THE GALL MIDGES of CALIFORNIA Diptera: Itonididae (Cecidomyiidae)

BULLETIN OF THE CALIFORNIA INSECT SURVEY VOLUME 2, NO. 2

THE GALL MIDGES OF CALIFORNIA Diptera: Itonididae (Cecidomyiidae)

BY A. EARL PRITCHARD

(Division of Entomoloky and Parisitology. University of Califiirnin, Rrrkrley )

UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES 1953 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Editors: E. G. Linsfey, E. 0. Essig, S. B. Freeborn, R. L. Usinger

Volunie 2, No. 2, pp. 125-150, plate 40

Submitted by Editors, August 13, 1952

Issued February 27, 1953

Price, 35 cents

UNIVERSITY OF CALIFORNIA PRESS

BERKELEY AND LOS ANGELES

CALIFORNIA

CAMBRIDGE UNIVERSITY PRESS

LONDON, ENGLAND

PRINTED BY OFFSET IN THE UNITED STATES OF AMERICA The Gall Midges of California Diptera: Itonididae (Cecidomyiidae)

?he present summary of the gall midges known Lestrerniinae are distinguishable morphologically. from California is a preliminary account, since our The larvae of these midges are saprophagous or knowledge of these insects is scant. It is hoped, fungivorous, and their adult classification is on a however , that by bringing the California records sound taxonomic basis. Members of the itonidine of the group together, further biological investi- tribe Porricondylini have similar feeding habits, gations will be encouraged. but their classification is in a state of chaos. A list of the gall midge fauna of western ?he other itonidines feed for the most part on North America was published by Felt (1912) in which higher living plants. Their specific classification 28 species were recorded from California. Seventy- has been largely based on a knowledge of the host nine species are recorded at present, but at least and the type of gall formed. Ilowever, host speci- several hundred probably occur in the state. Those ficity cannot always be taken for granted, because species known from Oregon and Arizona are also in- Jensen (1946) recently showed that Contarinia cluded in this bulletin because they probably occur Zycopersici Felt may attack the flowers of seven in California. A number of species described from different genera of plants in five different fami- Utah, Colorado, New Mexico, and elsewhere may lies. A great deal of biological and taxonomic work occur also along the Pacific coast. is needed before the classification of the plant Most of our knowledge of gall midges in the feeders can be considered to be on a sound basis. far West is based on miscellaneous rearings or very The larvae of certain genera and species of small collections of adults. However, the early itonidines are predaceous on mites and insects. work of P. H. Timberlake in rearing the makers of Although the classification of these midges is a number of galls is worthy of special mention. largely based on morphological characters, it can It will be noted that members of the subfamily be regarded as only preliminary.

Key to Subfamilies Adults

1. Tarsus with proximal segment longer than second segment ...... Lestremiinae Tarsus with proximal segroent much shorter than second segment ...... Itonidinae

Larvae

1. Anus circular and located terminally on last segment ...... Lestremiinae Anus longitudinal and located on venter of last segment ...... Itonidinae

Subfamily Lestremiinae

The larvae of Lestremiine midges are saprophagous or fungivorous. They have been found in such habitats as the soil, decaying wood, leaf litter, manure, and mushrooms. Three tribes are known from the western United States. [1251 126 BULLETIN OF THE CALIFORNIA INSECT SURVEY Key to Tribes

1. wing with Vein MI+* forked ...... 2 Wing with vein simple ...... Micromyini 2. Wing with vein M3+4 arising from M; ocelli three ...... Catotrichini Wing with vein M3+4 free; ocelli tm or none ...... Lestremiini

Tribe Catotrichini This species was previously known from Wash- ington and Oregon (Pritchard, 1948). A single The tribe Catotrichini is the most general- female was collected at Oakland, February 4, 1952 ized of the Itonididae. It is based on a single (W. C. Eentinck). genu s . 'Ihe female, previously unknown, resembles the male closely. 'Ihere are 14 flagellar segments, each slender, with a slight distal neck, and irregularly clothed with short setae. Two sclerotized spermathecae are present. Only three species are known in the genus Catotricha. Two of these are North American, one occurring on the west coast. Tribe Lestremiini

cacott~cbusubobsoktu (Alexander) Nine genera are recognized in the tribe Les- tremiini, and most contain only several known Catocha subobsoleta Alexander, 1924, bsec. Inscit. species. However, many of the species are holarc- Mens., 12: 82. Qpe 6, Lmgmire Springs, Wash- tic or cosmopolitan in distribution, and further ington. collecting should increase the California list.

Key to Genera Males' 1. Antenna with 16 segments ...... 2 Antenna with 10 to 12 segplents ...... Anarete 2. Flagellum with tm crenulate whorls on each segment ...... 3 Flagellum with a single crenulate whorl on each segment ...... 4 3. Flagellum with digitate sensory processes on each proximal segutent ...... Anaretella Flagellum with sensory spines only on each proximal segment ...... Lestrenia 4. Wing with R, terminating beyond level of end of M ...... Gongroatast ix Wing with R, terminating just proximal to level op&d of G ...... Allarete

Femalesa 1. Flagellum with sensory spines only near the distal end of each segment ...... 2 Flagellua with digitate senwry processes distally on each segment ...... Anaretel la 2. Flagellum with sements parallel sided, each with the distal stem differentiated ...Les trenia Flagellmwith segments obovate, and distal stuns scarcely evident ...... s.. 3. Wing with medial fork parallel sided medially ...... 4 Wing with =dial fork gradually divergent from origin ...... Anare te 4. Ovipositor with penultimate segment of lamella twice as long as terminal segplent ...Parare t e Ovipositor with last two segpents of lanella sube-al in length ...... A1 1arete

'The male of Parorcte is unknown. befemale of Gongromcrrtix is unknown. THE GALL MIDGES OF CALIFORNIA 127 Genus Lestremia Macquart

'Ihree species of Lestremia are known in North America. Two of these are holarctic, but the third is known only from the eastern United States.

Key to Species Males

1. Distiforceps with two teeth distally ...... cinerea Distiforceps with a spine distally ...... Zeucophaea Females

1. Flagellum with stem of third segment broader than long ...... cinerea Flagellum with stem of third segment longer than broad ...... leucophaea

Lestremia cinerea Macquart bekann. eur. zweifl. Insekt., 1: 288. Type from Europe. Lestremia cinerea Macq., 1826, Insect. Dipt. Nord. The male genitalia are distinctive in that the Fr., pt. 1, Tipul., p. 173. Type from France. distiforceps bear a single spine distally, and the ?he distiforceps of the aedeagus are distinc- basiforceps have no proximoventral lobe. tive in that each bears two teeth terminally. L. leucophaea is of frequent occurrence in L. cinerea is common in Europe and North Europe and across the northern United States. America, the adults being found principally in the A single female has been collected in California: shade of trees and on windows. Pritchard (1951) Berkeley, February 15, 1952 (W. C. Bentinck). recorded collections from the San Francisco Bay region where the adults are found mostly during the fall and spring after rains. Genus Amwetella Enderlein New records of this species from California include: 16, 6 $9, Willits, February 5 and 25, 1952 (W. C. Bentinck); 18, Niles, February 7, 1952 (W. There are two widespread species of Anare- C. Bentinck); 2 $9, Drake's Beach,December 9, 1951 tella that are known from the Pacific coast. A (C. D. MacNeill); 19, Walnut Creek, November 27, third North American species is known only from 1951 (D. G. Denning). the state of Washington. Only the female of A. defecta is known for certain. Lestremia leucopbaw (Meigen) The male distiforceps taper distally in the two species listed here; they are broadly expanded Sciara Zeucophaea Meigen, 1818, Syst. Beschr. distally in the species known only from Washington.

Key to Species Males 1. Basiforceps with a setose enlargement inside near distal end ...... spiraeina Basiforceps without a setose enlargement ...... defecta Anaretelkz defecta (Winnertz) A. defecta is found frequently in Europe and Lestremia defecta Winnertz, 1870, Verh. 2001.-bot. North America. Pritchard (1951) recorded this spe- Ges. Wien, 20: 33. Type from Germany. cies from the San Francisco Lby area. 128 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Anaretelkz spirakna (Felt) Alkzrete barberi (Felt)

Catocha spiraeina Felt, 1907, Bull. N. Y. State Lestrernia barleri Felt, 1908, &11. N. Y. State Mus., 110: 102. Albany, Kew York. Mus., 124: 310. Type 9, Las Vegas, New Mexico. A. spiraezna is known from scattered locali- Females of this species were recorded from ties in North America and Europe. The first record Burney Falls, California (Pritchard, 1951). The of this species from California is from Oakland, male is unknown. February 24, 1952 (W. C. Bentinck). Genus Pararete Pritchard Genus GongromaJtix Enderlein The genus Pararete was based on a species that This genus may be recognized by having €t5 very is known only by the female. It would be very de- long, reaching the level of the end of M,. The fe- sirable to find a male for complete characteriza- male is unknown. tion of the genus. One species is known from the western states.

