UNIVERSITY OF ILLINOIS LIBRARY AT URBANA CHAMPAIGN BIOLOGY J) 0.5 &1* HKLDIANA Zoology BIOLOGY LIBRARY BURRILL HW.L NEW SERIES, NO. 28 FEB 0 7 1986 Revision of the Genus Phyllotreta Chevrolat of America North of Mexico Part I. The Maculate Species (Coleoptera: Chrysomelidae, Alticinae) Eric H. Smith F 16 UNIVERSITY OF ILLINOIS URBANA-CHAMPAIQN December 31, 1985 Publication 1364 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Information for Contributors to Fieldiana General: Fieldiana is primarily a journal for Field Museum staff members and research associates, although manuscripts from nonaffiliated authors may be considered as space permits. The Journal carries a page charge of $65 per printed page or fraction thereof. Contributions from staff, research associates, and invited authors will be con- sidered for publication regardless of ability to pay page charges, but the full charge is mandatory for nonaffiliated authors of unsolicited manuscripts. Payment of at least 50% of page charges qualifies a paper for expedited process- ing, which reduces the publication time. Manuscripts should be submitted to Dr. Timothy Plowman, Scientific Editor, Fieldiana, Field Museum of Natural History, Chicago, Illinois 60605-2496, USA. Three complete copies of the text (including title page and abstract) and of the illustrations should be submitted (one original copy plus two review copies which may be machine copies). No manuscripts will be considered for publication or submitted to reviewers before all materials are complete and in the hands of the Scientific Editor. Text: Manuscripts must be typewritten double-spaced on standard- weight, 8'/2- by 11-inch paper with wide margins on all four sides. For papers longer than 100 manuscript pages, authors are requested to submit a "Table of Contents .'" a "List of Illustrations," and a "List of Tables." In most cases, the text should be preceded by an "Abstract" and should conclude with "Acknowledgments" (if any) and "Literature Cited." All measurements should be in the metric system. The format and style of headings should follow those of recent issues of Fieldiana. For more detailed style informa- tion, see The Chicago Manual of Style (13th ed.), published by The University of Chicago Press, and also recent issues of Fieldiana. In "Literature Cited," authors are encouraged to give journal and book titles in full. Where abbreviations are desirable (e.g. . in citation of synonymies), authors consistently should follow Botanico-Periodicum-Huntianum and TL-2 Taxonomic Literature by F. A . Stafleu & R. S . Cowan (1976 et seq.) (botanical papers) or Serial Sources for the Biosis Data Base (1983) published by the BioSciences Information Service. References should be typed in the following form: Croat, T. B. 1978. Flora of Barro Colorado Island. Stanford University Press. Stanford, Calif. , 943 pp. Grubb, P. J.. J. R. Lloyd, andT. D. Pennington. 1963. A comparison of montane and lowland rain forest in Ecuador. I. The forest structure, physiognomy, and floristics. Journal of Ecology, 51: 567-601. Langdon, E. J. M. 1979. Yage among the Siona: Cultural patterns in visions, pp. 63-80. In Browman. D. L., and R. A. Schwarz, eds.. Spirits, Shamans, and Stars. Mouton Publishers, The Hague, Netherlands. Murra, J. 1946. The historic tribes of Ecuador, pp. 785-821. In Steward, J. H., ed.. Handbook of South American Indians. Vol. 2, The Andean Civilizations. Bulletin 143, Bureau of American Ethnology, Smithsonian Institution. Washington. DC. Stolze, R. G. 1981. Ferns and fern allies of Guatemala. Part II. Polypodiaceae. Fieldiana: Botany, n.s., 6: 1-522. Illustrations: Illustrations are referred to in the text as "figures" (not as "plates"). Figures must be accompanied by some indication of scale, normally a reference bar. Statements in figure captions alone, such as " x 0.8." are not acceptable. Captions should be typed double-spaced and consecutively. See recent issues of Fieldiana for details of style. Figures as submitted should, whenever practicable, be 8'/2 by 11 inches (22 x 28 cm) and may not exceed 11'/: by l6'/2 inches (30 x 42 cm). Illustrations should be mounted on boards in the arrangement you wish to obtain in the printed work. This original set should be suitable for transmission to the printer as follows: Pen and ink drawings may be originals (preferred) or photostats; shaded drawings should be originals, but within the size limitation; and photo- stats should be high-quality, glossy, black and white prints. All illustrations should be marked on the reverse with author's name, figure number(s), and "top." Original illustrations will be returned to the author upon publication unless otherwise specified . Authors who wish to publish figures that require costly special paper or color reproduction must make prior arrangements with the Scientific Editor. Page Proofs: Fieldiana employs a two-step correction system. Each author will normally receive a copy of the edited manuscript on which deletions, additions, and changes can be made and queries answered. Only one set ol page proofs will be sent. All desired corrections of type must be made on the single set of page proofs. Changes in page proofs (as opposed to corrections) are very expensive. Author-generated changes in page proofs can only be made if the author agrees in advance to pay for them. ■ttifi FIELDIANA Zoology NEW SERIES, NO. 28 Revision of the Genus Phyllotreta Chevrolat of America North of Mexico Part I. The Maculate Species (Coleoptera: Chrysomelidae, Alticinae) Eric H. Smith Division of Insects Department of Zoology Field Museum of Natural History Present Address: Orkin National Service Department 2170 Piedmont Road NE Atlanta, Georgia 30324 Accepted for publication October 10, 1983 December 31, 1985 Publication 1364 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1985 Field Museum of Natural History Library of Congress Catalog Card Number: 84-63122 ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Contents List of Illustrations Abstract 1 1-3. Acknowledgments 1 Introduction 2 Methods 2 4-6. Classification Subfamily Alticinae 4 7. Genus Phyllotreta 4 8 . Key to the Maculate Species of Phyllotreta 1 9. Species Descriptions 10. Phyllotreta arcuata Smith, n.sp 13 11. Phyllotreta armoraciae (Koch) 14 Phyllotreta attenuata Smith, n.sp 15 12. Phyllotreta bipustulata (Fabricius) 17 13. Phyllotreta bisinuata Smith, n.sp 18 14-16. Phyllotreta conjuncta Gentner 19 Phyllotreta constricta Smith, n.sp 21 17. Phyllotreta decipiens Horn 23 18. Phyllotreta denticornis Horn 24 19. Phyllotreta dolichophalla Smith, n.sp. . 25 20-22. Phyllotreta emarginata Smith, n.sp. ... 26 Phyllotreta lepidula (LeConte) 28 23. Phyllotreta liebecki Schaeffer 29 24. Phyllotreta oblonga Chittenden 30 25. Phyllotreta oregonensis (Crotch) 31 26. Phyllotreta ramosa (Crotch) 33 27. Phyllotreta ramosoides Smith, n.sp. . . . 35 Phyllotreta robusta (LeConte) 36 28 . Phyllotreta spatulata Smith, n.sp 37 29. Phyllotreta striolata (Fabricius) 38 30. Phyllotreta undulata (Kutschera) 41 Phyllotreta utana Chittenden 42 31. Phyllotreta utanula Smith, n.sp 44 32. Phyllotreta zimmermanni (Crotch) 46 33. Species incertae sedis 34. Phyllotreta alberta Chittenden 48 Phyllotreta obtusa Chittenden 48 35. Phyllotreta perspicua Chittenden 48 36. Literature Cited 48 37-42. Illustrations 51 Tables 141 43. Appendix 44. List of Abbreviations 145 45. Specimens Examined 145 Indices 46. Beetle Names 166 47. Plant Names 167 Male and female 5th abdominal ster- num and label of lectotype of /Viy/- lotreta armoraciae 51 Aedeagus and spermatheca, with structures labeled 52 Phyllotreta arcuata 53 Phyllotreta armoraciae 54 Phyllotreta attenuata 55 Phyllotreta bipustulata 56 Phyllotreta bipustulata , color pat- tern variation 57 Phyllotreta bisinuata 58 Phyllotreta conjuncta 59 Phyllotreta conjuncta, color pat- tern variations 60 Phyllotreta constricta 61 Phyllotreta decipiens 62 Phyllotreta denticornis 63 Phyllotreta denticornis , color pat- tern variations 64 Phyllotreta dolichophalla 65 Phyllotreta emarginata 66 Phyllotreta lepidula 67 Phyllotreta liebecki 68 Phyllotreta liebecki, color pattern variation 69 Phyllotreta oblonga 70 Phyllotreta oregonensis 71 Phyllotreta oregonensis , color pat- tern variation 72 Phyllotreta ramosa 73 Phyllotreta ramosoides 74 Phyllotreta robusta 75 Phyllotreta robusta, color pattern variation 76 Phyllotreta spatulata 77 Phyllotreta striolata 78 Phyllotreta striolata, color pattern variations 79 Phyllotreta undulata 80 Phyllotreta utana 81 Phyllotreta utana, color pattern variation 82 Phyllotreta utanula 83 Phyllotreta zimmermanni 84 in 48. Phyllotreta zimmermanni color pat- 176-179 tern variation 85 49-64. Antennae of Phyllotreta arcuata , P. armoraciae , P. attenuata , P. bipus- 1 80- 1 83 tulata, P. bisinuata, P. conjuncta, P. constricta, and P. decipiens 86 65-82. Antennae of Phyllotreta denti- 184-187 cornis, P. dolichophalla, P. emar- ginata, P. lepidula, P. liebecki , P. oblonga, P. oregonensis, and P. 188-191 ramosa 87 83-99. Antennae of Phyllotreta ramo- soides , P . robusta , P . spatulata , P. 192 striolata, P. undulata, P. utana, P. utanula, and P. zimmermanni ... 88 193 Aedeagus of Phyllotreta arcuata 89 Aedeagus of Phyllotreta armora- ciae 90 194 Aedeagus of Phyllotreta attenuata 9 1 Aedeagus of Phyllotreta bipus- 195 tulata 92 Aedeagus of Phyllotreta bisinuata 93 196 Aedeagus of Phyllotreta conjuncta 94 Aedeagus of Phyllotreta constricta 95 197 Aedeagus of Phyllotreta decipiens 96 Aedeagus of Phyllotreta denti- 198 cornis 97 Aedeagus of Phyllotreta dolicho- 199 phalla 98 200 Aedeagus of Phyllotreta emargi- nata 99 201 Aedeagus of Phyllotreta lepidula 100 202 Aedeagus of Phyllotreta liebecki 101 Aedeagus of Phyllotreta oblonga 102 203 Aedeagus of Phyllotreta oregon- ensis 103 204 Aedeagus of Phyllotreta ramosa. . 104 Aedeagus of Phyllotreta ramo- 205 soides 105 Aedeagus of Phyllotreta robusta. . 106 206 Aedeagus of Phyllotreta spatulata 107 207 Aedeagus of Phyllotreta striolata 108 Aedeagus of Phyllotreta undulata 109 208 Aedeagus of Phyllotreta utana ... 110 209 Aedeagus of Phyllotreta utanula 111 210 Aedeagus of Phyllotreta zimmer- 211-213 manni 112 168-171. Spermathecae of Phyllotreta arcu- 214-216 ata, P. armoraciae , P. attenuata, and P. bipustulata 113 217 1 72- 1 75. Spermathecae of Phyllotreta bisinu- ata, P. conjuncta, P. constricta, and P. decipiens 114 100- -102 103- -105 106- -107 108- -110 111- -113 14-115A 116- -118 119- -121 122- -124 125- -126 127- -129 130- -132 133- -135 136- -138 139- -141 142- -144 145- -146 147- -149 150- -152 153- -155 156- 158 159- -161 162- 164 165- -167 Spermathecae of Phyllotreta denti- cornis, P. dolichophalla, P. emar- ginata, and P. lepidula 115 Spermathecae of Phyllotreta lie- becki, P. oblonga, P. oregonensis, and P. ramosa 116 Spermathecae of Phyllotreta ramo- soides, P. robusta, P. spatulata, and P. striolata 117 Spermathecae of Phyllotreta un- dulata, P. utana, P. utanula, and P. zimmermanni 118 Distribution of Phyllotreta arcuata and P. armoraciae 119 Distribution of Phyllotreta attenu- ata, P. bipustulata, and P. bisinu- ata 120 Distribution of Phyllotreta con- juncta 121 Distribution of Phyllotreta con- stricta 122 Distribution of Phyllotreta deci- piens 123 Distribution of Phyllotreta denti- cornis and P. dolichophalla 124 Distribution of Phyllotreta emar- ginata and P. lepidula 125 Distribution of Phyllotreta liebecki 126 Distribution of Phyllotreta oblonga and P. oregonensis 127 Distribution of Phyllotreta ramosa 128 Distribution of Phyllotreta ramo- soides 1 29 Distribution of Phyllotreta robusta and P. spatulata 130 Distribution of Phyllotreta strio- lata 131 Distribution of Phyllotreta undu- lata and P. utana 132 Distribution of Phyllotreta utanula 133 Distribution of Phyllotreta zimmer- manni 134 Phyllotreta alberta 135 Phyllotreta obtusa 136 Phyllotreta perspicua 137 Antennae of Phyllotreta alberta, P. obtusa, and P. perspicua 138 Spermathecae of Phyllotreta al- berta, P. obtusa, andf. perspicua 139 Histograms representing color pat- tern variation of 14 samples of Phyllotreta striolata 140 List of Tables 1 . Measurement ranges: head ratios, pronotum, elytra, and entire beetle 141 2. Antennal segment measurements: length/ width; total length of antenna 142 3. Comparative morphological data for native maculate nearctic species of Phyllotreta with respect to 5th antennal segment and aedeagal dorsal washboard 144 Revision of the Genus Phyllotreta Chevrolat of America North of Mexico Part I. The Maculate Species (Coleoptera: Chrysomelidae, Alticinae) Abstract Members of the genus Phyllotreta are small flea beetles (1.8-3.6 mm long), most of which inhabit open areas of disturbed or cultivated vegetation, but a few of which inhabit woodlands. They feed almost exclusively on the plant family Cruciferae, and several species are of economic importance as pests of cultivated crucifers. This is a revision of the maculate species of Phyl- lotreta Chev. known to occur in America north of Mexico. The aedeagus and spermatheca, the male and female antennae, and the elytral color pattern are il- lustrated for each of the 27 species. A key to the species is presented. The biology for each species has been summarized and the distribution is given. Nine new species of Phyllotreta are described: P. arcuata, P. attenuata, P. bisinuata, P. constricta, P. dolichophalla, P. emarginata, P. ramosoides, P. spatulata, and P. utanula. Two new synonyms are given: P. aequalis Hatch (=- P. denticornis Horn) and P. amphicornis Chit- tenden (= P. denticornis Horn). In addition, new synonyms of seven former varieties/variations (= subspecies) include: P. armoraciae var. biplagiata Chittenden (= P. armoraciae (Koch)); P. decipiens var. ordinata Chittenden (= P. decipiens Horn); and P. sinuata var. discedens Weise, P. sinuata var. monticola Weise, P. vittata var. artivitta Chittenden, P. vittata var. lineolata Chittenden, and P. vittata var. vernicosa Chittenden (all five = P. striolata (Fabricius)). Lectotypes are designated for six species: P. ar- moraciae (Koch), P. decipiens Horn, P. denticornis Horn, P. lepidula (LeConte), P. liebecki Schaeffer, and P. oregonensis (Crotch). Neotypes are designated for three species: P. bi- pustulata (Fabricius), P. striolata (Fabricius), and P. zimmermanni (Crotch). Three species, known only from females, are con- sidered incertae sedis: P. alberta Chittenden, P. ob- tusa Chittenden, and P. perspicua Chittenden. The male of P. oblonga Chittenden is described for the first time. Two of the new species, P. arcuata and P. bisinuata, have the wings reduced or vestigial. Acknowledgments Many individuals and institutions have assisted by loaning specimens and allowing examination of col- lections under their care. I would like to thank the following individuals in particular: Larry Burgess (ACSS); H. G. Wylie (ACWM); Lee H. Herman, Jr. (AMNH); W. Wayne Moss and David C. Rentz (ANSP); Melville Hatch (BMUW); Hugh B. Leech (CASC); Terry N. Seeno (CDAE); John A. Chemsak (CISC); Claude Chantal (CLCC); Edward C. Becker (CNCI); L. L. Pechuman (CUIC); the late David C. Miller (DCMC); Phillip J. Clausen (DEFW); Brett C. Ratcliffe (DEUN); W. J. Hanson (EMUS); Edward U. Balsbaugh, Jr. (EUBC and SDSU); the late Henry Dybas (FMNH); Robert E. Woodruff (FSCA); G. H. Nelson (GHNC); George C. Wallace (ICCM); Milton W. Sanderson (INHS); the late Jean L. Laffoon (ISUI); the late Joe Schuh; Charles L. Hogue and Roy R. Snelling (LACM); Larry Burgess (ACSS); John F. Lawrence, Jan Scott, and Margaret Thayer (MCZC); Roland L. Fischer (MSUC); David A. Young (NCSU); N. M. Downie (NMDC); Charles A. Triplehorn (OSUC); Paul Oman (OSUO); Edward E. Simons (PADA); Carlo Brivio (PIME); Ke Chung Kim (PSUC); Arwin Provonsha (PURC); E. Phil Rouse (UADE); George Ball (UASM); Saul Frommer SMITH: REVISION OF PHYLLOTRETA CHEVROLAT (UCRC); F. W. Wood (UMDC); Eben A. Osgood, Jr. (UMDE); W. R. Enns and E. G. Riley (UMRM); Richard E. White (USNM); Lutz J. Bayer (UWEM); the late Vernon M. Kirk (VMKC); Michael Kosztarab (VPIC); William J. Turner (WSUC); Jan Edelmann (UZMC); Gerhard Scherer (ZSBS); and F. Hieke (Zoological Museum of Humboldt University, Berlin, DDR). 