the specimens seen from Val Verde County have reduced apical markings. Another variable feature of T. bicinctus is the coloration of the elytra in pre¬ served specimens. All specimens collected during this study have bright orange elytra, but older specimens tend to have the elytra more testaceous. An examina¬ tion of all material available for study reveals that as specimens age, the elytra gradually change in color, eventually becoming testaceous. Specimens examined (116). — Texas: Garza Co., Justiceburg, 4 mi. E, 1 May 1974 (D. E. Eoster), 1 d (TTCC). Eor additional records, see Eoster (1973). Ornatus Group Description. — Small to medium sized, robust; antenna with club more than twice as long as broad, apex obliquely rounded in side view; prothorax finely punctate and sparsely clothed with fine hairs; elytra with sides expanded at posterior one-fourth, apices rounded, surface finely and indistinctly punctate. Species included. — T. ornatus, T. nexus, T. nutalli, T. bimaculatus, and T. oregonensis. Trichodes ornatus Say (Figs. 7, 8, 9, 10) Trichodes ornatus Say, 1823:189; Klug, 1842:340; Spinola, 1844:327; Le Conte, 1849:18, 1859:120; Desmarest, 1860:table 19, fig. 2; Le Conte, 1866:55; Dike, 1875:820; Horn, 1876:231; Brodie, 1888:214; Horn, 1891:6; Wickham, 1895:249; Schenk- ling, 1906:286; Snow, 1907:176; Wolcott, 1909:99, 1910a:372; Wickham and Wolcott, 1912:59; Boving and Champlain, 1920:612; Linsley and Usinger, 1934: 105; Balduf, 1935:108; Wade, 1935:110; Moore, 1937:39; Linsley and MacSwain, 1943:589; Wolcott, 1944:58; Linsley and MacSwain, 1946:18; Barr, 1950:508, 1952:188; Nye and Bohart, 1952:6; Barr, 1961:109; Krombein, 1967:376; Parker and Bohart, 1968:1; Stephen et al., 1969:127; Barr, 1969:18; Ekis and Gupta, 1971 :60; Waters, 1971:1. Trichodes ornatus VdiX. obsolet us V^oXcoii, 1944:59. Description. — Male. Small to large sized, slender to robust; clothed with short and long, recumbent to erect, whitish to fulvous hairs; head, thorax, scutellum, venter, and legs dark metallic blue or greenish, sometimes with an aeneous luster, tibia and tarsus sometimes testaceous; antenna testaceous to piceous; elytra yellow or red with dark metallic blue or atropurpureus subbasal, medial, and apical bands, umbones dark. Head finely punctate and densely roughened; front broadly impressed between eyes and above clypeus; antenna with club twice as long as broad in side view (Fig. 7); clypeus testaceous to dark brown, glabrous; labrum dark brown, surface finely, sparsely punctate, front margin broadly, shal¬ lowly emarginate; galea short, no longer than one-half distance between eyes 30 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Fig. 7. — Male genitalia and antennal club: a, Trichodes oregonensis, dorsal view; b, Trichodes oregonensis, lateral view; c, Trichodes oregonensis— ornatus group, lateral view. across front. Pronotum with sides abruptly constricted at posterior one-fourth; finely, sparsely punctate and densely, rugosely roughened, moderately densely clothed with suberect and erect fine hairs. Scutellum sparsely punctate and densely roughened, disc flattened or sometimes medially impressed, hind margin rounded. Elytra about two and one-fourth or two and one-half times longer than humeral width, finely sparsely punctate and densely roughened with umbones roughened, sides expanded at posterior one-fourth, apices rounded. Mesostemum finely, sparsely punctate and densely, rugosely roughened; moderately densely clothed with long, recumbent and suberect, fine hairs. Metafemur not swollen disproportionately. Abdomen with sclerites one through five sparsely punctate and finely rugose; fifth sternite with hind margin deeply, arcuately emarginate; sixth sternite elongate, disc basomedially glabrous, lateral margin nearly straight, hind margin broadly, shallowly emarginate; eighth tergum longer and narrower than sixth sternite, disc shallowly impressed, impression clothed with moderately dense, suberect hairs, hind margin semicircularly rounded and deflexed at middle. Aedeagus arcuate; lateral lobes abruptly constricted before the bluntly rounded FOSTER — REVISION OF NORTH AMERICA TRICHODES 31 Fig. 8. — Male genitalia: a, Trichodes ornatiis hartwegianus, dorsal view; b, Trichodes ornatus hartwegianus, lateral view; c, Trichodes nexus, dorsal view; d, Trichodes nexus, lateral view. apices in lateral view, apices hooked and convergent in dorsal view; median lobe laterally compressed, apex ovoid (Fig. 8). Female. Similar to male except: abdomen with hind margin of fifth stemite broadly, very shallowly emarginate or truncate; sixth stemite transverse, disc feebly convex or flattened, hind margin broadly, very shallowly emarginate; eighth tergum with disc medially impressed, impression clothed with scattered, suberect hairs, hind margin broadly, shallowly emarginate. Discussion. — T. ornatus is a highly variable polytypic species occurring com¬ monly throughout much of western North America. It can be separated from other species of the ornatus group on the basis of characters presented in the key. The recurved apices of the lateral lobes of the aedeagus of this species provide a further means of identification. The more than 10,000 specimens of this species assembled and examined dur¬ ing the study reveal that within the aforementioned variation, a clear pattern exists. Five large composites of similar local populations are apparent, each oc¬ cupying a climatically, topographically, and geographically definable region. 32 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Similarities among populations and intermediates between adjacent popula¬ tions within each geographic region suggest that gene flow is operative at varying levels, thus uniting local populations, allowing them to function as a more or less cohesive evolutionary unit sharing a common gene pool. It is also apparent that gene flow between the five large composites is restricted. The apparent restricted gene flow between adjacent composites and the rela¬ tively free genetic exchange within each, coupled with a degree of anatomical and geographic distinctiveness, make it advisable to consider them subspecies. Each has reached the potential, if future selective pressures dictate, of evolving to the point of satisfying the biological species concept as illucidated by Mayr (1963). Despite their distinctiveness and apparent reproductive isolation in some areas, each of the five large composites shares zones of intergradation with one or more counterparts. Two types of intergradation are apparent. Along some major eco- tones, for example the high plateau of southeastern Wyoming and the transitional zone between the Mojave and Great Basin deserts, hybrid zones are characterized by populations comprised chiefly of intermediate forms between the two sub¬ species involved. Variation within transitional populations is not noticeably greater than that found in either parental stock. Such hybrid zones are suggested to result from restricted gene flow across ecotonal areas accompanied by the evolution of increasingly distinctive gene pools on either side. Mayr (1963) terms hybrid zones of this type “primary” because there is no evidence of past geo¬ graphic isolation and subsequent reunion. Such an interpretation does not neces¬ sitate, but certainly does not preclude, the possibility of past geographic separa¬ tion. Other zones of hybridization between subspecies are obviously secondary, having resulted from the reunion of former isolates that have at some past time been isolated long enough to acquire a measure of anatomical distinctiveness. Hybrid populations in such cases characteristically exhibit features of both con¬ tributing subspecies and a higher level of variation than exists in either contrib¬ utor. Hybridization between T. o. douglasianus and T. o. tenuosus in southern California and between the former subspecies and T. o. hartwegianus in the mountain ranges of northern Oregon clearly meet this criterion. Local populations within subspecies exhibit various levels of distinctiveness. In some instances, local populations that now hybridize with adjacent populations clearly have been isolated in the past and have gained a noticeable measure of anatomical distinctiveness. Because such populations are small and localized, the probability that they eventually will be swamped by the effects of hybridization are infinitely greater than for the larger subspecies level composites already dis¬ cussed. Such local populations are herein considered as infrasubspecific variants of the larger elements within which they occur. Variation. — Variation following several patterns is noted in different sub¬ species; these include phenotypic response to local conditions, local semi-isolates, and color dimorphism. Phenotypic response to local conditions is evident in T. o. douglasianus in which the occurrence of three distinctive phenotypes can be correlated with the elevations at which they occur throughout the Sierra Nevadas and coastal FOSTER — REVISION OF NORTH AMERICA TRICHODES 33 Table 1. — Elytral coloration o/ Trichodes ornatus adults reared from larvae maintained under controlled temperature and relative humidity. 4°C 20°C 36°C Room temperature Species 50% 75%. 50% 75% 50% 75% T. o. bonnevillensis No. larvae used 2 2 2 2 2 2 2 No. adults emerged 0 1 1 0 0 0 1 No. yellow elytra 0 0 0 0 0 0 0 No. red elytra 0 1 1 0 0 0 1 T. o. hartwegianus No. larvae used 8 8 8 8 8 8 8 No. adults emerged 3 1 0 2 0 1 No. yellow elytra 3 1 0 2 0 1 2 No. red elytra 0 0 0 0 0 0 0 mountain ranges of California. Another response to local conditions with respect to elytra patterns and vestiture is evident in all subspecies. Specimens from cool, damp habitats tend to have darkly pigmented elytral bands more broadly expanded and have darker hairs on the head, pronotum, venter, and legs than do specimens from warmer, drier areas. Local populations or groups of local populations in some subspecies, particu¬ larly in T. o. bonnevillensis and T. o. tenuosus, have existed as isolates at some past time in geologic history. During their isolation, certain of these populations apparently evolved rapidly, allowing the expression at high frequencies of other¬ wise uncommon phenotypes. The features most conspicuously involved are color and elytral patterns. Such phenomena occurring in small isolates are discussed by Mayr (1963) and Dobzhansky (1970). Color dimorphism is expressed in T. o. tenuosus and T. o. bonnevillensis, in which forms with both red and yellow elytra exist at varying frequencies in local populations. Rearing experiments conducted as a part of this study (see Table 1) suggest that the base coloration of the elytra is not a response to either the larval food source or to the conditions under which larvae develop and pupate. There is some evidence that red forms have some selective advantage in warm, arid habitats; in some instances, they predominate under other habitat conditions. For example, many local populations that exist under warm, arid conditions such as those found in Death Valley and the lower Colorado River Valley, along with local populations from cool, mesic mountainous areas of southeastern Idaho and western Wyoming, are predominantly red. Populations from the Snake and Creek mountain ranges of eastern Nevada, in an ecological situation somewhat similar to that of the aforementioned mountainous areas, have only yellow forms (Fig. 9). There is no evidence that the frequency of yellow and red forms varies clinally. Rather, it appears that red forms have comprised at some past time a large percentage of individuals found in many semi-isolated, local populations 34 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Eig. 9. — Percentage of red forms in local populations of Trichodes ornatus bonnevillensis and Trichodes ornatus tenuosiis. throughout the heterogeneous desert regions as evidenced by the high frequency of red individuals presently found in the Bagdad, Charleston Mountains, Pioche, Elba-Basin Pass, and Jackson populations (Fig. 9). Correspondingly, those peripheral populations of both desert subspecies that hybridize with adjacent “all yellow” subspecies exhibit lower frequencies of red forms than do popula- FOSTER— REVISION OF NORTH AMERICA TRICHODES 35 tions nearer the center of either subspecies range. A few peripheral populations, for example those at Wadsworth and Jackson, have retained a high percentage of red forms exhibiting a high degree of distinctiveness. In some instances, as in California and southwestern Utah, a gradual increase in the frequency of red forms is evident as one moves from the area of hybridization toward the center of the subspecies range. The occurrence of yellow forms in low percentages throughout most of the epicenters of the ranges of T. o. bonnevillensis and T. o. tenuosus and the absence of red forms from nondesert subspecies suggest a possible adaptive value for red forms in desert areas or, conversely, a disadvantage for yellow forms. In con¬ junction with these possibilities, it is interesting to note that throughout the desert regions local, predominantly yellow populations occur either in cool, relatively damp, high altitude habitats or in peripheral areas where the influence of ad¬ jacent subspecies is expressed. Another highly variable feature of T. ornatus adults is size. Whereas the size ranges exhibited among subspecies as well as between males and females are distinct, a great deal of variation exists within the individual parameters of each. Size variation in response to larval food reserves has been demonstrated in the laboratory. Individuals provided with only one alfalfa leafcutter bee cell were smaller at the time they constructed their pupal chambers than were individuals supplied with two leafcutter cells. The broad host range of T. ornatus, which in¬ cludes hosts of varied sizes and nesting habits, apparently contributes to the marked size variation exhibited by the species. In all five subspecies, there exists a tendency for specimens from northern parts of their ranges or from higher ele¬ vations to be relatively larger, with a gradual size reduction in progressively more southerly or lower elevation populations. Anatomical characters. — Five subspecies of T. ornatus are recognized. Tax¬ onomic characters employed are limited to external anatomical features of the adult. No means have been found for separating larvae of various subspecies. Furthermore, present knowledge of biologies and host associations offers no char¬ acters useful at the subspecies level. The anatomical characters most beneficial for separating subspecies include body shape, size, elytral markings, coloration, and vestiture, or some combination of these. Body size and shape. Whereas size is quite variable, it nevertheless falls with¬ in a distinctive range in some subspecies and provides a useful taxonomic char¬ acter. The shape of the lateral elytral margins is also distinctive. Both characters are most useful in separating T. o. hartwegianus and T. o. tenuosus from other subspecies. Elytral markings. The nature of the dark subbasal, postmedial, and apical bands on the elytra provide characters valuable in subspecies recognition even though the bands exhibit marked infrasubspecific variation, particularly in T. o. bonnevillensis. Principal features of value are the degree of constriction in the postmedial transverse band, shape of the subbasal band, and the degree to which the anterior angles of the subbasal band are produced to the sutural vitta. The 36 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY extent to which the dark bands attain and are produced along the lateral margins of the elytra and, conversely, the degree to which the yellow or red maculations attain the elytral margin are useful character states. Another feature associated with the dark elytral bands is the degree to which they are expanded and, consequently, the portion of the elytra covered by them. Specimens of T. o. hartwegianus have in excess of 50 per cent of the elytral sur¬ face covered by the dark bands, whereas in T. o. tenuosus and some specimens of T. o. bonnevillensis less than 25 per cent of the elytral surface is covered by the dark bands. Coloration. A conspicuous feature of T. ornatus is the occurrence of two color forms in the Sonoran, Mojave, and Great Basin deserts. The presence of both red and yellow forms facilitates the separation of T. o. tenuosus and T. o. bonnevillensis from other subspecies. Vestiture. Whereas the nature and coloration of hairs on the head and pronotum are not distinctive enough to serve as primary characters, they are of value in a supporting role. T. o. tenuosus is characterized by fine, sparse, whitish hairs. Those of other subspecies are more numerous, coarser, and frequently darker in color. Key to Subspecies o/Trichodes ornatus Say 1. Elytra yellow or red with metallic blue or greenish bands usually covering less than 50 per cent of surface, middle and subapical yellow or red maculations usually complete to lateral margin . 2 Elytra yellow with dark blue metallic markings covering more than 50 per cent of surface, middle and subapical yellow maculations never complete to lateral margin; northern Rocky Mountain states and Pacific Northwest, extending north to Alaska (Fig. 10) . . hartwegianus 2. Elytra yellow (never red) with blue or atropurpureus bands; coastal and mountainous regions of California, Colorado Plateau, and the Rocky Mountains of Colorado and New Mexico . 3 Elytra red or yellow with metallic blue or verdigris bands; Sonoran, Mojave, and Great Basin deserts . 