THE GRASSHOPPERS OF CALIFORNIA (Orthoptera: Acridoidea) An endemic California grasshopper, Morsea californicus tamalpaisenais Rehn and Hebard, male. (Courtesy of E. S. Ross.) BULLETIN OF THE CALIFORNIA INSECT SURVEY # VOLUME 10 THE GRASSHOPPERS OF CALIFORNIA (Orthoptera: Acridoidea) BY H. F. STROHECKER (University of Miami, Coral Gables, Florida) WOODROW W. MIDDLEKAUFF (Department of Entomology and Parasitology, University of California, Berkeley) AND DAVID C. RENT2 (Department of Entomology and Parasitology, University of California, Berkeley) UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES 1968 BULLETIN OF THE CALIFORNIA INSECT SURVEY Advisory Editors: J. N. Belkin, R. M. Bohart, J. R. Douglas, R. L. Doutt, C. A. Fleschner, E. G. Linsley, E. I. Schlinger, R. L. Usinger VOLUME 10 Approved for publication March 10, 1967 Issued March 8, 1968 Price, $4.50 UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES CAMBRIDGE UNIVERSITY PRESS LONDON, ENGLAND 01968 BY THE REGENTS OF THE UNIVERSITY OF CALIFORNIA PRINTED BY OFFSET IN THE UNITED STATES OF AMERICA CONTENTS Introduction ............... 1 Bionomics ............... 2 Distribution .............. 4 Morphology .............. 6 Preservation of Specimens ........... 10 Acknowledgments ............. 10 Systematic Treatment ............ 12 Superfamily Acridoidea ........... 12 Key to California Families and Subfamilies ...... 12 Family Tetrigidae ............ 13 Family Eumastacidae ........... 15 Family Tanaoceridae ........... 19 Family Acrididae ............ 21 Subfamily Romaleinae .......... 22 Subfamily Cyrtacanthacridinae ........ 24 Subfamily Acridinae ........... 66 Subfamily Oedipodinae .......... 81 Literature Cited .............. 133 Index ................137 V THE GRASSHOPPERS OF CALIFORNIA (Orthoptera: Acridoidea) BY H. F. STROHECKER, WOODROW W. MIDDLEKAUFF AND DAVID C. RENT2 INTRODUCTION ALTHOUGHGRASSHOPPERS possess many attributes The last quarter of the nineteenth century brought which should make them sought after by entomolo- many papers from the pens of Samuel Scudder and gists, they have never attracted the large number of Cyrus Thoaas. These two workers were joined in collectors and students which have been drawn to the eighties by Lawrence Bruner but it was 1897 such other groups as the butterflies and beetles. before the first adequate treatments of even limited Knowledge of North American grasshoppers grew groups of North American grasshoppers were pub- very slowly from the time of Linnaeus to the last lished. Scudder‘s Revision of the Melanopli and decade of the eighteenth century, during which period Jerome McNeill’s Revision of the Truxalinae remain most descriptions of Nearctic species were from the fundamental works. pens of European entomologists. During the period 1825 to 1865, the American authors Thomas Say, Understandably the study of California grasshop- Thaddeus Harris, Samuel Haldeman, Asa Fitch and pers was far behind The first report based on Cali- P. R. Uhler published descriptions of a number of fornia acridids was the description by Stal in 1861 of species and several genera. Nevertheless, Scudder‘s two species. From 1876 into the twentieth century catalogue of 1868 listed fewer than seven presently Scudder, Thomas and Brunner described species and recognized species of Acridoidee from America north genera based on material collected by army and of the Rio Grande. geological survey expeditions. Saussure at Geneva and Bruner in Vienna received material from Ameri- In 1873 Cyrus Thomas essayed a synopsis of ca. Economic damage caused by grasshoppers had North American Acrididae, a work valuable ROW for attracted attention and several works on their de- it original descriptions, but otherwise more likely to structive activities were published in the last two confuse than to direct a student. A rapid perusal of decades of the nineteenth century. C. F. Baker and the section of this work devoted to the region north James Grinnell cdlected California Orthoptera early of the Rio Grande shows 145 specific names con- in this century, and a rudimentary account of Cali- sidered, of which quite a few are now regarded as fornia acridids may be extracted from Bruner’s synonyms. This work of Thomas, while dealing with volume of the BioZogia Centrali-Americana R&n and species, is not at all comparable to the CZassificafion Hebard published in 1909 the results of extensive of North American Coleopfera presented ten years collecting in California, and material taken by them later by Leconte and Horn. on subsequent trips has been reported in part in a number of papers. ‘These studies were financed in part by funds from Only recently, however, has a list of California Regional Research Project (W-371,a cooperetive project with Federal and