Pararete elongata (Felt) Gongromastix epista Pritchad Lestrernia elongata Felt, 1908, Bull. N. Y. State Congrornast ix epista Pritchard, 1951, Univ. Calif. Mus., 124: 310, Type 9, Argus Mountains, Publ. Ent., 8: 253. tiolotype male, American California. River , Washington. The single known specimen was collected in The occurrence of this species in California May, 1891. is based on a single male collected on the car window at Santa Cruz, November 18, 1947 (A. E. Pritchard) . Genus Anarete Haliday

Genus Allarete Pritchard Adults of the genus Anarete differ fran others of the tribe Lestremiini in that they may be col- Females belonging to the genus Allarete are lected in much drier habitats, and the males may sometimes collected at lights. Ihe only specimen form large swarms. Nothing is known of the larval that has been reared was from cow manure. One of habits. the two known species occurs in California, and it Nine species are known from North America. may be recognized by having the fork of the medial The males are very similar, and the females are vein only twice as long as the stem. even more difficult to determine.

Key to Species Males 1. Wings whitish ...... 2 Wings pale brownish or grayish ...... 3 2. Claw acuminate distally ...... johnsoni Claw with a denticulate expansion distally ...... busck i 3. Claw with a mall lateral tooth near tip ...... anepsia Claw simple ...... 4 4. Enpodium nuch longer than claws ...... rubra Empodium approximately as long as claws ...... corni

Anarete johnsoni (Felt) sylvan i a. A. johnsoni may be recognized by havink the Microcerata johnsoni Felt, 1908, hll. N. Y. State wings white, the claw of the male acuminate dis- h~.,124: 310. Tyne 8, Philadelphia, Penn- tally, and the distiforceps of the male without a THE GALL MIDGES OF CALIFORNIA 129 setose swelling on the inner side. wings, four palpal segments, and the male empo- This is a very commn species throughout the dium approximately as long as the claws. United States, and it also occurs in Europe. This species is known from across the north- Pritchard (1951) recorded collections from the ern United States and from Europe. It occurs in San Francisco Bay region. Oregon and Washington and should be found in northern California.

Anarete buscki (Felt) Anarete anepsia Pritchard Microcerata buscki Felt, 1915, Proc. U. S. Natl. Mus., 48: 198. Type 6, Raracoa, Chba. Anarete unepsia Pritchard, 1951, Univ. Calif. Publ. The distal, denticulate expansion of the male Ent., 8: 264. Type c3, Orinda, California. claw is distinctive. A. anepsia may be recognized by having a small This species has been recorded from Cuba, tooth near the end of the claw of the male. Puerto Rico, and Minnesota. The new record of two Two collections have been made of this species 66 from Molino Basin, Santa Rita Mts., Arizona, in California: Tilden Regional Park near Orinda, November 5, 1951 (C. D. MacNeill), at light, indi- and San Antonio Valley near Livermore. cates that the distribution is very wide, probably including California. Tribe Micromyini Anarete rubra Kieffer

Anarete rubra Kieffer, 1906, Ann. Soc. Sci. Brux- Adults of micromyine midges resemble those elles, 30: 33. Types 6 and 9, Germany. belonging to the Lestremiini in that they are A. rubra may be recognized by having the wings collected most commonly in shaded, damp local- brownish, the male empodium much longer than the ities, particularly after rains in the fall and claws, and the male flagellum with the distal seg- spring in California. Larvae may be found com- ments subglobular. monly in decaying plant material in these local- This species is known in California from ities, and a taxonomic study of the immature &scent City (Pritchard, 1951). stages is needed. Fifteen genera are recognized in the tribe Micromyini, all but two being known from North Anarete comi (Felt) America. Four additional genera have been proposed on larval characters alone. A number of Micromyia corni Felt, 1907, Bull. N. Y. State Mus., species have been collected that are undescribed, 110: 102. Type 6, Albany, New York. and the California fauna has been very inade- A. corni may be recognized by having brownish quately sampled and studied. Key to Genera Males 1. Wing with vein R1 at least three times as long as R, ...... 2 n Wing with vein R, not more than twice as long as Rs ...... 3 2. Flagellum with platelike sensory processes dstally on proximal segments .... Campylomyza Flagellum with sensory spines only at distal ends of proximal segments .... Cordylomyia3 3. Genital rod present ...... 4 Genital rod absent ...... 5 4. Distiforceps with a terminal spine or cusp ...... Tetraxyphus, Polyardis Distiforceps without a terminal spine ...... Bryoaryia3 5. Flagellum with 12 segments, the stems long ...... 6 Flagellum with 8 to 10 segments, the stems short ...... Mycophila 6. Costa ending well beyond Rs ...... Aprionus 3 Costa ending at or slightly beyond Rs ...... ~erornyia’

These genera have been collected in California, but the species are undescribed. 130 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Females

1. Wing with vein R, at least three times as long as R, ...... 2 Wing with vein R, not more than twice as long as R, ...... *. .3 2. Flagellum with a sensory collar distally on each segment ...... Campy I ony za flagellum with sensory spines only near distal end of each segment .....Cordy loayial 3. Costa reaching more than halfway of distance betmeen ends of €$ and M ...... 4 Costa ending abruptly at tip of Rs or reaching slightly beyond ...... - ~erornyia~ 4. Spermathecae two ...... 5 Spermatheca one ...... 7 5. Flagellum with two platelike sensory processes distally on each segment ..... Mycophila Flagellum with four or more platelike sensory processes distally on each segment or withawl-like or rodlike sensory processes ...... 6 6. Empodiumas longas claws ...... Polyardis Empodium rudimentary or not more than one-half as long as claws . . - . - - . Aprionus4 7. Flagellum with two slender or digitate sensory processes distally on each segment . . 6 Bryomyia4 Flagellum with four disclike sensory processes distally on each segment .... Te tr azyphus

Genus Campylomym Meigen

Adults of Campylomyza are often found on windows, as well as in agricultural areas. The males are known to swarm. Records, as well as observations in California, indicate that the larvae are found principally in the soil.

Key to Species Males

1. Tegpen with lamellate loops dorsodistally: distiforceps with inner, distal end some- what produced ...... fusca Tepn with an anteriorly directed pair of angulations medidistally; distiforceps without extensian of inner, distal end ...... flavipes

Females 1. Ovipositor with a small, geminate, median plate ventrally in ninth segment .....fusca Ovipositor without a median plate in ninth segment ...... flavipes

These genera have been collected in California, but the species are undescribed. THE GALL MIDGES OF CALIFORNI2 13 1

Campylomyza j?av;Pes Meigen Europe. Tlie first collection from California is from Santa Csuz, April 15. 1947 (A. E. Pritchard). Cmpylomyza f lavipes Meigen, 1818 , System Ekschr. kkann. Eur. Zweifl. Insekt., 1: 101. Type 9, from Europe. Genus Tetraxyphus Kieffer C. flavipes is found frequently in North America and Europe. The only California collection Te traxyphus has page priority over Xy lopr iona, is from Calistoga, April 13, 1947 (A. E. Pritchard). and the revised international rules of zocl ogical nomenclature make page priority mandatory. Camjylmyza fusca Winnertz 'Ihe genus is heterogeneous, and a review is needed in connection with Polyardis and Monardia. The males are all similar, but the females differ Canpylomyza fusca Winnertz, 1870, Verh. 2001. -bot. markedly. Two species that have been-assigned to ' Ges. Wien, 20: 12. Type 6, from Europe. the generic concept are known from the western C. fusca is common in both North America and states.

Key to Species

1. Palpus four segmented ...... ater Palpus three segmented ...... toxicodendr i

Tehmypbus der (Meigen) distinctive among the tribe because the distiforceps each bear a strong distal spine that is one- Canpylornyza ater Meigen, 1804, Klassif. 1, p. 40. half as long as the segment. Type 8, Germany. "his species appears to be widespread in North Edwards ( 1938) adopted Meigen' s name for this America and Europe. Several specimens were reared species, and he redescribed it dequately. The four from cow dung in Illinois. palpal segments together with the broad but taper- 'Ihe following collections are the first from ing sensory plates on the flagellum will separate California: 1 $, Berkeley, February 19, 1947 (A. E. atet from related species of Tetraxyphus. Pritchard), on window; and 18, 1 9, San Francisco, The first record from North America is 1 8, kember 27, 1948 (A. E. Pritchard), in greenhouse; Oakland, California, November 11, 1951 (W. C. 3 68, 5 9$, January 21, 1952 (J. h. MacSwain), Bentinck). reared from horse hg.

Tetraypbus toxicodendri (Felt), new combination Genus Polyczrdis Pritchard

Ccrrnpylonyta toxicodendri Felt, 1907, Bull. N. Y. 'Iwo species of the holarctic genus Polyardis State Mus. , 110: 98. Type 9, Albany, New York. are here recognized from the western United States. T. toricodendri differs from related species Several other species are known from eastern North by having only three palpal segments. The male is Punerica.

