MEMOIRS OF THE AMERICAN ENTOMOLOGICAL SOCIETY NUMBER 48 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) By RAYMOND J. GAGNE PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY AT THE ACADEMY OF NATURAL SCIENCES PHILADELPHIA 2008 The American Entomological Society Philadelphia Founded 1859 Incorporated 1862 PUBLICATIONS COMMITTEE Daniel Otte Jorge Santiago-Blay Norman E. Woodley The offices, library, and meeting rooms of the Society are at The Academy of Natural Sciences of Philadelphia PUBLICATIONS OF THE SOCIETY TRANSACTIONS OF THE AMERICAN ENTOMOLOGICAL Society are published in annual volumes of quarterly numbers. Subscription prices: United States $30.00 per yearly volume (four issues); foreign countries $34.00 per volume; U.S. and foreign members of the society $15.00 per volume. Entomotocicat News is published bimonthly except July-August. Subscription prices: $30.00 per annual volume; foreign countries $34.00 per volume; U.S. and foreign mem- bers of the society $15.00 per volume. Memoirs oF THE AMERICAN EnTomotocicat Society are published irregularly when a suit- able monograph is submitted and accepted for publication. Prices vary with size; see back cover for list of available numbers. Please address all inquiries relative to publications and other society matters to: The American Entomological Society The Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA 19103 USA e-mail: AES@acnatsci.org Please visit the society’s internet website at: http: //www.acnatsci.org /hosted /aes / MEMOIRS OF THE AMERICAN ENTOMOLOGICAL SOCIETY NUMBER 48 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) By RAYMOND J. GAGNE S << Ny igg ny, “ny, ' PLAT TMNNN ANNE LTT PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY AT THE ACADEMY OF NATURAL SCIENCES PHILADELPHIA 2008 Norman E. Woodley Editor Issued 15 September 2008 Composition by Anne M. Landgraf Brooklyn BookWorks Brooklyn, Michigan PRINTED IN THE UNITED STATES OF AMERICA by Sheridan Books Chelsea, Michigan TABLE OF CONTENTS MrtLOG CHOI Creare rte el ceacte eveiteteteste cuatro -vovevens crstlate mucuaeleue ceatereyeieue, oeueteuererers ausreamichs 2 pes ET CKOLICS Me see RNR erica tee RR aCe ere Rete Re te rei vel arene Felt anes eke escent astateweroceiel 3 MaterialsiandiMethodSinrarrscgrcte si ictersicr Weir oiee eicteics sisisuecis races shoe sors eeceg sais 5 GenusiGaryemyiggheltee re rte Te ree ee er ey ick era eos cee toe elaerer 7 Generic eserip thom cyssreen te cpserscre stax reyevohel romeesreliens ssh steteveite susyaves ysis ti cceuehols situcee tees 8 ING SIgHLELES), oro aa ain GRO OS oG GR Mere Ob. Ae Mohd ig CATE EB G Souler: © iter bedi Aart auc eI eRe rar oats 9 Intrageneric Morphology and Relationships ................... 0:5 e eee e eee eee 11 [MO Ofny Goovcop dap daoHops nang ben ooddagood ORD ODOU OU GDR OOnR aOR aD adece coer 14 RIOSte Specificity per verses ures tapers er Gentine ees castes gtecespersrsahars: ave niciekoners a emir 16 Distrib utionte terse: cies cots eee rene race tee stetaters tn Siete sco s fa eusis eee mer eo stee ele revagels 16 IMGTEUNG IEEE OES pootpad bots obo oa bind 6 ond 6 bUbI00.d 6 Bou Ho Dod do bonantHoeooue 19 Speciesuireatmentsi rae eee eer EEC et ericae cantik racers 20 Genus Caryadiplosis Gagné, New Genus ...........6 00006 Sr ER aeine arte 88 GenericidescripHomapee tence Rie ace eae hie seein etetaeh ce tarstenel release. tns 88 SpeciesilreatmentSyapss cramer arin GME Eee PRP re ee II fC L Yep eta eksisee veh 89 Genus ContariniaRondaniterr rere reiki aot errr rr enenrere 91 Speciestireatmentsiarmpii miter yore yeasterrccne ere sesieeerscorierstcey siecerare rays ceases 91 GenusiGliaspilotalGagneyiINewsGenusierar tyke eis sates: clays oe eiclsieirale | eveleie sees cine 93 Cenericidescriptlomnwacery russe cecyoererals sliver iebsl coos keer iene anes euerelanesors steiner eran 93 Genus) harmandiolaidettensarta. fers seine seis ari isi) nee ee eee 95 Genus!Parallelodiplosisiubsaamnentyr sei ihe tev ieksietcher-e- seth ote verve duets oleh aisiyssetees 96 Key to Hickory Galls Caused by Cecidomyiidae ................... ccc cece 99 Key to Mature Larvae of Cecidomyiidae on Hickories ..................0000000- 102 JNA AION MISe Fania aocobod kab edons dois Bacoud CoD uE ap ans DoD DUC OsMdaU cates arc 104 Pitera tres Cited. ieee cestery- roe, ststey no era os etietey sdenere euonevencietseede ai scstescuey suatevavanearaiovate iar suans- 104 Bi runes vl = 36 Zeanrwr cc.) weer Persiren creates men = nepeteteee ewstetore ones chsh cSvel ac ciea fall lke aes 108 IIA S acer ered tee cc tone eh ccioneurtemeasecy seniors isis ear sve rena atcusteneac evenesz yore Torerchoicin gaa. suays seevaveust 146 ... The moon makes no protest. It rolls what