1 would like to thank the following persons who have reviewed the manuscript and offered valuable suggestions: Edward U. Balsbaugh, Jr., Donald J. Borror, Laurnet LeSage, G. Allan Samuelson, Charles A. Triplehorn, Barry D. Valentine, Richard E. White, and John A. Wilcox. I am particularly in- debted to Barry Valentine, under whose guidance the original research was completed, and to G. A. Sam- uelson and R. E. White for critically reviewing this present version. It is with pleasure that I acknowledge John A. Wilcox's never-ending encouragement, the sharing of his vast knowledge of the chrysomelids, and his friendship over the years. I gratefully acknowledge and thank Marlene Hill- Werner for the handsome habitus drawings and the elytral color pattern variation drawings. I wish to thank Mary E. Lydon for her assistance, especially in proofreading. Most of the research was done while I was a gradu- ate student in the Department of Entomology at Ohio State University. The resulting dissertation was sub- mitted in partial fulfillment of the requirements for my Ph.D. degree to the Graduate School of the Ohio State University in 1973. At Ohio State University, partial financial support was received in the form of an NSF Summer Traineeship for graduate teaching assistants for the summer of 1971 and as a Dissertation Year Fellowship from the Graduate School; these are grate- fully acknowledged. Additional research was done after I joined the staff of Field Museum of Natural History. I am grateful to both of these institutions and their personnel for providing me with working space and material aid. Introduction Phyllotreta is a cosmopolitan flea beetle genus of about 150 species, of which about 44 occur in the Nearctic Region. Two of the nearctic maculate spe- cies, P. armoraciae and P. striolata, are regarded as introductions from Eurasia. Also, a third Eurasian species, P. undulata, is included, but its establish- ment in the United States is questionable. This partial revision of the genus Phyllotreta in- cludes only the maculate species known to occur in America north of Mexico. A key to these species is presented. The biology for each species has been sum- marized and the distribution is given. This revision is based on a study of about 13,100 specimens; in addi- tion, the types of most of the immaculate species, including their male genitalia, were studied. The genus Phyllotreta was proposed by Chevrolat (1837) in the third edition of the DeJean Catalogue of Coleoptera and included 14 species, two occurring north of Mexico. Crotch (1873), in the first American publication of a comprehensive nature treating the North American chrysomelids, described all the new species he recognized in the LeConte and Horn col- lections. Chapuis (1875) was the first to present a complete classification of the Alticinae. He placed Phyllotreta in the group "Aphthonites " which was later considered of tribal rank. Horn (1889) mono- graphed the Alticinae of America north of Mexico, the only time this has been done. He slightly altered Chapuis' classification and placed Phyllotreta , along with Longitarsus, Glyptina, and Aphthona, in the group "Aphthonae" which is, with the addition of Palaeothona (? = Lupraea ), now considered the tribe Aphthonini. However, there currently is no gen- erally accepted tribal classification for the Alticinae. Horn (1889) presented the first really comprehen- sive revision of Phyllotreta. The most recent revision was by Chittenden (1927), and in it, 17 new species and five new varieties or subspecies were described. Even though 16 of his new species are valid, unfortunately, his key is poor and his descriptions are somewhat inaccurate and inadequate. He studied nei- ther the aedeagi nor the spermathecae; the former provide the critical characters used for the separation of many species. This genus contains many species of economic im- portance (see the Biology section for the genus and for each species). Biological work has recently been done and is being continued on several species by research- ers at Cornell University (see Root & Tahvanainen, 1969; Feeny et al., 1970; Tahvanainen, 1972; and Hicks, 1972). J. K. Nielsen of Denmark is now the most active researcher/publisher of Phyllotreta bio- logical studies, especially with respect to host plant interrelationships (see Nielsen, 1977, 1978a, 1978b; and Nielsen et al., 1977, 1979). Methods Measurements For the key, comparative measurements were made at 16x for the beetle length (whole beetle, one mea- FIELDIANA: ZOOLOGY surement), at 64 x for antennal segments and stripe width, and at 160X for the spermatheca. All mea- surements, including those below, were made using a Leitz Model TS binocular dissecting microscope with ocular grid (1 mm divided into 40 parts). Each species description includes those specimens representing both the shortest and longest beetles. Table 1 gives a comparative chart of the measurement ranges for Nos. 1-3 and 5-6 below, and Table 2 gives a comparative chart of the antennal segments. The measurements were made in the following manner: 1. Beetle length: sum of elytral length (see No. 6 below) and distance measured along midline from pronotal base to most anterior part of frontal carina; measured at 64 x . 2. Beetle width: Measured on a line perpendicular to elytral suture at greatest breadth (about elytral mid- length); measured at 64 x. 3 . Interocular distance/maximum diameter of eye (IOD/MDE): narrowest separation of eyes on dorsal aspect of head; and maximum diameter of eye, max- imum dimension of eye in lateral view; both measured at 160X. 4. Antennal segment length/width: measurements are given in ocular grid units and represent maximum length or width; only half units are estimated, if di- mension was only slightly more or less than a unit, this is indicated by a plus or minus sign; measured at 160X. 5. Pronotal length and width: length measured on dorsal midline from basal to anterior edge, and width measured on a line perpendicular to pronotal midline at greatest breadth; measured at 64 x. 6. Elytral length and width: length measured along dorsal line from base of scutellum along suture to apex of elytra; for width, see No. 2 above; measured at 64x. 7. Length of aedeagus: measured dorsally or lat- erally from base of basal foramen to apex; measured at 160X. 8 . Length of spermatheca: measured in lateral view from most anterior part of sclerotized portion of sper- mathecal duct to posteriormost extent of pump; mea- sured at 160x. 9. Length of spermathecal receptacle: measured in lateral view from most anterior part of receptacle to base of pump (transverse line where sclerotization markedly changes intensity). Genitalia Preparation The beetle was first softened, the abdomen re- moved and treated with KOH, and the genitalia dis- sected out and studied. The genitalia were placed in glycerine in a small polyethylene genitalia vial (males) or mounted on a card beneath the specimen (females). For details about dissection and prepara- tion techniques, see Smith (1979a). Illustration Preparation Within each set of illustrations, all comparable structures were drawn at the same magnification; i.e., all antennae were drawn at the same magnification and each one is therefore comparable with all other antennae. The beetle habitus drawings, as well as those of the antennae and sterna, were made with the aid of a Wild M5 binocular dissecting microscope equipped with a camera lucida. For the habitus drawings, the elytra and pronotal basal margins were drawn viewed per- pendicularly to the elytral disk, and the pronotum and head were drawn from the angle where all four pro- notal corners were in focus at once. These were put together by matching the basal pronotal angles and using the basal margin drawn from the former orientation. Male genitalia were placed in a drop of glycerine in a shallow circular depression slide (if necessary, 1 or 2 grains of white sand was used for positioning). The general features were drawn using a Ken-A- Vision technical model microprojector, and the details were added using a Leitz TS binocular dissecting micro- scope. The spermathecae were drawn with the same microprojector, or if the spermathecal gland was not clearly visible, a camera lucida-equipped phase- contrast Wild M20 compound binocular microscope was used for drawing the spermathecal gland. Distribution Maps The maps represent the distribution of specimens personally examined. The localities are recorded in two ways. When a locality within a state or province is known, it is represented by a solid symbol (dot, square, etc.) at this locality. If only a state or pro- vincial locality is known (and no other locality within that state/province is known), it is represented by the same symbol but in open configuration, and it is placed in approximately the center of the state/ province. Systematic Formats Key Format — The key to species is artificial and dichotomous, but an explanation is needed about the order of characters within the couplet. If color pattern SMITH: REVISION OF PHYLLOTRETA CHEVROLAT will separate the species, it is given first because it is easiest, though it may vary in reliability. Color is followed by morphological characters, principally of the antennae and genitalia. The male genitalia provide the most reliable characters. Species Format — The descriptions or rede- scriptions are based on holotypes, lectotypes, or neo- types, with the exception of P. undulata for which the location of the type is unknown. All descriptions of variations are placed in parentheses, but no variation is given for punctation, and measurement ranges are summarized in Table 1. In the Biology section under host plants of adults, an asterisk is used to indicate records which are probably hosts, i.e., plant species belonging in the Cruciferae, Capparidaceae, Lim- nanthaceae, or Tropaeolaceae. The Habits section in- cludes a summary of the dates of collection. A detailed listing of the specimens examined is located in the Appendix. The four-letter abbreviations denoting the location of the specimens are listed in the Appendix under List of Abbreviations. Classification Subfamily ALTICINAE Illiger Head rounded, more or less inserted into prothorax; antennae inserted between inner margins of eyes, usu- ally short, often dilated toward tip, rarely pectinate or long and filiform. Pronotum transverse, almost as wide as elytra, less often square or subrectangular; mostly punctate. Elytra usually oblong oval, always covering tergites except rarely with pygidium ex- posed. Prosternum more or less convex between coxae, very rarely narrow and coxae contiguous. Legs of medium length, rather robust; metafemora always more or less swollen, grooved beneath, almost always with a large extensor apodeme; tibiae most often grooved on outer face; tarsi short, dilated, last seg- ment sometimes globose or swollen; claws appen- diculate, rarely bifid or simple. For a key to the genera of the Alticinae, see Arnett (1963). Genus Phyllotreta Chevrolat Phyllotreta Chevrolat, 1837:391. Type-species: Chrysomela brassicae F. (Designated by Chevrolat, in d'Orbigny, 1845:6; clarification by White, 1970.) Orchestris Crotch, 1873 (not Kirby, 1837):65. Type-species: Chrysomela nemorum nemorum L. (Synonymy by LeConte, 1878:615.) Tanygaster Blatchley, 1921:26-27. Type-species: Tanygaster ovalis Blatchley (by original designation and monotypy). (Synonymy by Smith, 1979b:359.) Major References— Crotch (1873), Horn (1889), Blatchley (1910), Duckett (1920), Chittenden (1927), Beller & Hatch (1932), Heikertinger & Csiki (1939), Wilcox (1954), Hatch (1971), and Balsbaugh & Hays (1972). Diagnosis — Small (less than 3.7 mm), shape elon- gate oblong, glabrous, completely dark or each ely- tron with a median pale stripe or 1 or 2 pale marks. In addition, antenna 1 1-segmented, slender (males of some species with segment 5 and often also 4 dilated); postantennal tubercles obsolete; pronotal width about 2 x length; elytra confusedly punctate; procoxal cavi- ties open; metatibial apical spur short, simple; 1st metatarsal segment much less than Vi tibial length, 5th tarsal segment not swollen apically; claws simple. Description — Small, 1.79-3.62 mm long, 0.88-1.85 mm wide; elongate oval to oblong oval; moderately convex to subdepressed; usually glabrous above but lightly pubescent ventrally with vesture short, fine, and pale, antennal segments pubescent with vesture sparse on basal segments to moderate on apical segments; dark, but legs, antennae, and/or ely- tra may have pale pattern. Head — Dark; covered by prothorax to just behind eyes; eyes prominent, not emarginate; postantennal tubercles obsolete, reduced to flat spots without sharp superior delineation; frons usually evenly punctate; vertex punctation quite variable in size and spacing within a species so variation not given in species de- scriptions. Antennae 1 1-segmented, about Vi body length, slender with 1st and apical segments slightly wider (males of some species with 1 or more of seg- ments 4-7 dilated, usually 5 and often also 4 dilated). Maxillary palpi with last segment about as long as preceding, elongate conical. Prothorax — Width about 2 x length, somewhat narrower apically; moderately convex, without dis- tinct antebasal or lateral impressions; basal margin adjacent to hind angles transverse, not oblique; punc- tures coarsest and closest on disc, usually less coarse and more widely spaced laterally (puncture size and spacing varies considerably within most species such that it is not diagnostic, so punctation variation is not included in species descriptions); procoxal cavities open. Elytra — Oblong oval, moderately convex to sub- depressed; entirely dark or each elytron with a median pale stripe or 1 or 2 pale marks; confusedly punctate with punctures coarsest and closest on disc, usually less coarse and more widely spaced laterally and in apical V2-V3 (puncture size and spacing varies enough FIELDIANA: ZOOLOGY within most species such that it is not diagnostic, so punctation variation is not included in species de- scriptions); humeri never prominent. Wings — Normally with metathoracic wings well developed, but both males and females of P. arcuata with wings vestigial (sometimes well developed) and P. bisinuata with wings reduced in males and well developed (sometimes vestigial) in females. Legs — Metafemora greatly enlarged, saltatorial; tibiae normal, without a tooth above followed by a ciliate emargination; apical spurs inserted medially on apex, short, with a single point, 1st metatarsal seg- ments much less than 14 tibial length, 5th metatarsi simple, not swollen apically; claws simple with, at most, a very small basal swelling. Normal color se- quence: trochanters pale, femora darkest, tibiae inter- mediate, tarsi pale (often with apical segments darker), all articulations paler than adjoining seg- ments; rarely with a different sequence. When "nor- mal color sequence" is used in a species description, it means the sequence described here. Abdomen — Fifth sternum sexually dimorphic, simple in outline in females (fig. 1), but males (fig. 2) with an apical median lobe formed by median Vs of apical margin set off by 2 narrow, moderately deep emarginations (pygidium fits this 3-part configura- tion), median lobe usually concave with concavity extending toward but ending before sternal base, male 5th sternum usually with a median linear impression extending part of or sometimes entire length; females sometimes with a median linear or oblong impression in apical 14 of 5th sternum, or sometimes extending to base; both sexes usually with shallow lateral prebasal impressions. Dark, often with apical margins of sterna 1-4 paler or darker, male usually with apical 14—14 of 5th paler. Male Genitalia — Figures 4, 5; terminology modified from that used by Sharp & Muir (1912) and Powell (1941). Aedeagus a single slender tube arched dorsoventrally; nearly uniform in width. Basal foramen narrower in basal half, almost entirely open ventrally. Tegmen ventral, attached medially at about midpoint of basal foramen; slender, usually about 14— lA total length of aedeagus and bifurcate at apical 14—14, the branches wrapped around median lobe but not meeting dorsally. Median lobe usually slightly tapering to apex which, in dorsal view, is usually rounded or acute, but may be emarginate or truncate; orifice dorsal, located preapically (in lateral view, on a plane oriented about 30° from longitudinal axis of median lobe), oblong in outline, occluded by a trans- parent area reinforced by 3 longitudinal sclerotized strips; sometimes, dorsally and/or ventrally with a "washboard" which consists of a median series of short transverse, parallel, heavily sclerotized, evenly spaced ridges, which usually start postbasally to basal foramen and extend onto orifice base, a ventral wash- board may be interrupted medially by a longitudinal concavity resulting in 2 parallel rows of "teeth"; usu- ally with a ventral concavity or impression, at least in apical lA. Female Genitalia — Figure 6; terminology mod- ified from that used by Samuelson (1966, 1973). Spermatheca, in lateral view, with pump of uniform width, about 14-% length of receptacle, usually more heavily sclerotized in apical XA , apex bluntly rounded. Receptacle (lateral view) usually cylindrical to elon- gate oval, slightly constricted premedially, but may be elongate pear-shaped or oblong; usually with a ring collar at junction with pump; uniformly sclerotized; in dorsal view, laterally compressed, shallowly C- shaped. Spermathecal duct (lateral view) usually at- tached to anterior end of receptacle premedially (in dorsal 14), rarely on dorsum (figs. 157, 186), usually paralleling receptacle outline anteriorly and ventrally to just beyond gland valve attachment; usually con- stricted at anterior end of gland valve attachment, sometimes swollen just before constriction; usually gradually tapering posteriorly beyond gland valve at- tachment, uniformly sclerotized to where this taper- ing ceases, then not visibly sclerotized (most of un- sclerotized portion of duct usually broken off during dissection). Gland valve (lateral view) globose, mod- erately developed, prominent; spermathecal gland at- tached on ventral posterior surface (often broken off during dissection or not visible except with phase- contrast or interference microscope). Sexual Dimorphism — This is exhibited in overall size (females usually slightly larger), 5th abdominal sternum (see section on abdomen above), and in mid- dle antennal segments of several species (see section on head above). Immature Stages — The egg, larva, and pupa have been described for only 3 species included in this revision: P. armoraciae by Chittenden (1917), P. striolata by Shimer (1869) and Riley (1885a), and P. zimmermanni by Riley (1885b). All 3 species have the larva and pupa illustrated, but only P. armoraciae has the egg drawn. The descriptions of Riley and Chit- tenden are fairly detailed, but Shimer's is sketchy. Adults of other species have been reared from larvae, but no descriptions have been published of the imma- ture stages. Both P. armoraciae and P. striolata are Eurasian introductions, and numerous European au- thors have treated their immature stages. Biology — Most members of this genus are confined to the Cruciferae, but a few feed on the closely related Capparidaceae, Limnanthaceae, and Tropaeolaceae. It has been demonstrated that these beetles are attracted by the mustard oils and mustard SMITH: REVISION OF PHYLLOTRETA CHEVROLAT oil glucosides found in these plants (Gornitz, 1953; Feeny et al., 1970; Matsumoto, 1970). In the temperate zone, a generalized life cycle would be as follows: adults overwinter; mating and egg-laying occur in early spring, the eggs are laid on or close to the part of the plant upon which the larvae feed (some are leaf miners, few are stem miners, and others are root feeders); the prepupal and pupal stages are spent in an earthen cell in the ground; and the adults emerge in late summer and are active until late fall. The larvae are leaf miners (P. constricta, P. lie- becki, P. oregonensis, and P. zimmermanni), stem miners (P. armoraciae, restricted to the petiole and midrib), or root feeders (P. striolata ). The transition from leaf to stem feedling has been observed for P. zimmermanni (Chittenden, 1927, p. 21), and from stem to root feeding in P. armoraciae (Chittenden, 1917, p. 10). Contrary to the habits of North Ameri- can species, worldwide, the majority of known larvae are stem and root borers or root feeders (Frost, 1924). Among our North American species, P. ar- moraciae, P. conjuncta (recorded as P. bipustulata ), P . decipiens , P . oregonensis , P . ramosa , P . striolata , and P. zimmermanni have been irregularly of economic importance. Recently, several researchers at Cornell University have been working on such biological aspects as phe- nology and microhabitat selection (Tahvanainen, 1972), the influence of vegetation diversity on popu- lation ecology (Tahvanainen, 1972), host plant specificity (Feeny etal., 1970), and the role of tempo- ral hosts on seasonal development of the crucifer fauna (Root & Tahvanainen, 1969). In 1979, Wilcox published a list of host plants for the Chrysomelidae of northeastern North America in which he incorporated original data from my Ph.D. dissertation and the manuscript I sent him for inclu- sion in the North American Beetle Fauna Project; due to an oversight, my dissertation was not included in the bibliography. This involves six of my new host plant records recorded under four species (P. bi- pustulata, P. liebecki, P. ramosa, and P. robusta). In the interest of documenting these new records, I have left these entries in my New Collection Records sec- tions, but have put them in brackets. Discussion — Characters and Previous Revi- sions— It should be pointed out that, with the excep- tion of the antennal morphological characters, most of the external morphological and color characters used by previous workers are highly variable. In particular, this includes the texture, punctation, and luster of the head, pronotum, and elytra, the color of the antennae and legs, and the elytral pattern for most species. Contrary to Chittenden (1927), the sexes may be easily distinguished by the apical margin of the 5th abdominal sternum (see section on abdomen above). The aedeagi show the most reliable characters for separation of species, whereas the spermathecae are usually reliable only for separation to groups of closely related species because of their variability in shape. The male antennae provide very reliable char- acters for most of the species, particularly those which exhibit sexual dimorphism. Several species exhibit reduction in the size of pale stripes or marks to the extent that they may be indis- tinct or completely absent. Therefore, it is possible that representatives of these maculate species may be found among the thousands of undetermined immacu- late specimens, the subject of Part II of this revision. It should be noted that I use the term "alutaceous" to mean "covered with minute cracks, like the human skin" as defined by Toreo-Bueno (1962, p. 9). Many previous workers have used this term to describe the texture condition which I call granulate, meaning peb- bly or like sandpaper. Chittenden's 1927 revision of Phyllotreta deserves a cautionary comment concerning his treatment of the maculate species. Excluding P. armoraciae , his key to species is based almost completely on male charac- ters, yet four of his new species (P. alberta, P. ob- longa, P. obtusa, P. perspicua) which are separated by "male" characteristics were known to him only from females that he incorrectly sexed. Also, Hatch's 1971 review of Phyllotreta for the Pacific Northwest deserves a note of caution. A fair number of the speci- mens which I have examined that bear his deter- mination label are misidentified. Therefore his listed localities should be accepted provisionally. Nomenclature and Name — The authorship of the genus Phyllotreta has been credited to several people, but White (1970) has clarified this situation, Chevro- lat (1837) being the correct author. Also, several spe- cies have been cited as the type-species, but Chevrolat (1845, in d'Orbigny) satisfactorily designated Chrysomela brassicae Fabricius (1787) as the type- species (White, 1970); it is now a synonym of P. exclamationis (Thunberg, 1774). Crotch's un- fortunate adoption of Kirby's name Orchestris for this genus has been adequately refuted by LeConte (1878, p. 615). The name Phyllotreta comes from the Greek phyllo- meaning leaf, and treta meaning to perforate; leaf perforation is the characteristic flea beetle dam- age. The accent is on the penult syllable (based on the Greek vowel 17, which is long): Phyllotreta. Relationships — The genus Phyllotreta is most closely related to Longitarsus, Glyptina, Aphthona, FIELDIANA: ZOOLOGY and Palaeothona, the other members of the Aph- thonini. This tribe is characterized by the following: small size, oval form, 11 -segmented antennae, open procoxal cavities, pronotum without any antebasal impression, mesosternum always visible, metatibiae not sulcate or sinuate on outer edge, 1st metatarsal segments long and slender, 5th metatarsal segments slender, and claws simple. Phyllotreta can be sepa- rated from these four genera on the basis of the follow- ing: metatibiae not grooved on the outer edge, but slightly excavated near the tip and the spur inserted medially, the 1st metatarsal segments not much more than one-third tibial length at most, the elytra con- fusedly punctate, each often with a pale stripe or marks, and the postantennal tubercles obsolete. Superficially, species of Phyllotreta might be con- fused with those of Systena (Chevrolat, 1837), Epi- trix (Foudras, 1860), and Chaetocnema (Stephens, 183 1 ), but in these three genera, the procoxal cavities are closed instead of open. In addition, species of Epitrix are usually pubescent instead of glabrous, those of Chaetocnema are more robust and their meta- tibiae have a preapical tooth followed by a ciliate emargination, and species of Systena are usually much longer (about twice as long), if they have pale stripes these are simple and straight, and the antennae are simple, with unmodified segments. Phylogeny — It is difficult to propose phylogenetic relationships of this cosmopolitan genus after study- ing in detail only about half the nearctic species. However, one observation seems worthy of note: the native nearctic maculate species form two distinct groups. These groups can be separated on the basis of antennal segment modifications and ratios and the dor- sal washboard character of the aedeagus; for details, see Table 3. To be specific, in Group I, the male 5th antennal segment is enlarged (one species has seg- ment 5 elongated only) and the aedeagus is without a dorsal washboard, whereas in Group II, the male 5th antennal segment is normal (one species has segment 5 slightly elongated only) and the aedeagus has a dorsal washboard. In addition, the length ratio of the 5th to 6th antennal segments in Group I averages 3.14 for males (lowest ratio is 2.0) and 1.57 for females (three ratios of 1.38), versus in Group II the ratio averages 1 .07 for males (one ratio of 1 .5) and 1 .05 for females (one ratio of 1.22). For both groups, as noted above, there is one spe- cies each which has antennal segment 5 elongated only. These two species exhibit distinctive extremes in ratios in their respective group; i.e. , in Group I, this species represents the lowest ratio for males, and its female is one of three females with the lowest ratio, and in Group II, this species represents the largest ratio for both sexes. One might speculate that these species represent a transition between the two groups with respect to modification of the 5th antennal segment. It should be pointed out that the maculate species do not constitute a natural group. This division of the genus is one of convenience. Key to the Maculate Species of Phyllotreta of America North of Mexico 2(1) 2' 3(2) 3' 4(3') 4' 5(4) 5' Elytra maculate, with pale marks or stripes 2 Elytra immaculate, entirely dark brown to black; may have metallic luster but no pale markings Revision of these species to be published as Part II Male, 5th abdominal sternum with an apical median lobe whose margin is surrounded by apical margin of pygidium (fig. 2), median lobe concavity not attaining sternal base 3 Female, apex of 5th abdominal sternite evenly rounded or smooth (fig. 1), apical xh sometimes with a median linear or oblong impression 32 Each elytron pale yellow, outlined in brown or black with sutural margin slightly wider (fig. 8); northern xh U.S. and adjacent Canada east of Rocky Mts. and Pacific Northwest armoraciae (Koch) p. 14 Elytra brown to black, with pale stripes or marks 4 Each elytron with 1 or 2 pale marks 5 Each elytron with a median pale stripe 12 Antennal segment 6 with a sharp anteroventrally directed process (fig. 66); each elytron with an indistinct postbasal and/or preapical yellowish/reddish brown mark (fig. 20); Pacific Coast states. (in part) denticornis Horn p. 24 Antennal segment 6 simple, with no process; each elytron with 1 indistinct small preapical yellowish/ reddish brown mark or with 2 yellowish marks 6 SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 6(5') Each elytron with 1 small preapical reddish brown to yellow mark (fig. 18); Rocky Mts. and westward, Texas decipiens Horn p . 23 6' Each elytron with 2 yellowish marks 7 7(6') Antennal segment 5 about 2 x length of 6 and enlarged (fig. 89); elytral preapical mark incurved apically (figs. 40-42); widespread (in part) striolata (F). p. 38 7' Antennal segments 5 and 6 subequal in length, 5th simple; elytral preapical mark with sutural margin straight, apex not incurved (figs. 10-11, 15-16); east of Rocky Mts 8 8(7') Each elytron with both pale marks tapering toward each other (fig. 15); east of Rocky Mts (in part) conjuncta Gent. p. 19 8' Each elytron with only apical pale mark at most slightly tapered toward basal mark (figs. 10-11, 16) 9 9(8') Less than 2.3 mm long or greater than 2.6 mm long 10 9' Between 2.3 and 2.6 mm long 11 10(9) Less than 2.3 mm long; east of Rocky Mts (in part) conjuncta Gent. p. 19 10' Greater than 2.6 mm long; east of Great Plains (in part) bipustulata (F.) p. 17 1 1(9') Aedeagus with apex in lateral view at most gradually curved ventrad (fig. 115), lateral margin of apical orifice not reaching midlength of median lobe (in part) conjuncta Gent. p. 19 1 V Aedeagus with apex in lateral view curved at an angle ventrad (fig. 109), lateral margin of apical orifice extending at least to midpoint of median lobe (in part) bipustulata (F). p. 17 12(4') Elytral stripe of almost uniform width, without lateral or terminal dilations, slightly incurved apically (figs. 7, 12, 19, 35); 5th antennal segment expanded bilaterally, dorsoventrally flattened (fig. 49) 13 12' Elytral stripe with 1 or 2 lateral dilations and/or terminal dilations (figs. 9, 24); 5th antennal segment may be enlarged or expanded 17 13(12) Antennal segment 6 with a sharp anteroventrally directed process (fig. 66); Pacific Coast states .... (in part) denticornis Horn. p. 24 13' Antennal segment 6 simple, without a process 14 14( 13') Antennal segments 7- 1 1 darkest, in contrast to paler basal segments; antennal segment 5 with venter longitudinally concave only on longitudinal Vi furthest from other antenna; southern California arcuata n.sp. p. 13 14' Antennal segments 5-11 darkest at least dorsally; antennal segment 5 with venter longitudinally concave medially or flat 15 15(14') Antenna with at least segments 1-5 paler on venter, segment 2 often pale; antennal segment 5 with venter flat; northern California spatulata n.sp. p. 37 15' Antennal segments 2-3 pale, others dark; antennal segment 5 with venter longitudinally concave medially 16 16(15') Antennal segment 4 length about 2 x maximum width (fig. 72); metathoracic wings normal, well developed; aedeagus in lateral view slightly sigmoid (fig. 131); California, ?New York lepidula (LeC.) p. 28 1 6' Antennal segment 4 length and width subequal (fig. 57); metathoracic wings reduced to less than elytron length; aedeagus in lateral view with a very abrupt, strong bend premedially (fig. 1 12); California. bisinuata n.sp. p. 18 17( 12') Elytral pale stripe greatly expanded over most of elytral width, at least in apical xh (expanded portion rarely only slightly paler than background color) reaching lateral, posterior, and sutural margins apically (figs. 26, 33) 18 17' Elytral pale stripe at most moderately expanded preapically, never covering most of preapical area, stripes never reaching apical margins 19 18(17) Antennal segment 5 with a long pointed apical prolongation (fig. 85); northern Vi U.S. and adjacent Canada robusta LeC. p. 36 Antennal segment 5 with apical Vi at most moderately expanded laterally (fig. 75); Great Plains and eastward, ?California liebecki Schffr. p. 29 19(17') Antennal segment 4 moderately and 5 strongly expanded bilaterally, dorsoventrally flattened (figs. 94, 98) 20 FIELDIANA: ZOOLOGY 19' Antennal segment 4 and 5 simple, or if 5 enlarged it is essentially cylindrical (figs. 58, 81, 89) 25 20(19) Antennal segments 2-5 pale, in distinct contrast to darker segments 7-11; segment 6 may be pale or dark 21 20' Antennal segments 2-4 pale and/or dark, segments 5-11 dark 22 21(20) Length over 2.8 mm (about 3 mm); aedeagus in lateral view slightly sigmoid with apex almost straight dorsally, and strongly swollen medially on venter (fig. 160); California, Nevada, Oregon, Utah, Washington utana Chttn. p. 42 21' Length less than 2.8 mm (about 2.6 mm); aedeagus in lateral view with dorsum gradually arched, apex curving lightly ventrad, and venter only very slightly swollen medially (fig. 163); Pacific Northwest, Nevada, Utah utanula n.sp. p. 44 22(20') Antennal segment 5 with a distinct concavity in basal V* anteriorly or to outside, 5th much wider than 4th (fig. 98); apex of aedeagus with a narrow median emargination (fig. 165), moderately curved ventrad in lateral view (figs. 166-167); widespread zimmermanni (Crotch) p. 46 22' Antennal segment 5 evenly flattened ventrally although slightly concave, 5 subequal in width to 4 (figs. 61, 70, 79); apex of aedeagus entire (figs. 116, 139) or if emarginate, then ventrally straight or flat in lateral view (figs. 127-128) 23 23(22') Elytral stripe width at middle distinctly less, usually much less, than distance from stripe to suture (fig. 24); apex of aedeagus with a narrow median emargination (fig. 127), straight or flat ventrally in lateral view (fig. 128); Oregon, northern California emarginata n.sp. p. 26 23' Elytral stripe width at middle about equal to or usually greater than distance from stripe to suture (figs. 17, 29), rarely less but if less, then apex of aedeagus entire, with no median emargination (figs. 1 16, 139) 24 24(23') Pronotal disc punctures usually more widely separated than elytral disc punctures; aedeagus abruptly narrowed in apical Vi and apex in lateral view distinctly directed ventrad (fig. 118); east of Rocky Mts. and west of Mississippi River constricta n.sp. p. 21 24' Pronotal and elytral disc punctures usually separated by about same distance; aedeagus gradually narrowed toward apex which is essentially straight in lateral view (fig. 140); Rocky Mts. and westward oregonensis (Crotch) p. 31 25(19') Elytral stripe apical dilation with inner margin straight, apex not incurved (figs. 13-14); east of Rocky Mts (in part) conjuncta Gent. p. 19 25' Elytral stripe apex strongly incurved (figs. 23, 31) 26 26(25') Antennal segments 4 and especially 5 slightly to moderately enlarged in diameter, essentially cylindri- cal, 5th subequal to or 2 x longer than 6th (figs. 89, 92) 27 26' Antennal segments 4 and 5 simple; segments 4-6 usually subequal in length, or if 5th is longer than 4 or 6 it is not enlarged (figs. 77, 83) 28 27(26) Antennal segment 5 at least 2 x length of 6th, 5th uniformly enlarged for most of its length (fig. 89); elytral stripe constricted in median xh , width medially usually much less than distance from stripe to suture (figs. 36-39); widespread (in part) striolata (F.) p. 38 27' Antennal segment 5 subequal to 6th, 5th distinctly enlarged in apical W-% and tapering to base (fig. 92); elytral stripe gradually somewhat narrower in median Vi , width medially often almost equal to distance from stripe to suture (fig. 43); Maryland undulata Kutsch p. 41 28(26') Elytral stripe with median Vi very narrow, of uniform width, postbasal lateral dilations with posterior margin almost at 90° angle to stripe, apical Vs of stripe abruptly widened, appearing notched basally on outside (figs. 31-32) 29 28' Elytral stripe with median VS not of uniform width, gradually widening basally into postbasal lateral dilation and apically into wider apical VS, sometimes with a lateral notch Vi from apex (figs. 9, 23, 28) 30 29(28) Antennal segment 5 longer than 4 or 6, 4 and 6 subequal in length (fig. 83); longer, about 2.5 mm; aedeagus in lateral view moderately and evenly arched dorsoventrally with apex gradually curved ventrad (fig. 146); Pacific Northwest, Northwest Territories ramosoides n.sp. p. 35 SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 29' 30(28') 30' 31(30') 31' 32(2') 32' 33(32') 33' 34(33) 34' 35(34) 35' 36(35') 36' 37(34') 37' 38(37) 38' 39(38') 39' 40(39) 40' 41(39') 41' 42(37') Antennal segments 4-6 subequal in length (fig. 81); shorter, about 2.2 mm; aedeagus in lateral view only very slightly arched dorsoventrally with apex bent ventrad at about 45° angle (fig. 143); Pacific Coast states, Nevada, ?New York ramosa (Crotch) p. 33 Elytral stripe with inner margin subparallel to suture except incurved apically, its width medially much less than distance from stripe to suture, and no basal dilation (fig. 28); antennal segment 5 about 1 .25 x length of 6 (fig. 77); Canada, New Hampshire, Oregon, Wisconsin oblonga Chttn. p. 30 Elytral stripe with inner margin sinuous, stripe width medially almost equal to distance from stripe to suture, stripe with basal dilation (figs. 9, 23); antennal segments 