4 3. Elytra with postmedial dark band strongly constricted on either side near sutural vitta; pubescence usually fulvous; California and western Oregon (Fig. 10) . douglasianus Elytra with postmedial dark band only slightly constricted on either side of sutural vitta; pubescence usually whitish; Colorado Plateau and Rocky Mountains (Fig. 10) . . . ornatus 4. Small and slender, elytra subparallel; hairs of head and pronotum whitish; Sonoran and Mojave deserts (Fig. 10) . tenuosus Large and robust, elytra distinctly expanded at posterior one-third; hairs of head and pronotum often dusky; Great Basin Desert and Snake River plains (Fig. 10) . . bonnevillensis Trichodes ornatus hartwegianus White, new status Tricliodes hartwegianus'Whhe, 1849:60; Cockerell, 1893:329; Schenkling, 1906:286. Trichodes ornatus, Le Conte, 1869:371; Anderson, 1914:57; Bush, 1914:60; Stace Smith, 1930:23. Description. — Medium to large sized, robust; clothed with whitish hairs; head, thorax, scutellum, venter, and legs dark metallic blue. Elytra about two and one- FOSTER— REVISION OF NORTH AMERICA TRICHODES 37 fourth times longer than humeral width; sides expanded at posterior one-third; surface yellow with broad, more or less transverse, dark metallic blue subbasal, postmedial, and apical bands covering more than 50 per cent of elytral surface, expanded along lateral margins and continuous with one another, subbasal band with front angles often confluent with sutural vitta, postmedial band with a feeble constriction on either side of sutural vitta. Length: male, 7.1 to 12.8; fe¬ male, 8.8 to 1 4.2. Type locality. — White incorrectly listed the type locality as “California”; the locality label on the type specimen reads “Rocky Mountains.” This specimen is indicative of populations occurring in the northern Rockies, the Pacific Northwest, and western Canada. Type material. — Type, female, British Museum (Natural History), London, England. Distribution. — Extreme western Nebraska and South Dakota west to Washing¬ ton, western Canada, and Alaska (Fig. 10). Discussion. — T. o. hartwegianus is separated easily from other subspecies of T. ornatus by the predominance of blue coloration on the elytra, usually covering more than 50 per cent of the surface, and by its large size and robust form. It is also unique in having the elytral margins entirely blue except for the front angles, whereas the other four subspecies have additional areas of yellow or red along the lateral margins. Throughout the range of T. o. hartwegianus, there is a tendency for specimens found at high elevations to have the dark coloration of the elytra greatly ex¬ panded. Some specimens from such localities have the yellow areas on the elytra reduced to small, isolated maculations. Conversely, there is a tendency for reduction of dark pigmentation and an expansion of yellow coloration in lower, drier climes. Specimens from the Columbia Basin of Washington exemplify the latter condition by having the yellow and dark areas about equal. Despite its distinctiveness, T. o. hartwegianus cannot be separated with cer¬ tainty from all local populations of T. o. douglasianus with which it shows a very complex band of hybridization in the Cascade and coastal mountain ranges of extreme southern Washington and northern Oregon. The pronounced topographic relief of the region complexes the situation, casting further doubt on the precise affinities of some local populations. Populations throughout most of the region are intermediate between the subspecies. Populations from Corvallis, Benton County, Oregon, and immediate adjacent localities are typical of this condition, with 57 per cent of the specimens exhibiting expanded dark elytral markings similar to the typical hartwegianus. However, the pronotal hairs are fulvous, the lateral margins of the elytra are not uniformly dark, and the body is only moder¬ ately robust, all features reminiscent of douglasianus. Eighteen per cent of the specimens from the area closely resemble douglasianus, whereas 25 per cent ap¬ pear similar to hartwegianus. A few localized populations found at higher elevations throughout central and northern Oregon, for example the Alsea Mountain, Benton County, popula¬ tion, have specimens resembling hartwegianus. Similar populations occur at Deer 38 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Fig. 10. Distribution of subspecies of Trichodes ornatus: T. o. hartwegianiis, diamonds; T. o. douglasianus, diagonal lines; T. o. ornatus, horizontal lines; 7. o. honnevillensis, stippled; 7. o. tenuosus, squares. Creek, Deschutes County; O’Brien, Jackson County, and in northeastern Jo¬ sephine County. Conversely, some populations, especially along the eastern side of the Cascades in northern Oregon and southern Washington, very closely re¬ semble douglasianus. For example, 80 per cent of the material examined from Mt. Adams and along Satus Creek, Yakima County, Washington, resemble douglasianus, with reference to the elytral pattern, coloration, and pronotal pubescence. FOSTER— REVISION OF NORTH AMERICA TRICHODES 39 T. o. hartwegianus intergrades with T. o. bonnevillensis in southeastern Oregon and southwestern Idaho and with the nominate subspecies in Albany, Goshen, and Platt counties of southeastern Wyoming. The former situation will be dis¬ cussed under T. o. bonnevillensis. In the latter case, inadequate specimens pre¬ clude discussion. However, from the few specimens available, it appears that populations in the area are intermediate between the two subspecies. Specimens examined (1637). — See Foster (1973). Trichodes ornatus douglasianus White, new status Trichodes doiiglasianus While, 1849:60; Cockerell, 1893:329; Schenkling, 1906:286. Trichodes ornatus, Essig, 1934:724; Linsley, 1942:164; Parker and Bohart, 1968:1 (in part). Description. — Small to medium sized, robust; clothed with whitish to fulvous hairs; head, thorax, scutellum, venter, and legs blue or greenish, sometimes with an aeneous luster, tarsi and tibial apices sometimes testaceous. Elytra about two and one-fourth times longer than humeral width; sides expanded at posterior one- third; surface yellow with atropurpureus subbasal, postmedial, and apical bands covering less than 50 per cent of elytral surface and seldom expanded along lateral margin, subbasal band oblique and narrow, attaining the lateral margin and with its front angles broadly separated from the sutural vitta, postmedial band transverse, broad at lateral margin with an abrupt, narrow constriction at the middle on each elytron. Length: male, 5.2 to 12.1; female, 7.1 to 13.7. Type locality. — “California.” Type material. — Type, female, British Museum (Natural History), London, England. Distribution. — Along the Pacific coast from central Oregon to northern Baja California and extending eastward to the Great Basin, Mojave, and Sonoran deserts (Fig. 10). Discussion. — T. o. douglasianus is found in a wide range of habitats from sea level to high elevations in the Sierra and Cascade mountains. Throughout its range, the subspecies is typified by the occurrence of three distinctive pheno¬ types. Despite such variation, the subspecies is recognized easily because it is the only one with yellow elytra in which the postmedial atropurpureus band is con¬ stricted abruptly and narrowly at the middle of each elytron to form a characteris¬ tic transverse hourglass shape. Phenotypes of T. o. douglasianus are understood most easily when they are considered with respect to the elevation at which they occur. The common phenotype found at elevations below 2500 feet is small, has the head, thorax, and abdomen dark metallic blue, and is clothed with whitish hairs. At middle elevations from 2500 to 5000 feet, most individuals tend to be large, have the head, thorax, and abdomen greenish with an aeneous luster, and are clothed with fulvous hairs. For example, 94 per cent of the specimens studied from Grass Valley, Nevada County, California (2600 feet), 95 per cent of the specimens seen from Quincy, Plumas County, California (3400 feet), and 81 per cent of the material from Meadow Lake, Plumas County, California (4000 feet), have greenish bodies with an aeneous luster and are clothed with fulvous hairs. 40 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY In the southern coastal mountains of California as well as in the northern reaches of the subspecies range, similar situations exist. Ninety-one per cent of material from Tanbark Flat, Los Angeles County, and all of the specimens examined from Grass Lake, Siskiyou County (5000 feet), have aeneous-fulvous coloration. At high elevations in the Sierra and Cascade mountains, the aeneous-fulvous coloration occurs less frequently, being replaced by smaller individuals with dark blue head, thorax, and abdomen, and clothed with whitish hairs. Sixty- seven per cent of the specimens from Conner’s Summit, Nevada County (7000 feet), and 87 per cent of the specimens from near Sonora Pass, Tuolumne County (9000 feet), are of this phenotype. A few specimens from high elevations in the Sierra and Cascade mountains have the dark blue elytral bands greatly expanded, leading to possible confusion with T. o. hartwegianus. Such specimens are smaller and are clothed with hairs not nearly as white as those found in the northern subspecies. Zones of hybridization exist between T. o. douglasianus and three adjacent subspecies. It hybridizes with T. o. hartwegianus in west-central Oregon, with T. o. bonnevillensis along the Nevada-Califomia border southwest of the Warner Mountains and again in the area around Topaz Lake and Lake Tahoe, and with T. o. tenuosus in southern Mono and northern Inyo counties, northern Kern County, and along the San Jacinto and Santa Rosa mountains of southern Cali¬ fornia. Areas of hybridization are discussed under the three above mentioned subspecies. Specimens examined (3568). — See Foster (1973). Trichodes ornatus ornatus Say Trichodes ornatns Say, 1823:189; Le Conte, 1878:472; Cockerell, 1893:329, 1898:155; Town¬ send, 1895:44. Trichodes ornatns var. ohsoietnsV^o\coii, 1944:59. Clerns ornatns Packard, 1 877:813. Description. — Medium sized, robust; clothed with whitish or fulvous hairs; head, thorax, scutellum, venter, and legs dark metallic blue or greenish with an aeneous luster, tarsi and sometimes tibial apices testaceous. Elytra about two and one-fourth times longer than humeral width; sides expanded at posterior one- fourth; surface yellow with dark metallic blue or atropurpureus subbasal, post- medial, and apical bands covering less than 20 per cent of elytral surface and moderately expanded along lateral margin with postmedial and apical bands con¬ tinuous in some specimens, subbasal band oblique, narrow or moderately broad, seldom attaining lateral margin and with its front angles at least narrowly sep¬ arated from the sutural vitta, postmedial band broad with a feeble constriction on either side of sutural vitta. Length: male, 6.9 to 1 1.4; female, 7.5 to 13.3. Type locality. — “Arkansa Territory.” A review of Major Stephen Long’s ex¬ pedition along the eastern slope of the Colorado Rockies (Goetzmann, 1959) shows that members of the party followed the Arkansas River west into the area of Royal Gorge and on toward Salida, Colorado. It is only in this segment of their recorded route that members of the party could have entered the presently estab- FOSTER— REVISION OF NORTH AMERICA TRICHODES 41 lished range of the species. It seems very likely that the type of T. ornatus was col¬ lected from that area. The type locality of T. o. var. obsoletus is the Sierra Ancha Mountains, Ari¬ zona. Type material. — The type of T. ornatus is unknown, presumably having been destroyed along with many of Say’s other types. Holotype, male, of T. o. var. obsoletus, FMNH. Distribution. — Rocky Mountain ranges of Colorado, New Mexico, and ex¬ treme northern Mexico, west to the Great Basin and Sonoran deserts (Fig. 10). Discussion. — Characters of the elytral pattern offer easy separation of T. o. ornatus from the other two subspecies with yellow elytra. The elytral margins of the nominate subspecies have alternating yellow and dark areas. The postmedial dark band is broad with only a slight constriction at the middle on each elytron. T. o. hartwegianus has the lateral margins of the elytra entirely blue except for the front angles, whereas in T. o. douglasianus the postmedial elytral band is con¬ stricted narrowly at the middle on each elytron. Most specimens of T. o. ornatus are typical of those found along the Arkansas River in the Colorado Rockies. They are medium sized, have metallic blue bodies, legs, and elytral markings, and are clothed with whitish hairs. The only notable exception to the typical condition is found in the Sierra Ancha Mountains of east- central Arizona and extending north and east from that area into extreme south¬ eastern Utah, southwestern Colorado, and northwestern New Mexico. Popula¬ tions from these areas frequently have a high percentage of specimens with body coloration greenish with an aeneous luster, atropurpureus elytral markings, a prominence of reddish coloration on the tibia and tarsi, and clothed with fulvous hairs. This condition occurs in all specimens studied from the Navajo Mountains of San Juan County, Utah, and Navajo County, Arizona; in 94 per cent of the specimens from the Sierra Ancha Mountains and 84 per cent from the White Mountains of Arizona; in 81 per cent of the specimens from Jemez Springs and Fort Wingate, Sandoval County, and in both specimens seen from the Tsankawi Indian Ruins, Santa Fe County, New Mexico; and in 66 per cent of the specimens from the Cortez vicinity, Montezuma County, Colorado. It was from the Sierra Ancha population that the holotype of T. o. var. obsole¬ tus was collected by D. K. Duncan. In his original description, Wolcott (1944) emphasized the indistinct nature of the dark elytral markings, a feature not found in other representatives from that population. Two other specimens collected by Duncan and bearing the same collection data as the holotype have similarly faded elytral markings. It seems likely that this material and the holotype have been exposed to some foreign substance, resulting in their discoloration. Aside from the features of the elytra, Wolcott’s description is indicative of the populations just discussed. However, the disjunct, isolated, and restricted distributions of these populations, the occurrence of high frequencies of the aeneous-fulvous forms only in mountainous areas, and their occurrence at lower frequencies in intermediate habitats dominated by more typical forms of T. o. ornatus as ex¬ emplified in New Mexico by the Jemez Springs, Wingate, and Cortez populations 42 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY make it advisable to consider them local variants of that subspecies and to treat T. o. var. ohsoletus as a junior synonym. Two zones of overlap occur between T. o. ornatus and adjacent subspecies. These are discussed under T. o. hartwegianus and T. o. honnevillensis. Specimens examined (963). — See Foster (1973). Trichodes ornatus tenuosus Foster, replacement name, new status Trichodes tenelliis Le Conte, 1859:72; Snow, 1907:176. Trichodes ornatus, Davidson, 1896:25; Parker and Bohart, 1968:1 (in part). Trichodes ornatus var. tenellus Horn, 1891:7; Schenkling, 1906:286; Wolcott, 1910a:373; Wickham and Wolcott, 1912:59; Davis, 1932:82; Wolcott, 1944:55. Trichodes ornatus tenellus Linsley and MacSwain, 1943:592. Description. — Small sized, slender; clothed with whitish hairs; head, thorax, scutellum, venter, and legs metallic blue, verdigris in some specimens, tarsi and tibial apices sometimes testaceous. Elytra about two and one-half times longer than humeral width; sides subparallel; surface red or yellow with dark metallic blue or verdigris subbasal, postmedial, and apical bands covering much less than 50 per cent of elytral surface and never expanded along the lateral margin, sur¬ face band either narrow and oblique, reduced to isolated maculations, or absent, its front angles always widely separated from sutural vitta, postmedial band very strongly constricted and sometimes interrupted on either side of sutural vitta. Length; male, 4.9 to 9. 1 ; female, 6.0 to 1 1 .4. Type locality. — Fort Yuma, Imperial County, California. Type material. — Type, female, MCZC. The type specimen bears a gold disk = “California.” Distribution. — Sonoran and Mojave deserts of California, southern Nevada, western Arizona, and northern Sonora, Mexico (Fig. 10). Discussion. — T. o. tenuosus is one of the most easily recognized subspecies because of its small size and distinctive parallel-sided, slender form. Infrasub¬ specific variation except for color and size is relatively slight. In some populations from the central part of the subspecies range, 84 to 100 per cent of the individuals have red elytra. Nearly all such specimens are clothed with whitish hairs. Less isolated populations or those nearer zones of hybridization exhibit an increase in the frequency of individuals with yellow elytra and dusky colored hairs. Three zones of hybridization between T. o. tenuosus and adjacent subspecies is apparent. Two of these are shared with T. o. douglasianus in southern Cali¬ fornia; one along the eastern side of the San Jacinto and Santa Rosa mountains and in the area of Anza-Borrego State Park, and the second in northern Kern County. In both instances, intermediate populations appear more variable than adjacent populations of either contributing subspecies. Specimens from both zones are quite variable in size as well as coloration of elytra and vestiture. The frequency of red forms found in hybridizing populations along the eastern side of the two mountain areas ranges from 18 to 31 per cent, whereas 47 per cent of the specimens of the Kern County hybrid bands are red. Forty-four per cent of the specimens from these zones have whitish vestiture, 26 per cent have fulvous F.OSTER— REVISION OF NORTH AMERICA TRICHODES 43 vestiture, and 30 per cent are intermediate between the extremes. The third zone of overlap is discussed under T. o. bonnevillensis. The proposal of tenuosus as a replacement name for tenellus Le Conte is neces¬ sitated by Say’s (1835) use of tenellus for a variety of his Trichodes verticalis = Phyllobaenus verticalis (Say). Le Conte’s tenellus is thus a junior primary homo¬ nym and must be rejected. Etymology. — The replacement name tenuosus refers to the distinctive slender form of this subspecies. Specimens examined (2354). — See Foster (1973). Trichodes ornatus bonnevillensis Foster, new subspecies Trichodes ornatus. Tanner, 1928:273; Horning and Barr, 1970:31. Trichodes nntalli, Knowlton, 1930:37; Barr, 1961:109 (misidentifications). Description. — Medium sized, robust; clothed with dusky or whitish hairs; head, thorax, scutellum, venter, and legs dark metallic blue, tarsi and tibial apices sometimes testaceous. Elytra about two and one-fourth times longer than humeral width; sides expanded at posterior one-third; surface red or yellow with dark metallic blue subbasal, postmedial, and apical bands covering less than 50 per cent of elytral surface and moderately expanded along lateral margin with post- medial and apical bands continuous in some specimens, subbasal band either broad and transverse, narrow and oblique, reduced to isolated maculations, or absent, postmedial band usually broad with a feeble constriction on either side of sutural vitta, appearing narrow or incomplete in a few specimens. Length: male, 6.1 to 12.0; female, 6.3 to 13.2. Type material. — Holotype 6 and allotype, Austin, Lander Co., Nevada, 12 mi. E, 16 July 1969 (D. E. Foster and R. L. Penrose), TTCC. Paratypes are de¬ posited in the collections of AMNH, CASC, FMNH, NMNH, TTCC, CISC, UICM, and in the personal collections of W. F. Barr and D. E. Foster. Distribution. — Great Basin Desert and Snake River plains of Idaho, Oregon, Nevada, and Utah (Fig. 10). Discussion. — The dimorphic coloration shared by T. o. bonnevillensis and its southern desert counterpart, T. o. tenuosus, makes them easy to separate from the other subspecies of T. ornatus, none of which has red forms. T. o. bonnevillensis is distinguished from T. o. tenuosus by its robust form and the expanded sides of its elytra. The range occupied by T. o. bonnevillensis has had an interesting geologic history involving the building of north-south mountain ranges in Nevada and Utah and the development and subsequent drying of the Bonneville and Lahontan lake systems (Blackwelder, 1948). As a result of varied local topography and climatic conditions supplemented