it sees into the scene it illumines, and lends its old weight- . * afloat and paper thin and mottled with maria— to what men have thrown up as it once beamed benign on Crusader castles, fern swamps turning into coal, and the black ocean when no microbe marred it. —From Condo Moon, by John Updike MEMOIRS OF THE AMERICAN ENTOMOLOGICAL SOCIETY NUMBER 48 The Gall Midges (Diptera: Cecidomyiidae) of Hickories (Juglandaceae: Carya) By RAYMOND J. GAGNE Systematic Entomology Laboratory, PSI Agricultural Research Service, U.S. Department of Agriculture c/o Smithsonian Institution MRC-168 P.O. Box 37012 Washington, D.C. 20013-7012 _USA (e-mail: raymond.gagne@ars.usda.gov) ABSTRACT—This monograph treats the 63 species of gall midges known to feed on North American hickories (Carya: Juglan- daceae). The distribution and history of hickories are outlined. Once widely dispersed across the Holarctic Region during the Eocene epoch and later, hickories are now limited in range to eastern North America and a smaller area in southeastern Asia. Fifty-six species belong to the single genus Caryomyia that is known exclusively from hickories. Each is univoltine and forms a characteristic, complex leaf gall in which their larvae eventually pupate the following spring. These species exhibit a great num- ber of striking biological and morphological traits. Six other species in four other genera are responsible for one complex and five simple galls on the leaves or nut husks of hickories, while one other is an inquiline in galls formed by Caryomyia spp. Morphol- ogy, relationships, distribution, and biology of each of the gall midge taxa are treated in turn. Keys for the identification of hick- ory galls made by gall midges and to larvae of cecidomyiids on hickories are provided. Nineteen of the 63 species were previ- ously named. Fifteen of them are redescribed and the remaining four are relegated to junior synonymy. One species is renamed and a new genus is described for another. Already described species are: Caryomyia ansericollum Gagné, new name for Caryomyia caryaecola Felt; Caryomyia antennata Felt; Caryomyia caryae (Osten Sacken) (= Caryomyia arcuaria (Felt), new synonym); Caryomyia caryaecola (Osten Sacken); Caryomyia cynipsea (Osten Sacken); Caryomyia holotricha (Osten Sacken) (= Caryomyjia caryae (Felt), new synonym); Caryomyia inanis Felt; Caryomyia persicoides (Osten Sacken); Caryomyia sanguinolenta (Osten Sacken); Caryomyia tubicola (Osten Sacken); Caryomyia thompsoni (Felt) (= Caryomyia consobrina Felt, Caryomyia similis Felt, new synonyms); Gliaspilota gluti- nosa (Osten Sacken), new combination; Harmandiola nucicola (Osten Sacken), new combination; and Parallelodiplosis caryae (Felt). Two new genera, Caryadiplosis and Gliaspilota, are described, as are 49 new species, as follows: Caryadiplosis biconvexa, Caryadiplo- sis venicola, Caryomyia aggregata, Caryomyia albipilosa, Caryomyia arcuata, Caryomyia asteris, Caryomyia biretta, Caryomyia caminata, Caryomyia cilidolium, Caryomyia conoidea, Caryomyia cucurbitata, Caryomyia deflexipili, Caryomyia echinata, Caryomyia eumaris, Cary- omyia flaticrustum, Caryomyia glauciglobus, Caryomyia glebosa, Caryomyia guttata, Caryomyia hirtidolium, Caryomyia hirtiglobus, Cary- omyjia inclinata, Caryomyia inflata, Caryomyia lenta, Caryomyia levicrustum, Caryomyia leviglobus, Caryomyia marginata, Caryomyia melicrustum, Caryomyia ovalis, Caryomyia procumbens, Caryomyia purpurea, Caryomyia recurvata, Caryomyia shmoo, Caryomyia sphe- rica, Caryomyia spiniglobus, Caryomyia spinulosa, Caryomyia stellata, Caryomyia striolacrustum, Caryomyia striolata, Caryomyia subu- lata, Caryomyia supina, Caryomyia tuberculata, Caryomyia tuberidolium, Caryomyia tumida, Caryomyia turbanella, Caryomyia turbinata, Caryomyia urnula, Caryomyia viscidolium, Contarinia bulliformis, and Contarinia cucumata. 2 THE GALL MIDGES (DIPTERA: CECIDOMYITIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) INTRODUCTION Cecidomyiidae, the gall midges, are a large family of over 6,000 described species worldwide (Gagné 2004 & subsequently). They are of great economic and ecological interest because at least two-thirds of their number feed during their lar- val stage on flowering plants, some of agricultural and horticultural importance. Many cecidomyi- ids, by means of their larval salivary secretions, cause their hosts to produce galls that surround the larvae and produce their food. Except for the more economically important pests, however, Ce- cidomyiidae remain one of the least well known families of Diptera. A case in point is the present monograph on the gall midges of hickory. The number of species treated in this study is aug- mented from the 14 previously known