Key to Species

1. Palps with four segnents ...... nwnotheca Palps with three segrnents ...... kas loens is

Polyardis kasloetzsis (Felt) State Mus., 124: 314. Type $, Kaslo, British Columbia. Cainpylomyza kasloensis Felt, 1908, Bull. N. Y. P. kasloensis has been previously known from 132 BULLETIN OF THE CALIFORNIA INSECT SURVEY a single female from western Canada. The male is previously been known from North America only from distinctive in having 14 flagellar segments. The a female collected in British Columbia, Canada. tegmen of the aedeagus is also distinctive in that California records are as follows: 3 d'z. 2 $9, it is long and narrows strongly on the distal half Berkeley, February 14, March 4, and March 10, 1947 to form a sclerotized, angulate apex. (A. E. Pritchard), on window; 1 $? Lagunitas, April California records are as follows: 4 22, 27, 1947 (A. E. Pritchard); 1 9, Orinda, March 3, Berkeley, February 14, March 4, and April 4, 1947 1947 (A. E. Pritchard). (A. E. Pritchard) on window; 2 GS,Orifida, March 3, and April 6, 1947 (A. E. Pritchard); and 1 8, Wal- nut Creek, April 6, 1947 (A. E. Pritchard). Genus Mycophikz Felt

The genus Mycophila differs from most other Polyardis monotheca (Edwards) micromyine midges in that its members are paedo- genic, the mature larvae giving birth to larvae of Monardia monotheca Edwards, 1938, Proc. Roy. Ent. the next generation when conditions are favorable. SOC. Lond., 7(ser. B): 241. Type 9, from Four species are known of which two have been England. found in California. These two are distinctive Monotheca differs from all other species of in having the empoclium one-half as long as the Polyardis by having four palpal segments. It has claws.

Key to Species Females

1. Flagellun with seven segments, the seventh compound ...... fungicola Flagellum with nine segments, the ninth compound ...... barnesi

Mycophila barnesi Edwards fornia. This species, the genotype, is known from a Mycophila barnesi Edwards, 1938, Proc. Roy. Ent. male and female reared in September, 1897, from SOC.Lond. (ser. B), 7: 254. Type 8, Litch- mushrooms in Marin County. worth, England. M. barnesi has been known only from the type series from England, collected on windows and Subfamily Itonidinae reared from mushrooms and manure. %e occurrence of this species in California The subfamily Itonidinae is very large, con- is base6 on 16, 1 $! reared January 27-February 4, taining more than 400 genera that are still consid- 1947 (A. E. Pritchard), from larvae found on paper ered valid. The more generalized members are flowerpots in a nursery near Half Moon Bay. saprophytic, and certain of the more specialized species are funpivorous. Some speoies are predaceous or even parasitic on mites or insects. tlow- Mycophila fiingicola Felt ever, most of the species feed on higher, living plants and often produce characteristic galls. A Mycophila fungicola Felt, 1911, Jour. N. Y. Ent. few are inquilines in these galls or the galls of Soc., 19: 33. Lectotype 6, San Rafael, Cali- other insects.

Key to Tribes

1. Wing with R, absent ...... 2 Wing with Rs present ...... Porricondylini THE GALL MIDGES OF CALIFORNIA 133

n 2. Antenna with flagellar segments of male cylindrical, without long distal stems ...... $ Antenna with flagellar segments of male with long distal stems ...... 4 3. Wing with costa setose; vein Rs wll removed from costa and reaching distal end of wing ...... Asphondy 1 in i Wing with costa clothed with scales; vein R, approximate to costa and joining it before end of wing ...... L~s iopter in i 4. Flagellum of male with only a proximal node on each segment ...... 01 igotrophini Flagellum of male with each segment binodose ...... 5 5. Flagellum of male with nodes subequal, the distal node of each segmnt with a single circwnfilun ...... Con tar ini in i Flagellum of male with the nodes unequal, the distal node of each segment having two circwnfili ...... Itonidini

Tribe Porricond ylini the winter rains, but no species has been recorded 'Ihe larvae of species belonging to the tribe from the state. Only four species are recorded Porricondylini are saprophagous or fungivorous. from the West, two from Arizona and two from Adults are found in well-shaded, damp woodlands. Oregon (each being described from a single speci- A number of undetermined specimens have been men). Each species represents a separate genus that taken along the coastal area of California during may be distinguished by the following key.

Key to Genera

1. R8 appearing as a cross vein ...... 2 R, appearing continuous with R, ...... Porricondyla 2. Flagellum with circumfila; spermathecae absent ...... 3 Flagellum with horseshoe-shaped sensoria; spermathecae present ...... Winnertzia 3. Wings broad; present ...... Hormosomyia Wings long and narrow; M,, absent ...... Colpodia

Genus Witznertziu Rondani Genus Hormosomyiu Felt

Thirteen species of Winnertzia have been "he genus Hormosomyia is based on a single described from North America of which all but two species. were based on a single individual. No attempt has sexes, and it is impossible been made to associate Hmosomyia oregonensis Felt to make determinations from the literature. An unidentified species has been collected at Berkeley. The only published record from the West Hormosomyia oregonensis Felt, 1919, Ent. News, is from Arizona. 30: 220. Type 6, Forest Grove Oregon. This very interesting species is known from a single male from Oregon. It should occur in Winnertzia arizonensis Felt California.

Winnertzia arizonensis Felt, 1908, Bull. N. Y. Genus Co&odia Winnertz State Mus., 124: 421. Type 9, Williams, Arizona. Nineteen North American species have bean pro- This species is known only from the type posed in the genus Colpodia, each based on a single female. specimen. Accurate identification is impossible i.74 BULLETIN OF THE CALIFORNIA INSECT SURVEY

from the descriptions. A number of specimens of POr+icondyla barberi Felt colpodia have been taken near Berkeley. Porricondyla barberi Felt, 1908, Ball. N. Y. State Mus., 124: 418. Type 6, Williams, Arizona. Copodia colei Felt This species is known only from the type male.

Colpodia colei Felt, 1919, Ent. News, 30: 223. Type 6, Forest Grove, Oregon. Tribe Asphondylini ?he species is known from a single male. Asphondyline midges may be recognized by the long, threadlike antennae. The adults are often Genus Porricondyla Rondani large and heavy-bodied, and the long, needlelike, protrusible ovipositor is characteristic of Cali- A large number of species have been described fornia genera. in the genus Porricondyla, and a revision of the The larvae are plant feeders, breeding pri- group is needed before identifications can be made. marily in the buds or fruits of a wide variety of A single species has been recorded from the West. plants, and they produce monophthalmous galls.

Key to Genera

1. Palpus with 4 segments ...... Schizomyia Palpus with less than 4 segments ...... Asphordylia

Genus Schizomyia KieEer whether they have one, two, or three palpal segments. Otherwise, their identification is based In addition to having a fourth palpal segment, primarily on the host and the gall. Schtzornyra may he distinguished frcm Asphondyl ia Ly the characteristic hypopygium and ovipositor. Schizomyra is world-wide in distribution. Aspbondylia adenostema Felt Eleven species have been described from North her- ica, of which one is from the West. Several unde- Asphondylia adenostoma Felt, 1916, Jour. iv. Y. Ent. scribed species have been reared in California. SOC., 24: 177. Type Laurel Canyon near fiol lywood, California. lhis midge was reared in June, 1922, from an Schizomyia macrofika (Felt) apparently normal seed of Ademstoma fasciculaturn.

Asphondylia rnacrofila Felt, 1907. N. Spp. Cecid. Aspbondylia arizonensis Felt 11, p. 16. Types 6 and $2, Los Angeles and Alameda, Califorcia. Asphondylia arizonensrs Felt, 1907, I\r. Spp. Gxid. lhis species is apparently common on Am- 11, p. 13. Types 6 and $!, Fort Grant, Ari- srnckza lycopsoides along the California coast, zona. the palls causing distorted flower heads. Adults A, arizonensis was described as producing emerge in May, and it is a question where they fruitlike galls on prickly pear cactus, Opuntia sp. oviposit since Amsinck ra nearly disappears at this Later, Felt (1916) recorded adults as being reared time on the San Francisco peninsula. in March, 1886, from cactus seed pods at Los An- geles, California, and he also recorded the species from Colorado. Genus Asphondylia hew Aspbondylia artemisiae Felt

Asphondylia is a very large cosmopolitan ge- Asphondylia artemisiae Felt, 1908, hll. N. Y. nus. A nuder of species have been described from State hlus., 124: 377, Type 9, F'ort Grant, North America that may be grouped according to Arizona. THE GALL MIDGES OF CALIFORNIA 135

'his species was described from adults reared Fnt. Soc., 20: 151. Type 8, Puente Hills near from woolly bud galls on Artemisia SP. Felt (1916) Whittier, California stated that the types were reared in July, 1882, ?his species was reared in October, 1911, from from galls received from Ft. Grant, Arizona. a cabbagelike, woolly, bud gall on Diplacus longi- f lorus. It has not since been recognized. Asphondylia auripikz Felt Asphondylia dondiae Felt Asphondylia auripila Felt, 1907, N. Spp. Cecid, a, p. 14. Types d' and 8, Tucson, Arizona. Asphondylia dondiae Felt, 1918, Jour. Econ. Ent. , A. auripila was originally reared from an un- 11: 381. Type 6, Point Fermine, California. described type of gall on Larrea divaricata. Ihe This species is known from a single male Larrea of the hbjave Desert is considered by some reared in April, 1918, from a globose, sessile, botanists to differ from that in South America, and leaf deformation of sea blite, Dondia or Sue& they refer to it as L. tridentata. A study of the multiflora. Sueda multiflora is a Chilean plant, several pall midges on this host is needed. and the California host is probably Sueda californica pubescens. Asphondylia bea Felt Asphondylia enceliae Felt Asphondylia bea Felt, 1925, Pomona Jour. Eht. sol., 17: 19. Types 6 and $, Claremont, California. Asphondylia enceliae Felt, 1912, Jour. N. Y. Ent. A. bea produces inconspicuous flower galls in Soc., 20: 152. Type 9, Puente Hills near the floral head of Senecio douglasii. The types Wh i t t i e r , Ca 1i f o rni a. were reared in December and January. According to A. enceliae is known only from the original Needham (1925) there appears to be a continuous females reared in February, 1911, from bud galls on succession of broods throughout the year. Encelia ca lifornica.