by temporary physical barriers provided by the lake system, separate local populations apparently developed. These popula¬ tions, which are presently connected by intermediate forms, collectively com¬ prise the large panmictic center of the subspecies. Populations throughout most of the central region are characterized by a pre¬ dominance of red forms, by pronotal hairs that appear dusky in 82 per cent of 44 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY the specimens studied, and by the narrow, oblique, and complete nature of the subbasal elytral band, a feature occurring in 99 per cent of the specimens. T. o. honneviltensis hybridizes with all four of its fellow subspecies. A narrow zone of secondary intergradation between it and T. o. hartwegianus occurs along the northern edge of the Snake River plains in Butte, Blaine, Camas, and Ada counties, Idaho. Specimens from Butte and Blaine counties are indicative of the situation found along the entire hybrid band. Forty-one per cent of the specimens examined had dark elytral margins indicative of T. o. hartwegianus, whereas 22 per cent were typical of T. o. bonnevillensis. The remaining 37 per cent are inter¬ mediate. The frequency of red individuals is 14 per cent, much lower than the frequencies noted for the large panmictic center of T. o. bonnevillensis. Although T. o. bonnevillensis and T. o. douglasianus show little tendency toward hybridization, a few populations along the Califomia-Nevada border between Topaz Lake and the Warner Mountains have a high percentage of yellow forms, whereas the elytral markings of most specimens are similar to those of T. o. bonnevillensis. Along the eastern side of the Great Basin Desert a complex area of hybridiza¬ tion occurs. Populations from northeastern Utah exhibit marked variation re¬ sulting from both inter and infrasubspecific hybridization. T. o. bonnevillensis populations from both the central Great Basin area and southeastern Idaho, along with elements of T. o. ornatus from western Colorado are involved. Because of the occurrence of red forms, which range from 1 8 to 64 per cent, these popula¬ tions are considered to be T. o. bonnevillensis. Three local populations of T. o. bonnevillensis have attained a level of dis¬ tinctiveness that may warrant their recognition as separate subspecies. However, their localized nature, low population densities, and the degree to which they intergrade with adjacent populations make it advisable to discuss each of these separately as an extreme local variant of T. o. bonnevillensis. Pioche Population, Lincoln County, Nevada. The Pioche population occurs along the east-facing slope of the Highland range, an area characterized by a juniper-sage community. The population is distinguished by a reduction of the dark elytral markings, a narrow and somewhat linear shape, and the complete absence of yellow forms. Of the 29 specimens from the Pioche locality examined, seven lack the subbasal elytral band, and 14 have the band very faintly and in¬ completely indicated. Eight specimens have very narrow but complete subbasal bands. The subbasal and postmedial bands are never produced to the lateral margins. The postmedial dark band appears narrow and discontinuous in four specimens, constricted medially on each elytron but complete in 1 1 specimens, and broad and complete in 14 specimens. There is strong evidence of hybridization between the Pioche population and adjacent populations in Washington County, Utah. A tendency toward reduction of the dark elytral bands and a more linear form is expressed in six of 22 speci¬ mens examined from that county. The tendency toward the reduction of dark elytral markings is not restricted to the Pioche population. Similar phenotypes are found in widely disjunct por- FOSTER— REVISION OF NORTH AMERICA TRICHODES 45 lions of the subspecies range, as well as in widely separated localities within the range of T. o. tenuosus. All of these cases are restricted to the occurrence of isolated specimens that comprise only a fraction of their local populations. There is no reason to associate such occurrences with the Pioche population or to assume a close genetic relationship among the populations involved. Elba-Basin Pass Population, Cassia County, Idaho. This population occurs in a high altitude mountain meadow situation characterized by an aspen-sage community. Of the specimens examined, 98 per cent have red base coloration of the elytra and most have the dark elytral bands noticeably expanded relative to other populations of T. o. bonnevillensis. Twenty-four per cent of the specimens have the subbasal dark band expanded to the point that it appears transverse, a feature that has led to misidentification of some specimens as T. nutalli. Hy¬ bridization between the Elba-Basin Pass and adjacent populations is evidenced by specimens examined from eight miles west of Elba and from Silent City of Rocks, Cassia County. Twenty per cent of the specimens from the former locality have the subbasal band broad and transverse, whereas specimens from the latter locality do not exhibit the trait. Also, the frequency of yellow forms, 33 and 30 per cent respectively, is intermediate between the Elba-Basin population and that found in neighboring populations. The Elba-Basin Pass population is unique among local populations found in higher elevations in the northern Great Basin region. However, some populations found throughout most of the mountainous regions of southeastern Idaho are somewhat similar in that they exhibit a higher than normal frequency of red forms; occasional specimens throughout this region appear similar to the pheno¬ type expressed in the Elba-Basin Pass population. Western Wyoming Population. Populations occurring west of the Wind River Mountains, from central Lincoln County, Wyoming, north into Yellowstone Park, are quite distinctive. They are characterized by large size, robust form, constant color pattern, and the occurrence of only red forms. The dark elytral bands on most specimens resemble those of typical T. o. bonnevillensis. A most interesting attribute of these populations is a narrow zone of overlap with T. o. hartwegianus that shows no evidence of intergradation. Apparently, the two subspecies have achieved isolating mechanisms sufficient in this area to preclude hybridization. Etymology. — The name bonnevillensis is used with reference to the Bonneville lake system, the history of which has greatly influenced the current distribution and variability of this subspecies. Specimens examined (1626). — Among the specimens examined, 45 are designated as primary and secondary types as follows: Holotype d and allotype, Nevada: Lander Co.: Austin, 12 mi. E, 16 July 1969 (D. E. Foster and R. L. Penrose). Forty-three paratypes, Nevada: Lander Co.: Austin, 12 mi. E, 16 July 1969 (D. E. Foster and R. L. Penrose), 1 9; Austin, 8 mi. E, 16 July 1969 (D. E. Foster and R. L. Penrose), 2 dd; Austin Summit, 2.5 mi. E, 6 June 1960 (J. W. MacSwain), 2 dd, 1 9; (R. W. Thorp), 1 d; Austin Summit, 6 June 1960 (E. G. Linsley), 1 9. Idaho: Owhyee Co.: Grasmere, 6 mi. SE, 6 August 1965 (L. S. Hawkins, Jr.), 1 9; Marys Creek, 17 July 1957 (R. A. MacKie), 1 9; (G. B. Hewitt), 1 d; 3 May 1969 (A. R. Gittins), 1 d; Murphy Hot Springs, 5 mi. NE, 7 July 1953 (T. B. O’Connell), 1 d, l9; 46 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Murphy Hot Springs, 20 June 1965 (W. F. Barr), 1 d, 2 99; Cassia Co.: City of Rocks, 5 June 1969 (D. E. Foster), 3 dd, 4 99; Elba, 9 mi. W, 16 July 1964(R. L. Westcott), 2 dd; Elba, 8 mi. W, 5 June 1969 (D. E. Foster), 5 dd, 8 99. Utah: Box Elder Co.: Kelton, 29 June 1969 (G. F. Knowiton), 1 9; 30 June 1969 (G. F. Knowiton and W. J. Hanson), 2 dd; Kelton Pass, 24 October 1972 (G. F. Knowiton), 1 9. For additional records, see Foster (1973). Trichodes nexus Wolcott (Figs. 8, 11) Trichodes nexus Wolcott, 1910a:372; Wickham and Wolcott, 1912:59; Wolcott, 1944:55; Barr, 1950:507. Description. — Male. Small, robust; clothed with short and long, suberect and erect, very fine, whitish hairs; head, thorax, scutellum, venter, and legs very dark metallic blue sometimes with a greenish cast, abdomen with lateral and hind margins sometimes reddish, tibia and tarsus becoming increasingly reddish towards apex; antenna testaceous or reddish; elytra yellow with broad, blackish subbasal, medial, and apical bands, umbones dark, umbonal spot produced to front margin. Head sparsely punctate and densely roughened; front impressed above clypeus on either side; antenna with club about twice as long as broad; clypeus testaceous or glabrous; labrum black with broadly, arcuately emarginate front margin testaceous, surface finely, sparsely punctate; galea short, no longer than one-half distance between eyes across front. Pronotum with sides abruptly constricted at posterior one-fourth; finely, sparsely punctate and densely, rugosely roughened, moderately densely clothed with suberect and erect, fine hairs. Scutellum sparsely punctate and densely roughened, disc flattened, hind margin arcuate. Elytra about two and one-fourth times longer than humeral width, finely, sparsely punctate and densely roughened with umbones roughened, sides expanded at posterior one-fourth, apices rounded. Mesosternum finely, sparsely punctate and densely, rugosely roughened, moderately densely clothed with long, recumbent and suberect, fine hairs. Metafemur not swollen dispropor¬ tionately. Abdomen with sternites one through five sparsely punctate and finely roughened; fifth sternite with hind margin deeply, arcuately emarginate; sixth sternite elongate, disc convex with basomedial surface glabrous, lateral margin nearly straight, hind margin broadly, triangularly emarginate; eighth tergum narrower and longer than sixth sternite, disc shallowly, longitudinally impressed, impression covered with dense, suberect hairs, hind margin rounded and deflexed at middle. Aedeagus feebly arcuate with lateral lobes produced into long, slender, bluntly rounded apices in lateral view, parallel in dorsal view; median lobe laterally compressed, apex ovoid (Fig. 8). Length: 7.4 to 9.0. Female. Similar to male except: abdomen with hind margin of fifth sternite truncate; sixth sternite transverse, disc slightly convex, hind margin rounded; eighth tergum with disc medially impressed to hind margin, impressed area clothed with scattered, suberect hairs, hind margin broadly, shallowly notched. Length: 7.8 to 9.2. Type locality. — San Jose del Cabo, Baja California. Type material. — Holotype, female, FMNH. FQSTER— REVISION OF NORTH AMERICA TRICHODES 47 Fig. 11. — Distribution of Trichodes nexus, stippled; Trichodes hitnaculatiis, shaded; Trichudes oregonensis, horizontal lines. Distribution. — Lower Baja California (Fig. 1 1). Discussion. — The remote and inaccessible range of T. nexus has contributed to the lack of specimens and information necessary for a comprehensive study of variation within the species and establishment of its relationships with other species of the ornatus group. T. nexus seems to be fairly consistent in appearance. However, Wolcott (1910a) discusses variation in coloration of abdominal sternites in which the basic coloration ranges from blue to green, with the lateral margins of the basal segments and sometimes the entire fifth and sixth segments appearing reddish. [ 48 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY T. nexus most closely resembles T. ornatus in elytral coloration and color pat¬ tern. Both species have yellow elytral coloration and an oblique subbasal elytral band. Elytral bands of T. nexus are much broader than the alternate yellow areas, which appear as narrow, oblique maculations. Yellow forms of T. ornatus have the dark elytral bands narrower than the yellow areas. The umbone spots of T. nexus are produced to the front elytral margin, whereas those of T. ornatus are never produced anteriorly. T. nexus occurs sympatrically with T. p. peninsularis, but not with T. ornatus. The allopatric but adjacent occurrence of T. nexus and T. ornatus further sug¬ gests that they are related more closely to one another than to the other species of the ornatus group. Specimens examined (5). — See Foster (1973). Trichodes nutalli (Kirby) (Figs. 12, 13) Clerns nutalli Kirby, 1818:394. Trichodes nutalli Klug, 1842:337; Le Conte, 1849:18; Chapin, 1917:31; Wolcott, 1944:60; Knuil, 1951:300; Dillon and Dillon, 1961;279; Ekis and Gupta, 1971:61. Trichodes humeral is Spinola, 1 844:3 1 7. Trichodes nuthalli, Spinola, 1844:317. Trichodes nuttalli, Horn, 1876:231; Wickham, 1892;141, 1895:294; Wolcott, 1910u:374, 19106:852; Wickham and Wolcott, 1912:59; Leng, 1915:254; Boving and Champlain, 1920:635; Carr, 1920:6; Balduf, 1935:108; Chagnon, 1935:173. Trichodes apivorus, Emmons, 1854:91; Lugger, 1 899: 149 (misidentifications). Description. — Male. Small sized, robust; clothed with short to long, suberect and erect, fine, whitish hairs; head, thorax, scutellum, venter, and legs dark metal¬ lic blue; antennal funicle testaceous to dark brown, club dark brown to piceous; elytra reddish orange with atropurpureus subbasal, medial, and apical trans¬ verse bands, umbones dark, umbonal spot produced to front margin. Head densely punctate with numerous punctations confluent; front impressed above clypeus on either side; antenna with club about twice as broad as long; clypeus dark testaceous and glabrous; labrum dark brown, surface finely, sparsely punctate, front margin emarginate; galea short, no longer than one-half distance between eyes across front. Pronotum with sides gradually constricted behind middle, finely, sparsely punctate and densely roughened, clothed with moderate¬ ly dense, suberect to erect, fine, whitish hairs. Scutellum sparsely punctate and densely roughened, disc feebly convex or flattened, sometimes with a medial impression extending to the arcuate hind margin. Elytra about two and one- fourth times longer than humeral width, finely, sparsely punctate and densely roughened with umbones roughened, sides expanded at posterior one-fourth, apices rounded. Mesosternum finely, sparsely punctate and densely, rugosely roughened; moderately densely clothed with long, recumbent and suberect fine hairs. Metafemur not swollen disproportionately. Abdomen with sternites one through five sparsely punctate and finely roughened; fifth stemite with hind margin deeply, arcuately emarginate; sixth sternite elongate, disc convex with basomedial surface glabrous, lateral margin feebly arcuate, hind margin slightly FO.STER— REVISION OF NORTH AMERICA TRICHODES Fig. 12. — Male genitalia: a, Trichodes niitalli, dorsal view; b, Trichodes nutalli, lateral view; c, Trichodes himaculatus, dorsal view; d, Trichodes hinuiculatus, lateral view. arcuate or truncate; eighth tergum narrower and longer than sixth stemite, disc shallowly, longitudinally impressed, impression covered with moderately dense, suberect hairs, hind margin rounded and deflexed at middle. Aedeagus feebly arcuate with lateral lobe gradually constricted, arcuately curved to the rounded apices in lateral view, divergent in dorsal view; median lobe laterally compressed, apex ovoid (Fig. 12). Length; 7.0 to 9.4. Female. Similar to male except: abdomen with hind margin of fifth stemite truncate; sixth stemite transverse, disc feebly convex, sometimes medially im¬ pressed to semicircularly rounded hind margin; eighth tergum with disc medially impressed to hind margin, impressed area clothed with scattered, suberect hairs, hind margin broadly, triangularly notched. Length; 7.2 to 10.5. Type locality. — “North America.” Type material. — Type, female, British Museum (Natural History), London, England. Distribution. — Southern Canada and northern United States east of the Conti¬ nental Divide (Fig. 13). Discussion. — Many features of T. nutalli, especially size, coloration, and color pattern, exhibit little variation and rival the consistency found in T. 50 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY oregonensis and T. bimaculatus. This is especially remarkable considering the large and varied geographic range inhabited by the species. Furthermore, it seems likely that larval host specificity is involved with subsequent adult size and that the larval biology of T. nutalli is similar to that of T. oregonensis. In the eastern part of its range, T. nutalli occurs sympatrically with T. a. borealis-, their separation has been discussed under that taxon. On the high plains and along the eastern slope of the Rocky Mountains, T. nutalli and T. ornatus occur sympatrically. In the area of sympatry, they can be separated by either of two easily observable elytral characters. T. nutalli has the subbasal band transverse and its umbonal spot produced to the front elytral margin, whereas the subbasal bands appear oblique in T. ornatus and the umbonal spots never attain the front margin of the elytra. Separation of sympatric populations is facilitated further by the occurrence of only yellow forms of T. ornatus. Reports recording the occurrence of T. nutalli in Idaho have found their way into the literature (Barr, 1961). An exhaustive examination of both species has shown the invalidity of such records, but has revealed that some specimens of T. o. bonnevillensis from southern Idaho and western Wyoming superficially resemble T. nutalli. The nature of the umbone spots (already discussed) and the distinctiveness of the lateral lobes of the aedeagus allow positive identification. Specimens examined (1636). — See Foster (1973). Trichodes bimaculatus Le Conte (Figs. 11, 12) Trichodes bimaculatus Le Conte, 1867:63; Wolcott, 1910«:374, 1944:55; Barr, 1952:188, 1961:109. Trichodes hisignatiis Horn, 1876:231, 1891:7; Wolcott, 1910a:374; Barr, 1952:188, 1961: 109. FOSTER— REVISION OF NORTH AMERICA TRICHODES 51 Description. — Male. Small to medium sized, moderately robust; clothed with short to long, suberect and erect, very fine, whitish hairs; head, thorax, scutellum, venter, and legs dark metallic blue or nearly black; antennal funicle testaceous at base, becoming dark toward apex with club dark brown; elytra dark metallic blue or nearly black with a large subrectangular to rounded maculation near middle of either elytron, umbones dark. Head densely punctate with numer¬ ous punctations confluent; front shallowly impressed above clypeus on either side; antenna with club more than twice as long as broad; clypeus testaceous and glabrous; labrum dark brown, surface finely, sparsely punctate, front margin truncate or shallowly emarginate; galea short, slightly shorter than one-half distance between eyes across front. Pronotum with sides gradually constricted behind middle; finely, sparsely punctate and densely roughened, clothed with scattered, suberect and erect, fine hairs. Scutellum sparsely punctate and rough¬ ened, disc flattened, hind margin semicircularly rounded. Elytra about two and three-fourths times longer than humeral width, finely, sparsely punctate and densely roughened with umbones roughened, sides expanded at posterior one- fourth, apices rounded. Mesosternum finely, sparsely punctate and densely rugosely roughened, clothed with widely scattered, long, suberect hairs. Meta¬ femur not swollen disproportionately. Abdomen with stemites one through five sparsely punctate and finely roughened; fifth sternite with hind margin deeply, arcuately emarginate; sixth sternite elongate, disc convex with basomedial surface glabrous, lateral margin arcuate, hind margin feebly arcuate; eighth tergum nar¬ rower and longer than sixth sternite, disc deeply, longitudinally impressed, im¬ pression bearing a row of dense, suberect hairs along either side, hind margin rounded and deflexed at middle. Aedeagus feebly arcuate with lateral lobes gradually constricted to rounded apices in lateral view, subparallel in dorsal view; median lobe laterally compressed, apex ovoid (Fig. 12). Length: 7.4 to 9.6. Female. Similar to male except: abdomen with hind margin of fifth sternite truncate; sixth sternite transverse, disc feebly convex, hind margin arcuate with a small triangular notch at middle; eighth tergum with disc medially impressed to hind margin, impressed area clothed with scattered suberect hairs, hind margin truncate with a deep triangular notch at middle, not deflexed. Length: 8.3 to 10.3. Type locality. — “California and Oregon”; the type specimen bears a label stating “Or.” A type locality for T. bisignatus was not designated. Type material. — Holotype, female, MCZC. Horn apparently did not designate a type for T. bisignatus. Distribution. — Coastal mountain ranges of Lake, Mendocino, and Humboldt counties, California (Fig. 1 1). Discussion. — At the time of its description, Le Conte aligned T. bimaculatus with T. nutalli on the strength of prothoracic similarities. The prothorax of T. bimaculatus appears similar not only to that of T. nutalli, but also to T. oregonen- sis. Similarities in the shape of the lateral lobes of the aedeagus support the idea of a close relationship among the three species. Still further, the small range of size variation found within each of the species suggests certain life history simi¬ larities. 