from hick- ories to 63, 49 of them new to science. The first notice of hickory gall midges was published in 1862 by Charles R. von Osten Sacken. A European amateur dipterist, he lived in the United States between 1856 and 1877, first as Secretary of the Russian Legation in Washington, later as Consul General of Russia in New York, and finally as a retired person before returning permanently to Europe (Osten Sacken 1904). His 1862 paper presented the first introductory sketch and synopsis of North American Cecidomyiidae. In that long article, still a delight to read for the simple, straightforward prose, Osten Sacken de- scribed and named many species of gall midges he found in the vicinity of Washington, DC. Among his discoveries was a suite of eight species from hickories. Seven of them form complex, de- tachable leaf galls and belong to the genus Cary- omyia. The eighth species causes a sticky leaf spot and is placed here in a new genus, Gliaspilota. Os- ten Sacken reared the adult stage from only one of the detachable galls; all the remaining species were described chiefly from their galls, with a line or two on the larvae of some. In 1870 Osten Sacken described a ninth species, placed here in Harmandiola, from swellings on hickory nut husks and associated larvae that he collected in Rhinebeck, New York. Additional species from hickory were named between 1907 and 1921 by Ephraim Porter Felt, distinguished State Entomologist of New York from 1898 to 1928. Felt was involved with many statewide insect problems, including mosquitoes and shade tree insects, but as a major side interest while State Entomologist and afterwards he named 1080 species of gall midges from North America and elsewhere. That number is phenom- enal, even taking into account the standards of the day, but the accomplishment left him little time to reexamine the body of work as it grew. He nonetheless left to posterity an overall well- mounted and labelled collection of type material as well as a major work on galls, Plant Galls and Gall Makers (Felt 1940). The book is somewhat dated, but still useful and unique in that it treated all of the North American insect and mite gall makers known to 1940, including what are still the more noticeable and common gall makers of our fauna. Felt acknowledged Osten Sacken’s species and named nine more from hickory, all from the Albany, New York, vicinity except one from Massachusetts. Two species were based on adults netted on hickory (Felt 1907) and the re- maining seven were reared from galls (Felt 1908, 1909, 1921), but his association of the reared species with particular galls wavered in consecu- tive papers. Felt (1909) proposed the name Cary- omyia for the hickory gall midges and later (Felt 1921) gave an overview of the included species. Four of Felt’s nine species are treated as junior synonyms here, and another, a junior homonym, is renamed. Notices of some of these Osten Sacken and Felt species were published subsequently, including summaries in Felt (1940), the posthumous Felt (1958) (as revised by Remington & Remington), and Gagné (1989), but only one further study, by Bertram W. Wells (1916), treated the cecidomyiid- caused hickory galls in a comprehensive manner. Fascinated by gall architecture while a botany stu- dent and later assistant professor at Ohio State University, Wells published several papers on galls. In his 1916 paper, he made effective draw- ings, whole and in section, of 28 kinds of ce- cidomyiid hickory galls that he collected, mainly in southeastern Ohio. His two plates are repro- duced here as Figures 1-33. Wells’s drawings of galls in longitudinal section demonstrated the im- portance of internal gall structure, including wall thickness, any intrusions from the exterior, and how each is attached to the leaf. He attempted to assign Osten Sacken’s and Felt’s species names to the galls but refrained from coining new names himself because he believed that task should be done by those studying the insects (Wells 1916). Wells subsequently moved to North Carolina RAYMOND J. GAGNE 3 where he distinguished himself in the field of plant ecology. Following his 1916 paper and the review by Felt (1921), essentially nothing new has been published on hickory gall midges for these last eighty-seven years. Table 2, pp. 17-18, lists the 63 species of Ce- cidomyiidae known to feed on American hicko- ries, all but one responsible for specific galls. They belong to six separate genera and show remark- able morphological diversity and a variety of life strategies. The genera are as follows: Caryomyia. This genus comprises 56 species found exclusively on hickories. Each species forms a distinctive complex leaf gall that usually