Asphondylia brevicuuda Felt Asphondylia gurryae Felt

Asphondylia brevicauda Felt, 1907, N. Spp. Ckcid. Asphondylia garryae Felt, 1934, Pomona Jour. Ent. XI, p. 14. Type 8, Fort Yuma, Arizona. Zool., 26: 34. Types d' and 8, Mt. Konocti, A. brevicauda is known from a single female Lake County, California. collected at Fort Yuma, Arizona. Ihe adult re- The types were reared in May, 1933, from a sembles A. auripila and differs from all other bud gall on Garrya frewntii. members of the genus by having the palpus one segmented. The type was described as differing from Asphondylia iutegrayolke Felt auripila by being smaller, but it is probably a synonym of that species. Asphondylia intcgrifoliae Felt, 1908, Bull. N. Y. Asphondylia wuotbi Felt State Mus. 124: 376. Type $, Los Angeles County, California. Asphondylia ceanothi Felt, 1908, h11. N. Y. State 'he type was reared in April, 1886, from a hs., 124: 377. Type 9, Oakland, California. subglobular flower gall of Rhus integrifolia. Felt A. ceanothi was described from a female reared (1916) regarded as conspecific specimens that were in June, 1888, from a large, rather loose bud gall reared in June, 1914. from distorted flower buds on C'eanothus (identified as C. velutinus) from of Rhus trilobata, near Salt Lake City, Utah. Oakland. Felt subsequently (1934) identified as this species midges reared in July, 1933, from a flower bud gall on Ceanothus cuneatus, in Lake ' Asphondylia optuniue Felt County , Ca 1i f ornia . Asphondylia opuntiae Felt, 1908, all. N. Y. State Asphotidylia diphd Felt Mus., 124: 377. Type of unknown sex, Sinton, Texas. Asphondylia diplaci Felt, 1912. Jour. N. Y. State Felt (1940) indicated that A. opuntiae differs 136 BULLETIN OF THE CALIFORNIA INSECT SURVEY from A. arizonensis in that it breeds in swollen come enlarged and distorted, and they are sometimes leaves of Opuntia cactus rather than in swollen a localized pest in Arizona and New Mexico. fruit. Felt (1916) indicated that the type of opuntiae was from Texas, and he further recorded it Tribe Lasiopterini from Colorado, Arizona, and from Los Angeles, Cali- fornia (reared in April, 1908); an infested “lob&’ Members of the tribe Lasiopterini are stocky of the plant was illustrated. ?he adult of A. opwz- and densely covered with scales that often form tiae was regarded as being very similar to A. ari- a distinctive color pattern. Unfortunately, most zonensis, and it is probable that the two are of the scales ani theircoloration are lost in slide synonymous. preparation, and a part of each series reared should be pinned on minuten nacieln.‘ Larvae of the lasiopterine midges are plant Aspbondylia photiniae Pritchard feeders. Three genera have been recognized from the Asphondylia photiniae Pritchard, 1952, Pan-Pac. Pacific coast states, but it is probable that only Jbt., 28: 16. Holotype 6, Palo Alto, Califor- one of these is valid. The palpal characters used nia. by Felt to separate Lasioptera and Asteromyia are The type series was reared from distorted admitted by the describer of the latter genus to fruit of toyon, Photinia arbutifolia. Adults intergrade, although the gall-making habits of the emerge in April and May. two proposed genera have general differences. The venational characters that were defined to separate Neolasioptera and Lasioptera are not always Aspbondylia websteri Felt recognizable among the types of the species described and assigned to the two genera by Felt. Asphondylia websteri Felt, 1917, Jour. Econ. Fht., The present article, however, is not revisional, 10: 562. Types 8 and 9, Tempe, Arizona. and no new generic synonymy is proposed. It is Webster (1912) gave an account of this spe- merely pointed out that the distinctions of the cies. Larvae cause the seed pods of alfalfa to be- following key will probably not prove to be valid.

Key to Genera

1. Palpus with three or four segments ...... 2 Palpus with one segment ...... Asteronyia 2. Vein M3+4 indistinct or absent ...... Lasioptera Vein M3+4 present ...... Neolasioptera

Genus Lusioptera Meigen are of ten avail ahle for reared material . Six species have been described from Califor- Larvae of the genus Lasioptera feed character- nia, and five are known from Arizona. An unde- istically in the stems of herbaceous and woody scribed species was reared by E. S. lbss from pass plants, producing more or less nodular swellings. at Santa Cruz, and a numlier of specimens have been lhere is usually a single generation annually, the takeLi by sweeping near Berkeley. pupal stagp occurring in the gall and the adults emerginf in the spring or early summer. The geniis Lasioptera, including related genera Lusioptera arizonensis Felt of doubtful validity, contains more than 150 species described from North America. Their identification is not possible from the descriptions of Lasioptera arizonensis Felt, 1908, Bull. N. Y. their coloration and the few morphological charac- State Mus., 124: 325. Type 6,Oracle, Arizona. ters that have been considered. Ilowever, each spe- L. arizonensis was o’escribed from a male cies is believed to be host specific, and names reared in May, 1889, from the stems of Senecio ari- THE GALL MIDGES OF CALIFORNIA 137 zonensis. There is no plant named S. arizonensis, species of oak, and it is known only from the the reference probably being to S. arizonicus. types.

Lusioptera cussiae Felt Lusioptwa spinulae Felt

Lasioptera cassiae Felt, 1909, Jour. Econ. Ent. , Lasioptera spinulae Felt, 1908, b11. N. Y. State 2: 287. Cotypes 6 and 2, Fort Huachuca, Ari- Mus., 124: 325. Types 6 and 9, Fort Huachuca, zona. Arizona. This species is known only from the type L. spinulae is known only from the types series, reared in May, 1883, from stem galls of reared in June, 1883, from stem galls on an unknown the sensitive plant referred to as Cassia nic- host . titans. The host identification was erroneous (being an eastern species), and it probably represents Lusioptera tibialis Felt C. leptadenia.

Lusioptera diplaci Felt Lasioptera tibialis Felt, 1914, Insec. Insc. Menst., 2: 119. Types 6 and 2, La Jolla, Cali- forni a. Lasioptera diplaci Felt, 1912, Jour. N. Y. Ent. This species is known only from the types that Soc., 20: 151. Type 6,Whittier, California. were reared in April, 1914, from stem galls of L. diplaci is known only from the two type Pent s ternon ant i rrh ino ides. males, reared from stem gal 1s on Dip lacus Zongi- f lorus. Lasioptera umbetzae Felt

Lasioptwa howardi Felt Lasioptera uerbenae Felt, 1912, Jour. N. Y. Ent. SOC., 20: 150. Types 6 and 8, Whittier, Calif. Lasioptera howardi Felt, 1921, Bull. Brook. Fnt. ?his species is known only from the types Soc., 15: 94. Type 9, Mount Diablo, Cali- reared in November, 1910, from flower stem galls fornia. of Verbena prostrata, at Whittier, California. , L. howardi was described from specimens reared from “scrub” oak on Mt. Diablo and also from Martinez, California, in 1883. Sixty-eight years Genus Neolasioptera Felt later, there are no further records of this species. The genus Neolas ioptera was differentiated from Lasioptera on a basis of having M3+4.present and GJ~absent. The distinction is so difficult to Lasioptera lupini Felt appreciate that many of Felt’s types do not appear to correspond with the generic distinction, and Lasioptera lupini Felt, 1908, Bull. N. Y. State European workers have not accepted it for the Mus., 124: 326. Type $, California. paleearctic species. The only known specimen of this species was The only species from the western United reared from an undescribed gall on lupine. A stem States that were ascribed to this genus by Felt gall occurs on Lupinus arboreus in the San Fran- are the following: cisco Bay area, and it is probably the work of this species. Adults have not been reared, however. Neolasioptera hirsuta Felt

, Lasioptera qfierdJoriae Felt Neolasioptera hirsuta Felt, 1908, Bull. N. Y. Lasioptera querciflorae Felt, 1908, Bull. N. Y. State Mus., 124: 331. Type 9, Fort Huachuca, State Mus., 124: 325. Type 9, Fort Grant, Arizona. Arizona. N. hirsuta was described from a specimen This species was reared in June, 1882, from that was reared in May, 1883, from a stem gall on larvae feeding on the flowers of an undetermined an unknown host. It has not since been recognized. 138 BULLETIN OF THE CALIFORNlA INSECT SURVEY

Neolasioptera trimera Felt Asteromyii grindeliue (Felt)