52 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Beyond the level at which the three species are related, T. bimaculatus exhibits an even closer relationship to T. oregonensis. Both species have very similar coloration and color patterns. They have similar limited geographic ranges and both occupy high altitude localities typified by broken woodlands with grass- covered, south-facing slopes. Despite its close relationship to T. oregonensis, T. bimaculatus exhibits very distinctive diagnostic characters. T. bimaculatus appears elongate with the elytra about two and two-thirds times longer than humeral width. By contrast, T. oregonensis appears short and chunky with the elytra no more than two and one- fourth times longer than their humeral width. Elytral maculations offer another means of easily separating the species. T. bimaculatus has large elytral macula¬ tions, the diameter of each greater than one-half the width of the elytron. The outer edge of the maculation attains the lateral margin of the elytron or is sepa¬ rated from it by a distance less than one-half its diameter. The elytral maculations of T. oregonensis are small, with each covering less than one-half the width of the elytron. The outer edge of the maculation is separated from the lateral margin of the elytron by a distance nearly equal to its own diameter. Specimens examined (27). — See Foster (1973). Trichodes oregonensis Barr (Figs. 7, 11) Trichodes oregonensis Barr, 1952:186, 1961:109. Description. — Male. Small, robust; clothed with short to long, suberect and erect, very fine, whitish hairs; head, thorax, scutellum, venter, and legs dark metallic blue, verdigris in a few specimens; antennal funicle dark brown, its under surface sometimes testaceous, club dark brown or piceous; elytra dark metallic blue or verdigris with a small, obliquely transverse, orangish maculation near middle of each elytron, umbones dark. Head densely punctate with numerous punctations confluent; front shallowly impressed above clypeus on either side; antenna with club about twice as long as broad; clypeus dark testaceous to brown, surface glabrous; labrum dark brown, surface finely, sparsely punctate, front margin feebly arcuate; galea short, no longer than one-half distance between eyes across front. Pronotum with sides gradually constricted behind middle; finely, sparsely punctate and densely roughened, moderately densely covered with suberect and erect, fine hairs. Scutellum sparsely punctate and densely roughened, disc flattened, sometimes with a medial impression extending to the arcuate hind margin. Elytra about two and one-fourth times longer than humeral width, finely, sparsely punctate and densely roughened with umbones roughened, sides ex¬ panded at posterior one-fourth, apices rounded. Mesostemum finely, sparsely punctate and densely, rugosely roughened, moderately densely clothed with long, recumbent and suberect fine hairs. Metafemur not swollen disproportionately. Abdomen with stemites one through five sparsely punctate and finely roughened; fifth sternite with hind margin deeply, arcuately emarginate; sixth sternite elon¬ gate, disc convex with basomedial surface glabrous, lateral margin feebly arcuate, hind margin arcuate; eighth tergum narrower and longer than sixth sternite, disc FOSTER— REVISION OF NORTH AMERICA TRICHODES 53 shallowly, longitudinally impressed, impression covered with dense, suberect hairs, hind margin rounded and deflexed at middle. Aedeagus feebly arcuate with lateral lobes gradually constricted and apices rounded in lateral view, subparallel in dorsal view; median lobe laterally compressed, apex ovoid (Fig. 7). Length; 6.5 to 8.7. Female. Similar to male except: abdomen with hind margin of fifth stemite truncate; sixth stemite transverse, disc feebly convex, hind margin arcuate with a small triangular notch at middle; eighth tergum with disc medially impressed to hind margin, impressed area clothed with scattered, suberect hairs, hind margin broadly, triangularly notched. Length: 7.5 to 9.6. Type locality. — Lake of the Woods, Klamath County, Oregon. Type material. — Holotype, male, no. 8337, Department of Entomology, CASC. Distribution. — Coastal mountain ranges of southern Oregon and extreme northern California (Fig. 1 1). Discussion. — T. oregonensis was not described until 1952. Specimens belong¬ ing to it were theretofore recognized as extreme variants of T. ornatus (Horn, 1891; Wolcott, 1944). T. oregonensis resembles some specimens of T. ornatus from higher elevations in the Pacific Northwest, but differs markedly in colora¬ tion from those T. ornatus populations with which it occurs sympatrically. T. ornatus specimens from within the range of T. oregonensis have yellow elytra with atropurpureus subbasal, medial, and apical bands. Specimens of T. ornatus from the Pacific Northwest that look similar to T. oregonensis can be dis¬ tinguished by their pale anterior elytral angles. The anterior angles of T. oregon¬ ensis are dark metallic blue. T. oregonensis, along with T. bimaculatus and T. nutalli, are the least variable species in our fauna. The only noticeable variation occurring in T. oregonensis is body color, which usually appears dark metallic blue, but ranges to verdigris in a few specimens. Remarkable consistency associated with larval host specificity is noted in the size range of T. oregonensis. The food supply afforded T. oregonensis by the egg pods of its grasshopper host apparently guarantees suf¬ ficient food reserves to insure attainment of consistent size. Species of Trichodes thus far associated with multiple hosts exhibit a much greater range of sizes as exemplified by T. ornatus. Separation of T. oregonensis from its sister species has been fully discussed under T. bimaculatus. Specimens examined (325). — See Foster (1973). Larval Taxonomy Description. — Mature larva. Body digitiform, widest at third or fourth abdomi¬ nal segment, tapered gradually anteriorly and posteriorly; membranous areas salmon to scarlet colored, densely covered with short to long yellowish hairs, sclerites reddish brown; ampullae not developed; spiracles annuliform. Head capsule compact, subquadrangular, about twice as broad as long in dorsal view (Fig. 14), epicranium tumescent, gena flat or tumescent, surface smooth to 54 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY faintly rugose; frons triangular, as broad as long with surface roughened behind front margin and epicranial suture incomplete; stemmata subequal in size and shape, five on either side, arranged in an anterior row of three and a posterior row of two with posterior pair positioned about equidistantly above and below level of dorsal member of anterior row; antenna composed of basal membrane and three segments, ratio between basal, middle, and apical segments 2: 1.5:1, basal segment glabrous, middle segment with apical appendix less than one-sixth as long as apical segment and three subapical hairs shorter than apical segment, apical segment bearing a medial hair about equal in length to that segment; clypeus smooth with disc flat; labrum smooth with disc bearing a medial, trans¬ verse tumescence and with front margin arcuate, truncate, or shallowly emargin- ate; mandible longer than broad in dorsal view, inner margin with prominent retinaculum, maxilla and labium well developed, maxilla from cardo base to apex longer than gula (10:7); gula slightly longer than epicranial suture, narrow¬ est at posterior one-fifth and expanded in front of middle. Thorax with large paired sclerites covering most of protergal surface, smaller meso and metanotal sclerites sometimes present; prosternum bearing a well -developed medial sclerite, about four times as long as broad. Abdomen with ninth segment posteriorly rounded, its dorsoposterior surface bearing a pair of short, subconical, slightly hooked, dorsally produced urogomphi. Discussion. — The only previous larval characterization of Trichodes (Boving and Champlain, 1920) was based solely on the larva of T. ornatus. Eight species of North American Trichodes, bibalteatus, bicinctus, nutalli, oregonensis, ornatus, peninsularis, simulator, and oresterus, were examined and serve as a basis for the description and diagnosis presented herein. Subsequent discussion pertains only to mature larvae. Trichodes larvae are distinguished by a combination of characters not shared by other genera. On the basis of general appearance and biology, Trichodes might be confused with Lecontella, the larvae of which also frequent bee and wasp nests. Both have similarly shaped body, head capsule, and urogomphi. They are separated most readily on the basis of body color and stemmatal number. Tri¬ chodes larvae are salmon to scarlet colored, and have the five stemmata present on either side of the head arranged in two rows, whereas Lecontella larvae are cream colored, and have only three stemmata on either side. Trichodes also might be confused with any one of several other genera, including Enoclerus, Thanasimus, Phlogistosternus, Thaneroclerus, or Chariessa, all of which have the same number of stemmata arranged in a similar fashion. Trichodes is the only presently known representative among these North American genera that has a chunky, salmon to scarlet-colored body, with the basal plate of the ninth tergum lacking, and the urogomphi small and hook shaped. It is further distinguished by the shape of the head capsule, which appears subrectangular in lateral view. Enoclerus, Thanasimus, and Thaneroclerus, genera with orangish or pinkish body coloration, might be mistaken for Trichodes. However, their head cap¬ sules are distinctively elongate and wedge shaped in lateral view. FOSTER— REVISION OE NORTH AMERICA TRICHODES 55 Relationship Among Species Mature larvae of eight species, including representatives of all species groups, were examined. Larval characteristics support the arrangement of species groups already presented. They particularly reaffirm the distinctiveness of the ornatus group. Members of the ornatus group have the body and head covered with rela¬ tively fewer and shorter hairs than do members of the other groups. The head capsule in the ornatus group has the genal areas slightly swollen in dorsal view whereas in other groups it appears more flattened. Larvae belonging to the bibalteatus and peninsularis groups cannot be dis¬ tinguished at the group level. Anatomical Characters The larval portion of this study is based on limited material, making it im¬ possible in most instances to assess infraspecific variation. For this reason, the characters and their states used herein should be applied in combination with one another. Identifications should be based on total anatomical similarity in col¬ laboration with biological and distributional information. Labrum. — The shape of the front labial margin may be emarginate as in T. oregonensis, transverse as in T. simulator, or feebly arcuate, a condition found in some specimens of T. ornatus. Each of these three character states applies to more than one species, but can be used satisfactorily in combination with other characters. Mandible. — Characters found on the mandibles of some species, for example, the number of hairs on the mandibles of T. oregonensis and T. nutalli and the position of the retinaculum in T. oresterus, are of primary value in that they separate the species on which they occur from all other known species. In other instances, these characters are of secondary value and their states must be used in combination with other character states. Maxilla. — The number and position of setae on the cardo base, stipes base, and palpiger are somewhat variable. They are useful nevertheless as supporting char¬ acters and are covered in the mature larval description under each species. Labium. — The number and position of setae found on the submentum, men- tum, and prementum are also variable but nevertheless are useful as supple¬ mentary diagnostic characters. The disposition of these setae is characterized for mature larvae of each species in the following descriptions. Gula. — The number of hairs on the gula and their arrangement provide useful characters for the separation of several species. Key to Mature Larvae 1. Head bearing scattered, relatively short hairs, most of which are much shorter than one- third head capsule width; genae slightly swollen (Fig. 14) . 2 Head covered with erect hairs, many of which are longer than one-third the head capsule width; genae not swollen (Fig. 14) . 4 2. Gula with two hairs behind front margin (Fig. 16); mandible with one or two hairs behind middle . 3 56 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Gula with four or five hairs (Fig. 16); mandible with three to seven hairs on disc (Fig. 15) . ornatus 3. Mandible with two hairs behind middle of outer surface (Fig. 16) . oregonensis Mandible with a single hair behind middle of outer surface (Fig. 16) . nutalli 4. Gula with three or more hairs . 5 Gula with two or fewer hairs . 7 5. Gula with all hairs in front of middle; mandible only slightly longer than broad in dorsal view . 6 Gula with one or two hairs behind middle; mandible at least one and one-half times as long as broad in dorsal view (Fig. 15) . simulator 6. Gula with four to six hairs irregularly arranged (Fig. 16); mandible with several hairs on disc (Fig. 15) . hibalteatiis Gula with four hairs arranged in a diamond shape behind front margin (Fig. 16); mand¬ ible with one large hair in front of base (Fig. 15) . hicinctus 7. Gula with a single, medial hair behind front margin (Fig. 16); mandible with retinaculum at middle of inner margin (Fig. 15) . peninsularis Gula with a pair of hairs behind front margin (Fig. 16); mandible with retinaculum be¬ hind middle of inner margin (Fig. 15) . oresterus Trichodes ornatus Say (Figs. 14, 15, 16, 17) Description. — Mature larva. Length, 11-15; head capsule width, 1.5-1. 9. Body digitiform, broadest at third or fourth abdominal segment; membranous areas salmon to bright red, densely covered with yellowish hairs of short to moderate length, sclerites reddish brown. Head capsule almost twice as broad as long in dorsal view, genal area slightly swollen (Fig. 14), epicranium and frons smooth to finely, shallowly rugose, gena smooth; stemmata subequal in size and shape with anterior row entire but nearly contiguous, posterior pair distinctly separate and positioned about equidistantly above and below level of dorsal member of anterior row; antenna comprised of basal membrane and three seg¬ ments, basal and middle segments subcylindrical, middle segment with three sub- equally spaced, subapical hairs around its circumference and with its apical surface lying at a 70-degree angle to its longitudinal axis and bearing a conical appendix less than one-sixth as long as apical segment, apical segment cylindrical and slender, apex bearing three stiff, erect, minute hairs equidistantly placed around the margin and a single hair subequal to length of segment at center; clypeus with surface smooth and disc flat; labrum with surface smooth, with disc tumescent, anterior margin transverse to slightly arcuate; mandible longer than broad in dorsal view with middle area of dorsolateral surface bearing three to seven stiff, irregularly positioned hairs, outer margin strongly arcuate, inner mar¬ gin with retinaculum nearer apex than base (Fig. 15); maxilla with bases of cardo and stipes platelike and heavily scleritized, cardo base bearing a single hair near inner angle, stipes base bearing a cluster of five or six hairs, palpiger lightly scleritized, bearing four hairs in a transverse row; labium membranous, sub- mentum bearing four or five irregularly positioned hairs, mentum glabrous, prementum bearing four or five hairs in a transverse row in front of hind margin, palpiger bearing two transversely arranged hairs; gula slightly longer than epi¬ cranial suture with sides expanded in front of middle, surface smooth, bearing FOSTER— REVISION OF NORTH AMERICA TRICHODES 57 Fig. 14. — Head capsule of mature larva, dorsal view: a, Trichodes ornatiis hartwegiant/y, b, Trichodes simulator. four or five irregularly placed hairs (Fig. 16). Prothorax with well-developed dorsal plate, its ventral surface bearing a single long, narrow median sclerite; meso and metanota immaculate or occasionally bearing small, poorly developed paired sclerites; metathoracic legs almost as long as head capsule width. Abdo¬ men with ampullae of eighth segment undeveloped; ninth segment with uro- gomphi much shorter than head capsule in dorsal view, projecting dorsally and slightly curved forward before apices in lateral view, surface glabrous, roughened around base and becoming smooth toward slightly divergent apices (Fig. 17). Discussion. — Examination of larvae belonging to all subspecies except T. o. ornatus has revealed no morphological differences upon which to base their separation. Variation manifest in a number of characters, including setal arrange¬ ment on mandibles, maxillae, labium, and gula, presence or absence of meso and metathoracic tergites and basal plate of the urogomphi, is not presently under¬ stood. Although all five larval instars are somewhat similar, they can be separated on the basis of size, aided to some extent by morphological features. Allometric growth is noted with respect to several structures, including legs, mandibles, and urogomphi, and is especially apparent when the first instar is compared to later stages. The first instar is distinguished from later stages by its slender, parallel- 58 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Fig. 15. — Mandibles of mature Trichodes larvae, dorsal view: a, T. ornatus hartwegianiis', b, T. oregonensis', c, T. nutalli\ d, T. hihalteatus; e, T. hicinctus\ f, T. simidator, g, T. peninsularis horni; h, T. oresterns. sided body, relatively long legs, and large urogomphi. Long hairs, particularly on the genae, pleural body regions, and urogomphi, further distinguish the first instar larva. Later instars share a greater degree of morphological similarity but are distinguished by head capsule measurements. No attempt has been made to compare each instar of T. ornatus with those of other species. Comparisons are drawn only between the mature larvae of avail¬ able species. Mature larvae of T. ornatus are distinguished from other known species by the combination of characters presented in the key. Separation from species with which it occurs sympatrically in various parts of its range, T. nutalli, T. oregonensis, T. simulator, and T. peninsularis, is discussed in greater detail under those species. The reader is referred to Boving and Champlain (1920) and Linsley and Mac- Swain (1943) for additional drawings of both larvae and pupae of this species. Specimens examined (68). — Various localities in Arizona, California, Idaho, Nevada, Oregon, and Utah. Description of Other Life Stages of Trichodes ornatus Egg. — Length, 1.61-1.75; maximum width, 0.46-0.50. Shape elongate and slightly curved, thickest at middle, one end slightly smaller and more pointed than the other. Surface smooth and shiny, creamy yellow, becoming orangish with age. FOSTER— REVISION OF NORTH AMERICA TRICHODES 59 / rr / 1 1 // e f 1 g Fig. 16. — Gulae of mature Trichodes larvae, ventral view: a, T. ornatiis hartwe^ianiis\ b, T. nutaUv, c, T. bihalteatus', d, T. hicinctiis; e, T. peninstdaris horni\ f, T. sinitdator, g, T. oresterus. First instar. — Length, 2.2-4. 