4-6 subequal in length 31 Elytral stripe with basal dilation small and not attaining midlength of postbasal lateral dilation; inner margin somewhat incurved (fig. 23); antennal segments 2-5 pale, 6-7 transitional, 8-11 dark; aedeagus longer, about 1.4 mm, in lateral view apex distinctly directed ventrad (fig. 126); California dolichophalla n.sp. p. 25 Elytral stripe with basal dilation large and extending beyond midlength of postbasal lateral dilation, inner margin subparallel to suture (fig. 9); antennal segments 2-4 pale, 5 transitional, 6- 1 1 dark; aedeagus shorter, about 1.2 mm, in lateral view apex almost straight (fig. 107); Colorado, South Dakota attenuata n.sp. p. 15 Each elytron pale yellow, outlined in brown or black with sutural margin slightly wider (fig. 8); northern V2 U.S. and adjacent Canada east of Rocky Mts. and Pacific Northwest armoraciae (Koch) p. 14 Elytra brown to black with pale stripes or marks 33 Each elytron with 1 or 2 pale marks 34 Each elytron with a median pale stripe 43 Each elytron with 1 pale mark 35 Each elytron with 2 pale marks 37 Each elytron with a postbasal pale mark (fig. 21); Pacific Coast states (in part) denticornis Horn p. 24 Each elytron with a preapical pale mark 36 Elytral mark usually incurved (fig. 18); shorter than 2.5 mm (about 2-2.4 mm); spermathecal recep- tacle shorter, distinctly tapered in apical Ys to a moderately developed apical ring collar (fig. 175); Rocky Mts. and westward, Texas decipiens Horn p. 23 Elytral mark not incurved (fig. 22); longer than 2.5 mm (about 2.6 mm); spermathecal receptacle longer, gradually tapering apically to a slightly developed ring collar, receptacle very slightly constricted medially on venter (fig. 176); Pacific Coast states ... (in part) denticornis Horn p. 24 Antennal segments 5 and 6 subequal in length (figs. 56, 60); east of Rocky Mts 38 Antennal segment 5 about 1 .5 x length of 6th (figs. 67, 90) 42 Each elytron with both pale marks tapering toward each other (fig. 15); east of Rocky Mts (in part) conjuncta Gent. p. 19 Each elytron with only apical pale mark at most slightly tapering toward basal mark (figs. 10-11, 16) 39 Less than 2.4 mm long or greater than 2.7 mm long 40 Between 2.4 and 2.7 mm long 41 Less than 2.4 mm long; east of Rocky Mts (in part) conjuncta Gent. p. 19 Greater than 2.7 mm long; east of Great Plains (in part) bipustulata (F). p. 17 Spermathecal duct attached dorsally , distance to gland valve attachment very short, gland valve located in anterior aspect (fig. 173); east of Rocky Mts (in part) conjuncta Gent. p. 19 Spermathecal duct attached anteriorly, distance to gland valve attachment much greater, gland valve located in ventral aspect (fig. 171); east of Great Plains (in part) bipustulata (F). p. 17 Elytral marks much larger and wider, with dilations, incurved (figs. 40-42); shorter than 2.5 mm (about 2-2.3 mm); spermathecal receptacle shorter, distinctly tapered in apical V3 to a moderately developed apical ring collar, pump longer than V2 receptacle length (fig. 187); widespread (in part) striolata (F). p. 38 10 FIELDIANA: ZOOLOGY 42' Elytral marks small and narrow, without dilations, not incurved (fig. 20); longer than 2.5 mm (about 2.6 mm); spermathecal receptacle longer, slightly narrowed apically with a slightly developed ring collar, pump much shorter (fig. 177); Pacific Coast states (in part) denticornis Horn p. 24 43(33') Elytral stripe of almost uniform width, without lateral or terminal dilations, slightly incurved apically (figs. 7, 12, 19, 25, 35); antennal segment 5 about 1.5 X length of 6 44 43' Elytral stripe with 1 or 2 lateral dilations and/or terminal dilations; antennal segment 5 from subequal to 1 .5 x length of 6th 48 44(43) Elytral stripes usually indistinct with borders blending into background color, somewhat pale reddish brown; antennal segment 4 about 1 .5 x length of 6th (fig. 67); Pacific Coast states (in part) denticornis Horn p. 24 44' Elytral stripes usually with borders clearly delimited, straw yellow; antennal segments 4 and 6 subequal in length 45 45(44') IOD/MDE ratio greater then 1.75 (about 1.8); spermathecal receptacle unique (fig. 186), basal end tapering to a blunt predorsal point, rather abruptly and strongly swollen ventrally in basal xh and elongate, almost parallel-sided in apical !4, ring collar moderately developed; northern California. spatulata n.sp. p. 37 45' IOD/MED ratio less than 1.70; spermathecal receptacle not as described above (figs. 168, 172, 179) 46 46(45') IOD/MED ratio less than 1.45 (1.33-1.38); spermatheca more robust, receptacle length about 2.7 x width and basal and apical width subequal (fig. 179); California, ?New York lepidula (LeC.) p. 28 46' IOD/MED ratio greater than 1.45 (1.50-1.63); spermatheca more elongate, receptacle length about 3.5 x width (fig. 172) or if length about 2.7 x width, then basal width greater than apical width (fig. 168) 47 47(46') Spermathecal receptacle with dorsal indentation opposite midpoint of ventral basal swelling (fig. 172), receptacle length/width ratio about 3.5, basal and apical width subequal (fig. 172); California. . . bisinuata n.sp. p. 18 47' Spermathecal receptacle without such an indentation (fig. 168), receptacle length/width ratio about 2.8, basal width 1.25 x apical width (fig. 168); southern California arcuata n.sp. p. 13 48(43') Elytral stripe greatly expanded over most of elytral width at least in apical lA, reaching lateral, posterior, and sutural margins (figs. 26-27 ', 33-34) 49 48' Elytral pale stripe at most moderately expanded preapically, never covering most of preapical area, stripes never reaching apical margins 50 49(48) Elytral median pale stripe with a moderate postbasal lateral dilation and usually greatly expanded laterally only in apical 16, leaving median 56 of stripe narrow (figs. 26-27); Great Plains and eastward, ?California liebecki Schffr. p. 29 49' Elytral stripe with a strong postbasal lateral dilation and greatly expanded in apical xh-xh, leaving usually much less than median % of stripe narrow (figs. 33-34); northern lh of U.S. and adjacent Canada robusta LeC. p. 36 50(48') Elytral stripe with postbasal lateral dilation at most very small (fig. 43); Maryland undulata Kutsch p. 41 50' Elytral stripe with postbasal lateral dilation distinct and well developed (figs. 17, 23, 32, 47) ... 51 51(50') Elytral stripe with median xh very narrow, of uniform width, postbasal lateral dilations with posterior margin almost at 90° angle to stripe, apical Vi of stripe abruptly widened, appearing notched basally on outside (figs. 31-32) 52 51' Elytral stripe with median xh not of uniform width, gradually widening basally into postbasal lateral dilation and apically into wider apical xh , sometimes with a lateral notch !6 from apex (figs. 13, 28-29, 44, 47) 53 52(51) Antennal segment 5 longer than 4 or 6 (rarely subequal), 4 and 6 subequal (fig. 84); longer, about 2.5 mm; spermathecal sclerotized duct between receptacle attachment and gland valve, with length about 1 .5 x maximum width (fig. 184); Pacific Northwest, Northwest Territories ramosoides n.sp. p. 35 SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 11 52' Antennal segments 4-6 subequal in length (fig. 82); shorter, about 2.2 mm; spermathecal sclerotized duct between receptacle attachment and gland valve with length about 3.5 X maximum width (fig. 183); Pacific Coast states, Nevada, ?New York ramosa (Crotch) p. 33 53(51') Elytral stripe width medially almost equal to or usually greater than distance from stripe to suture (figs. 9, 17, 29, 45) 54 53' Elytral stripe width medially much less than distance from stripe to suture (figs. 13, 28, 30, 44) 57 54(53) Antennal segments 5 and 6 subequal in length (fig. 54); Colorado, South Dakota attenuata n.sp. p. 15 54' Antennal segment 5 about 1.5 x length of 6 (figs. 62, 80, 95) 55 55(54') Antennal segments 2-5 pale, 6 pale/intermediate/dark, 7-11 dark; spermathecal receptacle almost parallel-sided, base not swollen (fig. 189); California, Nevada, Oregon, Utah (in part) utana Chttn. p. 42 55' Antennal segment 2 pale, 3 pale/intermediate, 4 pale/intermediate/dark, 5- 1 1 dark, segment 5 never pale; spermathecal receptacle with ventral constriction about midlength, base swollen (figs. 174, 182) 56 56(55') IOD/MED ratio 1 .25- 1 .44; spermathecal receptacle strongly constricted medially on venter (fig. 174); east of Rocky Mts. and west of Mississippi River constricta n.sp. p. 21 56' IOD/MED ratio 1 .44- 1 .56; spermathecal receptacle only slightly constricted medially on venter (fig. 182); Rocky Mts. and westward (in part) oregonensis (Crotch) p. 31 57(53') Antennal segments 5 and 6 subequal in length (figs. 60, 69) 58 57' Antennal segment 5 at least 1 .4 x length of 6, if 5 only 1 .25 x length of 6 then elytral stripe with no basal dilation and inner margin of stripe subparallel to suture except incurved at apex (fig. 28) . . 59 58(57) Elytral stripe with apical inner margin straight, apex not incurved (figs. 13- 14); smaller, less than 2.75 mm long (usually less than 2.4 mm); spermathecal receptacle oval, duct attached on dorsum of receptacle with distance to gland valve attachment very short (fig. 173); east of Rocky Mts (in part) conjuncta Gent. p. 19 58' Elytral stripe incurved apically (fig. 23); larger, about 3 mm long; spermathecal receptacle unique (fig. 177), with normal duct and gland valve attachments; California, Oregon dolichophalla n.sp. p. 25 59(57') Elytral stripe with inner margin subparallel to suture except incurved at apex and stripe with no basal dilation (fig. 28); antennal segment 5 about 1.25 x length of 6 (fig. 78); Canada, New Hampshire, Oregon, Wisconsin oblonga Chttn. p. 30 59' Elytral stripe with inner margin sinuous, if subparallel to suture, then stripe moderately incurved basally (fig. 210); antennal segment 5 at least 1.4 x length of 6 60 60(59') Elytral stripe with basal terminal dilation strongly dilated and about same size as postbasal lateral dilation (figs. 24, 29-30, 36-39, 182) 61 60' Elytral stripe with basal terminal dilation at most only slightly dilated in comparison to postbasal lateral dilation (figs. 44, 46-48, 208-210) 63 61(60) IOD/MED ratio less than 1 .30 (1 . 1 1- 1 .28); spermathecal pump longer than Vi receptacle length (fig. 187); widespread (in part) striolata (F.) p. 38 61' IOD/MED ratio greater than 1 .40 (1 .44- 1 .76); spermathecal pump much less than Vi receptacle length (figs. 178, 182) 62 62(61') Spermathecal receptacle shorter (0.25 mm), more robust with maximum length about 2.6 x width(fig. 178); Oregon, northern California emarginata n.sp. p. 26 Spermathecal receptacle longer (0.27 mm), more elongate with maximum length about 3.1 x width (fig. 182); Rocky Mts. and westward (in part) oregonensis (Crotch) p. 31 63(60') Spermathecal receptacle with maximum width basally and apically subequal, ring collar indistinct to very slightly developed (figs. 189, 216) 64 Spermathecal receptacle with maximum width greater basally (figs. 191, 214-215) or apically (fig. 190), ring collar slightly to strongly developed 65 64(63) Elytral stripe sinuous, with basal dilation (figs. 44-45); Nevada, Oregon, Utah, Washington (in part) utana Chttn. p. 42 FIELDIANA: ZOOLOGY 64' Elytral stripe almost parallel to suture except for apices, with no basal dilation (fig. 210); Oregon perspicua Chttn. p. 48 65(63') Spermathecal receptacle with basal Vi wider than apical xh (figs. 191, 214-215), ring collar slightly or strongly developed 66 65' Spermathecal receptacle with basal V2 narrower than apical V2, ring collar moderately developed (fig. 190); Pacific Northwest, Nevada, Utah utanula n.sp. p. 44 66(65) Spermathecal receptacle with slightly developed ring collar (figs. 214-215) 67 66' Spermathecal receptacle with strongly developed ring collar (fig. 191); widespread zimmermanni (Crotch) p. 46 67(66) Beetle longer, about 2.8 mm; spermathecal receptacle longer (0.28 mm), basal Vi more strongly and abruptly swollen on venter (fig. 214); Alberta alberta Chttn. p. 48 67' Beetle shorter, about 2.2 mm; spermathecal receptacle shorter (0.24 mm), basal V2 less strongly and gradually swollen on venter (fig. 215); Colorado obtusa Chttn. p. 48 Phyllotreta arcuata Smith, NEW SPECIES. Figures 7,49-50, 100-102, 168, 192. Holotype: Male, deposited at Los Angeles County Mu- seum of Natural History. Type locality: Griffith Park, Los Angeles, California. Diagnosis — Each elytron with the median stripe simple, of uniform width and without lateral or termi- nal dilations, slightly incurved toward suture apically; male 4th and 5th antennal segments expanded bilaterally, dorsoventrally flattened, 5th with venter longitudinally concave on inner Vz only; antennal seg- ments 2-6 pale, in contrast to darker segments 7-11; wings usually vestigial. Description of Holotype (species variation, ex- cluding punctation, in parentheses) — Figure 7; elon- gate oval, length 2.50 mm (6*2.28-2.50, 92.18- 2.58), width 1.05 mm (6*1.05-1.12, $1.02-1.20); head and pronotum black with slight metallic luster, elytra brownish black, each with a median straw- yellow stripe. Head: Black; vertex finely granulate basally to slightly roughened anteriorly (entirely finely granulate), moderately punctate with punctures separated by less than to 2 x their diameter, mostly by about 1 diameter; interocular distance/maximum diameter of eye, 1.63 (61.50-1.63, 91.50-1.63). Antennae: Figures 49-50; segments 4 and 7 about 2 x length of 6, 5 about 2 x length of 4; segments 4 and 5 expanded bilaterally and dorsoventrally flattened; segment 5 distinctly longest, with venter longitudinally concave on inner V2 only; see Table 2 for segment length/width, length 1.50 mm (allotype: length 1.45 mm; simple); antennae various shades of brown, basal 6 (5) segments paler with dorsum of 1 darker, none (6) intermediate, 7-11 darkest. Pronotum: Length 0.45 mm (60.42-0.45, 90.38- 0.48), width 0.75 mm (60.72-0.78, 90.72-0.78); black, texture slightly roughened (finely granulate); moderately to coarsely punctate, punctures separated by less than to equal their diameter, mostly by about 1 diameter. Elytra: Length 1.80 mm (61.60-1.85, 9 1.62- 1.88), width 1.05 mm (6 1.05- 1.12, 91.02- 1.20); brownish black (dark brown), median stripe as in Figure 7; texture slightly roughened (usually smooth); coarsely punctate in basal V2, moderately punctate in apical V2 , punctures separated by less than Vi to equal their diameter, mostly by less than 1 diameter. Legs [metatarsi with left missing tar- someres 2-5, right missing 5th]: Normal color se- quence. Abdomen: 5th sternum with apical median lobe moderately concave, concavity oval in its exten- sion to within V3 of sternal base, with a median linear impression extending from sternal base and ending preapically (6: median linear impression ending XA from base; 9 : simple in outline, usually with apical V2 shallowly concave, usually with median linear im- pression in apical V2); black with apical V3 paler except along midline (dark brown to brownish black). Male — Aedeagus (figs. 100-102); length approxi- mately 0.80 mm; moderately arched dorsoventrally, of rather uniform width; apex in dorsal view very bluntly rounded with shallow, moderately wide me- dian emargination, in lateral view gradually but strongly curved ventrad, end bluntly rounded; ventral view as in Figure 168. Female — Spermatheca (fig. 168); length approxi- mately 0.35 mm; receptacle elongate, ring collar slightly developed; pump long. Type Material — The holotype, allotype, and 1 paratype ( 9 ) are at the LACM; 2 paratypes (16, 19) are at the USNM; 2 paratypes (16, 1 9) are in my collection, EHSC; 4 paratypes (16, 3 9) are at the CASC; and 1 paratype (6) is at the FMNH. The holo- type is mounted on a point and the pin bears 2 labels (descending order): "Griffith Pk LA 4.4.25" and my holotype label. The allotype bears the same locality label and my allotype label. The paratype at the LACM and the 2 each at the EHSC and USNM have the same locality label as the holotype; all 4 paratypes SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 13 at the CASC have the same 2 labels (descending or- der): "Santa Marguerita [sic] Calif. IV-26-1933" and "R. HOPPING COLLECTION". The paratype at the FMNH has 3 labels (descending order): "Hullville, VI:9:31 Calif." "Col. by H. Dorn" and "Pres. by E. Liljeblad". Biology — Host plants and immature stages are un- known. Habits — Adults have been collected in southern California during May and June. Distribution — Figure 192; P. arcuata has been found only in Hullville, Los Angeles, and Santa Mar- garita, California. Specimens Examined — Total 12, see Type Mate- rial above. Discussion — Relationships — Phyllotreta arcuata is most similar to P. lepidula and P. bisinuata and less similar to P. utanula and P. utana. Phyllotreta arcuata shares having a simple median elytral stripe with P. lepidula, P. bisinuata, P. spatulata, and the striped specimens of P. denticornis. The antennal character of the 5th segment dorsoventrally flattened is shared with P. utana, P. utanula, P. oregonensis, P. emarginata, P. constricta, and P. zimmermanni , but P. arcuata shares with P. utana and P. utanula the additional feature of having antennal segments 2 through 6 pale in contrast to 7 through 1 1 dark. The apex of the aedeagus in dorsal view is very similar in P. arcuata, P. lepidula, and P. bisinuata, but the lateral view will separate these since P. arcuata is evenly arched dorsoventrally, whereas P. lepidula is essentially straight, and P. bisinuata is strongly sigmoid. Etymology — The specific name arcuata comes from the Latin arcus- meaning arch, and refers to the moderately and evenly arched aedeagus as seen in lateral view. Phyllotreta armoraciae (Koch). Figures 8, 51-52, 103-105, 169, 192. Haltica armoraciae Koch, 1803. Entomol. Hefte, 2:75-76; table 3, fig. 6. Lectotype (here designated): Male, at Zoological Mu- seum of Humboldt Univ., Berlin, DDR. Type locality: ?; lectotype is from Hungary. Phyllotreta armoraciae: Foudras, 1860. Ann. Soc. Linn. Lyon, 6(ser. 2):344. Haltica vittata Stephens, 1831 (not Fabricius, 1801). Illus. British Entomol., Mandibulata, 4:297. (Syn- onymy by Weise, 1888:865.) Phyllotreta armoraciae var. biplagiata Chittenden, 1927. Entomol. Amer., 8(n.s., no. 1): 16. NEW SYNONYMY. Holotype: Male, USNM type #28792, at United States National Museum. Type locality: Green Bay, Wisconsin. Diagnosis — Each elytron pale, outlined in brown or black with sutural margin slightly wider; antennae simple. Description of Lectotype (species variation, ex- cluding punctation, in parentheses) — Figure 8; elon- gate oval, length 2.82 mm (6*2.48-3.38, 93.08- 3.62), width 1.35 mm (6*1.30-1.62, 91.60-1.85); head and pronotum black with slight metallic luster (no luster), elytra dark brown, each elytron with a very wide median straw-yellow stripe covering most of surface. Head: Black (brownish black); vertex finely granulate basally to roughened anteriorly (finely granulate, finely alutaceous basally to rough- ened anteriorly, finely alutaceous), moderately punc- tate with punctures separated by 1-2 x their di- ameter, mostly by about 1 diameter; interocular distance/maximum diameter of eye, 1.30 (61.30- 1.50, 91.36-1.67). Antennae: Figures 51-52; seg- ments 4-6 subequal in length, simple; see Table 2 for segment length/width, length 1.76 mm (9 paralecto- type: length 1.95 mm; simple); antennae various shades of brown, basal 3 paler, 4-11 (4 intermediate, 5-11) darker. Pronotum: Length 0.50 mm (60.42- 0.60, 90.55-0.62), width 0.92 mm (60.85-1.05, 9 1.05-1.15), black (brownish black); texture finely granulate to slightly roughened (finely granulate, slightly roughened); moderately punctate, punctures separated by less than Vi to equal their diameter, mostly by about 1 diameter. Elytra: Length 2. 