Neolasioptera trimera Felt, 1911, Jour. Lon. Ent., Asteromyia grindeliae Felt, 1912, Jour. N. Y. ht. 4: 484. Type 2, Fort Grant, Arizona. Soc., 20: 149. 'lype 8, Santa Barbara, Cali- "he single female was reared from a stem gall fornia. on sun f 1ower . A. grindeliae was originally known from a male reared in July, 1911, from a blister leaf gall on Neolasioptera mimuli Felt Grindelia robusta at Santa Barbara. It was later (Felt, 1916) recorded on a basis of adults bred in October, 1915, from similar galls on Grindelia Neolasioptera mimuli Felt, 1908, Bull. N. Y. State cuneifolia at Millbrae. Variations were noted be- Mus., 124: 332. Type 9, Alamecia, California. tween add ts from the two collections. "his species is known from a single female reared from the stem of Miaulus aurantiacus in Tribe Oligotrophini November, 1885. Oligotrophine midges may be recopnized by having Rs absent and the flagellar segments of the male each with only a proximal node and a distal Genus Astwomyis Felt stem. 'Ihe group has been divided into two tribes by Felt and certain other workers on a basis of whether the claw is simple or else bears a proximo- Asteromyia is distinguished from Lasioptera by ventral tooth (the latter being the Dasineurini). having one or two (rarely three) palpal segments Certain genera, however, have a tooth only on the rather than three or four. The distinction is anterior legs, and its development may vary con- scarcely worthy of generic consideration. However, si derably. the larval habits of Asteromyia differ from those Except for the genus Coccidomyia, all the of Lasioptera in a general way inasmuch as they lsembers of the Oligotrophini feed on higher living usually form leaf blisters rather than stem galls. plants.

Key to Genera

1. Claws on all legs simple ...... * .2 Claws on leg I, and usually I1 and 111, with a proximal tooth ...... 5 2. Palpus fmr-segmented ...... 3 Palpus with one or two segments ...... 4 3. R, reaching tip of wing ...... Phytophaga R5 reaching costa well before tip of wing ...... Janetiella 4. Ovipositor slender; male with distiforceps plump ...... Rhopalornyia Ovipositor broadly triangular; male with distiforceps slender except proximally . . Walshornyia 5. Pal pus four-segmented ...... 6 Palpus two-segroented ...... a 6. R5 reaching tip of wing ...... Rhabdophaga R, reaching costa well before tip of wing ...... 7 7. Wing membrane with macrotrichiae slender ...... Uasineura Wing membrane with scales ...... Phaenolauthia 8. Antenna 18-segmented ...... Diarthonomyia Antenna 12-segmented ...... Cocc idorny i a

Genus Pbytopbagu Rondani alis Rondani of which the first is the type of the genus- (not cerealis which is generally regarded as "he genus Phytophaga originally included two a synonym of destructor Say). Salicina is regarded species, salicina (accredited to Degeer) and cere- by European workers as belonging to either the THE GALL MIDGES OF CALIFORNIA 139

genus Rhabdophaga or Dasineura. At any rate it ap- Genus Jaizetielh Kieffer pears that the American usage of Phytophaga is in error, and that the generic name Mayetiola should be employed for these species as it is for similar Janet iel la closely resembles Phytophuga except species in Europe. However, no generic change is that vein % unites with the anterior margin of the proposed in this review of the several species wing. A single species from western North herice known from California. was provisionally referred to this genus.

Pbytopbaga callfurnica (Felt) Janetielk-z siskiyou Felt

Mayetiola californica Felt, 1908, eUll. N. Y. State Janetiella siskiyou Felt. 1917, Jour. N. Y. Ent. Mus., 124: 370. Types d' and 2, Alameda County, SOC., 25: 194. Type $, 1 mile north of Siski- California. you National Forest, along south fork of Co- P. culifornica is known only from the type quille River, Oregon. specimens reared in February from an undescribed The seeds of Chamaecyparis lawsoniana were re- gall on currant, Ribes rnenziesii (Felt, 1916). ported to be deformed by larvae of this species, pupation occurring in cocoons on the sides of the seeds. One female was reared in January, 1917. Fbytopbaga destructor (Say)

Cecidomyia destructor Say, 1817, Jour. Phil. Acad, Genus Rhopalomyia Riibsaamen Nat. Sci., 1: 45. Types 6 and 9, northern and middle states. Rhopalomyia resembles Phytophugc except that The status of the Hessian fly in California only one or two palpal segments are present. The was summarized by Packard (1928). It has been known ovipositor is slender and extensile. from California since 1879, and it also occurs in Members of the genus feed on higher, living Oregon and Washington. This pest is only of local- plants, usually psoducing galls of the flower buds. ized importance in the far West. Ten species have been described that are from Cali- Wade (1934) presented an extensive, annotated fornia or probably so. bibliography of P. destructor, more than 1,200 There are a large number of species of Rhopa- references being included. lomyia described from North America, and their clzssification has been based primarily on the number of antennal segments, which is variable. Fbytophaga occidentalis Felt The number of palpal segments and the relative lengths of the node and the stem of the flagellar Phytophaga occ identat is Felt, 1926, Pomona Jour. segments are the only other morphological charac- Fht. Zool., 18: 79. Types 6 and 8, Berkeley, ters of importance that have received considera- Cali forni a. tion. The host relationships are of paramount The types of P. occidentalis were reared in importance for identification at the present. February, 1925, -from unenlarged branches of Sal ix SP- Rhopalomyia baccharis Felt

Rhopalomyia ba-charis Felt, 1908, hll. N. Y. State Mus., 124: 364. Types 6 and Q, from undis- Fbytopbaga tetradymia Felt closed locality. R. bacchar is was originally reared from stem galls of Baccharis pilularis, in December, 1885. Phytophaga tetradyaia Felt, 1925, Pomona Jour. Stem galls on this host are common on the pros- Ent. Zool., 17: 15. Types 6 and ?, Claremont, trate form of this plant in the sand dunes of the California. San Francisco peninsula. Types were reared in March, 1925, from a woolly, polythalmous bud gall on Tetradymiu Rhopalomyia bigelm*odes Felt comosa. Rhopalomyia bigelovioides Felt, 1908, Bull. N. Y. State Mus., 124: 366. Types CP and 9, Los 140 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Angel es, California. Eht. Soc., 24: 189. Type 8, Whittier, Cali- 'Ihe galls from which the types were reared fornia. in February, were reported to be deformed seeds R. erigerontis is known only from the speci- of Bigelouia sp., later identified (Felt, 1917) as men reared in April, 1913, from a gall on a plant (Bige lovia graueolens) = Chrysothamnus nauseosus. provisionally identified as Erigeron fragi lis.

Rhopalomyia califonaim Felt Rhopalomyh grindelhe Felt fihopalomyia californica Felt, 1908, Bull. N. Y. Rhopalomyia grindeliae Felt, 1916, Jour. N. Y. Ent. State Mus. , 124: 364. Types 6 and 9, Alameda Soc., 24: 186. Types 3 and $, Millbrae, Cali- County, California. fornia. Felt (1916) reported that the types had been R. grindeliae is known only from the types reared in February, 1904, from Baccharis sp.; also that were reared in October, 1915, from apparently from Alameda Co. , in September, presumably 1885, uninodified flower heads of tarweed, Grindelia cu- and in April, 1886 (from €lowers) of B. pilularis. ne if0 1 ia. Felt (1940) considered this species to form flower bud or leaf galls, single or clustered, on Bw- charis pilularis. 'Ihe galls are comnon in the San RbopalmyLa lonicera Felt Francisco Bay region. Rhopalomyia lonicera Felt, 1925, Pomona Jour. Ent. Zool., 17: 15. Type Claremont, California. Rhopalomyia muziana Felt 6, 'he type was reared in February, 1925, from an irregular bud gall on Lonicera subspicata. Rhopalomyia cruziam Felt, 1908, h11. N. Y. State Mus., 124 366. Type S, Santa Cruz Mountains, California. Rhopaloqva saluhe Felt This species is known only from specimens reared before May, 1890, from flower galls on Soli- Rhopalomyia saluiae Felt, 1916, Jour. N. Y. Ent. dago sp. SOC., 24: 184. Types 8 and q, Puente Hills near Whittier, California. H. saluiae forms conical leaf galls. "he types Rhopalomyia enceliae Felt were from Salvia californica and S. niuea, the adults being reared in May, 1912. Felt (1916) also Rhopalomyia enceliae Felt, 1916, Jour. N. Y. Ent. recorded rearings in January from Alpine, Califor- SOC., 24: 183. Type $?$, from unspecified lo- nia. Similar galls are comnon on Salvia mellifera, cality. on Mount Diablo. R. enceliae types were reared in April, 1913, from I- robose, thick-walled bud galls that some- Limes were confluent and distorting the stem. The Rhopalomyh truncata Felt host, Encelia sp., was mentioned by Felt (1940). Rhopalomyia truncata Felt, 1908, Bull. N. Y. State Mus. , 124: 365. Type a, bsAngeles, Califor- Rhopalomyia ericameriae Felt nia. A single male, collected at bs Angeles, Cali- Rhopalomyia ericameriae Felt, 1916, Jour. N. Y. fornia, was made the type of this species. Eht. SOC., 24: 190. Types 6 and 9, Puente Hills near Hhittier, California. R. ericanericae was originally reared in April, Genus Wahhomyia Felt 1913, from a small rosette gall on Ericameria palmeri. It has not since been recognized.