1; head capsule width, 0.22-0.26. Body elongate, slender, sides subparallel; membranous areas cream colored, sparsely covered with stiff, long hairs, each segment bearing a very long hair at middle on either side, sclerites reddish brown. Head capsule only slightly broader than long in dorsal view (Fig. 17) with epicranium, frons, and gena smooth; stemmata ar¬ ranged in two clumps on either side with individual stemma indistinguishable; antenna as in mature larvae except appendix of second segment nearly one-half as long as apical segment and median distal hair of apical segment about equal in length to entire antenna including basal membrane; clypeus and labrum similar to those of mature larvae; mandible much longer than broad in dorsal view, outer margin strongly arcuate, inner margin with retinaculum about one-half way between apex and base; maxilla, labium, and gula similar to those of mature larvae. Prothorax with large, paired dorsal plates covering nearly its entire surface (Fig. 17); meso and metanota each with smaller paired sclerites, each sclerite bearing a distinct setal pit near its center; metathoracic legs more than twice as long as head capsule width. Abdomen with eighth segment bearing a pair of well-developed ampullae, one on either side just above the spiracles; urogomphi, including large basal plate, longer than head capsule in dorsal view (Fig. 17), projecting posteriorly and slightly curved dorsally before the apices 60 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Fig. 17. — Trichodes ornatus hartwegianiis larvae: a, urogomphi of mature larva, dorsal view; b, urogomphi of mature larva, lateral view; c, head and thorax of first instar, dorsal view; d, urogomphi of first instar, dorsal view; e, urogomphi of first instar, lateral view. in lateral view (Fig. 17), surface glabrous, roughened around base and becom¬ ing smooth toward the slightly divergent apices. Second instar. — Length, 3. 5-7. 5; head capsule width, 0.49-0.58. Body similar to mature larvae. Head capsule transverse in dorsal view (3:2), otherwise similar to mature larvae; stemmata arranged as in mature larvae but with stemma of an¬ terior row contiguous; antenna similar to mature larvae except appendix of mid¬ dle segment one-fourth as long as apical segment; mouthparts and gula similar to those of mature larvae. Thorax as in mature larvae. Abdomen similar to mature larvae except basal urogomphal plate always present, sharply defined, and cover¬ ing most of ninth tergum, urogomphi including basal plate subequal to length of head capsule in dorsal view. Third instar. — Length, 7.0-15.0; head capsule width, 0.70-0.74. Similar to mature larvae except urogomphi including basal plate nearly as long as head cap- FOSTER— REVISION OF NORTH AMERICA TRICHODES 61 sule in dorsal view, basal plate always present but varying from lightly scleritized with margins poorly defined to heavily scleritized with margins sharply defined. Fourth instar. — Length, 9.0-15.0; head capsule width, 0.94-0.98. Very similar in appearance to mature larvae. Pupa. — Salmon colored. Head projecting ventrally and nearly obscured by pronotum in dorsal view; vertex bearing scattered, stiff, short hairs; frons glabrous; clypeolabral area altus with surface coarsely, shallowly rugose and glabrous; eyes well developed; antennae projecting back along sides of pronotum above forelegs; mouthparts distinct. Thorax well developed, prothorax sub- cylindrical, surface coarsely rugose, clothed with scattered, short, stiff, cream colored hairs; meso and metanota coarsely rugose, glabrous, metanotum longer than mesonotum. Abdomen with tergites and sternites distinct, glabrous; sternites one through five with hind margins transverse; sixth sternite with hind margin rounded, surface transversely impressed at middle; sixth tergite with hind margin arcuate, slightly emarginate at center and visible in ventral view; paired caudal processes arising from an inflated basal process inserted near lateroposterior margin of sixth tergite, conical, glabrous, and apparently six segmented, segments one through five poorly delineated, sixth segment distinct and smaller in cir¬ cumference than other segments. Trichodes oregonensis Barr (Figs. 15, 16) Description. — Mature larva. Length, 11-13; head capsule width, 1.2- 1.4. Body similar to T. ornatus in shape, color, and vestiture. Head similar to T. ornatus except labrum with front margin broadly, shallowly emarginate; mandible a little longer than broad in dorsal view with dorsolateral surface bearing two stiff, long hairs behind middle, outer margin arcuate, inner margin with retinaculum nearer apex than base (Fig. 15); maxilla with stipes base bearing a single long hair near middle, and palpiger bearing three or four hairs; labium with submentum bearing a single hair near center, prementum bearing six hairs arranged in two clusters of three hairs each; gula with two long, suberect hairs near middle (Fig. 16). Discussion. — Mature larvae of T. oregonensis are most easily confused with those of T. ornatus because of overlapping size range, similar mandibular shape, and sympatric occurrence. The number of hairs on the gula provide the easiest means of distinguishing larvae of T. oregonensis from those of T. ornatus. Specimens examined (\1). — Oregon: Jackson Co., Ashland, 16 mi. NE. Trichodes nutalli (Kirby) (Figs. 15, 16) Description. — Mature larva. Length, 12-13; head capsule width, 1.3-1. 4. Body similar to T. ornatus in shape, color, and vestiture. Head similar to T. ornatus except labrum with front margin transverse or feebly emarginate; mandibles a little longer than broad in dorsal view with dorsolaterad surface bearing a single long, stiff hair behind middle, outer margin arcuate, inner margin with retinacu- 62 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY lum nearer apex than base (Fig. 15); maxillae with stipes base bearing two trans¬ versely positioned hairs; gula with two transversely positioned hairs in front of middle (Fig. 1 6). Discussion. — The close relationship between T. nutalli and T. oregonensis as indicated on the basis of adult morphology and biological similarity is substanti¬ ated by the similarity of mature larvae, as indicated by the number of mandibular setae and number and placement of setae on the maxilla, labium, and gula. T. nutalli occurs sympatrically with T. apivorus, the larvae of which remain unknown, and with T. ornatus. It can be separated from the latter species by the number of hairs on the gula and on the dorsolaterad surface of the mandible. Specimens examined O). — North Dakota: Ransom Co. Trichodes bibalteatus Green (Figs. 15, 16) Description. — Mature larva. Length, 22; head capsule width, 2.7. Body digiti- form, broadest at third abdominal segment; membranous areas salmon to bright red, densely covered with short and long yellowish hairs, sclerites reddish brown. Head capsule almost twice as broad as long in dorsal view, gena flattened; epi- cranium, frons, antennae, and stemmata similar to T. ornatus\ labrum with front margin broadly, shallowly emarginate; mandible longer than broad in dorsal view with disc of dorsolaterad surface bearing a cluster of seven stiff, short to long hairs, outer margin arcuate, inner margin with retinaculum nearer apex than base (Fig. 15); maxilla with cardo base bearing seven irregularly positioned hairs as in T. ornatus\ gula with four to six hairs irregularly arranged in a longi¬ tudinal row down the center (Fig. 16). Discussion. — T. bibalteatus can be separated from all known species except T. bicinctus and T. oresterus on the basis of distribution. It is most easily sepa¬ rated from those species by its large size and the arrangement of gular hairs. Specimens examined (3). — Texas: Brewster Co., Marathon, 23 mi. E, 2; Sutton Co., Juno, 19 mi. NE, 1. Trichodes bicinctus Green (Figs. 15, 16) Description. — Mature larva. Length, 14.7-15.2; head capsule width, 1.9. Body similar to T. bibalteatus in shape, color, and vestiture. Head similar to T. bibalteatus except labrum with front narrowly, shallowly emarginate; mandible slightly longer than broad in dorsal view with dorsolaterad surface bearing a single long, stiff hair behind middle, outer margin strongly arcuate, inner margin with retinaculum nearer apex than base (Fig. 1 5); maxilla with cardo base bearing two hairs, one positioned in front of the other at center of disc, stipe base with seven scattered hairs, palpiger bearing a transverse row of five hairs; labrum with submentum bearing a medial clump of eight irregularly positioned hairs, mentum with a transverse row of six hairs across its middle, prementum glabrous, palpiger bearing two transversely arranged hairs; gula with four hairs arranged in a diamond shape in front of middle (Fig. 16). FOSTER— REVISION OF NORTH AMERICA TRICHODES 63 Discussion. — Larvae assigned to T. hicinctus are associated on the basis of distribution, size, and anatomical distinctiveness. The most distinctive diagnostic feature of T. hicinctus is the arrangement of hairs on the gula. Specimens examined (2). — Texas: Garza Co., Justiceburg, 4 mi. E. Trichodes peninsularis horni Wolcott and Chapin (Figs. 15, 16) Description. — Mature larva. Length, 29; head capsule width, 2. 1-2.7. Body similar to T. bibalteatus in shape, color, and vestiture. Head similar to T. hihalteatus except frons rugose with a shallow, transverse depression in front of middle; labrum with front margin transverse; mandible distinctly longer than broad in dorsal view with dorsolaterad surface bearing a cluster of three to nine short to long, stiff hairs, outer margin arcuate, inner margin with retinaculum midway between apex and base and bearing a small pointed tooth midway be¬ tween retinaculum and base (Fig. 15); maxilla with cardo base bearing three transversely positioned hairs, stipes base bearing six irregularly positioned hairs and palpiger bearing a pair of transversely positioned hairs; labium with sub- mentum bearing two irregularly positioned hairs, prementum bearing a clump of four or five irregularly positioned hairs; gula with a single long hair arising from in front of middle (Fig. 16). Discussion. — T. p. horni is separated from other known species by the ap¬ pearance of the gula. It is the only known species in which the gula bears a single hair. Specimens examined O). — Arizona: Cochise Co., Portal, 1 mi. E. Trichodes simulator Horn (Figs. 15, 16) Description. — Mature larva. Length, 18-22; head capsule width, 2. 0-2. 2. Body similar to T. bibalteatus in shape, color, and vestiture. Head similar to T. bibal¬ teatus except labrum with front margin transverse; mandible more than one and one-half times longer than broad in dorsal view with basal one-half of dorso¬ laterad surface bearing a cluster of four to eight stiff, short to long hairs, outer margin with basal one-half more or less straight and anterior one-half arcuate, inner margin with large, anteriorly produced retinaculum a little in front of middle and a broad, low tooth midway between retinaculum and apex (Fig. 15); maxilla with cardo base bearing three transversely positioned hairs, stipes base bearing two clumps of three to five hairs each, palpiger bearing five to seven hairs irregularly positioned along a more or less transverse line; labium with sub- mentum bearing a pair of transversely positioned hairs, prementum with five to eight hairs positioned transversely across middle; gula bearing five irregularly positioned long hairs (Fig. 16). Discussion. — The only species occurring sympatrically with T. simulator is T. ornatus from which it can be distinguished usually on the basis of size. These species are separated positively by the shape of the mandible, which appears elongate in T. simulator and more wedge shaped in T. ornatus. T. simulator 64 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY is separated most easily from other species of similar size, T. peninsularis and T. hihalteatus, by the shape of the retinaculum and mandibular apex. Specimens examined (18). — Utah: Uintah Co., Vernal, 5; Iron Co., Cedar City, 3 mi. E, 13. Trichodes oresterus Wolcott (Figs. 15, 16) Description. — Mature larva. Length, 17; head capsule width, 1.94. Body similar to T. hihalteatus in shape, color, and vestiture. Head capsule similar to T. hihalteatus except frons with a pair of broad, shallow excavations positioned transversely at middle; labrum with front margin feebly arcuate; mandible more than one and one-half times longer than broad in dorsal view, with basal one-half of dorsolaterad surface bearing seven short to long hairs, outer margin with basal one-half more or less straight and anterior one-half arcuate, inner margin with large, obliquely rounded retinaculum nearer base than apex (Fig. 15); maxilla with cardo base bearing two transversely positioned hairs, stipes base bearing five irregularly positioned hairs, palpiger bearing a cluster of eight hairs; labium with submentum bearing four hairs, prementum with a cluster of 10 hairs near middle; gula bearing a pair of transversely positioned hairs behind front margin (Fig. 16). Discussion. — The larva treated here is associated with T. oresterus on the basis of distribution and anatomical distinctiveness. It is separated from sympatric species, T. hihalteatus and T. hicinctus, by the occurrence of only two gular hairs. Specimens examined (\). — Texas: Brewster Co., Marathon, 23 mi. E. Comparative Biology of North American Trichodes During the study, biological data on seven species of Trichodes were obtained. Of these, T. hihalteatus, T. oregonensis, T. ornatus, T. peninsularis, and T. simu¬ lator were studied in detail. Two species, T. hicinctus and T. oresterus, were observed, collected, and associated with larval hosts. Four species not studied in the field, T. apivorus, T. himaculatus, T. nexus, and T. nutalli, are discussed in light of available information and inferences drawn from related species. A com¬ plete list of flower visitation records for each species appears in Appendix 1. Composite Biology There is substantial biological diversity among the species of North American Trichodes. Nevertheless, species studied share a number of features that permit formulation of a generalized biology. T. oregonensis is excluded from the discus¬ sion, except by direct reference, because of its association with Orthoptera egg pods rather than Hymenoptera cells. What is known for each individual species is presented in a subsequent section under the biology of that species. Adults of all species are diurnal and occur on a variety of flowers (Appendix 1 ), which play a very important role in their life cycle. Flowers that serve as a site for feeding, mating, and, in some instances, oviposition are frequented during FOSTER— REVISION OF NORTH AMERICA TRICHODES 65 periods of activity and inactivity. Adults feed on pollen and a wide range of soft- bodied insects that are attracted to flowers. No instance of cannibalism was ob¬ served for any species, although Linsley and MacSwain (1943) reported post- copulatory cannibalism by females of T. ornatus. Mating occurs only on flowers. All seven species studied exhibit similar, un¬ complicated premating behavior. When a male locates a female, usually con¬ tacting her first with his antennae, he immediately attempts to mount from the rear. There is no evidence that precopulatory behavior serves as an isolating mechanism among sympatric species. Unlike mating, ovipositional behavior conforms to no generalized pattern. The ovipositional behavior of four species, T. bibalteatus, T. oregonensis, T. ornatus, and T. simulator, has been observed in the field. Comparisons among these and other species studied suggest that ovipositional behavior falls generally into three categories: T. bibalteatus and T. ornatus lay eggs on flowers frequented by hymenopteran hosts; T. simulator oviposits in or near nests of its hymen- opteran hosts; and T. oregonensis oviposits near the egg pod of its orthopteran host. The means by which Trichodes larvae reach their prey varies with the ovi¬ positional behavior. The larvae of T. bibalteatus and T. ornatus probably exhibit phoresia by attaching themselves to a bee or wasp and riding it back to the nest. Larvae of other species studied in detail seek their host after hatching. After arriving at the site of its food source, the first instar larva must enter the cell of its host while it is being provisioned. Once sealed into a cell, the Tri¬ chodes larva does not attempt to leave until it has devoured the contents. No evi¬ dence has been found to indicate that a first instar larva can gain entrance to a cell that has been sealed. Upon entering, the larva usually feeds to a limited extent on the provisions of the cell. When the host