10 mm (61.90-2.50, 92. 38-2. 78), widthO. 92 mm (6 1.30- 1.62, 91.60-1.85); dark brown (brown, usually brownish black), median stripe pattern as in Figure 8; texture slightly roughened (usually smooth); mod- erately to coarsely punctate, punctures separated by less than Vi to equal their diameter, mostly by less than 1 diameter. Legs: Normal color sequence except tibiae and tarsi same color (normal color sequence). Abdomen: 5th sternum with a shallowly concave api- cal median lobe, concavity extending to within ¥t of base; a median linear impression extending post- basally to base of median lobe, impression somewhat indistinct or interrupted (concavity moderate, linear impression distinct, extending to sternal base; 9 : sim- ple in outline, usually with a shallow narrow median elliptical impression extending from apex to within Vi of base or to base); brownish black, median lobe area paler (5th sternum with apical xh paler). Male — Aedeagus (figs. 103-105); length approxi- mately 1.06 mm; slightly arched dorsoventrally, al- most uniform in width, with a dorsal concavity ex- tending from about midpoint and ending preapically; apex in dorsal view with a very wide, moderately deep median emargination, in lateral view with a preapical dorsal swelling and a rounded end; ventral view as in Figure 105. 14 FIELDIANA: ZOOLOGY Female — Spermatheca (fig. 169); length approxi- mately 0.38 mm; receptacle oblong, ring collar wide and slightly developed; pump long. Type Material — The lectotype and 7 apparent paralectotypes (16*, 69) are in the Zoological Mu- seum of Humboldt University, Berlin, DDR. This series is accepted as the type series on the authority of Dr. F. Hieke of this museum. It should be pointed out that, within this series, only 2 specimens bear labels and 5 different methods of mounting are used. The syntype selected as lectotype was chosen because it was the first in the series and has 2 more detailed labels associated with it, and its mount is unique in the series. The lectotype was pinned through the right elytron with a short (23-24 mm long) silvery pin of about #5 thickness (now remounted on a point), and the pin bears 3 labels (descending order): "55898" "Armoraciae EH.* Crioc. Nasturtii Fab? Hung. Jr. Rhen. Koy Koch. " [see fig. 3] and my lectotype label. The paralectotypes are as follows: 29 , each mounted on a short white point supported by a thin silvery pin; 1 8 , pinned with a very thin silvery pin bearing a very small cream-colored to white square of paper below; 1 9 , pinned with a thin silvery pin bearing a very small cream-colored to white square of paper below; 1 9 , mounted on a short gray point supported by a moderate-size silvery pin and bearing the label "Halae. Erichs." [this could be Crisks]; 29, each mounted identically to the previous female but with- out the label. Biology — Host Plants, Adult — * Meerrettig (Co- chlearia Armoracia) [* horseradish, Armoracia rusticana (Lam.) Gaertn., = Armoracia Armoracia] (Koch, 1803, p. 75); * marsh cress (Nasturtium plaus- tre) [Rorippa islandica (Oeder) Borbas, = Radicula palustris] (Chittenden, 1895, p. 405); * cabbage, not observed feeding (Chittenden, 1917, p. 8); * radish, * turnip (Beirne, 1971, p. 109); *Raphanus sativus (Wilcox, 1979, p. 20). New Collection Records: al- falfa, from Sauk Co., Wis., 5 mi. N. LaValle, May 3, 1948, J.T. Medler Coll. (UWEM:1); DuPuits alfalfa, from Savage Farm, Ithaca, N.Y., 9 June 1967, C.U. Lot 946 (CUIC:1); *Raphanus, from Ithaca, N.Y., 30 July '30 (CUIC:3). Host Plants, Larva — Horse-radish (Chittenden, 1917, p. 2), Immature Stages — Egg, larva, and pupa described by Chittenden (1917). Habits — Larvae are stem miners on the basis of plant anatomy, being restricted to the petiole and mid- rib of horseradish leaves; rarely may they mine the roots (Chittenden, 1917). Adults cause either the typ- ical shot-hole leaf damage or gouge pits in the midrib and petiole (Chittenden, 1917). Adults of P. ar- moraciae have been collected in the northern half of the United States and adjacent Canada east of the Rocky Mountains from mid-April (as early as mid- February in the midwest) until mid-November and in Idaho in mid-August. The adults are long-lived, they overwinter, and then the females, with periodical mating, lay eggs until early August (Chittenden, 1917). In Wisconsin, larvae from eggs laid in late May emerge as adults in late July or early August (Chittenden, 1917). The most complete North Ameri- can biological account of this species is by Chittenden (1917). Distribution — Figure 192; P. armoraciae has been collected mainly in the northern half of the United States and adjacent Canada east of the Rocky Mountains but also in southern Missouri and northern Idaho and Oregon. It is an introduced species and was first collected in 1893 at Chicago, Illinois, by Chit- tenden (Chittenden, 1895). Specimens Examined — Total 1,121, see Type Ma- terial above and the Appendix. Discussion — Nomenclature — Koch (1803) de- scribed this species in Haltica, apparently from a series of eight specimens. Foudras (1860) transferred it to Phyllotreta. In 183 1 , Stephens described Haltica vittata which was synonymized by Weise (1888); this is not the Fabrician vittata described in Crioceris in 1801. Chittenden (1927) described the variety P. a. biplagiata which is to be treated as having subspecific rank according to the Rules of Zoological Nomen- clature, article 45 (d)(i) and 45 (e)(i). However, this subspecies is based on a unique specimen which is merely one that has the median pale stripe somewhat irregularly interrupted by various shades of brown. This makes the overall color somewhat darker, but otherwise, it is a typical P. armoraciae . Therefore, I have synonymized this subspecies. Relationships — Phyllotreta armoraciae shows no close similarities with any other species in this genus found north of Mexico. Phyllotreta attenuata Smith, NEW SPECIES. Fig- ures 9, 53-54, 106-107, 170, 193. Holotype: Male, deposited at United States National Museum.* Type locality: Cottonwood, South Dakota. Diagnosis — Each elytron with a median pale stripe whose large basal dilation extends beyond midlength of postbasal lateral dilation, inner margin of basal dilation subparallel to suture, medially stripe usually wider than distance from stripe to suture; antennal segments 4-5 simple, male antennal segments 6-11 * USNM has discontinued their type numbering system. SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 15 darkest in contrast to paler basal segments; aedeagus in lateral view with apex very slender, almost straight. Description of Holotype (species variation, ex- cluding punctation, in parentheses) — Figure 9; elon- gate oblong, length 2.58 mm (6*2.20-2.62, 92.30- 2.65), width 1.18 mm (61.05-1.20, 91.02-1.30); head and pronotum black with slight metallic luster, elytra brownish black (rarely with slight metallic lus- ter), each with a median straw-yellow stripe. Head: Black (rarely dark reddish brown or brownish black); vertex finely granulate basally to smooth anteriorly (usually finely granulate basally to slightly roughened anteriorly), moderately punctate with punctures sepa- rated by less than to 2 x their diameter, mostly by less than 1 diameter; interocular distance/maximum di- ameter of eye, 1.33 (61.30- 1.44, 91.33-1.56). Antennae: Figures 53-54; segments 5 and 6 equal in length, slightly longer than 4 and slightly shorter than segment 7; segments 4 and 5 simple, cylindrical; see Table 2 for segment length/width, length 1.46 mm (allotype: length 1.55 mm; simple); antennae various shades of brown, basal 4 (3) segments paler, 5 (4-5 or none) transitional, 6- 1 1 (4- 11 or 5- 1 1 ) darkest. Pronotum: Length 0.50 mm (60.40-0.50, 90.45- 0.50), width 0.82 mm (60.75-0.85, 90.75-0.85); black; texture finely granulate; coarsely punctate with moderate punctures interspersed, punctures separated by less than xh to equal their diameter, mostly by less than 1 diameter. Elytra: Length 1.88 mm (61.60- 1.95, 91.62-2.00), width 1.18 mm (61.05-1.20, 9 1 .02- 1 .30); brownish black (dark brown or black), median stripe as in Figure 9; texture smooth (usually smooth to slightly roughened or entirely slightly roughened); coarsely punctate with very coarse punc- tures interspersed in basal Vi, punctures separated by less than Vi their diameter. Legs: Normal color sequence. Abdomen: 5th sternum with an apical me- dian deeply concave lobe, concavity oblong and ex- tending to within xh of sternal base, with a median linear impression extending from sternal base to base of concavity (usually extending to base of median lobe, sometimes interrupted; 9: simple in outline, usually with a shallow longitudinal median impres- sion in apical xh ); black (brownish black) with median concavity paler (5th sternum unicolorous; 9 : usually with apical xh paler). Male— Aedeagus (figs. 106-107); length approxi- mately 1.20 mm, moderately arched dorsoventrally, of rather uniform width except slightly narrower in apical '/3 in dorsal view and very slender in apical !4 in lateral view; with a dorsal washboard; apex in dor- sal view acute, gradually tapering to a blunt point, in lateral view very narrow, ending in a sharp point. No ventral impression observed. Female — Spermatheca (fig. 170); length approxi- mately 0.37 mm; receptacle elongate, moderately swollen basally and tapering posteriorly to a wide, moderately developed ring collar. Type Material — The holotype, allotype, and 3 paratypes (26, 19) are at the USNM, 7 paratypes (1 6 , 69 ) are at the SDSU, and 2 paratypes (16,19) are in my collection, EHSC. The holotype is mounted on a point and the pin bears 3 labels (descending order): "Cottonwood S.D. June 9, 1954 H.C. Severin Coll. " and my and the USNM's type labels. The allo- type bears the same locality-date label and my and the USNM's type labels. The 7 paratypes at the SDSU have the following locality-date labels: same as holo- type (29); same as holotype, but date is July 9, 1954 (16); "Midland S.D. June 9, 1954 H.C. Severin Coll. " (1 9 ). The 3 paratypes at the USNM are labeled as follows (descending order): "Rocky Ford Col 26 August 15" "Radicula palustris" "H O Marsh Collector" "Chittenden No 1578" "oregonensis Phyllotreta Chttn. Ent Am 1927" (16, 19); "Rocky Ford Col 29 Aug 11" "Sugar Beets" "H O Marsh Collector" and "oregonensis Phyllotreta Chttn. Ent Am 1927" (16). The 2 paratypes (16, 19) in my collection (EHSC) are labeled with the same locality- date label as the holotype. Biology — Host Plants, Adult — * Marsh cress (Radicula palustris) (Chittenden, 1923, p. 135; re- corded as P. oregonensis) (USNM: 2); sugar beets, from Rocky Ford, Col. 29 Aug 11, H O Marsh Collector, Phyllotreta oregonensis Chttn. Ent. Am. 1927 (USNM:1). Immature Stages — Unknown. Habits — Adults have been collected in South Da- kota from late May until mid-August and in Colorado in late August. Distribution — Figure 193; P. attenuata has been collected in Colorado and South Dakota. Specimens Examined — Total 14, see Type Mate- rial above. Discussion — Relationships — Phyllotreta attenu- ata is most similar to P. dolichophalla and the speci- mens of P. conjuncta which have the postbasal and preapical marks connected. These three species share the following: antennal segments 4 and 5 simple, seg- ments 5 and 6 subequal in length, and the aedeagi very similar in form. The aedeagi differ mainly in length: that of P. dolichophalla is longest, that of P. attenu- ata is intermediate in length, and that of P. conjuncta is shortest; also, the apex in lateral view is almost straight in P. attenuata, whereas in P. conjuncta and P. dolichophalla it is distinctly directed ventrad. However, the spermathecae of these three species are very different: in P. conjuncta, the receptacle is oval 16 FIELDIANA: ZOOLOGY and robust, with the ring collar only slightly devel- oped and the spermathecal duct attached dorsally; in P. dolichophalla , the receptacle is elongate, strongly swollen in the basal two-thirds, and rather uniformly narrow in the apical one-third, with the ring collar strongly developed and the spermathecal duct at- tached predorsally on the base; and in P. attenuata, the receptacle is moderately swollen basally and ta- pered posteriorly, with the ring collar moderately de- veloped and the spermathecal duct attached pre- dorsally on the base. Phyllotreta attenuata and P. dolichophalla share the additional character of having the elytral stripe very wide medially, whereas in P. conjuncta the stripe is narrow medially. Phyllotreta attenuata might be confused with P. oregonensis because the elytral stripe of each is wide medially. However, P. attenuata has antennal seg- ments 5 and 6 subequal in length, 5 simple and cylin- drical, whereas P. oregonensis has antennal segment 5 about one and one-half times the length of 6, and in the male, segment 5 is dorsoventrally flattened. Etymology — The specific name attenuata comes from the Latin attenuat meaning thin and refers to the apical one-fourth of the aedeagus which is very nar- row in lateral view. Phyllotreta bipustulata (Fabricius). Figures 10-11, 55-56, 108-110, 171, 193. Crioceris bipustulata Fabricius, 1801. Systema Eleuther- atorum, 1:464. Neotype (here designated): Male, USNM type #100635, deposited at United States National Museum. Type locality: "Carolinae"; neotype is from Myrtle Beach, South Carolina. Orchestris bipustulata: Crotch, 1873. Proc. Acad. Nat. Sci. Philadelphia, 25:66. Phyllotreta bipustulata: Horn, 1889. Trans. Amer. Ento- mol. Soc, 16:300. Diagnosis — Each elytron with a postbasal and pre- apical pale mark, sutural margin of preapical mark straight, not incurved; antennal segments 4 and 5 sim- ple and cylindrical, segments 5 and 6 subequal in length; aedeagus and spermatheca distinctive. Description of Neotype (species variation, ex- cluding punctation, in parentheses) — Figure 10; elon- gate oblong, length 2.68 mm (6*2.38-3.00, 92.48- 3.08), width 1.22 mm (61.12-1.35, 91.10-1.45); head, pronotum, and elytra black with slight metallic luster, each elytron with a postbasal and preapical straw-yellow mark. Head: Black (often dark brown or brownish black); vertex finely granulate basally to slightly roughened anteriorly (entirely finely granu- late), moderately punctate with punctures separated by less than ¥x to equal their diameter, mostly by less than xh a diameter; interocular distance/maximum diameter of eye, 1.40(61.56-1.75, 91.56-1.60). Antennae: Figures 55-56; segment 4 shorter than 5, segments 5 and 6 subequal (equal) in length, segments 6 and 7 subequal (equal) in length; segments 4 and 5 simple and cylindrical; see Table 2 for segment length/width, length 1.49 mm (alloneotype: length 1 .44 mm; simple); antennae various shades of brown, basal 4 (1-6, usually 1-4; 1st rarely darker) paler, 5 (sometimes 3-4, 5-7, or none) transitional, 6-11 (usually 4—11 or 8-11) darkest. Pronotum: Length 0.60 mm (6 0.50-0.62, 9 0.52-0.68), width 0.85 mm (60.82-0.95, 9 0.82-0.98); black (rarely dark brown or brownish black); texture finely granulate (rarely finely alutaceous or roughened); coarsely punctate, punctures separated by less than Vi to equal their diameter, mostly by less than 1 diameter. Elytra: Length 1.88 mm (61.78-2.02, 91.82- 2.20), width 1.22 mm (61.12-1.35, 91.10-1.45); black (often brownish black), color pattern as in Fig- ure 10 (often as in fig. 11); texture slightly roughened (sometimes smooth to slightly roughened); coarsely punctate, punctures separated by less than V2 to equal their diameter, mostly by less than 1 diameter. Legs [metatarsi with left and tarsomeres 3-5 of right miss- ing]: Normal color sequence (often with tibiae and tarsi same color). Abdomen: 5th sternum with an apical median deeply concave lobe, concavity oblong and extending to within !6 of sternal base (oval in its extension to midpoint), with a median linear impres- sion (no impression) from sternal base to base of con- cavity (9: simple in outline, usually with a shallow median oblong impression in apical V3); black (brownish black, rarely dark brown) with apical V2 of 5th sternum paler (entirely dark or with median lobe to apical xh of 5th sternum paler). Male — Aedeagus (figs. 108-110); length approxi- mately 1 .27 mm; moderately arched dorsoventrally, of rather uniform width except slightly narrowed pre- apically in dorsal view, slender and tapering toward apex in apical V2 in lateral view; with a dorsal wash- board; apex in dorsal view acute, bluntly rounded, in