?he genus Walshomyia resembles Hhopalomyia Rhopalomyia mgerontis Felt except that the male and female genitalia are distinctive. Hhopalomyia erigerontis Felt, 1916, Jour. N. Y. A single species is known in the genus. THE GALL MIDGES OF CALIFORNIA 14 1

Walshomyia juniperina Felt The types of this species were reared in Feb- ruary, 1925, from nearly normal willow buds. H'aZshomyia juniperina Felt, 1908, Bull. N. Y. State Mus., 124: 360. Types 6 and 9, New Indria, Cali fomi a. Rbabdopbaga occidentalis Felt This unique species has been known only from types reared in June, 1884, from the fruit of Juni- Rhabdophaga occidentazis Felt, 1908, Bull. N. Y. perus californica. Several specimens were reared State MUS., 124: 353. Type 6, Santa Clara from Gavilan, Riverside County, in May, 1939 (P. H. County, California Timber 1ake 1. The single male of R. occidentalis was collected in June. Nothing is known of its biology. Genus Rhabdophaga Westwood Genus Dasineura Rondani Members of the genus Khabdophaga are characteristic of willow, particularly in the eastern Rondani' s original spelling Dasineura was United States. Their identification is very diffi- later emended to Dasyneura, and the latter spelling cult at the present, and the western species are is prevalent in most of the literature. The orig- mostly unidentified. inal name is here used because Rondani clearly Some species of Rhabdophaga occur on hosts wished it to be spelled that way and because the other than willow, and the genus resembles Phyto- emendation has not been submitted to the Interna- phaga except for having the claw unidentate. tional Comnission of Zoological Nomenclature. The classification of the North American spe- The genus contains a very large number of species of Rhabdophaga that was presented by Felt cies, most of which produce leaf galls. Some, (1915) is based on the number of antennal segments however, feed in the buds or seeds. lheir classifi- which is admittedly variable, the comparative cation in North America has been based primarily length of the flagellar stems of the male, and the on the number of antennal segments, which varies, termination of vein R ?he biological observations and the shape of the flagellar segments. Identi- 5. of the midges and their- individual hosts and galls fication must be based on reared material. are of more significance for identification at the present. Dasinewa califot.lica Felt

Rbabdopbaga cnlifmnica Felt Dasyneura californica Felt, 1908, hll. N. Y. State Mus. 124: 347. Type 9 from Alameda, Calif- Rhabdophaga californica Felt, 1908, Bull. N. Y. omia. State Mus., 124: 353. Type 6, Santa Clara D. californica was originally reared in Feb- County, California. ruary, 1886, from an undescribed type of bud gall R. californica is known from a single male on willow, Salix californica. This species has collected in June. Nothing is known of its host not since been recognized. or life history. Dasineura kgwninicokz (Linmer) Rbabdopbaga elymi Felt Cecidomyia leguninicola Lintner, 1879, Canad. Eht., Hhabdophaga elymi Felt, 1909, Jour. Econ. Ent., 2: 121. Described from larvae from New York. 2: 289. Type 9, Alameda, California. D. ZeguninicoZa was originally recognized as a R. elymi is known only from a single female serious pest of red clover throughout the northern reared in January, 1891, from (Elyw aneticanus) = part of the eastern United States and eastern Elymus glaucus. Canada. The larvae feed within the flowers and pre- vent the development of seeds. Mature larvae drop Rbabdopbaga essigi Felt to the ground and pupate in the soil. There are two or three generations annually. Barnes (1946) Rhabdophaga essigi Felt, 1926, Prmona Jour. Ent. has sumnarized the biological data on this midge. Zool., 18: 79. Types 6 and 9, Alviso, Cali- The clover-seed midge is recorded from British fornia. Columbia, Washington, and Oregon in the West, but 142 BULLETIN OF THE CALIFORNIA INSECT SURVEY it is not yet known from California. Diarthronomyia cbtysantbemi Ahlberg

Diarthronomyia chrysanthemi Ahlberg, 1939, Fnt. Dasineura lupini Felt Tijriskr., 60: 274. No types designated. The chrysanthemum gall midge was known for Dasyneura lupini Felt, 1916, Ent. News, 27: 413. many years as D. hypogaea LaW. Alberg showed that Types d' and 9, Buana Vista Park, San Fran- Felt erred in this identification. cisco, California. D. chrysanthemi makes small bullet-shaped D. lupini was originally reared in May from galls on thO leaves and stems of cultivated chrys- an irregularly fusiform, polythalmous, presumably anthemum. Essig (1916) studied their biology in stem gall on a large, perennial species of lupine. California. Greenhouse infestations in San Fran- A bud gall is common on Lupinus arboreus along the cisco and Los Angeles and outdoor infestations in San Francisco peninsula. the extensive comnercial fields of Sen Mateo and Santa Clara counties have been common. This midge was considered a very serious pest until chemi- Genus Pbuenokzutbiu Kieffer cals were recently developed that give excellent control. Edwards (1937) showed that Lasiopteryx as de- termined by Felt (not Lasiopteryx Stephens) is the same as Phaenolauthia. At least in the type of the Diartbronomyha f?occosa Felt enus, the anterior claws only are bifid. Members of this small genus are believed to be either gall makers or inquilines. A single species Diarthronomyia floccosa Felt, 1916. Jour. N. Y. is known from the West. Ent. Soc., 24: 195. Type 9, Kdge between Mill Valley and Muir Woods, Marin County, Cali- forni a. Pbaenokrutbk arizonensis (Felt), new combination The type female was reared in November, 1912, from large, woolly, lateral, bud galls on Artemisia Lasiopteryx arizonensis Felt, 1916, Bull,. N. Y. californica. Felt also noted that a similar gall State Mus. , 175: 195. Types 6 and 9, Prescott, was found at Santa Ehrbara, California. Arizona. This species is known only from adults reared in April, 1911, from Phylloxera galls on wild-grape Diartbronomyk occidentalis Felt leaves; and from adults collected in August, 1910. Diarthronomyia occidentalis Felt, 1912, Jour. N. Y. ht. Soc., 24: 194. Types d' and 9, Sweet Water Genus Diarthrotzomyia Felt Dam, San Eego County, California. Specimens were originally reared in Septem- ber, 1912, from small, oval, grayish galls on the Diarthronomyia contains several species of leaves of Artemisia heterophylla and also from gall-making midges of which one is of economic im- flower buds on this host taken in the Puente Hills portance. Four species have been recognized in near Whittier, California. Felt further considered California, their identification being based on the as being produced by this species bla.ckish galls on host. the underside of leaves of A. tridentata at Salt Lake City, Utah.

Dkrtbronomyha catifmica Felt Genus Coccidomyia Felt Coccidomyia contains tuo species, both North Diarthronomyia californica Felt, 1912, Pomcma Jour Ent. Zool., 4: 732. Types 6 and 9, Clarmont, American and both predaceous on members of the Coc- California. coidea. One species is known from the West. The types of D. californica were reared in March, 1912, from brownish- or reddish subconical Cocczidornyia Ctii Felt . galls on the underside of the narrow leaves of Arteaisia californica. Coccidomyia erii Felt, 1912, Jour. N. Y. Fht. Soc., THE GALL MIDGES OF CALIFORNIA 143

20: 147. Types 6 and 9, Casitas Pass road near (1950) showed that this group is worthy of tribal Carpenteria, California. recognition. The tribal name was originally pro- Adults were reared in November, 1911, from posed by Enderlein (1936). larvae, presumably predaceous on the mealybug (Eriurn lichtensioides) = Amnostheriua lichtensi- Members of the Contariniini feed on higher oides (Cockerell) , infesting Artemisia californica. living plants. fungi, mites, and insects. It is noteworthy that the genus Contarinia is not known Tribe Contariniini in the West. 'his genus contains such well-known pests as the sorghum midge, the grape-blossom On a basis of cytological studies, White midge, and the pear midge.

Key to Genera

1. Palpus four segmented ...... 2 Palpus three segnented ...... Zeuxidiplosis 2. Costa of wing with scales; pedicel of antenna with a prcmimodorsal tooth .....Endaphis Casta of wing setose; pedicel of antenna without tooth ...... "hecodiplosis

Genus ThecOrtipfosisKieffer Thecodiplosis uruschneriae Felt

The generic name Thecodiplosis was employed 'Zhecodiplosis zauschneriae Felt, 1912, Jour. N. Y. by Felt for those species allied to but not con- Ent. Soc., 20: 152. Types 6 and 9, Puente generic with Contarinia. Most, if not all, species Hills near Whittier, California. that he assigaed to this genus belong elsewhere. This species is known only from the types Their proper disposition can be made only in con- reared in December, 1910, from rosette galls on nection with revisional studies. Zauschneria cal ifornica. Two Western species have been provisionally assigned to this genus. Genus Endaphis Kieffer