larva is nearly fully grown, the first instar Trichodes larva punctures the host’s epidermis and feeds on the ensuing exudate. By the time the host larva and the contents of its cell have been consumed, the Trichodes larva has molted twice. The third instar larva then may do one of several things: it may remain in the host cell; it may burrow into the surrounding substrate and con¬ struct a pupal cocoon; or it may burrow into adjacent cells and continue feeding. All Trichodes species with which larvae have been associated have five instars, the first three of which are feeding instars. The fourth instar apparently does not feed, but rather prepares the pupal cell and lines it with an oral secretion. The fifth instar orients itself with its head toward the exit hole. Evidence for this gen¬ eral sequence is supplied by the usual occurrence of three cast skins in the host cells frequented by the Trichodes larva and by the presence of the other two cast skins in the pupal chamber. Each species can be regarded as either spring or fall occurring. Spring species, including the ornatus and bibalteatus groups and T. oresterus, occur as adults during the spring or early summer and are univoltine. They overwinter commonly as fourth instar larvae within their already constructed pupal cell. Autumn spe¬ cies, T. simulator and T. peninsularis, appear as adults in the late summer or fall 66 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY and have a two year life cycle. The first winter is spent as a small larva, which then undergoes major development the following spring. The second winter and following summer is spent as a fourth instar; pupation does not occur until late in the second summer. In all species, the fourth instar larva serves a similar function irrespective of whether a one or two year life cycle is involved. In both instances, it is a non¬ feeding diapause stage of development that transcends the winter preceding pupation. Laboratory observations indicate that exposure to cold is neces¬ sary to break diapause in spring species. Individuals held at room temperature during the winter spend one or more additional years as nonfeeding larvae and sometimes pass through an additional instar before pupating. Similar observations on longevity under laboratory conditions have been reported (Essig, 1934; Lins- ley and MacSwain, 1943, 1946). Within the framework of the generalized biology just presented exist features unique to individual species or species groups. These are best exemplified by summarizing the biology of each species. Peninsularis Group Biological similarities among T. simulator, T. peninsularis, and T. oresterus support the concept of the peninsularis group as recognized on the basis of external morphology. Sex ratios of all three species, correlated with observations on T. simulator and T. peninsularis, suggest that males spend most of their time on flowers, whereas females leave for long periods while searching for ovi- positional sites. Trichodes simulator Horn This species occurs as scattered local populations throughout the Colorado Plateau region of western Colorado, eastern Utah, northwestern New Mexico, and northeastern Arizona, and in extreme southwestern North Dakota. A requisite for the occurrence of the species is the presence of vertical streamside clay banks containing nests of bee species that serve as larval hosts. Stephen et al. (1969) suggested that T. simulator is restricted to anthophorine bees. During this study larvae have been taken from cells of Anthophora occidentalis and also from the cells of an unidentified megachilid bee, a secondary nester in old anthophorine burrows. An examination of T. simulator adult behavior reveals the nature of the restricted host range. During the day, adults can be found on a number of autumnal flowers in the immediate vicinity of the larval habitat. A sur¬ vey of such plants over a 24-hour period shows a differential behavior. Males remain on the blossom throughout the day, whereas females spend the night on flowers but are absent from them during most of the day, returning only oc¬ casionally for short periods of time. Table 2 shows the male-female ratio on flowers at the Cedar City study site during a 24-hour period. During the time females are away from the blossoms, they can be observed flying along the stream bed and crawling over the surface of vertical banks along the stream in search of ovipositional sites. The female inspects burrows by enter- FOSTER— REVISION OF NORTH AMERICA TRICHODES 67 Table 2. — Nunther of adult male and female Trichodes on flowers at hourly intervals. Time Species Sex 0800 0900 1000 1 100 1200 1300 1400 1500 1600 1 700-0800 Trichodes d 7 7 9 8 10 4 12 9 8 8 hihalteatus 9 5 5 6 10 5 7 8 7 9 8 Trichodes 6 56 48 41 47 53 49 52 44 47 37 ornatus 9 44 52 59 43 47 51 58 56 53 63 Trichodes d 48 50 47 55 46 51 39 54 58 49 oregonensis 9 52 50 53 45 54 49 61 46 42 51 Trichodes d 33 34 36 41 30 27 37 29 28 37 peninsularis 9 39 30 6 4 5 4 4 12 27 30 Trichodes d 14 14 16 12 12 18 11 15 14 16 simulator 9 12 10 3 2 0 2 0 4 10 15 ing head first, sensing when burrows are being provisioned. During this in¬ spection, only the head and prothorax are in the burrow. The female responds to an acceptable burrow by turning around and backing into it, where she remains for several minutes while ovipositing. Excavation of burrows visited by females has not revealed where eggs are deposited. However, two possibilities present themselves: females may oviposit at some point along the burrow, or they may oviposit directly in open cells. In the case of the former, newly hatched larvae would have to seek open cells, because inspection under the microscope of in¬ fested cells reveals that they are sealed. There is no evidence that T. simulator larvae enter the cell after it has been sealed. Adults are present from late July through mid-September. Presumably, winter is passed either as an egg or small larva. Cells excavated 4 June 1970 revealed that larvae had reached the second instar and were feeding within the closed cell. Cells taken 26 June 1970 contained third instar larvae that had devoured or nearly devoured the contents of each cell. After consuming the cell contents, the larva may enter an adjacent cell to feed, construct an earthen pupal chamber in the soil adjacent to the burrow, or pupate within the host cell. Examination of burrows in early June revealed that some larvae were feeding in the bee cells, whereas others were in their pupal chambers. These observations led to the fol¬ lowing conclusions about the life cycle. Eggs are laid in late summer and au¬ tumn, and the first winter is passed as either an egg or first instar larva within the closed cell of the host bee. Throughout the first spring and summer, the larva feeds within the closed cell of its host, during which time it passes through three instars and enters the fourth. The fourth instar larva constructs a pupal chamber in which it passes the second winter, spring, and summer before pupating. Trichodes peninsularis horni Wolcott and Chapin Observations were made in Cochise County, Arizona, where adults of the sub¬ species occur from late July through September. Specimens have been collected from a number of flowering plants (Appendix 1). The most frequently visited 68 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY flower in the study area was Baileya miiltiradiata Harv. and Gray. Adults of both sexes frequent flowers; their frequencies, relative to one another (Table 1), are similar to those presented for T. simulator, suggesting that females oviposit away from flowers. Although oviposition has not been observed, females have been observed crawling over the surface of fence posts as if in search of ovipositional sites. Another circumstance suggesting that females may oviposit in such situa¬ tions is that all larvae collected during this and a previous study (Krombein, 1967) were taken from the cells of four species of bees (Appendix 2) in block traps tied to fence posts. Numerous stick traps placed in the ground in the study area were uninfested. During the Krombein study, traps were placed in the field during March and were retrieved in May. Because adults of T. p. horni are present only during the autumn, it seems plausible that oviposition occurs at sites where bees are prone to nest, that eggs may overwinter, and that larvae seek their hosts’ cells the follow¬ ing spring. In this case, larvae apparently would have to enter open cells while they are being provisioned. Neither larva reared by Krombein completed its life cycle in a single year, sug¬ gesting a condition similar to T. simulator in which the life cycle normally spans two years. Trichodes oresterus Wolcott Adults are present from late April through July and have been observed in the field on two occasions. A single larva associated with this species was col¬ lected from a cell of Megachile brevis Say in a stick trap 23 miles east of Mara¬ thon, Brewster County, Texas. From these limited observations, it is apparent that ovipositional behavior differs in one respect from that of the other species in the peninsularis group. Its larva was taken from a stick trap placed in the ground, whereas neither of the other species use this type of ovipositional site. Nevertheless, the ovipositional behavior of T. oresterus is similar in one respect to that already presented for other members of the peninsularis group, as reflected in male-female sex ratios. Whereas populations encountered in the field have been of such low numbers that sex ratios could not be obtained, one can gain some insight into sex ratios by looking at museum collections, if the assumption is made that most collections are from flowers. Sixty-nine per cent of all T. oresterus specimens studied were males. Similarly high percentages occur in col¬ lections of the two aforementioned species but not in collections of T. ornatus, which is known to oviposit on flowers, the implication being that T. oresterus may seek the nesting site of its host much as T. simulator and T. p. horni do. Bibalteatus Group Members of the bibalteatus group, as far as is known, oviposit on flowers and prey upon hymenopteran hosts. T. bibalteatus oviposition has been observed and larval hosts of T. bibalteatus and T. bicinctus are known. Nothing specific is known of the biology of T. apivorus. FOSTER— REVISION OF NORTH AMERICA TRICHODES 69 Trichodes bibalteatus Le Conte Studies were conducted in Brewster, Garza, Kimble, Sutton, Terrell, and Val Verde counties, Texas, during May and June of 1971, 1972, and 1974. Adults frequent a variety of flowers (Appendix 1). Throughout the season, males are more numerous than females, as suggested by the sex ratio of preserved speci¬ mens. Sixty-six per cent of all specimens examined were males. The observed sex ratio might be interpreted as indicative of an ovipositional behavior similar to that found among members of the peninsularis group. How¬ ever, oviposition on flowers of Ratibida columnaris (Sims) D. Don. was observed on 16 and 23 May 1974 at the Texas Tech University Center, Junction, Kimble County. A 24-hour census (Table 1) and observations of females marked with florescent paint revealed that females do not leave the flowers for long periods. Differential occurrence cannot be explained by ovipositional behavior. During 1970, stick traps were set out in May and were retrieved in late August. Three larvae were taken from cells of two megachilid bees (Appendix 2). Two larvae were preserved; the third pupated and emerged during the last week of May 1971 after overwintering as a fourth instar within its pupal chamber. Trichodes bicinctus Green This species is restricted to the Rolling Plains, Trans-Pecos, and Edward’s Plateau regions of Texas and western Oklahoma. It occurs sympatrically with T. bibalteatus and T. oresterus. T. bicinctus exists in scattered, small populations and at low densities. It is far less common than T. bibalteatus in areas where local populations coexist. Adults have been recorded from several species of flowering plants (Appendix 1), including R. columnaris, which is also the preferred host of T. bibalteatus in areas of sympatry. Relative frequency of males and females on flowers could not be determined because of low population densities. However, the ratio between males and fe¬ males was nearly equal in preserved material, with males comprising 5 1 per cent of the specimens studied. Two fourth instar larvae associated with T. bicinctus were found in burrows of Microanthophora sp. four miles east of Justiceburg, Garza County, Texas, on 22 March 1974. Both were enclosed in pupal chambers constructed within host cells. Trichodes apivorus Germar This species occurs along the Atlantic and Gulf coasts where adults are pres¬ ent from March through May in the southern part of the range and from June to August in the more northerly portion. Adults frequent a variety of flowers as in¬ dicated by specimen labels (Appendix 1). No collections or field observations were made, and no larvae were available for study. A letter was sent to api- culturists working within the range of the species inquiring about its possible association with honeybees. No positive responses were received, suggesting that the name “apivorus” is not indicative of the larval host. 70 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY References to T. apivorus as a predator in nests of bees and wasps (Boving and Champlain, 1920; Balduf, 1935) were inferred apparently from the specific name or from what was known of Trichodes biology in general. Fifty-one per cent of all specimens examined during the study were males. Ornatus Group Unifying features among biologies of this species group are apparently few in view of the diversity exemplified by T. ornatus and T. oregonensis. Despite ap¬ parent ovipositional differences, both species exhibit approximately equal male- female frequency on flowers. Evidence presented herein suggests that biologies of T. bimaculatus and T. nutalli are similar to that of T. oregonensis. Trichodes oregonensis Barr Two features of this species, its restricted distribution and lack of variation, make it especially interesting. It is limited to habitats above 5000 feet in the coastal mountain ranges of southwestern Oregon and extreme northwestern California. Adults show relatively little size variation, especially when compared to T. ornatus, with which it occurs sympatrically. Both distribution and size are related to the specificity between T. oregonensis and its larval host, a slant-faced grasshopper, Chloealtis aspasma Rhen and Hebard. The known range and habitat occurrence of T. oregonensis fit almost exactly those of its host as presented by Gurney et al. (1964). Limited size variation among adults of T. oregonensis is a response to the constant quantity of food available to the developing larva. During its development, a larva normally consumes the contents of only a single grass¬ hopper egg pod. Because most pods contain eight eggs, the food resource avail¬ able to all larvae is essentially the same. Adults are present from early June to August and can be found on a number of flowering plants (Appendix 1). T. oregonensis depends on flowers, much as other species do, to provide a source of food, a resting place, and a means of bringing potential mates together. Nothing is known of the precise manner in which females oviposit, although they evidently oviposit at or near the egg pods of the host. C. aspasma eggs are found only in exposed, pithy stems of dead elderberry, Sambucus spp. Conse¬ quently, stick traps similar to those used for other species can be used as artificial nesting sites for the grasshopper and ultimately yield larval specimens of T. oregonensis. Means by which the larva enters the egg pod were not observed, although scars along side the egg pod plugs suggested that larvae burrow via that route. Larvae feed on eggs during the late summer and autumn, at which time the entire con¬ tents of a single egg pod is consumed. By the time the larva has consumed the con¬ tents of the egg pod, it has passed through three instars. The fourth instar larva constructs a pupal chamber directly below the egg pod gallery, pushing the ex¬ cavated wood fiber into the gallery as it works. The anterior end of the gallery then is sealed with the same type of brown leathery material used by other species FOSTER— REVISION OF NORTH AMERICA TRICHODES 71 to line their entire pupal chamber. The fourth instar overwinters in the pupal chamber, molts to a fifth instar in late May or June, and pupates soon after. All larvae thus far reared have reached maturity in a single year, suggesting that a single year life cycle is normal for the species. Some information concerning the relative abundance of T. oregonensis has been gathered during the study. Because C. aspasma oviposits in Sambucus, and because an increase in the availability of ovipositional sites associated with the invasion of Sambucus occurs after extensive logging, subsequent population in¬ creases of both insects could be expected. Such seems to be the case in areas where T. oregonensis was studied. A survey of northern Jackson County, Oregon, in an area that had not been logged recently produced only an occasional speci¬ men, whereas adults were numerous in the study area, a recent clearcut. A similar situation exists with reference to the type locality. At the time the type series was collected in 1951, the species was quite common (Jacques R. Heifer, personal communication). Today the area is covered by regrowth timber, and Sambucus plants are no longer present in the area. During the past four years, no specimens of T. oregonensis have been found at the type locality. It is of interest to note that only one North American clerid, Aulicus terrestris Tinsley, has been reported as a predator of grasshopper eggs (Tinsley, 1936). Rees (1973) failed to record any such relationship. Trichodes bimaculatus Te Conte This species, which occurs in the coastal mountain ranges of both northern and southern California, was not observed during the study. Although no biological information is available, some interesting inferences can be drawn based on dis¬ tributional information and what is known of the biology of its sister species, T. oregonensis. In their study of Chloealtis, Gurney et al. (1964) gave distributional infor¬ mation for three closely related West Coast species assigned to the genus. Their distributional data showed that the localities recorded for T. bimaculatus fall within the main portion of the range of the two species, C. dianae Gurney et al. in northern California and C. gracilis McNeill in southern California. It is probable that T. bimaculatus, which exhibits a restricted distribution and relatively slight size variation similar to that discussed for T. oregonensis, may be associated with Chloealtis and possibly with both C. dianae and C. gracilis in different parts of its range. A response to logging similar to that discussed under T. oregonensis may occur for T. bimaculatus. If so, it might explain