dehisces between spring and fall (Figs. 36-164). Larvae remain in their galls over winter and pu- pate the following spring. Caryadiplosis. This is a new genus that appears to be the closest known relative of Caryomyia. It contains two new species that make simple galls, one a circular, biconvex leaf blister (Figs. 167-168), the other a vein or petiole swelling (Figs. 165-166). When fully fed, larvae exit the galls and drop to the soil where they overwinter and then pupate the following spring. Contarinia. To this large group in its broad sense are added two new species known only from their larvae. One species forms a complex sac-like leaf gall (Figs. 172-173) in spring on sev- eral hickories, the other causes a bubble leaf blis- ter (Figs. 170-171) on pecan. Gliaspilota. This new genus is described for Ce- cidomyia glutinosa (Osten Sacken). Larvae of this species live freely on mucilaginous leaf spots that appear on the underside of new leaves in spring (Figs. 175-177). Fully-fed larvae soon drop to the soil to pupate and then emerge as adults the fol- lowing year. Harmandiola. Until recently this group com- prised several gall-formers on poplars and chest- nuts. It now includes one other species, Harmandi- ola nucicola (Osten Sacken), from hickory. Larvae form swellings on nut husks (Figs. 178-180). Fully grown larvae drop to the soil to overwinter. Parallelodiplosis. A number of species in this genus live as parasitoids in galls formed by other gall midges and cynipids. One, Parallelodiplosis caryae (Felt), lives in galls of various species of Caryomyia (Fig. 48). This species may have more than one generation per year. In addition to the taxa listed above, undeter- mined species of two additional genera were found. These genera are included in the larval key provided farther on but are not treated further in this paper except to list them here, as follows: Lestodiplosis. This is a large worldwide genus of over 160 described species that feed as larvae on insects and mites (Gagné 2004). Larvae of sev- eral species were found during this study on the surface of hickory leaves or in galls caused by Caryomyia, Phylloxera (Phylloxeridae), and various groups of mites (Acarina). Larvae are shown in Figs. 48 and 53. These predaceous larvae and the few adults reared from them during the present study cannot now be identified with confidence without a separate revision of Lestodiplosis. Dasineura. This is the largest genus of gall midges, a catchall category with more than 460 described species (Gagné 2004). On one occasion in early May, 2005, two full-grown larvae were found closely appressed to an expanding leaflet of mockernut hickory. The larvae were not associ- ated with any apparent damage. THE HICKORIES Hickories belong to Carya, a genus of the wal- nut family (Juglandaceae) that contains 12 North American species including pecan. The present range of the genus encompasses the eastern half of temperate North America, including southern Ontario and Quebec, eastern United States, and isolated areas in the mountains of northeastern Mexico, as well as parts of southeastern Asia (Grauke 2003, Manchester 1987, Stone 1990). The 12 North American hickories form two sections, Apocarya with four species and Eucarya with eight species and one variety (Table 1, p. 4). Additional Carya spp. occur in southeast Asia, three to six species depending on the authority, and belong to a third section, Sinocarya (Stone 1997). No galls from Asian species have been reported and no sign of galls occurs on leaflets of their representa- tives in the herbarium of the National Museum of Natural History, Smithsonian Institution, Wash- ington, DC, so the present study treats only the gall midges of North American hickories. Hickories are now used chiefly for firewood or charcoal-making, but their tough, resilient wood Was once more commonly used for such things as spokes for wagon wheels and tool handles (Grauke 2003). Pecan, originally a Mississippi basin species, is now widely planted and culti- 4 THE GALL MIDGES (DIPTERA: CECIDOMYITDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Table 1. Scientific and common names of North American Carya species (from Stone 1997). The correct name for the section Eucarya is Carya, but Eucarya is used in this paper to avoid confusion with the generic name. Section Apocarya: C. aquatica (F. Michaux) Nuttall C. cordiformis (Wangenh.) K. Koch C. illinoiensis (Wangenh.) K. Koch C. palmeri Manning Section Carya or Eucarya: C. floridana Sargent C. glabra (Miller) Sweet C. laciniosa (F. Michaux) Loudon C. myristiciformis (F. Michaux) Nuttall C. ovata (Mill.) K. Koch C. ovata var. mexicana (Helmsley) W.E. Manning C. pallida (Ashe) Engler & Graebner