lateral view very narrow and distinctly directed ventrad, ending in a sharp point; ventral view as in Figure 110. Female — Spermatheca (fig. 171); length approxi- mately 0.37 mm; receptacle elongate, slightly swollen in basal half, with ring collar wide , strongly developed . Type Material — The neotype and alloneotype are at the USNM. The neotype is mounted on a point and the pin bears 4 labels (descending order): "Myrtle Beach, S.C. 21 March 1963 V.M. Kirk" "Washed up on beach" and my and the USNM's type labels. The SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 17 alloneotype has the same 2 locality-date labels plus my and the USNM's type labels. Biology — Host Plants, Adult — *Dentaria di- phylla Michx. (Tahvanainen, 1972, p. 125); *Carda- mine douglassii and * Dent aria laciniata (Wilcox, 1979, p. 8, 11). New Collection Records: [*Cardamine douglassi [sic] (Torr.) Britton, from Cedar Springs Ont. May 2 1945 A. A. Wood (CNCI:2)]; [* Dentaria laciniata Muhl. (feeding on), from OHIO: Franklin Co. Sharon Woods Metro. Park 29.V.75 E.H. Smith Lot No. 450 (EHSC:6)]; Mullien [sic] (Verbascum sp.; collected on), from College Park, Md. 4/29/19 W H White Coll (USNM:1). Immature Stages — Undescribed. Habits — The adults overwinter in the top soil close to the host plants (Hicks, 1972) and usually are active by April. Eggs are laid in the soil around the host plant (Hicks, 1972), and the adults decline in numbers thereafter and are rarely encountered after June. The larvae probably spend most of the summer feeding on the green rhizomes (Tahvanainen, 1972) and the adults appear again in the fall. In the Ithaca, New York, area, P. bipustulata is re- stricted to woodland inhabiting Dentaria diphylla, with the adults common on it until the leaves die off in late June. I have collected it feeding on Dentaria lacin- iata in rich woodlands in Franklin County, Ohio, and Cook County, Illinois, and Marion County, Missouri. Adults have been collected from early April to late June and again from mid-July until late August or September in northeastern United States (in early May in Iowa and in late August in Mississippi) and in Canada from early May until early July. Many data in the literature about this species are erroneous; for example, being a pest in past years (Hicks, 1972) and feeding on a wide variety of cruci- fers (Chittenden 1902, 1923, 1927; Hicks, 1972). These data should probably be associated with P. conjuncta; see its Biology section. Distribution — Figure 193; P. bipustulata has been collected in northeastern United States with the addition of Iowa and Mississippi and in Ontario and Quebec in Canada. Specimens Examined — Total 237, see Type Mate- rial above and the Appendix. Discussion— Nomenclature— Fabricius ( 1 80 1 ) described P. bipustulata in the genus Crioceris. In 1873, Crotch in error placed P. bipustulata in Orchestris (see LeConte, 1878, p. 165) and Horn (1889) placed this species in Phyllotreta. The Fabrician type is considered to be lost and, therefore, a neotype is designated here; for details see Smith, 1977. Relationships — Phyllotreta bipustulata is most similar to P. conjuncta and less so to P. dol- ichophalla and P. attenuata; for a discussion of these relationships see this section under P. conjuncta and for the last two, see this section under P. attenuata. Phyllotreta bipustulata has often been confused with specimens of P. striolata which have the elytral stripe reduced to a postbasal and preapical pale mark. However, in P. bipustulata the elytral preapical mark has the sutural margin straight, not incurved, whereas in P. striolata the preapical mark is incurved apically. Also, in P. bipustulata, antennal segment 5 is simple and segments 5 and 6 are subequal in length, whereas in P. striolata antennal segment 5 in the male is en- larged, and in both sexes segment 5 is about two times longer than segment 6. Phyllotreta bisinuata Smith, NEW SPECIES. Fig- ures 12, 57-58, 111-113, 172, 193. Holotype: Male, CASC type #14211, deposited at California Academy of Sciences. Type locality: Green Point, Humboldt Co., California. Diagnosis — Each elytron with the median pale stripe simple, of uniform width and without dilations; male antennal segments 4 and 5 moderately expanded bilaterally, dorsoventrally flattened, 5 moderately concave ventrally; antennal segments 5-11 darkest in contrast to paler basal segments; metathoracic wings reduced to less than elytron length; aedeagus in lateral view with a very abrupt, strong bend premedially. Description of Holotype (species variation, ex- cluding punctation, in parentheses) — Figure 12; elon- gate oblong, length 2.65 mm (8 2.65-2.75, 92.68), width 1.28 mm (6*1.28-1.30, 9 1 .42); head and pro- notum black with slight metallic luster, elytra dark brown, each with a median straw-yellow stripe. Head: Black; vertex finely granulate basally to slightly roughened anteriorly (finely alutaceous bas- ally), moderately punctate with punctures separated by less than to equal their diameter, mostly by less than 1 diameter; interocular distance/maximum di- ameter of eye, 1.44(6*1.44-1.53, 9 1.56). Antennae [left missing segments 9-11]: Figures 57-58; seg- ment 4 longer than 6 and equal in length to 7, segment 5 more than 2 x length of 6 and almost 2 x length of 4; segments 4 and 5 moderately expanded bilaterally, dorsoventrally flattened, 5 with venter deeply concave in basal Vi to outside; see Table 2 for segment length/ width, length 1.59 mm (allotype: length 1.51 mm; simple); antennae various shades of brown, basal 4 (9:5) segments paler, 5 (none) darkest, 6-11 darker IS FIELDIANA: ZOOLOGY than basal segments. Pronotum: Length 0.50 mm (60.50, 90.52), width 0.88 mm (60.85-0.88, 90.88); black; texture finely granulate to slightly roughened (entirely slightly roughened); coarsely punctate, punctures separated by less than V2 to equal their diameter, mostly by less than 1 diameter. Elytra: Length 1.95 mm (6*1.95-2.00, 91.95), width 1.28 mm (6*1.28-1.30, 91.42); dark brown, brownish black in basal Va (usually entirely brownish black), median stripe pattern as in Figure 12; texture smooth (smooth to slightly roughened); moderately punctate with coarse punctures interspersed basally, punctures separated by less than lA to equal their diameter, mostly by about 1 diameter. Legs: Normal color sequence. Abdomen: 5th sternum with an api- cal median deeply concave lobe, concavity tapering as it extends almost to midpoint, with a median linear impression extending postbasally to just inside con- cavity (extending from sternal base to base of median lobe, sometimes interrupted; 9: simple in outline, with a very shallow oval median impression in apical V3 ); brownish black (black) with apical Vs paler (with only median lobe paler; 9 : entirely dark). Male — Aedeagus (figs. 111-113); length approxi- mately 1.37 mm; strongly sigmoid or twice bent, of rather uniform width except median lobe strongly constricted just anterior to junction with basal piece in dorsal view, swollen medially in lateral view; apex in dorsal view broadly rounded, with a moderately deep and moderately wide emargination, in lateral view gradually tapering to a wide bluntly rounded end which is directed ventrad, with a small preapical lobe on venter which is the ventral median emargination flange; ventral view as in Figure 113. Female — Spermatheca (fig. 172); length approxi- mately 0.39 mm; receptacle elongate, of rather uni- form width, moderately arched dorsoventrally, ring collar moderately developed. Type Material — The holotype, allotype, and 1 paratype ( 9 ) are at the CASC, 2 paratypes (16, 19) are in my collection (EHSC), and 3 paratypes ( 9 ) are at the MCZC. The holotype is mounted on a point and the pin bears 5 labels (descending order): "Green Point Humboldt Co., Cal. VI-9-16" "F.E. Blaisdell Collector" "Blaisdell Collection" and my and the CASC's type labels. The allotype bears the same first 3 labels and then my and the CASC's type labels. The paratype at the CASC is labeled: "Gustine Merced Co. IV 14 -45 Cal" "B.E. White Collector" "BURDETTE E. WHITE Collection Calif. Acad. Sci. Accession 1967". The 2 paratypes at EHSC are labeled: 6 , same first 3 labels of holo- type; 9, "Rosarito Bch., L. Cal. IV-5-39" "Com- positae" and "CD. Michener". The 3 paratypes at the MCZC are all labeled "Cal" and "Liebeck Col- lection"; 2 have an additional label, one is "P. ulkei Horn" and the other is "Museum of Comparative Zoology". Biology — Host Plants, Adult — Compositae, see Type Material above. Immature Stages — Unknown. Habits — Adults have been collected in California from early April to mid- June. Distribution — Figure 193; P. bisinuata is known only from California. Specimens Examined — Total 8, see Type Material above. Discussion — Relationships— Phyllotreta bisin- uata is most similar to P. lepidula and less similar to P. arcuata, P. spatulata, and striped specimens of P. denticornis; for a discussion of these relationships see this section under P. lepidula. Etymology — The specific name bisinuata comes from the Latin bi- meaning two and the Latin sinua- meaning bend. This refers to the sigmoid shape of the aedeagus as seen in lateral view. Phyllotreta conjuncta Gentner. Figures 13-16, 59-60, 114-115A, 173, 194. Phyllotreta conjuncta Gentner, 1924. Entomol. News, 35(5): 168. Holotype: Male, at Michigan State University. Type locality: East Lansing, Michigan. Phyllotreta bipustulata form conjuncta: Chittenden, 1927. Entomol. Amer., 8(n.s., no. 1):34. Phyllotreta bipustulata var. conjuncta: Wilcox, 1954. Ohio Biol. Surv. Bull., 43:467. Phyllotreta bipustulata var. conjuncta: Balsbaugh and Hays, 1972. Auburn Univ. Agr. Exp. Sta. Bull., 441:172. Phyllotreta conjuncta: Wilcox, 1979. World Nat. Hist. Pub. New York. p. 7. Diagnosis — Each elytron with a postbasal and pre- apical pale mark or these marks connected, sutural margin of preapical mark or stripe apex straight, not incurved; antennal segments 4 and 5 simple and cylin- drical, segments 5 and 6 subequal in length; aedeagus and spermatheca distinctive. Description of Holotype (species variation, ex- cluding punctation, in parentheses) — Figure 13; elon- gate oblong, length 2.22 mm (61.95-2.52, 92.02- 2.70), width 1.08 mm (60.95-1.20, 90.95-1.22); head and pronotum black with slight metallic luster, elytra dark brown (usually brownish black with slight metallic luster), each elytron with a median straw- yellow stripe dilated basally and apically (usually re- duced to a postbasal and preapical straw-yellow SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 19 mark). Head: Black (sometimes dark brown or brownish black); vertex finely granulate (usually finely granulate basally to slightly roughened ante- riorly), moderately punctate with punctures separated by much less than to 2 x their diameter, mostly by less than 1 diameter; interocular distance/maximum diameter of eye, 1.22 (6 1.20-1.25, 91.25-1.44). Antennae: Figures 59-60; segment 4 shorter than 5, segments 5 and 6 equal (subequal) in length, segments 6 and 7 equal (subequal) in length; segments 4 and 5 simple and cylindrical; see Table 2 for segment length/width, length 1.41 mm (9: length 1.54 mm); antennae various shades of brown, basal 4 (3 or 5; 1 sometimes darker) paler, 5 (none, 4, 4-5, 5-6) transi- tional , 6- 1 1 (4 or 7 through 1 1 , usually 5 or 6 through 11) darkest. Pronotum: Length 0.42 mm (60.40- 0.50, 90.40-0.50), width 0.75 mm (6*0.62-0.82, 90.65-0.85); black (rarely brownish black); texture finely granulate (rarely finely roughened to granulate or finely alutaceous to roughened); moderately punc- tate, punctures separated by much less than to equal their diameter, mostly by less than 1 diameter. Elytra: Length 1.65 mm (61.35-1.88, 91.42- 1.95), width 1.08 mm (60.95-1.20, 90.95-1.22); dark brown (usually brownish black), color pattern as in Figure 13 (sometimes as in fig. 16, usually as in figs. 14-15); texture slightly roughened (sometimes smooth to slightly roughened); coarsely punctate, punctures separated by much less than to equal their diameter, mostly by less than 1 diameter. Legs: Normal color sequence except tarsi slightly darker than tibiae (often with tibiae and tarsi same color). Abdomen: 5th sternum with an apical median deeply concave lobe, concavity oblong and extending half way to sternal base (usually extending to within V3 of sternal base, sometimes with base broadly rounded), with a faint median linear impression (no impression) from sternal base to base of concavity (9 : simple in outline, usually with a shallow median oblong impres- sion in apical V3 ); brownish black (dark brown or black) with 5th sternum dark brown (basally usually same color as other sternites; concavity usually paler). Male — Aedeagus (figs. 114-115A); length ap- proximately 1 .02 mm; slightly arched dorsoventrally, of rather uniform width in dorsal view, slender and tapering toward apex in apical Vi and with a smooth dorsally directed preapical arch in lateral view; with a dorsal washboard; apex in dorsal view acute, evenly tapered to rounded tip, in lateral view very narrow and distinctly directed ventrad, ending in a sharp point; ventral view (fig. 115A) from specimen of similar size. The tip was broken off on holotype but was added from specimen of similar size. 20 Female — Spermatheca (fig. 173); length approxi- mately 0.26 mm; receptacle oval, very robust with ring collar slightly developed; spermathecal duct attached dorsally with distance to gland valve attach- ment very short. Type Material — The holotype is at the MSUC. It is mounted on a point and the pin bears 8 labels (de- scending order): "AG. COLL. MICH. 12 July, 1921" "AG. COLLEGE LOT 1541 SUB. 52" "L.G. Gent- ner Collector" "AG. COLLEGE LOT 1541 SUB. 142" "6" "TYPE conjuncta gentner [sic] L.G.G." "Phyllotreta conjuncta Gentner" and "Abdomen lost E.H. Smith '80". Biology — Host Plants, Adult — The following represent those records verified by me via label data: *Arabis, *Barbarea verna, *Barbarea vul., *Brassica kaber, clover (red), * cabbage, * cabbage (Chinese), * cress, * horseradish, *Lepidium foliage, * mustard (feeding on), *peppergrass, Prunus vir- ginium [sic] flowers, *Radicula palustris, * radish foliage, and *turnip (foliage, matting). New Col- lection Records: Alfalfa, from Hagerstown, Md. 27 Apr. '15 W E Pennington Collector (USNM:1), *Brassica Pe-Tsai, from Waupaca, Wis., 6-23-20 L.G. Gentner Collector (USNM:1); *Radicula Nas- turtium-aquaticum [= Nasturtium officinale R.Br.; watercress], fromUtica, 111., V-22-1946 F.G.Werner (INHS:4); smartweed, from Harrison Co., Ind. VI- 23-1934 D W Lattue collector (PURC:1); sphagnum moss, from Volo, 111., Oct. 27, 1943 Ross & Sand- erson (INHS:1); 3-seeded Mercury, from Kokomo, Ind. June 27, 1922 G.M. Stirrett Collector (CNCL1, PURC:1); Trifolium pratense , from Westville, Mont- calm Co., Mich. 31 July 1959 H.D. Niemczyk (MSUC:1); wheat, from Urbana, 111. May 21, 1885 (INHS:1); W. Clover, from Arlington, Va. 5-14-28 F W Poos Collector (USNM:1); willow, from Morgan Co., Ind. V-14 1932 Musgrave (PURC:1). Immature Stages — Unknown. Habits — Most data previously attributed to P. bi- pustulata should probably be associated with P. con- juncta for the following reasons: (1) P. conjuncta inhabits open sunny areas of disturbed vegetation, whereas P. bipustulata is restricted to moist wood- lands; (2) most literature host records are for plants inhabiting open sunny areas of disturbed vegetation; (3) almost all of the literature host records for P. bipustulata are actually associated with P. conjuncta when the identification of the specimens is checked; and (4) specimens of P. conjuncta are much more numerous in collections than are those of P. bipustulata (1,024 vs. 237); also, see below under Nomenclature. FIELDIANA: ZOOLOGY In central Ohio, I have collected P. conjuncta and a few P. zimmermanni along with more numerous P. striolata and P. cruciferae (Goeze) feeding on Brassica kaber along the roadside in midsummer. I have collected it along with more numerous P. strio- lata and P. cruciferae, both in northwestern Indiana in early August feeding on turnips and lettuce at the end of the turnip rows and in northeastern Illinois in mid-September feeding on Chinese cabbage. Adults have been collected from late March to late June and again from early July until late August, occa- sionally as late as mid-November. Distribution — Figure 194; P. conjuncta has been collected east of the Rocky Mountains primarily north of 35° north latitude with the addition of southern Louisiana and Florida and in Manitoba, Ontario, Que- bec, and Saskatchewan in Canada. Specimens Examined — Total 1 ,024, see Type Ma- terial above and the Appendix. Discussion — Nomenclature — Gentner (1924) de- scribed this species from a single specimen. There are no synonyms, but I have listed for completeness the citations to this species when it was considered a form or variety of P. bipustulata. In 1979, Wilcox inadver- tently published this change of status while incorpo- rating original data from my Ph.D. dissertation and the manuscript I sent him for inclusion in the North American Beetle Fauna Project. Several previous systematists probably based their published papers on P. conjuncta instead of the recorded P. bipustulata:B\alch\ey (1910) and Wilcox (1954), as indicated by the size range and abundance given; Duckett (1920), as indicated by the size range, abundance, and host plants listed; and Chittenden (1927), as indicated by the size range and host plants listed. Possibly, the record of Balsbaugh & Hays (1972) should also be for P. conjuncta. Relationships — Phyllotreta conjuncta is most sim- ilar to P. bipustulata, with which it is often confused. Externally, only if both postbasal and preapical pale marks taper toward each other (= P. conjuncta ), and otherwise, size (within limits) is distinctive. Males less than 2.3 mm and females less than 2.4 mm long are almost always P. conjuncta, whereas males greater than or equal to 2.5 mm and females greater than 2.5 mm are almost always P. bipustulata. This leaves males 2.3-2.4 mm and females 2.4-2.5 mm long which must be dissected for examination