Tbdiphis pini-radktae (Snow and Mills) 'Ihe type of the genus Endaphis is a European species that parasitizes aphids. An American Diplosis pini-radiatae Snow ad Mills, 1900, Ent. Endaphis 'is known from Arizona that was reared News, 11: 489. Tjps d and 9, Palo Alto, Cali- from eriophyid galls; and another species was fornia. reared from mite leaf galls in Peru. A fourth The larvae of T. pini-radiatae cause Monterey species was reared from dactyLopiid scales in pine needles to be stunted and swollen proximally, Ceylon. and the species is sometimes regarded as a serious pest. Pinus tuberculatus, P. wicata, P. sabini- ana, P. coulteri, and P. sylvestris are also re- Enhpbis americana Felt ported as hosts. 'he midge is common in the San Francisco Bay region. Endaphis avricma Felt. 1911, Ent. News, 22: 129. Snow and Mills noted that the adults emerge Type $, Prescott, Arizona. from about the middle of January until the first The only known female of this species was of March. Eggs are laid between the scales of ter- reared in September, 1910, from galls of (Eriophyes minal buds. The egg stage lasts about twu weeks. fraxinif lorae = Aceria fraxiniflorae on Fraxinus and the larva feeds in the needle for ten or ve lut ina. The species undoubtedly occurs in Cali- eleven mnths. Ihe pupal stage is formed, according fornia. to Snow and Mills, usually in a cocoon within the deformed buds, and it lasts three or four weeks; Genus Zeuxidiplosis Kieffer however, Compere (1915) found the pupal stage to occur in the soil. The genus Zeuxidiplosis contains a single spe- 14.4 BULLETIN OF THE CALIFORNIA INSECT SURVEY

cies that forms budlike leaf galls on certain mem- Tribe Itonidini bers of the genus Hypericum. It is native to Europe The tribe Itonidini is very large, and it is diverse in biological habits. Included species may Zeuxidiplosis gricrdi (Kieffer) be predators, fungus feeders, or gall makers. The binodulate flagellar segment, with the distal nod- ule long and bearing two circumfila, of the male is Diplosis giardi Kieffer, 1896, Bull. Soc. Ent. characteristic, but females are difficult to dis- France, p. 383. Types fram France. tinguish frinn the contariniines and oligotrophines. Specimens imported from Germany were reared Ihe male terminalia have been used more than else- in the insectary at Albany, California, and they where in the subfamily for the recognition of were released by J. K. Holloway in the spring of genera and species, and this sex is necessary for 1950, in northern California, as an aid in control ideriti fication. of Klamath weed, Hypericum perforatun. According to ‘Ihe genus Lestodiplosis, a very large and com- Holloway, in litt., the midges are established in man genus of predaceous midges, is unrecorded from the field. Cali fonia.

Key to Genera

1. Legs with all claws toothed ...... 2 Legs with posterior claws simple, at least ...... 5 2. Flagellum with distal node of each segment subdivided ...... -3 Flagellum with distal node of each segment entire ...... -4 3. Basiforceps each with a strong angulation proximally on inside ...... Youngomyia Basiforceps without angular projection ...... Cleodiplosis 4. Claw sharply curved at right angle ...... l’homasiniana Claw evenly and shallowly curved ...... Dicrodip losis 5. Claws of anterior legs toothed ...... 6 Claws of anterior legs simple ...... 7 6. Claw sharply curved at nearly right angle ...... Clinodiplosis Claw evenly and shallowly curved ...... Mycodiplosis 8. Wing with R5 reaching apex ...... 9 Wing with R5 uniting with costa well before tip of wing ...... Arthrocnodaz 9. Flagellum with several circumfilar loops of each segment much longer than others . Aphidoletes Flagellum with the circumfilar loops all similar in length ...... * .10 10. Palpus with four segments ...... ll Palpus with one segment ...... Monart hropalpus 11. Flagellum with nodes short and broad and bearing about 20 short circumfilar loops. . Ret inodiplosis Flagellum with nodes sJender and bearing canparatively few circumfilar loops . . . . Itonida

Genus Youngomyia Felt The only known adult was reared in June, 1886, from a subglobular gall on the underside of young Two members of the genus Youngomyia have been leaves of “Quercus pmila.’ The host was obviously reared from blossoms in the eastern United States. misidentified and probably refers to Quercus fraxi- lhe only species known from California produces nifolia. leaf galls. Genus CZeodiplosiS Felt

Youngomyz’u qumdnu Felt The genus Cleodiplosis has been known from a single species that is predaceous on whiteflies in Youngomyia quercina Felt, 1911, Jour. Econ. Ent., Panama. A second species is here transferred to the 4: 551. Type 6, Ibnner, California. genus. THE GALL MIDGES OF CALIFORNIA 145

Cleodiplosis koebelei (Felt), new combination galls are common in Marin County and San Mateo County, California, the midges obviously having a Silvestrina koebelei Felt, 1932, Pan-Pac. Ent., single genera tion annually . 8: 167. Types 6 and 9, Sydney, New South Wales (at Riverside, California). Koebelei cannot belong to the genus Silues- Genus Clinodiplosis Kieffer trim, because a cotype studied in the U. S. Na- tional museum has the claws strongly bent and each Most of the species that have been assigned with a long proximal tooth. to Clinodiplosis have been regarded as inquilines The larvae of C. koebelei were found feeding in the galls of itonidids, cynipids, or caterpil- on the citrophilus mealybug, Pseudococcus gahani lars. Larvae of the only species known from the Green. According to Harry S. Smith (in Felt, 1932) West are found in galls formed by a rust. this species was introduced and became established in California. Specimens were obtained from Stanley E. Flanders, collected in Riverside, November, Clindiplosis pucciniae Pritchard 1947, feeding on Pseudococcus adonidurn (Linnaeus). C1 imd ip 10s is pucciniae Pritchard, 1948, Pan-Pac . Ent., 24: 29. Holotype 3, Palo Alto, Califor- Genus DiM.odipZosis Kieffer nia. This species was reared in March, 1947, from The European genus Dicrodip losis, known only larvae feeding on Puccinia rust on Baccharis pilu- from the female, has been employed by Felt for a laris. small number of North American species that are allied to Thomsiniana. Felt?s generic concept is heterogeneous, inasmuch as he included both gall Genus Mycodiplosis Riibsaamen makers and predators. One of the species is known from western North America. The genus Mycodiplosis, as utilized by Felt, is heterogeneous. The type of the genus feeds on fungus; but the only species from California that Dicrodiplosis calfmica Felt has been referred to the genus feeds on spider mites. However, Felt?s generic reference must be Dicrodiplosis californica Felt, 1912, Jour. .N. Y. retained until revisional studies are undertaken. Ent. Soc., 20: 244. Types CP and 2, Riverside, California. Adults of this species were reared in 1893, Mycdiplosis acarivora (Felt) from Pseudococcus mealybugs on So lanua. Cecidoryia acarivora Felt, 1907, ht. News, 18: 242. Types 6 and 9, aula Vista, California. Genus Tbomasiniana Strand M. acarivora was reared from larvae found feeding on tbe spider mites (Tetranychus mytilos- Thomasiniana was proposed as a new name for pidus, misidentified) = Iletatetranychus citri the preoccupied Thorosia that was used for many (Md;regor), and Tetranychus sexurculatus Riley on years as a genus of gall midges. lemon. The type of the genus of Thomasiniana is a European pest of roses and related fruit trees. l?he species described from California was referred Genus Arthromodax Riibsaamen to this genus very provisionally. Arthrocnodax contains a number of species whose larvae are predaceous on spider mites, erio- Tbonrashiiaiia cnlifwica (Felt), new combination phyids, and probably other mites. Only two species are known from California. Thomasia californica Felt, 1914, Jour. N. Y. Ent. Soc., 22 132. Type 2, Lake Tahoe, California. ?he species is known only by a female reared Artbromdax occidentalis Pelt in August, 1913, from larvae producing midrib galls on the leaflets of Synphricarpos. Similar Arthrocnodox occidentalis Felt, 1912, Jour. Econ. 146 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Ent., 5: 402. Type 8, Berkeley, California. were assigned to the familiar generic names Cecido- This species is known from a single specimen myia and Diplosis that are now considered synonyms reared in July, 1912, from larvae preying on a of Itonida. However, Itonida is still rather heter- spider mite, Tetranychus sp. ogeneous, and it includes those midges that do not fall into better defined genera. Two species have been described from the West. Arthrmodax apiphila Felt

Arthrocnodnx apiphila Felt, 1907, N. spp. Cecid. Ztonida hopkinsi Felt 11, p. 20. Type 6, California Adults of this species were reared from de- Itonida hopkinsi Felt, 1911, Jour. Econ. Ent., bris and excrement in the bottom of a beehive, 4: 554. Type 9, San Francisco, California. and from brood comb affected by American foul This species is known from a single female brood from Tulare County. The comb was decaying that was reared from scolytid burrows in Pinus and heavily infested with fungus mites (Acaridae). attenuata. The generic reference of the midge is Felt (1921) also considered as A. apiphila obviousI y prov isiona 1. adult midges reared from a twig infested with Pulvinaria and a breeding jar containing the forest tent caterpillar, at Tacoma, Washington. Ztonida citrulli Felt Others were reared from a jar containing Viburnm leaves infested with an eriophyd. Itonida citrulli Felt, 1935, Bull. Brook, Ent. Soc., 30: 79. Types 6 and 9. Tuscon, Arizona. I. citrulli, an Arizona species that is not Genus Retimdzjdosis Kicffcr yet known from California, is regarded as an im- portant localized pest of watermelon. The larvae Members of the genus Retinodiplosis breed infest the tips of the vines causing them to characteristically in the resinous exudate of curl and die. pine trees in North America. Several species have been described from the eastern states and one from California. Genus M onarthropaZpus Riibsaamen