the relatively large number of collections made during the late nineteenth and early twentieth centuries at a time when the coastal mountain ranges of California were logged more extensively than they are today. Trichodes nutalli (Kirby) This very widely distributed eastern species was not studied in the field. Pub¬ lished information concerning its biology is limited to a few flower visitation records and general statements associating its larvae with nests of bees and wasps 72 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY (Balduf, 1935; Boving and Champlain, 1920; Knull, 1951). No larvae taken from bee nests were available for study, and no authenticated records associating this species with bees or wasps could be found, a surprising revelation in view of the common occurrence of the species and the number of biological studies that have been conducted on various Hymenoptera within its range. There is one published report of larvae of an unidentified beetle destroying eggs of Chloealtis conspersa Harris (Brusven, 1967). Although Brusven’s work concentrated on Kansas grasshoppers, it should be noted that the observation cited here was made in Ransom County, North Dakota (Merlyn A. Brusven, per¬ sonal communication). These proved to be Trichodes larvae and are associated with T. nutalli on the basis of distribution. The range of size variation among adult specimens throughout the range of T. nutalli is similar to that found in T. oregonensis. Because the food resource available to the larva has a direct effect on the ultimate size of the adult, there is reason to suspect that T. nutalli larvae are specific predators of Chloealtis egg pods in a manner similar to that described for T. oregonensis. A list of plant visitation records drawn from specimen labels and the litera¬ ture is presented in Appendix 1 . Trichodes nexus Wolcott This rarely collected Baja California species was not studied and has not received biological treatment by previous authors. Nothing is known of its biology except that adults have been collected only during October. Trichodes ornatus Say Many particulars of the biology of this species were published by Linsley and MacSwain (1943). The present study has provided a reaffirmation of much of their work and has added additional information. All five subspecies of T. ornatus have been observed and collected, but detailed biological information was ob¬ tained for only three. Observations were made on T. o. douglasianus 16 miles northeast of Ashland, Jackson County, Oregon, on T. o. bonnevillensis 1 1 miles south of Fairfield, Camas County, Idaho, and on T. o. hartwegianus on Central Grade, Nez Perce County, Idaho. No evidence has been found to suggest con¬ sistent biological differences among subspecies. The following discussion is a compilation of observations made on these subspecies. Adults of both sexes frequent a wide variety of flowering plants (Appendix 1) where they occur in about equal numbers (Table 1). In the study areas, they favor Achillea millefolium L., Eriogonum spp., and Ceanothus spp. Linsley and MacSwain (1943) observed that eggs are laid singly and de¬ posited within the floral heads of A. millefolium. During the present study, oviposition was observed on Eriogonum elatum (Dougl.) ex Benth. as well as on A. millefolium. With both plants, eggs are inserted deep within the floral process away from the staminate portion of the flower and are placed on the flowers not yet mature enough to display their pollen. It seems most probable that larvae FOSTER— REVISION OF NORTH AMERICA TRICHODES 73 hatch when flowers are attractive to bees and reach the cells of their hosts by phoretic means. During the study, larvae have been taken from the cells of a number of bees. These records, along with those recorded in the literature or made available by other workers, are presented in Appendix 2. First instar larvae usually occur singly within host cells. In only one instance has multiple infestation of a host cell been found. A cell of Anthocopa cope- landica (Ckll.) containing three first instar larvae was found in a stick trap from the Jackson County, Oregon, study site opened 1 1 July 1970. Linsley and Mac- Swain (1943) reported that first instar larvae do not feed until the host grub is nearly mature and then attack it. In several instances, larvae in cells of A. cope- landica. Megachile geniula Cresson, and Osmia sp. were observed to feed on the pollen ball before attacking the host grub. T. ornatus larvae preying upon cavity nesting hosts may consume the contents of several bee cells during their development. Larvae usually pass the first two in¬ stars in the cell into which they are introduced. As third instars, they often bur¬ row into adjacent cells. A single larva may feed on cells both above and below the one where its initial development took place. The number of cells destroyed by a single larva during the course of its development varies principally with size of the host, number of cells in the nest, and the type of nest structure. When nests, of a medium sized bee, M. gemula, are considered, each T. ornatus larva destroys one to five cells, with a mean of two. Cells of a smaller bee. Megachile pacifica Panzer, with similar nest structure are preyed upon more extensively; one to eight cells, with a mean of four, are destroyed. Variation evident within the amount of host material consumed in the above instances suggests that larvae often consume more resource than is necessary for their development. Fourth instar larvae appear by late summer or autumn. They exhibit a great deal of flexibility in pupal chamber construction. Larvae developing in block traps made of hardwood constructed their pupal chamber within the last host cell occupied by pushing frass and remaining cell contents into the opening through which the larva entered the cell. The cell is then sealed with the brown leathery material described by Linsley and MacSwain (1943). Some larvae occupying nests in elderberry stick traps constructed pupal chambers in the above manner. However, most burrowed into the pith adjacent to the gallery and constructed an oblong pupal chamber. The tunnel between the gallery and pupal chamber was plugged with excavated material, and the entire pupal chamber was lined with leathery material. Orientation of the pupal chamber to the main burrow is vari¬ able. In some cases it lies more or less perpendicular to the axis of the burrow, in which case the entrance tunnel is used as an escape route by the adult. In other cases when the pupal chamber lies more or less parallel to the burrow, the larvae excavate a secondary tunnel at the anterior end of this chamber, tunneling al¬ most through to the main burrow but leaving a thin partition of pith mixed with oral secretion that can be broken easily by the emerging adult. The second¬ ary tunnel may contain sawdust excavated from the pupal chamber. 74 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Winter is passed usually as a fourth instar larva within the pupal chamber, followed by molt into a fifth instar during the early spring. The fifth instar orients itself within the pupal chamber. After about two weeks, it pupates. Pupation is completed after 10 to 12 days, when adult emergence completes the annual life cycle. Larvae that attain the fourth instar before winter complete their life cycle in a single year. Those overwintering as small larvae spend at least one additional year before maturation. Laboratory rearings were conducted to ascertain the effects of various com¬ binations of temperature and realtive humidity during the fourth and fifth instars and the pupal stage on the elytral coloration of the ensuing adult. The results of this study (Table 2) suggest that elytral coloration is not directly influenced by either temperature or relative humidity. Laboratory and field observations suggest that exposure of fourth instar larvae to cold temperatures triggers maturation. Larvae maintained at room temperature remain in the fourth in¬ star two years or longer before finally pupating. Larvae reared to the fourth in¬ star at room temperature and then allowed to overwinter out of doors pupated and emerged as adults the following spring. Acknowledgments Special thanks are due those persons whose assistance was instrumental in the completion of the study. William F. Barr displayed continuing interest in the project and offered encouragement throughout, always making his library avail¬ able and examining type specimens not accessible to me. He read, criticized, and edited early drafts of the manuscript. Critical reviews were provided also by Lawrence E. O’Keeffe, Steward C. Schell, and Janice M. Gillespie. Determinations of bees were made by Frank D. Parker and Richard W. Rust. Grasshopper determinations were made by Ashland B. Gurney. John L. Strother provided plant determinations. Financial support for this research was provided by a National Defense Educa¬ tion Act Fellowship, a grant from The Society of the Sigma Xi, and by the Uni¬ versity of Idaho. Literature Cited Anderson, E. M. 1914. Insects collected in the Okanogan Valley, 1913. Proc. Entomol. Soc. British Columbia, 4;57. Arnett, R. H. 1960. The beetles of the United States. The American Entomological Institute, Ann Arbor, xii+ 11 12 pp. Arnett, R. H., and G. A. Samuelson. 1969. Directory of Coleoptera collections of North America (Canada through Panama). Purdue Univ., Lafayette, Indiana, vii+123 pp. Balduf, W. V. 1935. The bionomics of entomophagus Coleoptera. John S. Swift Co. Inc., New York, 220 pp. Barr, W. F. 1950. Contributions toward the insect fauna of Lower California. No. 12. Coleoptera: Cleridae. Proc. California Acad. Sci., (4)24(12):485-5 19. - . 1952. A new species of Trichodes from southern Oregon (Coleoptera:Cleridae). Proc. Entomol. Soc. Washington, 54:186-188. FOSTER— REVISION OF NORTH AMERICA TRICHODES 75 - . 1961. Family Cleridae. Pp. 105-112, in The beetles of the Pacific Northwest: part III: Pselaphidae and Diversicornia (M. H. Hatch, ed.), Univ. Washington Press, Seattle, ix + 503 pp. - . 1962. A key to the genera and classification of the North American Cleridae (Coleoptera). Coleopt. Bull., 16:121-127. - . 1969. The Buprestidae and Cleridae of the Nevada test site (Coleoptera). Great Basin Natur., 29:1 1-19. Blackwelder, E. 1948. The geological background, in The Great Basin: with emphasis on glacial and postglacial times. Bull. Univ. Utah, 38(20):3-15. Boving, a. G., and a. B. Champlain. 1920. Larvae of the North American beetles of the family Cleridae. Proc. U.S. Nat. Mus., 52:576-649. Brodie, W. 1888. List of Coleoptera collected by Mr. Bruce Bailey, in Kicking Horse Pass, Rocky Mountains, C. P. R., 1884. Proc. Canadian Inst. Toronto, 3:212-220. Brusven, M. a. 1967. Differentiation, ecology and distribution of immature slant-faced grasshoppers (Acrididae) in Kansas. Kansas Agr. Exp. Sta. Tech. Bull., 149:1-59. Bush, A. H. 1914. A trip up Mount Chem. Proc. Entomol. Soc. British Columbia, 4:58-60. Carr, F. S. 1920. An annotated list of the Coleoptera of northern Alberta. Alberta Nat. Hist. Soc., pp. 1-6. Champenois, a. 1900. Synopsis des especes palearctiques du genre Clerus Muller ( Trichodes Herbst). L’Abeille, 30: 1-46. Chagnon, G. 1935. Contribution a I’etude des Coleopteres de la Province de Quebec. Naturelle Canada, 62:167-176. Chapin, E. A. 1917. Miscellaneous notes on Coleoptera. Bull. Brooklyn Entomol. Soc., 12:29-31. Chittenden, F. H. 1890. Remarks on the habits of some species of Cleridae. Entomol. Amer., 6: 1 54-155. Clemens, W. A. 1916. The pine bark beetle. Bull. Cornell Univ. Agr. Exp. Sta., 28:1-155. Cockerell, T. D. A. 1893. The entomology of the mid-Alpine zone of Custer County, Colorado. Trans. Amer. Entomol. Soc., 20:305-370. - . 1898. Life-zones in New Mexico II. The zonal distribution of Coleoptera. Bull. New Mexico Agr. Exp. Sta., 28:137-179. CORPORAAL, J. B. 1948. Further notes on systematics and synonomy. Entomol. Berlin, 12:286-288. - . 1950. Coleopterorum catalogus, pars 23 (editio secunda) Cleridae. Uitgeverij Dr. Junk, The Hague, Netherlands, 373 pp. Crotch, G. R. 1870. The genera of Coleoptera studied chronologically (1735-1801). Trans. Entomol. Soc. London, 1:41-52. Curtis, J. 1824. British entomology; being illustrations and descriptions of the genera of insects found in Great Britain and Ireland: containing coloured figures from nature of the most rare and beautiful species, and in many instances of the plants upon which they are found. Part 1. London, 320 pp. Davidson, A. 1896. On the nesting habits of Anthidium consimile. Entomol. News, 7:22-26. Davis, A. C. 1932. A list of the Coleoptera of Fort Tejon, California. Bull. California Acad. Sci., 3 1 :75-87. Desmarest, E. 1860. In Chenu, Encyclopedic d'histoire naturelle de Coleoptera. Paris, 2:226-279, pi. 19. Dillon, E. S., and L. S. Dillon. 1961. A manual of common beetles of eastern North America. Row, Peterson and Co., Evanston, Illinois, viiiT 884 pp. Dobzhansky, T. 1970. Genetics of the evolutionary process. Columbia Univ. Press, New York, ix 4- 505 pp. 76 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Ekis, G., and a. P. Gupta. 1971. Digestive system of Cleridae (Coleoptera). Intern. J. Insect Morph, and Embryol., l(l):51-85. Emmons, E. 1854. A natural history of New York agriculture, V. Insects. Albany, New York, 156 pp. Escherich, K. 1893. Zun Kenntniss der Coleopterengattung Tr/c/;o<7e5 Herbst. Verhand. K. K. Zool. Botan. Ges., 43:1-57. Essie, E. O. 1934. Trichocles ornatus Say. J. Econ. Entomol., 27:724. Fabricius, j. C. 1775. Systema entomologiae, sistens Insectorum classes, ordines, genera, species, adjuctis synonymis, locis, descriptionibus, observationibus. Flensburgi et Lipsiae, 832 pp. - . 1801. Systema elevtheratorum, Kiliae. Lxxiv 4- 506 pp. Forbes, T. M. 1922. The wing-venation of the Coleoptera. Ann. Entomol. Soc. Amer., 15:328-352. Foster, D. E. 1973. The taxonomy and biology of the genus Trichodes Herbst of North America (Coleoptera:Cleridae). Unpublished Ph.D. dissertation, Univ. Idaho, 340 pp. Gemminger, M., and E. von Harold. 1869. Catalogus coleopterorum hucusque descrip- torum synonymicus et systematicus. 6: 1 609-1800. Germar, E. F. 1824. Insectorum species novae aut minus cognitae, descriptionibus illustratae. Halas, xxiv -h 624 pp. Gibson, A. 1917. The entomological record, 1916. 47th Ann. Rep. Entomol. Soc. Ontario 1916, pp. 149-155. Goetzmann, W. H. 1959. Army exploration in the American West. Yale Univ. Press, New Haven, xx-l- 509 pp. Green, J. W. 1917. A new Tr/c/iot/e^ (Cleridae: Coleoptera). Entomol. News, 28:367. Gurney, A. B., H. F. Strohecker, and J. R. Helfer. 1964. A synopsis of the North American grasshopper of the genus-group Chrysochraontes (Othoptera:Acrididae). Trans. Amer. Entomol. Soc., 89:1 19-137. Hamilton, J. 1895. Catalogue of the Coleoptera of southwestern Pennsylvania, with notes and descriptions. Trans. Amer. Entomol. Soc., 22:317-381. Hennig, W. 1966. Phylogenetic systematics. Univ. Illinois Press, Urbana, 263 pp. Herbst, J. F. W. 1792. Natursystem aller bekannten in-und auslandischen Insekter als eine Fortsetzung der von Biiffonschen Naturgeschichte. Kafer, 4:1-197. Hope, F. W. 1840. The coleopterist’s manual, part the third, containing various fami¬ lies, genera, and species of beetles, recorded by Linneus and Fabricius. Also descriptions of newly discovered and unpublished insects, part 3. London, 189 pp. Horn, G. H. 1876. Synopsis of the species of Cymatodera and Trichodes of the United States. Trans. Amer. Entomol. Soc., 5:220-232. - . 1880. Contribution to the coleopterology of the U.S., no. 3. Trans. Amer. Entomol. Soc., 5:139-154. - . 1891. Trichodes ornatus Say . Entomol. News, 2:6-8. - . 1894. Coleoptera of Baja California. Proc. California Acad. Sci., 4(2):302- 449. Horning, D. S., and W. F. Barr. 1970. Insects of Craters of the Moon National Monu¬ ment, Idaho. Univ. Idaho, Misc. Ser., 8:1-118. Kirby, W. 1818. A description of several new species of insects collected in New Hol¬ land by Robert Brown. Trans. Linn. Soc. London, 12:375-453. Kirk, V. M. 1969. A list of beetles of South Carolina. Part 1 — North Coastal Plains. South Carolina Agr. Exp. Sta. Tech. Bull., 1033:1-124. Klug, j. C. F. 1842. Versuch einer systematischen Bestimmung und Auseinandersetzung der Gattungen und Arten der Clerii, einer Insekten-Familie Aus der Ordnung der Coleopteren. Abhandl. Konig. Akad., pp. 259-397. Knowlton, G. F. 1930. Notes on Utah Coleoptera. Florida Entomol., 14:35-36. FOSTER— REVISION OF NORTH AMERICA TRICHODES 77 Knull, J. N. 1951. The checkered beetles of Ohio (Coleoptera:Cleridae). Ohio Biol. Surv. Bull. 42, 8:269-350. Krombein, K. V. 1967. Trap-nesting wasps and bees: life histories, nests, and associates. Smithsonian Press, Washington, D.C., vi + 570 pp. Lacordaire, J. T. 1857. Genera des Coleopteres du expose methodique et critique de tous les genres proposes jusquici dans cet ordre d’insectes. Paris, 4:1-579. Latreille, P. a. 1804. Histoire naturelle, generale et particulare des crustaces et des insects. Paris, 9: 1-445. Le Conte, J. L. 1849. Synopsis of the Cleridae of the United States. Ann. Lyceum Nat. Hist., New York, 5:9-35. - . 1858. Catalogue of the Coleoptera of the regions adjacent to the boundary line between the United States and Mexico. J. Acad. Nat. Sci. Philadelphia, 4:9-420. - , ED. 1859. American entomology. A description of the insects of North America by Thomas Say. A. E. Foote, New York, 2:xxi + 412 pp. - . 1863. List of Coleoptera of North America, part 1. Smithsonian Misc. Coll., 6(140):l-78. - . 1867. Descriptions of new Coleoptera chiefly from the Pacific slope of North America. Trans. Amer. Entomol. Soc., 5:43-72. - . 1869. List of Coleoptera collected in Vancouver’s Island by Henry and Joseph Matthews, with descriptions of some new species. Ann. Mag. Nat. Hist., ser. 4, 4:369-385. - . 1878. The Coleoptera of the alpine regions of the Rocky Mountains, part 2. U.S. Geol. Geogr. Surv., 5:447-480. Leng, C. W. 1915. Cocinella transversogiittata, Trichodes niitalli and Malachius aeneiis. J. New York Entomol. Soc., 23:254. - . 1920. Catalogue of the Coleoptera of America, north of Mexico. John D. Sherman Jr., Mount Vernon, New York, x + 470 pp. Linsley, E. G. 1936. Studies in the genus Aidicus Spinola (Coleoptera:Cleridae). Univ. California Publ. Entomol., 6:249-262. - . 1942. Notes on the habits of some beetles from the vicinity of Yosemite National Park. S. California Acad. Sci., 41: 164-166. Linsley, E. G., and J. W. MacSwain. 1943. Observations on the life history of Trichodes ornatus (Coleoptera: Cleridae), a larval predator in the nests of bees and wasps. Ann. Amer. Entomol. Soc., 34:589-601. — - . 1946. Longevity of Trichodes and Pelonium larvae. Pan-Pacific Entomol., 22:18. Linsley, E. G., and R. L. Usinger. 1934. Insect collecting in California. Pan-Pacific Entomol., 10:101-105. Lugger, O. 1899. Beetles injurious to fruit producing plants. Univ. Minnesota Agr. Exp. Sta. Bull., 66:1-250. Mayr, E. 1963. Animal species and evolution. The Belknap Press of Harvard Univ. Press, Cambridge, xiv + 797 pp. Moore, I. 1937. A list of the beetles of San Diego County, California. Occas. Papers San Diego Soc. Nat. Hist., 2:1-109. Nye, W. P., and G. E. Bohart. 