of their genitalia. The aedeagus of P. conjuncta, in lateral view, has the apex gradually curved ventrad and the lateral margin of the apical orifice not reaching mid- length of median lobe, whereas P. bipustulata has the apex curved ventrally at an angle and the lateral mar- gin of the apical orifice extending at least to midpoint of median lobe. For the females, P. conjuncta has the spermathecal receptacle oval and very robust, with the duct attached dorsally and the distance to the gland valve attachment very short, whereas P. bipustulata has the spermathecal receptacle elongate, with the usual anterior duct and ventral gland valve attachments . Phyllotreta conjuncta has frequently been confused with specimens of P. striolata which have the elytral stripe reduced to a postbasal and preapical pale mark. However, in P. conjuncta, the elytral preapical mark has the sutural margin straight, not incurved, whereas in P. striolata, the preapical mark is distinctly in- curved apically. Also, in P. conjuncta, antennal seg- ment 5 is simple and segments 5 and 6 are subequal in length, whereas in P. striolata, antennal segment 5 in the male is enlarged, and in both sexes segment 5 is about two times longer than segment 6. Phyllotreta constricta Smith, NEW SPECIES. Fig- ures 17, 61-62, 116-118, 174, 195. Holotype: Male, deposited at United States National Museum. Type locality: Rocky Ford, Colorado. Phyllotreta constricta Smith, 1979. In Riley and Enns, Trans. Missouri Acad. Sci., 13:77. Nomen nudum. Diagnosis — Each elytron with a median pale stripe with dilations which never meet at suture, stripe medi- ally equal to or usually wider than distance from stripe to suture; male antennal segment 5 dark, expanded bilaterally and dorsoventrally flattened, subequal to 4 in width, with venter flat; aedeagus in lateral view abruptly narrowed in apical Vi and apex distinctly directed ventrad. Description of Holotype (species variation, excluding punctation, in parentheses) — Figure 17; elongate oblong, length 2.62 mm (6*2.55-2.85, 92.80-3.10), width 1.22 mm (61.15-1.30, $ 1 .32- 1 .48); head and pronotum black with distinct metallic luster, elytra brownish black with slight me- tallic luster (no luster), each with a median straw- yellow stripe. Head: Black; vertex finely granulate basally to slightly roughened anteriorly (entirely finely granulate), moderately punctate with punctures separated by less than !4 to 2 x their diameter, mostly by less than 1 diameter; interocular distance/ maximum diameter of eye, 1.56 (61.40-1.56, 9 1.25-1.44). Antennae: Figures 61-62; segment 4 longer than 3, 6, or 7, segment 5 about 1 .4 x (usually 1.5 x ) length of 4 and more than 2 x length of 6, 6 distinctly shortest; segments 4 and 5 expanded bilaterally, dorsoventrally flattened; segment 5 dis- SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 21 tinctly longest, subequal to 4 in width, with venter flat; see Table 2 for segment length/width, length 1 .56 mm (allotype: length 1 .61 mm; simple); antennae var- ious shades of brown, basal 3 segments pale with dorsum of 1 and 3 darker (3 usually entirely pale), 4-5 darkest (4 sometimes only dark), segments 6- 1 1 dark (9:4-11 dark). Pronotum: Length 0.45 mm (6*0.42-0.50, 90.48-0.55), width 0.82 mm (6*0.78-0.90, 90.90-1.02); black; texture finely granulate to slightly roughened (usually entirely finely granulate); moderately punctate with fine and coarse punctures interspersed, punctures separated by less than xh to 3 x their diameter, mostly by about 1 diameter. Elytra: Length 1.88 mm (61.80-2.02, 91.90-2.32), width 1.22 mm (61.15-1.30, 9 1 .32- 1 .48); brownish black (dark brown laterally), median stripe pattern as in Figure 17; texture slightly roughened (smooth to slightly roughened, or smooth); coarsely punctate with several moderate punctures in- terspersed, punctures separated by less than Vi their diameter. Legs [right mesotarsus and metatarsi miss- ing tarsomeres 4-5]: Normal color sequence. Abdo- men: 5th sternum with an apical median moder- ately concave lobe, concavity tapering (moderately rounded) posteriorly and extending to midpoint, a me- dian linear impression extending from sternal base to median lobe (ending postbasally and/or extending to preapex, sometimes interrupted; 9 : simple in outline, usually with a median longitudinal impression in api- cal 2A); black (brownish black) with median lobe paler. Male — Aedeagus (figs. 1 16- 1 18); length approxi- mately 0.94 mm; slightly arched dorsoventrally in dorsal view, of rather uniform width, in lateral view abruptly narrowed in apical V2; apex in dorsal view bluntly rounded, in lateral view abruptly tapered to a ventrally directed, rounded end; ventral view as in Figure 118. Female — Spermatheca (fig. 174); length approxi- mately 0.44 mm; receptacle elongate, strongly con- stricted medially on venter, ring collar indistinct. Type Material — The holotype, allotype, and 16 paratypes ( 126 , 4 9 ) are at the USNM; 1 paratype (6 ) is at AMNH; 5 paratypes (6) are at CASC; 3 para- types (6) are at CNCI; 7 paratypes (46, 39) are at EHSC; 4 paratypes (36, 19) are at MCZC; and 5 paratypes (46, 1 9) are at SDSU. The holotype is mounted on a point and the pin bears 6 labels (de- scending order): "Rocky Ford Col 26 Aug 15" "Ra- dicula palustris" "H O Marsh Collector" "6" "ore- gonensis Phyllotreta Chttn. Ent Am 1927" and my and the USNM's type labels. The allotype bears the same labels as the holotype, except in the place of the sex label is "Chittenden No 1578" and my and the USNM's allotype labels. The paratypes all have a paratype label added. The 5 6 paratypes at CASC are as follows: 2, "Eldorado Springs, Colo VII-12-1939 J.W. Green"; 2, "Porvenir, N. Mex. Dr. A. Fenyes" "A. Fenyes Col- lection"; and 1, "Show Low, Ariz. IX. 11. 1941 O. Bryant". The 3 6 paratypes at CNCI: 1, "Elbow, Sask. 24. VII 1954 Brooks-Wallis"; 1, "Ft. Collins Colo." "TESTE 16067 Chittenden"; and 1, "Ft. Col- lins Colo 6.13.22" "Colo 28" "TESTE 16067 Chit- tenden". The 7 paratypes in EHSC are: 26 and 1 9 , "Childress Tex. July 8, 09" "On turnip" "H O Marsh Collector" and female has in addition "oregonensis Phyllotreta Chttn. Ent Am 1927"; 16 and 19, "Ft. Collins Colo. 7. 1 1.22" correct sex symbol label, "or- egonensis Phyllotreta Chttn. Ent Am 1927" and fe- male has in addition "Colo 25"; and 1 6 and 1 9 , same first 4 labels of holotype. The 5 paratypes at SDSU are as follows: 36 and 19, "Elk Point, S.D. June 27, 1946 J. A. Lofgren, Coll."; and 16, "Freeman, S.D. June 27, 1946 J. A. Lofgren, Coll.". The 16 paratypes at the USNM are as follows: 16 and 19, same 5 labels as allotype; 16, "Medicine Hat Alta 31.V.25 F.S. Carr" "6" "oregonensis Phyllotreta Chttn. Ent Am 1927"; 16, "3 mi. N Tularosa N.M. 7/2/29 L. alyssoides" "from VE Romney"; 26, "Fowler Col 9 June 04" "ESG Titus Collector" "Cleome" 1 with "6 " and the other without; 1 6 and 1 9 , "LARAMIE WYO. 6-14-93" "WICKHAM Collection 1933"; 16, "Ft. Collins Colo. 7.3.32" "Colo 26" "ore- gonensis Phyllotreta Chttn. Ent Am 1927"; 1 9 , "Ft. Collins Colo. 7.11.22" "9" "oregonensis Phyl- lotreta Chttn. Ent Am 1927"; 16, "Garden City Kans"; 26, 19, "Childress Tex. July 8, 09" "On turnip" "H O Marsh Collector" sex symbol label, "oregonensis Phyllotreta Chttn. Ent Am 1927"; 16, "Childress Tex. July 8, 09" "On turnip" "H O Marsh Collector"; 16, "Lincoln Neb June". Biology — Host Plants, Adult — *Cleome ser- rulatum [sic], * marsh cress {Radicula palustris), and * turnip (Chittenden, 1923, p. 135; recorded as P. oregonensis); see Type Material for label and col- lection data. Host Plants, Larva — Cleome serrulata (Chit- tenden, 1923, p. 135; recorded as P. oregonensis). These host records represent Chittenden's Fowler, Colorado, record of Cleome serrulata, his Chil- dress, Texas, and Garden City, Kansas, records of turnip, and part of his Rocky Ford, Colorado, record of Radicula palustris. Immature Stages — Undescribed. Habits — The larvae are leaf miners of Cleome ser- 22 FIELDIANA: ZOOLOGY rulata (Chittenden, 1923, p. 135). Adults have been collected in southern Saskatchewan and Alberta in late April and late May, in the central Great Plains from mid-June until late August, and in New Mexico and Texas in early July. Distribution — Figure 195; P. constricta has been collected in south central Canada, the central part of the Great Plains, southern New Mexico, and northern Texas. Specimens Examined — Total 42, see Type Mate- rial above. Discussion — Relationships — Phyllotreta con- stricta is most similar to P. oregonensis and is less similar to P. emarginata, P. zimmermanni , P. utanula, and P. utana. For a discussion of these re- lationships, see this section under P. oregonensis. Phyllotreta constricta might be confused with P. attenuata and P. dolichophalla on the basis of having the elytral stripe wide medially, but P. constricta has the male 5th antennal segment dorsoventrally flattened, whereas in P. attenuata and P. dol- ichophalla it is simple. Etymology — The specific name constricta comes from the Latin constrictus meaning drawn together or contracted. This refers to the aedeagus in lateral view being abruptly narrowed in the apical one-half and to the deep premedian emargination on the venter of the spermathecal receptacle. Phyllotreta decipiens Horn. Figures 18, 63-64, 119-121, 175, 196. Phyllotreta decipiens Horn, 1889. Trans. Amer. Ento- mol. Soc, 16:298-299. Lectotype (here designated): Male , MCZC type #32849, at Museum of Comparative Zoology, Harvard. Type locality: Oregon and Washington Territory; lec- totype is from Oregon. Phyllotreta decipiens var. ordinata Chittenden, 1927. Entomol. Amer., 8(n.s., no. 1):38. NEW SYN- ONYMY. Holotype: Male, USNM type #28814, at United States National Museum. Type locality: Elko, Nevada. Diagnosis — Each elytron with 1 small preapical reddish brown to yellow mark or elytra entirely dark; male antennal segment 5 dark, simple but slightly enlarged, about 2 x length of 6. Description of Lectotype (species variation, ex- cluding punctation, in parentheses) — Figure 18; ob- long, length 2.35 mm (6*1.85-2.40, 92.38-2.55), width 1.15 mm (6*0.92-1.15, 91.20-1.30); head and pronotum black with slight metallic luster (head no luster), elytra brownish black with slight metallic luster (no luster), each with a preapical yellowish (to reddish) brown mark. Head: Black; vertex finely granulate (to slightly roughened anteriorly), mod- erately punctate with punctures separated by less than to equal their diameter, mostly by less than 1 di- ameter; interocular distance/maximum diameter of eye, 1.22 (61.00-1.28; 91.22-1.38). Antennae [left missing segments 3-11]: Figures 63-64; seg- ments 4 and 7 equal (subequal) in length, longer than 6, segment 5 about 2 X length of 6; segment 5 simple, slightly enlarged in diameter; see Table 2 for segment length/width, length 1.49 mm (9: length 1.32 mm; simple); antennae various shades of brown, basal 3 segments paler, 4-6 darkest (4 intermediate or usu- ally 4-6 only dark), 7-11 dark (4-11 or usually 5- 1 1 darkest) (9 : basal 2 or 3 segments paler, 3 or 4 intermediate, 4-11 or 5-11 darkest). Pronotum: Length 0.45 mm (60.35-0.45, 90.42-0.48), width 0.75 mm (60.60-0.75, 9 0.60-0.80); black; texture finely granulate to slightly roughened (usually entirely finely granulate); coarsely punctate, punctures sepa- rated by less than xh to equal their diameter, mostly by less than V2 a diameter. Elytra: Length 1.68 mm (61.40-1.80, 91.75-1.90), width 1.15 mm (60.92-1.15, 91.20-1.30); brownish black (often black, rarely dark brown), color pattern as in Figure 18 (or elytra entirely dark); texture slightly rough- ened (often smooth); coarsely punctate, punctures separated by less than V2 diameter, moderate (to slight) serial tendency. Legs [right metatarsus missing tarsomeres 4-5]: Normal color sequence. Abdomen: 5th sternum with an apical median shallowly concave lobe, concavity oval and extending to apical V3 (oblong or tapering, extending to midpoint), with a median linear impression extending from midpoint to preapex (extending from sternal base to preapex, sometimes interrupted; 9 : simple in outline, usually with a shallow median oval impression in apical lA); black (often brownish black) with median lobe dark brown except median linear impression black (apical margin to apical % paler or entirely dark). Male — Aedeagus (figs. 1 19-121); length approxi- mately 0.79 mm; moderately arched dorsoventrally, of rather uniform width except slightly narrowed pre- apically in dorsal view, with a dorsal washboard; apex in dorsal view broadly acute (about 45°), lateral mar- gins almost straight, tip rounded, in lateral view grad- ually tapering in apical XA to very narrow apex, abruptly directed ventrad with tip pointed; ventral view as in Figure 121. Female — Spermatheca (fig. 175); length approxi- mately 0.32 mm; receptacle elongate oblong, wider basally and gradually tapering to posterior, mod- erately developed ring collar. SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 23 Type Material — The lectotype, allolectotype, and 8 paralectotype are at the MCZC. The lectotype is mounted on a point, and the pin bears 4 labels (de- scending order): "Or." "PARA-TYPE 3852.3" and my and the MCZC's type labels. The allolectotype is labeled: "W.T." "PARA TYPE 3852.2" and my and the MCZC's type labels. The paralectotype is labeled: "W.T." "LectoTYPE 3852". This lectotype label on the male designated paralectotype by me was added by a subsequent curator and is not a valid designation. The male selected as lectotype was chosen because it has one complete antenna, whereas the other male lacks antennae. Biology— Host Plants, Adult— * Radish, ♦tur- nip, sugar beet, and potato (Chittenden, 1927); the records for turnip and potato have been verified by me. New Collection Records: Artemisies [sic] sp., from Klamath Falls, Ore. (Algome) April 24, 1956 Joe Schuh, Coll. (AMNH:1); Solidago sp., from Creston,B.C. 2. VI. 1950 G. Stace Smith, C.A. Frost Collection 1962 (MCZC:1). Immature Stages — Unknown. Habits — Adults have been collected in the Pacific Northwest and Montana and Utah from late April until early August and again in late August and mid- October and in California in early July. No dates are associated with the other collection records. Distribution — Figure 196; P. decipiens has been collected mainly in the Pacific Northwest. The Texas record may be a case of mislabeling. Specimens Examined — Total 214, see Type Mate- rial above and the Appendix. Discussion — Nomenclature — Horn (1889) de- scribed P. decipiens from at least three specimens and noted that two of them had a preapical pale mark on each elytron, whereas the third was entirely black (it actually has a slightly paler, indistinct preapical mark). Chittenden (1927) named the color variation with the preapical pale marks P. decipiens var. ordi- nata. According to the Rules of Zoological Nomen- clature, article 45 (d)(i) and 45 (e)(i), this variety is to be treated as having subspecific rank. However, the aedeagi are the same, and the other characters all fall within the observed variation for this species. There- fore, I have synonymized this subspecies. The vast majority of P. decipiens specimens are immaculate. Therefore, they will be treated/included in Part II to be published later. Relationships — Phyllotreta decipiens is a rather distinctive species, except it is easily confused with those few specimens of P. striolata which have only a preapical pale elytral mark. Although P. decipiens often shows a tendency for serial elytral punctation, and the male 5th antennal segment of P. striolata is usually distinctly enlarged (about two times width of 6th segment), the most reliable way to separate these species is by their genitalia. For P. decipiens, the aedeagus has the lateral margin of the apical orifice extending less than half the length of the median lobe, whereas P. striolata has the lateral margin extending about three-fourths the length of the median lobe. The spermatheca of P. decipiens is longer and has the pump measuring less than one-half the receptacle length, whereas P. striolata has the pump greater than one-half the receptacle length. Phyllotreta denticornis Horn. Figures 19-22, 65-67, 122-124, 176, 197. Phyllotreta denticornis Horn, 1889. Trans. Amer. En- ' tomol. Soc, 16:297-298, fig. 19. Lectotype (here designated): Male, MCZC type #32850, at Museum of Comparative Zoology, Harvard. Type locality: California. Phyllotreta amphicornis Chittenden, 1927. Entomol. Amer., 8(n.s., no.l): 35-36. NEW SYNONYMY. Holotype: Male, USNM type #28802, at United States National Museum. Type locality: Wawawai, Washington. Phyllotreta aequalis Hatch, 1971. The beetles of the " Pacific Northwest. Part V:210. NEW SYNONYMY. Holotype: Male, at United States National Museum. Type locality: Bear Springs, Oregon. Diagnosis — Each elytron with the median pale stripe simple, of uniform width and without dilations, or reduced to a postbasal and/or preapical mark, or elytra entirely dark; male antennal segment 5 ex- panded bilaterally, dorsoventrally flattened, segment 6 with a sharp anteroventrally directed process. Description of Lectotype (species variation, ex- cluding punctation, in parentheses) — Figure 19; elon- gate oblong, length 2.65 mm (£2.45-2. 