The genus Monarthropalpus includes a single Retinodiplosis resinicoloides (Williams) species. Monarthropa&us hi(Laboulb'ene) Cecidonyia res inicoloides Williams, 1919, Ent. News, 20: 2. Types 6 and $, Palo Alto, Cali- Itonida buxi Laboulb&ne, 1873, Ann. SOC. Ent. fornia. France (ser 51, 3: 321. Types 6 and 9, Paris, Adults were observed by F. X. Williams in France. March and April , 1907, there being a single genera- The larvae of Y. buxi make blisterlike leaf tion annually. ?he larvae live gregariously in galls on the leaves of ornamental boxwood, Buxus masses of pitch on Monterey pine. Rpation occurs sempervirens, and the pupae are formed in the in a cocoon that protrudes from the exudate. galls. "here is a single generation annually, the adults emerging in April in the San Joaquin Valley (Harry S. Smith, 1915). An infestation was found Genus Ztonida Meigcn by the writer in 1949, in a nursery at Hayward, California. This midge is sometimes a serious Before 1900, a large number of the gall midges pest, but it is seldom encountered in California. THE GALL MIDGES OF CALIFORNIA 147

Host List of Phytophagous Gall Midges

Host Larval habitat Midge PMACEAE Pinus radiata pitch exudate Retinodiplosis resinicoloides Pinus spp. needle swelling Thecodip losis pini-radiatae CUPRESSACE AE Chaecyparis I wsoniana fruit Janet ie 11a siskiyou Juniperus cal ifornica fruit Walshomyia juniperina GRMEAE Elyw glaucus undescribed Rhabdophaga elymi Wheat, barley, rye stem Phytophaga destructor SALICACE AE Sal ix californica bud gall Dasineura californica Salix sp. nearly normal bud Rhabdophaga essigi FAGACEAE

Qaercus frazinifolia globul~~leaf gall Youngomyia quercina Qwrcus sp. flowers Lasioptera qusrcif loroe Qercus sp. undescribed Lasioptera homardi CHENOPODIACEAE Sueda californica globular leaf gall Asphondylia dondiae SAXIFRAGACEAE Ribes nenziesii undescribed Phytophaga cal ifornica ROSACEAE Adems toma f asc iculaturn seed Asphondylia odenostom Photinia arbut if0 lia fruit Arphondylia pkotinw L EGUMINOSAE Cassia leptadenia stem gall hsioptcra casies Lupinus sp. stem gall Larioptera iupini Lupinus spp. probably hd gall Dasineura lupini Trifolium pratense flower head Dasineura leguainicota hfedicago sativa distorted seed pods Asphondyl ia wbsteri ZYGQPHYLLACEAE Lwrea tr identata undescribed Asphondylia awipila BUXACEAE Buxus sempervirens blister leaf gall Mmar t hropulpus buxi ANACARDIACEAE Rhus integrifoLia flower gall Asphondyl ia integr ifo lia RHAMNACEAE Ceanothus SFP bud gall Asphondyl ia ceunothi HYPERICACEAE

Hypericum spp. budlike leaf gall Zcuxid ip 10s is g iord i 148 BULLETIN OF THE CALIFORNIA INSECT SURVEY Host Larval habitat Midge CACTACEAE Opuntia spp. fruit Asphondylia arixonensis Opuntia spp. leaves Asphondylia opuntiae ON AGRACEAE Zauschneria californica rosette gall necodip los is xauschner iae CUCURBITACEAE Cucumis me10 de f orned ti ps Itonida citrulli GARRYACEAE Carrya fremontii bud gall Asphondylia garryae BORAGIN ACE AE Ams inck ia 1ycopso ides flower gall Schizomy ia macro f i 1 a VERBENACEAE Verbena prostrata stem gall Lasioptera verbenae LAEIIATAE Salvia spp. conical leaf gall Rhopalomyia salviae SCROPHULARI ACE AE Pents tenon ant irrhino ide s stem gall Lasioptera tibialis Diplacus longif lorus woolly bud gall Asphondy 1 ia dip 1 ac i Diplacus longif lorus f 1over he ad Lasioptera diplaci stem Mimulus aurantiacus Neolasioptera mimuli CAPRI FOLIACEAE Lonicera subspicata bud gall Rhopalomyia lonicerae Symphoricarpos sp. midrib leaf gall Thomasiniana californica COMPOSITAE Gr inde 1 ia cuneifolia flower head Rhopa lomyia grinde 1 iae Grindelia robusta blister leaf gall Asteromyia gr inde 1 iae Ericaneria palseri rosette gall Rhopa lomy ia er icamer iae Wrysothamnus nauseosus deformed seed Rhopalomyia bige louioides Solidago sp. flower gall Rhop a 1omyi a c ruz iana Erigeron fragi lis undescribed Rhopalomyia erigerontis Baccharis pilularis bud or leaf gall Rhopalomyia cal ifornica Baccharis pilularis stem gall Rhopalomyia baccharis Encelia sp. bud gall Phopalonyia enceliae Chrysanthemum spp. conical leaf gall Diarthronomyia chrysanthemi Ar temisia ca lifornica conical leaf gall Diarthrononyia cal ifornica Artemisia ca1 ifornica woolly bud gall Diarthrononyia f loccosa Artemisia spp. oval leaf gall Diartkonomyia occidentalis Artemisia sp. woolly bud gall Asphondy 1 ia ar teais i ae Senecio arizonicus stem gall Lasioptera ar izonens is Senecio douglasii flower head Asphondyl ia bea Tetrtxlynia comsa bud gall Phytophaga tetradymia THE GALL MIDGES OF CALIFORNIA 149

Host List of Zoiiphagous Gall Midges Family of apparent host Apparent host Midge ACARINA Tetranychidae Metatetranychus citri Mycodip 10s is acar ivora Tetranychus sexraculatus @codiplosis acarivora Tetranychus sp. Arthrocdax occidental is Eriophyid- Aceria fraxiniflorae Endqhis amer icana Acaridae undetermined Arthrocnodax apiphila INSECTA Pseudococc idae Pseudococcus spp. Dicrodiplos is cal ifornica Pseudococcus spp. Cleodiplosis koebelei Amnostherium 1 ichtensioides Cocc idomyia er ii

Literature Cited

Barnes. H. F. Comstock Publishing Co. , Ithaca, New 1946. Gall midges of economic importance, York. 2. 160 pp. Crosby Lockwood and Son, Jensen, D. D. London. 1946. 'he identity and host plants of blos- Compere, Harold sm midge in Henaii (Diptera: hi&- 1915. Insect Notes. Mon. Bull. Calif. State myiidae: Contarinia). Proc. Hawaiian Hort. Cam., 4:574. ht. Soc., 12(3):525-534. Edwards, F. VI. Needham, James G. 1937. New records of British Cecidomyi- 1925. Observations on a flower gall of the idae (Diptera), with taxonomic notes chaparral. Pomona Jour. Ent. Zool., on certain genera. Ent. Mon. Mag. , 17: 17-20. lxxiii: 145-154. Packard, C. M. Enderlein, Gkther 1928. 'Ihe Hessian fly in California. U. S. 1936. Die Tiemelt Mitteleuropas, Band 6, Dept. Agric., Tech. Bull. 81:l-25. Lief 2, Insekten, Teil 3, pp. 59-75. Pritchard, A. Earl Essig, E. 0. 1951. Ihe North American gall midges of the 1916. The chrysrnthemnm gall-fly, Diarth- tribe Lestremiini; Ittmididae (Ceci- ronoryia hypogaea (F. Lb). Jour. domyiidae) ; Diptera. Univ. Calif, Econ. Ent., 9:461468. Publ. Ent., 8(6): 239-275. Felt, E. P. Smith, Harry S. 1912. 'Ihe gall midge fauna of western North 1915. The occurrence of the Furopean box- America. Pomona Coll. Jour. Ent., wood leaf miner in California. Mon. 4(3 ):753-757. h11. Calif. Hort. Com., 4:340-344. 1915. A study of gall midges 11. hll. N. Y. Wade, J. S. State Mus. 589:79-243. 1934. An annotated bibliography of the Hes- 1916. New western gall midges. Jour. N. Y. sian fly, Phytophaga destructor (Say), Eht. Soc., 24:175-1%. U. S. Dept. Agric., Misc. Publ., 1917. Key to American insect galls. Bull. 198: 1-100. N. Y. State Mus., 200:s-310. Webster, F. M. 1912. The alfalfa gall midge. U. S. Dept. 1921. A study of gall midges, VII. Bull. ofAgric., Bureau Fnt. Cir. 147. N.. Y. State Mus. , 231-232:81-280. White, M. J. D. 1934. Two western species of Asphondylia. 1950. Cytological studies on gall midges Pomona Jour. Ent. Zool,, 26:34. (Cecidomyidae). Univ. Texas Publ., 1940. Plant galls and gall makers, 364 pp. 5007: 1-80 150 BULLFTIN OF THE CALIFORNIA INSECT SURVEY

---.DlSTlFORCEPS

----..I0 th T E R 6 ITE ---IOth STERNITE ----e E W ITA L R 0 0

-----T E 8 Y E N

Explanation of Plate 40 a. Wing venation of a porricondyline midge. b. Male genitalia of a lestremiine midge.