1952. A larva of Trichodes ornatus from a pollen trap on a hive of honey bees. Pan-Pacific Entomol., 27:6. Packard, A. S. 1877. A list of Coleoptera collected in 1875, in Colorado and Utah. 9th Ann. Rep. U.S. Geol. Surv., pp. 811-815. Papp, C. S. 1960. The Cleridae of North America, part 1. Bull. S. California Acad. Sci., 59:76-88. Parker, F. D., and R. M. Bohart. 1968. Host-parasite associations in some twig-nesting Hymenoptera from western North America, part 2. Pan-Pacific Entomol., 44:1-6. Peterson, A. 1959. Entomological techniques. Edward Brothers, Ann Arbor, Michigan, v-l- 435 pp. 78 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Rees, N. E. 1973. Arthropod and nematode parasites, parasitoids and predators of Acrididae in America north of Mexico. U.S. Dept. Agr. Tech. Bull., 1460:1-288. Say, T. 1823. Descriptions of coleopterous insects collected in the late expedition to the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under the command of Major Long. J. Acad. Nat. Sci. Philadelphia, 3:1 39-2 16. - . 1825. Descriptions of new species of coleopterous insects inhabiting the United States. J. Acad. Nat. Sci. Philadelphia, 5:160-204. - . 1835. Descriptions of new North American coleopterous insects, and observa¬ tions on some already described. Boston J. Nat. Hist., l(2):151-203. Schaeffer, C. 1908. On new and known Coleoptera of the families Cocinellidae and Cleridae. J. New York Entomol. Soc., 16:125-135. Sc HENKLiNCi, S. 1903. Genera insectorum, Coleoptera, family Cleridae. 13:1-124. - . 1906. Die Cleriden des Deutschen entomologischen nationalmuseums, nebst Beschreibungen neuer Arten. Deutsche Entomol. Zeitschr., pp. 241-320. Sharp, D., and F. A. G. Muir. 1912. The comparative anatomy of the male genital tube in Coleoptera. Trans. Entomol. Soc. London, part 3. Entomol. Soc. Amer., Col¬ lege Park, Maryland, vi + 642 pp. Snow, F. H. 1907. List of species collected in New Mexico. Trans. Kansas Acad. Sci., 20:41-65. Spinola, M. 1844. Essai monographique sur les Clerites, Insectes Coleopteres. Geneva, 1:1-386. Stage Smith, G. 1930. Coleoptera of Cooper Mt., British Columbia, part 2. Mus. and Art Notes, 5:22-25. Stephen, W. P., G. E. Bohart, and P. F. Torchio. 1969. The biology and external morphology of bees. Oregon Agr. Exp. Sta., 147 pp. Sturm, J. 1826. Catalog meiner Insecten-Sammlung, Ester Theil, Kafer. Niirnberg, 207 pp. - . 1843. Catalog der Kaefer-Sammlung von Jacob Sturm. Niirnberg, 386 pp. Tanner, V. M. 1928. The Coleoptera of Zion National Park, Utah. Ann. Entomol. Soc. Amer., 21:269-281. - . 1934. The Coleoptera of Zion National Park, no. II, Utah. Ann. Entomol. Soc. Amer., 27:43-49. Thompson, W. R., and F. J. Simmonds. 1965. A catalogue of the parasites and predators of insect pests. Section 4, host predator catalogue. Commonwealth Agr. Bull., 198 pp. Townsend, C. H. T. 1895. On the Coleoptera of New Mexico and Arizona, including biologic and other notes. Canadian Entomol., 27:39-51. Ulke, H. 1875. Report upon the collections of Coleoptera. Rep. U.S. Geol. Surv. (Wheeler), 5:809-827. - . 1902. A list of the beetles of the District of Columbia. Proc. U.S. Nat. Mus., 25:1-57. Van Dyke, E. C. 1943. New species of West American Coleoptera. Pan-Pacific Entomol., 19:41-42. Wade, J. S. 1935. A contribution to a bibliography of the described immature stages of North American Coleoptera. USDA, ARS, Washington, D.C., 1 14 pp. Waters, N. D. 1971. Insect enemies of the alfalfa leafcutter bee and their control. Idaho Current Info. Ser., 163:1-4. White, A. 1949. Nomenclature of coleopterous insects in the collection of the British Museum, Part IV, Cleridae. British Mus. (Nat. Hist.), London, 68 pp. Wickham, H. F. 1892. Description of four- insect monstrosities. Canadian Entomol., 24:141-142. - . 1895. The Coleoptera of Canada. XIII. The Cleridae of Ontario and Quebec. Canadian Entomol., 27:247-253. FOSTER— REVISION OF NORTH AMERICA TRICHODES 79 - — . 1896. A list of some Coleoptera from northern New Mexico and Arizona. Bull. Lab. Nat. Hist. Univ. Iowa, 3:153-171. - . 1910. List of Van Duzee collection of Florida beetles. Bull. Buffalo Soc. Nat. Sci., 9:399-405. Wickham, H. F., and A. B. Wolcott. 1912. Notes on Cleridae from North and Central America. Bull. Lab. Nat. Hist. Univ. Iowa, 6:49-67. Wolcott, A. B. 1909. The Cleridae of the public museum of the city of Milwaukee. Bull. Wisconsin Nat. Hist. Soc., 7:93-102. - . 1910a. Notes on some Cleridae of Middle and North America with descrip¬ tions of new species. Field Mus. Nat. Hist. Publ. Zool. Ser., 7:339-401. - . 19106. Family Cleridae. Pp. 46, 852, in Coleoptera or beetles known to occur in Indiana (W. S. Blatchley, ed.), Indiana Dept. Geol. and Nat. Res. Bull., 1:1- 1386. - . 1944. The American species of Trichodes (Coleoptera: Cleridae). Pan-Pacific Entomol., 20:54-60. - . 1947. Catalogue of North American beetles of the family Cleridae. Fieldiana: Zool., 32:61-105. Wolcott, A. B., and E. A. Chapin. 1918. Notes on Cleridae. Bull. Brooklyn Entomol. Soc., 13:107-108. Zimmerman, S. 1971. Ergebnisse zoologischer Sammelrisen in der Tiirkel Gattung Trichodes Herbst (Cleridae: Coleoptera). Ann. Nat. Mus. Wien, 75:591-625. 80 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Appendix 1. — Flower visitation records for North American Trichodes. Trichodes apivoriis apivorus Castanea sp. Enpatoriiim sp. Ruhiis sp. Trichodes apivorus borealis Ceanothas sp. Ruhiis sp. Seriocarpus sp. Trichodes hihalteatas Acacia preggii Gray Acacia sp. Coreopsis tinctoria Nutt. Gossypiani sp. Heiianthas sp. Opuntia sp. Pithecolohiiou sp. Prosopis glandulosa Torn. Ratihida col a at nar is (Sims) D. Don. Ratihida sp. Rad heck ia sp. Verhesina encelioides (Ca\.) Gray Trichodes hicinctas Ratihida colaninaris (Sims) D. Don. Trichodes natalli Achillea niillefoliaia L. Apocy/uini sp. Asclepias sp. Aster sp. Chrysantheautai leacantheiuain L. Heiianthas sp. Heracleam inaxiaiani Bartr. MeiHotas albas Lam. Melilotas sp. PotentiUa recta L. Rosa sp. Rahas sp. Radheckia hirta L. Spiraea sp. T r ichod es or ego nensis Achillea inillefoliani L. Achillea sp. Brodiaea congesta Sm. Brodiaea hyacinthina {Lindi.) Baker Erigeron deca/nhens Nutt. Eriogonani elation Dougl. Eriogonain sphaerocephahon Dougl. Eriogonani anihellatton Torr. Eriogonani sp. Eriophyllani lanatani (Prush) Forbes Gnaphaliani heneolens Davids Madia elegans D. Don. PotentiUa draniniondii Lehm. PotentiUa glandulosa Lindl. Sanihacas caendea Raf. Trichodes ores ter as Asclepias sp. Haplopappas heterophyllas (Gray) Blake Haplopappas sp. Larrea tridentata (DC.) Cov. Prosopis sp. Taniarix pentandra Pall. Trichodes ornatas honneviUensis Achillea lanalosa Nutt. Achillea sp. Asclepias sp. Balsaniorhiza sagittata (Pursh) Nutt. Calochortas sp. Cardans sp. Chaniaehatiaria inillefoliani (Torr.) Maxin. Chrysothaninas naaseosas (Pall.) Britt Chrysothaninas sp. Cirsiani andalatani (Nutt.) Spreng. Eriogonani ovalifoliani Nutt. Eriogonani anihellatani Torr. Eriogonani sp. Fallagia paradox (D. Don.) Endl. Geraniani sp. Heiianthas sp. Iris sp. Juniper as sp. Malva sp. Medicago sp. Oenothera sp. Penstenion cyanthas Hook. Penstenion sp. Phacelia sp. PotentiUa spp. Rosa woodsii Lindl. Rahas niacropetaias Dougl. Sarcohatas verniicalatas (Wook.) Torr. Sol id a go sp. Sphaeralcea sp. Taraxacani officinale Weber Tetradyniia canescens DC. Tetrad}’ niia sp. Trichodes ornatas doaglasianas Achillea inillefoliani L. Achillea sp. Adenostonia sp. Arctostaphylos sp. Argenione sp. Asclepias sp. FOSTER— REVISION OF NORTH AMERICA TRICHODES 81 Baccharis sp. Brassica sp. Brodiaea sp. CalochortKs liiteus Dougl. Calochortiis splendens Dougl. Calochortus venustits Dougl. Calochortiis sp. Ceanothiis ciineatiis (Hook.) Nutt. Ceanothus parvifoliiis (Wats.) Trel. Ceanothiis sp. Cirsiiini sp. Clarkia williamsonii (Dur. and Hilg.) Lewis and Lewis Clarkia sp. Cryptantha nuiricata (H. and A.) Nels. and Macbr. Cryptantha sp. Datura sp. Eriodictyon sp. Erioponiiin coinpositiini Dougl. Eriogoniini latifoliiim Sm. Eriogoniini niidiini (Dougl.) S. Stokes Eriogoniini iinihellatiini Torn. Eriogoniini sp. Eriophyll inn confertiflorinn (DC.) Gray Eschscholzia californica Cham. Eschscholzia sp. Eiicnide iirens (Gray) Parry Cilia capitata Sims Cilia sp. Lapinas sp. Mentha palegiani L. Oenothera tanacetifolia T. and G. Pad as sp. Penstemon sp. Phacelia sp. Qaercas sp. Ranancalas sp. Rhaninas californica Esch. Rhaninas crocea Nutt. Rhaninas sp. Salvia colanihariae Benth. Sidalcea sp. Sonchas sp. Sphaeralcea sp. Stanleya pinnata (Pursh) Britton Syniphoricarpos sp. Trifoliiun sp. Verhascain thapsas L. Whipplea inodesta Torr. Wyethia sp. Trichodes ornatas hartwegianas Achillea millefoliani L. Achillea sp. Artemisia sp. Balsaniorhiza sp. Calochortiis elegans Pursh Calochortiis inacrocarpas Dougl. Ceanothiis sp. Chrysothainnas sp. Crepis sp. Daacas carota L. Eriogoniini sp. Lapinas sp. Medicago sp. Phacelia sp. Popalas tacainahacca Mill. Potent ilia sp. Rhus glabra L. Rosa sp. Sol id ago sp. Spiraea sp. Trifoliiun pratense L. Vida sp. Wyethia sp. Trichodes ornatas ornatas Achillea lanalosa Nutt. Artemisia sp. Asclepias capricorna Woodson Aster ahatas Blake Astragalus sp. Brodiaea spp. Cleome latea Hook. Dichelostemma sp. Erigeron sp. Eriogonam compositam Dougl. Eriogoniini e latum Dougl. Eriogonam latifoliiim (T. and G.) Nels. Eriogonam sphaerocephalam Dougl. Eriogonam amhellatam Torr. Eriophyllam lanatam (Prush) Forbes Erysiniam capitatam (Dougl.) Greene Cilia capitata Dougl. Cnaphaliam sp. Madia sp. Opantia engelmannii Salm-Dyck. Pe rider id ia sp. Potent ilia drammondii Lehm. Potentilla glandalosa Lindl. Potentilla sp. Pseadocyniopteras sp. Rhaninas sp. Samhacas caeralea Raf. Senecio sp. Sphaeralcea amhigaa Gray Sphaeralcea laxa Woot. and Standi. Stanleya elata Jones Tamarix sp. 82 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Thelespeniui suhnuiiuni Gray Tor'll is sp. Yiiccd sp. Trichocles onuittis leniiosus Acacia sp. Anisconui acaitlis Torr. and Gray Arpeiiione intenneiHa Sweet Artemisia sp. Asclepias asperala (Decne.) Woodson Asclepias erosa Torr. Asclepias sp. Atriplex canescens (Pursh) Nutt. Baileya nuihiradiata Harv. and Gray Bailey a sp. Cassia covesii Gray Cercidium florid am Benth. Cholla sp. Cleome jonesii (Macbride) Tidestrom Coreopsis higelovii (Gray) Hall Coreopsis sp. Croton californiciis Muell. Dalea schottii Torr. Echinocactiis acanthodes (Lemaire) Britt, and Rose Encelia farinosa Gray Encelia sp. Eriogonum fascicidatum Benth. Eriogonum inflatiini Torr. and Frem. Eriogonam sp. Eacnide areas Parry Geraea canescensTovr. and Gray Haplopapptis cooperi (Gray) Hall Hemizonia sp. Hyptis emoryi Torr. Larrea tridentata (Sesse and Moc.) Larrea sp. Lapinas sp. Machaeranthera canescens (Pursh) Gray Machaeranthera tortifolia (Torr. and Gray) Cronq. and Keck Malacothrix sp. Mentzelia involacrata S. Wats. Mohavea confertiflora (Benth.) Nanui demission Gray Opantia hasilaris Englem. and Bigelow Opantia sp. Palafoxia sp. Phacelia crenalata Torr. Plachea sp. Prosopis sp. Salix sp. Sarcohat as sp. Senecio sp. Sphaeralcea amhigaa Gray Spliaeralcea orcattii Rose Sphaeralcea sp. Stanley a elata Jones Tamarix sp. Vigil iera del to idea Gray Trichodes peninsalaris hasalis Boerhaavia wriglitii Gray Pectis papposa Harv. and Gray Prosopis jidiflora (Sw.) DC. Trichodes peninsalaris horni Asclepias sahverticiUata (Gray) Vail Baileya sp. Boerhaavia wriglitii Gray Eriogonam ahertianam Torr. Eriogonam sp. Gaillardia palchella Foug. Gatierrezia sarothrae (Pursh) Britt, and Rushy Gatierrezia sp. Haplopapptis sp. Helianthas annaas L. Hymenothrix wislizenii Gray Kallstroemia grand iflora Torr. Lepidiam tharheri Woot. Solidago altissima L. Verhesina encelioides (Cav.) Benth. and Hook. Trichodes simulator Achillea sp. Baileya sp. Chrysothamnas naaseostis (Pall.) Britt. Chrysothamnas viscidifloras (Hook.) Nutt. Cleome sp. Gatierrezia sarothrae (Pursh) Britt, and Rushy Senecio spartiodesTorr. and Gray Solidago sp. FOSTER— REVISION OE NORTH AMERICA TRICHODES 83 Appendix 2. — Hosts for larvae of North American Trichodes. Records other than those obtained directly from this study are indicated by source. Trichodes bibalteatus Megachilidae Megachile niontivaga Cresson 23 mi. E Marathon, Brewster Co., Texas Megachile policaris Say 23 mi. E Marathon, Brewster Co., Texas 19 mi. NE Juno, Sutton Co., Texas Apidae Microanthophora sp. Trichodes bicinctus 4 mi. E Justiceburg, Garza Co., Texas Trichodes nutalli Acrididae Chloealtis conspersa (Harris) Ransom Co., North Dakota M. A. Brusven, personal communi¬ cation Trichodes oregonensis Acrididae Chloealtis aspasma Rhen and Hebard Megachilidae Anthocopa copelandica (Cockerell) Hoplitis albifrons argentifrons (Cresson) Hoplitis fulgida (Cresson) Hoplitis hypoorita (Cockerell) Megachile pacifica Panzer Osmia sp. Proteriades bidlifacies Michener 16 mi. NE Ashland, Jackson Co., Oregon Elba-Basin Pass, Cassia Co., Idaho 12 mi. S Fairfield, Camas Co., Idaho Elba-Basin Pass, Cassia Co., Idaho 1 2 mi. S Fairfield, Camas Co., Idaho 12 mi. S Fairfield, Camas Co., Idaho 12 mi. S Fairfield, Camas Co., Idaho Parker and Bohart, 1968 Trichodes ornatus bonnevillensis Trichodes ornatus douglasianus Apidae Ceratina acantha submaritima Cockerell Kyburz, El Dorado Co., California Linsley and MacSwain, 1943 84 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Megachilidae Anthocopa copelandica (Cockerell) Ashmeadiella sp. Callanthidiiun illustre (Cresson) Dianthidium macswaini Timberlake Dianihidium pudicum provancheri Titus Dianthidium sp. Hoplitis alhifrons maura (Cresson) Hoplitis productus (Cresson) Hoplitis samhuci Titus Hoplitis sp. Megachile brevis onohrychidis Cockerell Megachile gemula Cresson Megachile pacifica Panzer Osmia sp. Vespidae Euodynerus formaminatus scutellaris (Saussure) Odynerus blandinus Cresson 16 mi. NE Ashland, Jackson Co., Oregon Mt. Diablo, Contra Costa Co., California Linsley and MacSwain, 1943 Mt. Diablo, Contra Costa Co., California Linsley and MacSwain, 1943 Mt. Diablo, Contra Costa Co., California Linsley and MacSwain, 1943 Antioch, Contra Costa Co., California Coarsegold, Madera Co., California Linsley, 1942 Strawberry, El Dorado Co., California Essig, 1934 16 mi. NE Ashland, Jackson Co., Oregon Mineral King, Tulare Co., California Linsley and MacSwain, 1943 Mineral King, Tulare Co., California Linsley and MacSwain, 1943 Banning, Riverside Co., California 18 mi. W Blythe, Riverside Co., California White Water, Riverside Co., California F. D. Parker, personal communication Davis, Yolo Co., California Linsley and MacSwain, 1943 16 mi. NE Ashland, Jackson Co., Oregon 16 mi. NE Ashland, Jackson Co., Oregon 16 mi. NE Ashland, Jackson Co., Oregon Banning, Riverside Co., California F. D. Parker, personal communication Mt. Diablo, Contra Costa Co., California Linsley and MacSwain, 1943 FOSTER— REVISION OF NORTH AMERICA TRICHODES 85 Pseudomasaris coquillet ti Rohwer Mt. Diablo, Contra Costa Co., California Linsley and MacSwain, 1943 Trichodes ornatus hartwegianus Megachilidae Hoplitis hypoorita (Cockerell) Hop I it is sambuci Titus Megachile pacifica Panzer Osmia spp. Central Grade, Nez Perce Co., Idaho Central Grade, Nez Perce Co., Idaho Touchet, Walla Walla Co., Washington Carl Johanson, personal communication 2 mi. E Samuels, Bonner Co., Idaho Central Grade, Nez Perce Co., Idaho Megachilidae Ashmeadiella sp. Trichodes ornatus ornatus 5 mi. N Las Cruces, Dona Ana Co., New Mexico F. D. Parker, personal communication Trichodes ornatus tenuosus Megachilidae Anthocopa sp. Ashmeadiella higeloviae (Cockerell) Ashmeadiella breviceps Michener Ashmeadiella sp. Diathidium consimile (Ashmead) Hoplitis biscutellae (Cockerell) F. D. Parker, personal communication 1 8 mi. W Blythe, Riverside Co., California Parker and Bohart, 1968 18 mi. W Blythe, Riverside Co., California Parker and Bohart, 1968 18 mi. W Blythe, Riverside Co., California Sand Canyon, 3 mi. W Brown, Kern Co., California F. D. Parker, personal communication Palm Springs, Riverside Co., California Davidson, 1896 Sand Canyon, 3 mi. W Brown, Kern Co., California 18 mi. W Blythe, Riverside Co., California Palo Verde, Imperial Co., California F. D. Parker, personal communication 86 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Osmia dare see ns Cockerell Osmia kincaidii Cockerell Osmia spp. Proteriades sp. Sphecidae Leptochilus washo Solierella sp. 20 mi. E Indio, Riverside Co., California Linsley and MaeSwain, 1943 20 mi. E Indio, Riverside Co., California Linsley and MaeSwain, 1943 F. D. Parker, personal communication Sand Canyon, 3 mi. W Brown, Kern Co., California Borrego Valley, San Diego Co., California White Water, Riverside Co., California F. D. Parker, personal communication Westgard Pass, Inyo Co., California F. D. Parker, personal communication Bonigo Valley, California F. D. Parker, personal communication Megachilidae Megachile brevis Say Trichodes oresterus 23 mi. E Marathon, Brewster Co., Texas Trichodes peninsularis horni Megachilidae Ashmeadiella meliloti meliloti (Cockerell) 1 mi. E Portal, Cochise Co., Arizona Krombein, 1967 Ashmeadiella occipitalis Michener 1 mi. E Portal, Cochise Co., Arizona Krombein, 1967 Ashmeadiella sp. 1 mi. E Portal, Cochise Co., Arizona Trichodes simulator Apidae Anthophora occidentalis Cresson 3 mi. E Cedar City, Iron Co., Utah Microant hop hora sp. Vernal, Uintah Co., Utah R. B. Selander, collector Megachilidae Megachile sp. 3 mi. E Cedar City, Iron Co., Utah > E r ■ I Copies of the following numbers of Special Publications of The Museum may be obtained on an exchange basis from, or purchased through, the Exchange Librarian, Texas Tech University, Lubbock, Texas 79409. No. 1 Watkins, L. C., J. K. Jones, Jr., and H. H. Genoways. 1972. Bats of Jalisco, Mexi¬ co, 44 pp., 3 figs . $1.00 No. 2 Krishtalka, L. 1973. Late Paleocene mammals from the Cypress Hills, Alberta, 77 pp., 21 figs . $2.00 No. 3 West, R. M. 1973. Review of the North American Eocene and Oligocene Apatemy- idae (Mammalia: Insectivora), 42 pp., 20 figs . $1.00 No. 4 Gardner, A. L. 1973. The systematics of the genus Didelphis (Marsupialia: Didel- phidae) in North and Middle America, 81 pp., 14 figs . $2.00 No. 5 Genoways, H. H. 1973. Systematics and evolutionary relationships of spiny pocket mice, genus Liomys, 368 pp., 66 figs . $7.00 No. 6 Northington, D. K. 1974. Systematic studies of the genus Pyrrhopappus (Com- positae, Cichorieae), 38 pp., 14 figs . $1.00 No. 7 King, M. E., and 1. R. Traylor, Jr., eds. 1974. Art and environment in native America, 169 pp . $5.00 No. 8 Pence, D. B. 1975. Keys, species and host list, and bibliography for nasal mites of North American birds (Acarina: Rhinonyssinae, Turbinoptinae, Speleognathi- nae, and Cytoditidae), 148 pp., 728 figs . $4.00 No. 9 Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. 184 pp., 62 figs . $5.00 No. 10 Baker, R. J., J. K. Jones, Jr., and D. C. Carter, eds. 1976. Biology of bats of the New World family Phyllostomatidae. Part 1., 218 pp . $6.00 No. 11 Foster, D. E. 1976. Revision of North American Trichodes (Herbst) (Coleoptera: Cleridae), 86 pp., 17 figs . $2.00