80, 92.55- 2.80), width 1.20 mm (£1.00-1.20, 91.20-1.28); head and pronotum black with slight metallic luster, elytra black with slight metallic luster (no luster), each with a somewhat indistinct, slightly paler median stripe. Head: Black; vertex finely granulate basally to slightly roughened anteriorly (entirely slightly rough- ened), moderately punctate with punctures separated by less than Ms to equal their diameter, mostly by less than Vi a diameter; interocular distance/maximum di- ameter of eye, 1.62 (£1.44-1.74, 91.56-1.75). Antennae: Figures 65-67; segment 4 longer than 3 or 7, shorter than 5, segment 5 more than 2 x length of 6; segment 5 bilaterally expanded, dorsoventrally flattened, venter concave in basal % ; segment 6 with a sharp anteroventral process , segments 7 and 8 with a short blunt apical process; see Table 2 for segment length/width, length 1.50 mm (9: length 1.51 mm; 24 FIELDIANA: ZOOLOGY simple); antennae various shades of brown, basal 2 (3) segments paler with dorsum of segment 1 dark, 3-6 (4-5 or none) darkest, 7- 1 1 (6- 11 or 4- 1 1) darker (9: basal 3 or 4 segments paler, 4-11 or 5-11 darker). Pronotum: Length 0.45 mm (6*0.42-0.48, 90.42-0.48), width 0.82 mm (60.75-0.85, 90.80- 0.82); black (rarely brownish black); texture finely granulate (finely granulate to slightly roughened, rarely finely alutaceous to finely granulate); coarsely punctate, punctures separated by less than xh their diameter. Elytra: Length 1.92 mm (6*1.75-2.02, 9 1.88-2.02), width 1.20mm (6 1.00- 1.20, 91.20- 1.28); black (dark brown, usually brownish black), median stripe indistinct, slightly paler than back- ground color as in Figure 19 (or usually reduced to a postbasal and/or preapical mark as in Figures 20-22, or elytra entirely dark); texture smooth (smooth to slightly roughened); moderately and coarsely punc- tate, punctures separated by less than % to equal their diameter, mostly by less than xh diameter. Legs: Nor- mal color sequence. Abdomen: 5th sternum with an apical median moderately concave lobe, concavity tapering and extending to midpoint (extension ending postbasally, oblong and extending to midpoint), with a median linear impression extending postbasally to apex (9 : simple in outline, usually with a shallow to moderately deep oval impression in apical xh ); black (usually brownish black) with concavity and apical margin paler (apical xh paler or entirely dark). Male — Aedeagus (figs. 122-124); length approxi- mately 0.93 mm; essentially straight except for basal piece slightly and apex strongly curved ventrad, of rather uniform width except apical XA slightly wider; apex in dorsal view very bluntly rounded with a shallow, moderately wide median emargination, in lateral view strongly and rather abruptly curved ven- trad with a small ventral median preapical lobe, of rather uniform width with end bluntly rounded; ven- tral view as in Figure 124. Female — Spermatheca (fig. 176); length approxi- mately 0.36 mm; receptacle elongate, slightly con- stricted medially on venter, gradually tapering to a slightly developed ring collar posteriorly. Type Material — The lectotype and paralectotype (6) are at the MCZC. The lectotype is mounted on a point and the pin bears 5 labels (descending order): "Cal" "LectoTYPE 3850" "P. denticornis Horn" and my and the MCZC's type labels. The paralec- totype pin bears 3 labels (descending order): "Cal" "PARA-TYPE 3850" and my type label. The cura- torially added lectotype 3850 label is not a valid designation. Biology — Host Plants, Adult — New Collection Records: Medicago sative [sic], from Gregory Sta., Ore. VI-3-1932 Jones-Gammon Collector (CDAE:1); ♦mustard, from Hornbrook, Cal. IV-27-1949 Col- lector H T Osborn (CDAE:3) and from San Miguel, Cal. 5 • 5 • 49 Keifer (USNM:1). Immature Stages — Unknown. Habits — Adults have been collected in California, Oregon, and Washington, from late April until early July. Specimens Examined — Total 48, see Type Mate- rial above and the Appendix. Distribution — Figure 197; P. denticornis is known only from California, Oregon, and Washington. Discussion — Nomenclature — Horn (1889) de- scribed P. denticornis from male specimens. In 1927, Chittenden described P. amphicornis , remarking that its antennae were similar to those of P. denticornis , but that the 6th antennal segment lacked the acute process; the acute process is present on the 6th anten- nal segment of his unique type, and the specimen is otherwise within the normal variation of P. denti- cornis. Hatch (1971) described P. aequalis from a striped specimen of P. denticornis and, not noting the acute process on the 6th antennal segment, placed it close to P. lepidula. The type of P. aequalis has a well-developed acute process on the 6th antennal seg- ment and is a normal, striped P. denticornis. There- fore, I have synonymized these two species with P. denticornis. Relationships — Phyllotreta denticornis is most similar to P. spatulata and less similar to P. arcuata, P. bisinuata, and P. lepidula, with which it shares having a simple elytral stripe and having the male 5th antennal segment dorsoventrally flattened. Phyl- lotreta denticornis and P. spatulata have very similar aedeagi, differing mainly in the overall shape in lat- eral view in which P. denticornis gradually and uni- formly tapers toward the preapical ventral bend, whereas P. spatulata has a moderate dorsal swelling at approximately the base of the apical orifice. In dorsal view of the apex, P. denticornis is shallowly emarginate, whereas P. spatulata is faintly truncate. Phyllotreta denticornis has a sharp process on the male 6th antennal segment which is unique. Phyllotreta dolichophalla Smith, NEW SPECIES. Figures 23, 68-69, 125-126, 177, 197. Holotype: Male, CASC type #14212, deposited at California Academy of Sciences. Type locality: Stinson Beach, Marin Co., California. Diagnosis — Each elytron with a median pale stripe with dilations basally somewhat and apicaily abruptly SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 25 incurved toward suture, but never meeting at suture; antennal segments 4-5 simple, male antennal seg- ments 6-11 darkest in contrast to paler basal seg- ments; aedeagus in lateral view with apex very narrow, directed ventrad. Description of Holotype (species variation, ex- cluding punctation, in parentheses) — Figure 23; ob- long, length 2.78 mm (6*2.66-3.16), width 1.30 mm (9 1.12-1.48); head and pronotum black with slight metallic luster, elytra brownish black, each with a median straw-yellow stripe. Head: Black; vertex finely granulate basally to slightly roughened anteriorly, finely to moderately punctate with punc- tures separated by less than lh to 1 .5 x their diameter, mostly by about 1 diameter; interocular distance/ maximum diameter of eye, 1.67 (91.37-1.67). An- tennae [left missing segments 4- 1 1]: Figures 68-69; segments 4-6 equal (subequal) in length, segment 7 longer than (equal to) 6; segments 4 and 5 simple, cylindrical; see Table 2 for segment length/width, length 1.68 mm (allotype: length 1.90 mm; simple); antennae various shades of brown, basal 5 (4) seg- ments paler, 6-7 (5-7) intermediate, 8-11 darkest. Pronotum: Length 0.58 mm (90.52-0.64), width 0.92 mm (90.85-1.04); black, texture slightly roughened to finely alutaceous (finely granulate to slightly roughened, smooth to slightly roughened); finely to moderately punctate, punctures separated by less than lh to 2 x their diameter, mostly by about 2 diameters. Elytra: Length 1.98 mm (9 1.88-2.31), width 1.30 mm (91.12-1.48); brownish black, me- dian stripe as in Figure 23; texture slightly roughened (smooth to slightly roughened); coarsely punctate, punctures separated by less than V2 to equal their diameter, mostly by less than V2 a diameter. Legs [left front leg missing tibia and tarsus, left middle leg and right hind leg missing tarsi]: Normal color sequence except tibiae and tarsi same color. Abdomen: 5th sternum with an apical median deeply concave lobe, concavity tapering and extending to within Vi of ster- nal base, with a median linear impression extending postbasally to base of median lobe, interrupted at base of concavity (9: simple in outline, with a median longitudinal impression in apical Vi to XA); black (brownish black with apical Vi of 5th sternum paler). Male — Aedeagus (figs. 125-126); length approxi- mately 1.31 mm; moderately arched dorsoventrally, of rather uniform width except gradually tapering toward apex in apical V3 in lateral view; apex in dorsal view acute with tip bluntly rounded, in lateral view very narrow, directed ventrad, ending in a sharp point. No ventral impression observed. Female — Spermatheca (fig. 177); length approxi- mately 0.42 mm; receptacle unique, basal Vz strongly swollen ventrally, apical Vi of rather uniform width, ring collar very wide, strongly developed. Type Material — The holotype and allotype are at the CASC. All the paratypes are female, with 1 at CASC, 2 at CDAE, and 1 at EHSC. The holotype is mounted on a point and the pin bears 5 labels (de- scending order): "Stenson's [sic] B Cal 4.11.33" "COL BY E J BLUM" "J.E. Blum Collection, gift of W.H. Nutting. Calif. Acad. Sci. Accession 1968" and my and the CASC's type labels. The allotype bears the same first 3 labels and then my and the CASC's type labels. The paratype at the CASC is labeled: "Alpine Dam Cal 9-33" and "E.S. Ross Collection". The 2 para- types at CDAE are labeled: one, "CAL. Santa Cruz Co. 2 m E Scotts Valley 3-IV-1972 W.H. Tyson", and the other, "Niles, Calif. Alameda Co. 14 • II • 1966" and "W.H. Tyson Collector". The paratype at EHSC is labeled: "Cal. Marin, Co. L. Lagunitas V- 10-59 D.C. Rencz". Biology — Host plants and immature stages are un- known. Habits — Adults have been collected in California from mid-February until mid-May. Distribution — Figure 197; P. dolichophalla has been collected only in California. Specimens Examined — Total 6, see Type Material above. Discussion — Relationships — Phyllotreta dolicho- phalla is most similar to P. attenuata and the speci- mens of P. conjuncta which have the postbasal and preapical marks connected; for a discussion of these relationships, see this section under P. attenuata. Phyllotreta dolichophalla might be confused with P. oregonensis on the basis of having the elytral stripe wide medially, but P. dolichophalla has antennal seg- ments 5 and 6 subequal in length, segment 5 simple and cylindrical, whereas P. oregonensis has antennal segment 5 about one and one-half times the length of segment 6, and in the male, segment 5 is dor- soventrally flattened. Etymology — The specific name dolichophalla comes from the Greek dolicho- meaning long and the Greek phallo meaning penis, which refers to the aedeagus being much longer than the average length for the genus. Phyllotreta emarginata Smith , NEW SPECIES . Fig- ures 24, 70-71, 127-129, 178, 198. Holotype: Male, deposited at American Museum of Natural History. Type locality: Mare's Egg Spring, Klamath Co., Oregon. 26 FIELDIANA: ZOOLOGY Diagnosis — Each elytron with a median pale stripe with dilations which never meet at suture, medially stripe width distinctly less than distance from stripe to suture; male antennal segment 5 dark, expanded bilat- erally and dorsoventrally flattened, subequal to 4 in width, with venter flat; apex of aedeagus in dorsal view with a narrow deep median emargination, straight ventrally in lateral view. Description of Holotype (species variation, ex- cluding punctation, in parentheses) — Figure 24; elon- gate oblong, length 2.75 mm (6*2.25-2.78, 92.50-2.80), width 1 .21 mm (60.95- 1 .32, 9 1.22- 1.50); head and pronotum black with slight metallic luster, elytra brownish black, each with a median straw-yellow stripe. Head: Black; vertex finely gran- ulate (finely granulate basally to slightly roughened anteriorly), moderately punctate with punctures sepa- rated by less than V2 to equal their diameter, mostly by less than 1 diameter; interocular distance/maximum diameter of eye, 1.39(61.30-1.50, 91.44-1.76). Antennae: Figures 70-7 1 ; segments 4 and 7 subequal (equal) in length, 6 about V2 length of 4, segment 5 about 2 x length of 4 and almost subequal to 4 in width; segments 4 and 5 expanded bilaterally, dor- soventrally flattened; segment 5 distinctly longest, with venter flat, not concave; see Table 2 for segment length/width, length 1.58 mm (allotype: length 1.40 mm; simple); antennae various shades of brown, basal 2 (9: 3) segments pale with dorsum of 1 dark, seg- ments 3 and 7-11 (sometimes 8-11; 9:4-11) dark, 4-6 (sometimes 4-7) darkest. Pronotum: Length 0.52 mm (60.40-0.52, 90.48), width 0.85 mm (60.72-0.92, 90.82-0.92); black; texture finely granulate; coarsely punctate with moderate punctures interspersed, punctures separated by less than V2 to equal their diameter, mostly by less than 1 diameter. Elytra: Length 1.90mm(6 1.42-2.02, 91.80-2.05), width 1.21 mm (60.95- 1.32, 9 1.22-1.50); brown- ish black, median stripe pattern as in Figure 24; tex- ture slightly roughened (smooth); coarsely punctate with moderate punctures interspersed, punctures sep- arated by less than V2 to equal their diameter, mostly by less than V2 a diameter. Legs [right metatarsus missing]: Normal color sequence. Abdomen: 5th sternum with an apical median lobe moderately con- cave, concavity tapering as it extends to within xh of sternal base, a median linear impression in middle V3 only (9: simple in outline, no median impression); black. Male — Aedeagus (figs. 127-129); length approxi- mately 0.90 mm; slightly sigmoid, of rather uniform width; apex in dorsal view abruptly triangular with tip broadly rounded each side of a median narrow mod- erately deep emargination, in lateral view almost straight ventrally, not directed ventrad, tip bluntly rounded; ventral view as in Figure 129. Female — Spermatheca (fig. 178); length approxi- mately 0.36 mm; receptacle elongate oblong with a slight dorsoventral arch, basal V2 slightly swollen, tapering to a slightly developed ring collar. Type Material — The holotype, allotype, and 9 paratypes (56, 49) are at the AMNH. One paratype (6) is at the CASC, 6 paratypes (36 , 3 9 ) are in my collection (EHSC), and 1 paratype (6) is at OSUC. The holotype is mounted on a point and the pin bears 3 labels (descending order): "Mare's Egg Spring, Ore.; Klamath County May 30, 1962 Joe Schuh, Collector" "Phyllotreta oregonensis 16067 Cr. M.H. Hatch '67" and my type label. The allotype bears the same first label and my type label. The paratypes at the AMNH are as follows: 26 and 1 9 , same as second pair cited for EHSC; 1 6 and 1 9 , same as third pair cited for EHSC; 1 6 and 1 9 , "Ore.; W. side of Upper Klamath Lk. July 30, 1969 Joe Schuh, Coll."; 16, "SpragueRiv., Ore. 5 mi. E. Bly June 16, 1957 Joe Schuh, Coll. "; and 1 9 , "Klamath Falls, Oregon Geary Ranch 6-21 1961 Joe Schuh, Collector". The paratype at the CASC is labeled: "Orinda, Cal Con Cos Co VI. 10. 1939" "B.E. White Collector," and "BURDETTE E. WHITE Collection Calif. Acad. Sci. Accession 1967". The 6 paratypes in my collection (EHSC) are labeled: 1 6 and 1 9 , same as holotype; 16 and 19, "Ore.; Klamath Co. Williamson Riv. Rch.; alfalfa & cruciferous weeds; 5-28-70 Joe Schuh, Coll. "; and 1 6 and 1 9 , "Crystal Cr., Ore. Upper Klamath Lk. May 30, 1960 Joe Schuh, Coll." The paratype at OSUC is labeled: "Chester Cal. VII- 18-56" and "D.J. & J.N. Knull Collrs.". Biology — Host Plants, Adult — Alfalfa and * cruciferous weeds, see Type Material above. Immature Stages — Unknown. Habits — Adults have been collected in California in mid-June and mid-July and in Oregon from late May until late July. Distribution — Figure 198; P. emarginata has been collected in California and Oregon. Specimens Examined — Total 19, see Type Mate- rial above. Discussion — Relationships — Phyllotreta emar- ginata is most similar to P. zimmermanni and is less similar to P. utana, P. utanula, P. oregonensis , and P. constricta, with which it shares elytral color pat- tern and having the male 5th antennal segment dor- soventrally flattened. However, the elytral stripe of P. oregonensis and P. constricta is usually much wider than that of the other 4 species. The male 5th antennal segment of P. zimmermanni has a basal con- SMITH: REVISION OF PHYLLOTRETA CHEVROLAT 27 cavity ventrally, whereas that of the other five species is evenly flattened ventrally; also antennal segments 2 through 5 are pale in P. utana and P. utanula, whereas in the other four species, antennal segments 2 and 3 are pale and/or dark and segments 5 through 1 1 are dark. Phyllotreta emarginata has the male 5th antennal segment subequal in width to the 4th, whereas P. zimmermanni has the 5th segment dis- tinctly wider than the 4th. The aedeagi of these six species separate into three distinct groups (based on dorsal view of apex) as follows: P. utana and P. utanula have the apex very broad and its margin mod- erately and broadly emarginate; P. constricta and P. oregonensis have the apex not broadened and its mar- gin entire; and P. emarginata and P. zimmermanni have the apex not broadened and its margin mod- erately but narrowly emarginate medially. However, P. emarginata has its apex in dorsal view abruptly triangular with its tip broadly rounded on each side of the median emargination and in lateral view almost straight, whereas P. zimmermanni has its apex in dor- sal view bluntly rounded with short premedian exten- sions and in lateral view the apex is directed ventrad. The spermatheca of P. emarginata has the receptacle slightly arched dorsally and tapering in the apical one- half to a slightly developed ring collar, whereas P. zimmermanni has its receptacle almost straight dorsally, of rather uniform width in apical one-half, and with a strongly developed ring collar. Also see this section under P. oregonensis and P. utana. Etymology — The specific name emarginata comes from the Latin emarginatus meaning notched at the apex, and refers to the apex of the aedeagus in dorsal view which has a narrow and moderately deep notch or emargination. Phyllotreta lepidula (LeConte). Figures 25, 72- 73, 130-132, 179, 198. Haltica lepidula LeConte, 1857. Report of insects col- lected on the survey. Washington, p. 68. Lectotype (here designated): Male, MCZC type #32851, at Museum of Comparative Zoology, Harvard. Type locality: San Jose or San Diego, California. Orchestris lepidula: Crotch, 1873. Proc. Acad. Nat. Sci. Philadelphia, 25:65-66. Phyllotreta lepidula: Horn, 1889. Trans. Amer. En- tomol. Soc, 19:294-295. Diagnosis — Each elytron with its median pale stripe simple, of uniform width and without dilations; male antennal segment 5 moderately expanded bilat- erally, dorsoventrally flattened, moderately concave ventrally; antennal segments 2-3 pale in contrast to darker 5-11; aedeagus in lateral view slightly sigmoid. 28 Description of Lectotype species variation, ex- cluding punctation, in parentheses) — Figure 25; elon- gate oval, length 2.60 mm (62. 60-2.75, 9 2.55- 2.72), width 1.22 mm (