Experiment Station workera participating in grasshoppers been published. The Zndex of the Arizona, Montana, Colorado, Idaho, Wyoming, and California Acn’doidea of California compiled by Thompson and 2 Bulletin of the California Znsecf Survey Buxton (1964) is a valuable guide to the systematic these insects usually matches closely the color of literature. the earth, in fact Faure (1932) demonstrated that some locusts reared on variously colored substrates changed color at molts to match the background. BIONOMICS Isely (1938a) reared grasshoppers of different colors by this technique and exposed them, on plots of dark The Orthoptera, while comprising only a fraction and light earth, to predation by chickens, a turkey, of the number of species to be found in such orders and native mockingbirds. His reports indicate definite as the Lepidoptera, Coleoptera and Diptera, must be survival value for background-matching coloration. judged as among the more successful groups of insects. Grasshoppers are the most ubiquitous, conspicuous, A stringent requirement for persistence of acridian and destructive group in the Orthoptera. populations is a suitable site for oviposition This site is generally moist but welldrained soil; loose or While mostly phytophagous, grasshoppers may on sandy soil seems to meet the requirements of the occasion consume a variety of organic matter. The greatest number of species. Some acridids regularly reported "cannibalism" is the eating of dead or molt- oviposit in cracked or decaying wuod and Criddle ing and helpless individuals rather than active preda- (1930) reported that partly dried cow dung, as well tion. A few instances of monophagy seem definitely as decaying wood, was used as a place of oviposition established, for example, acridids of the genus by the cow grasshopper, Neopodismopsis abdominalis Botitettix are confined to kea,but such a restricted Thos, in Manitoba. Such habits have not been re- food habit seems to be exceptional. corded for California acridids although Wilson and Keifer (1941) egg of Melanoplus All students of acridian ecology have recognized observed pods mar- the occurrence of particular species of grasshoppers ginatus in the crowns of alfalfa. According to Parker with certain plant associations but have mered as to (1957) human artifacts, for example, roadway the importance of specific feeding habits in restrict- shoulders and railway embankments, may increase ing the range of species of acridids Anderson (1964) the sites favorable for egg deposition. Edwards and Epp (1965) have recently presented the results of attributes to vegetative cover a definite influence on acridids both by its taxonomic composition and experiments on the effect of soil moisture and soil physical structure but points out, as did Isely pH on oviposition by Melanoplus SanguiNpes. Sand (19386) and others, that colonies of grasshoppers which is dry on the surface but moist at a depth of an inch was approximately as favored for egg deposition may be localized within large areas more or less covered by a favored food plant Gangwere ( 1961), as sand damp throughout; dry sand was used by few in summarizing extensive observations on feeding females A range of pH from 5 to 9 seemed without habits of a number of acridid species of eastern significance. North America, states that, The herbivores among Overlying local ecological conditions is the effect Orthoptera show definite constant preferences. They of temperature according to latitude and altitude usually specialize in a single category of foods, e.g, (Morse, 1889). In all ecological studies generaliza- grasses or forbs, but select widely within the category. tions should be tempered by recognition of the prob- . food probably plays only a minor part in the ability that a species of wide range exists as a series insects' habitat selection" (p. 134). of physiological "races" of different requirements or Isely (1944) tested the acceptability of various tolerance. Hastings and Pepper (1964) bring out this plants as food for some Texas Orthoptera and corre- problem in their study of populations of Aulocara lated mandibular structure with predominant con- in Montana sumption grasses or Acridinae generally tend of forbs. In preparing a site for oviposition the female grass- to grass feeding and Cyrtacanthacridinae to coflsump- hopper bends her abdomen into an arc to bring the tion of but mixed diets are common. forbs, oviposibor valves to a position perpendicular to the The band-winged locusts, long known as Oedipo- substrate. In this action the dorsal and ventral valves dinae, occur only in areas where there is considerable are brought together and thrust into the soil, then bare earth. These insects are seldom found on plants, spread apart. The scooplike