C. texana Buckley C. tomentosa (Poiret) Nuttall water hickory bitternut pecan Mexican bitternut scrub hickory pignut shellbark nutmeg hickory shagbark Mexican hickory sand hickory black hickory mockernut vated throughout southeastern United States and abroad for its nutmeat. Shagbark and white hicko- ries, especially north of the range of pecan, still have some value to people for their nutmeat. All are important to wildlife, particularly tree squir- rels (Manchester 1987). The name “Carya” is Greek for walnut. The common name “hickory” is derived from the American aboriginal word “pawcahiccora” for a mash made from the nutmeat. The word was shortened to hickory for the name of the tree by the early English colonists. “Pecan” is another Ameri- can aboriginal word meaning “hard-shelled nut.” Only one other genus of Juglandaceae, Juglans or walnut, occurs in North America. It is widely distributed throughout the temperate Holarctic Region. The only gall midges recorded from Juglans across its entire range are two undescribed eastern North American species that live as larvae in marginal rolls of leaflets, one a Dasineura on Juglans nigra, or black walnut, the other a Con- tarinia on Juglans cinerea, or butternut (Gagné 1989). Hickories are an ancient genus that was once much more widely dispersed. Carya-like pollen is known from the early Paleocene, about 60 million years before the present (bp), but the earliest rec- ognizable Carya fossil fruit is from mid-Eocene Europe about 45 million years bp (Manchester 1987, Stone 1990). Post-Eocene hickories extended across the Northern Hemisphere until after the mid-Miocene (ca. 15 million years bp), when the climate cooled and the individual elements of a once uniform Arcto-Tertiary forest began to sepa- rate and undergo extinctions (Wolfe & Leopold 1967). Fossil Carya fruit have been found in Alaska (Lower Miocene), Washington state (Miocene) and Colorado (Lower Oligocene), as well as in Eurasia (Middle Eocene to Pliocene) from central and eastern Europe, western Siberia, northern China, and Japan (Wolfe & Leopold 1967, Man- chester 1987). The present narrow range of the genus is due to extinctions during the uppermost Tertiary and Pleistocene (Manchester 1987). During the Pleistocene glacial maxima, hicko- ries and other deciduous trees were much more severely restricted in range than presently. Decid- uous angiosperm trees survived in small, isolated populations in the lower Mississippi valley and on the southern coastal plain among the more widespread conifers, as determined by pollen de- posits in lake beds (Davis 1983, Delcourt and Del- court 1984). When the glaciers of the most recent ice age began to recede about 12,000 years ago, the various tree species that survived gradually spread north on a species basis and not as com- RAYMOND J. GAGNE 5 munities, their individual paths dependent upon their particular refugia and individual strategies (Davis 1983). Moving from southwest to north- east, Carya expanded more slowly than some other genera, in part because of their heavy fruit; even so, hickories arrived early (10,000 bp) in the upper midwest at sites close to their present northern boundary, but were slow to cross the Ap- palachian Mountains, arriving in New England only about 5,000 years bp (Davis 1983). Populations of hickories in Mexico (Map 1) may not have been greatly affected by glaciation. Hickories there are part of an assemblage of de- ciduous trees generically similar to that of eastern North America. This representation of northern trees in Mexico appears to have been in place since the mid-Miocene when the cooling conti- nental climate allowed the deciduous forest to reach so far south (Graham 1999). Evidence for this is that pollen and spore deposits in Mexico show a late Cenozoic history of the assemblage’s presence there. In addition, it is unlikely that the soil between central Texas and the relict popula- tions in Mexico would have supported a rich de- ciduous forest more recently than the mid- Miocene (Graham 1999). These points will be of interest farther along when considering the rela- tive absence of galls on a Mexican population of Carya ovata var. mexicana. North American hickories vary greatly in dis- tribution. A few species, C. cordiformis, C. glabra, C. ovata, and C. tomentosa, range widely and occupy most of the combined range of the genus in Canada and the United States indicated in Map 1. The remaining species, except pecan, Carya illi- noiensis, which has been extensively planted for commerce, are restricted to small parts of the com- bined range (Elias 1980). MATERIALS AND METHODS Hickory names and identifications.—In dis- cussions that refer to both Carya and Caryomyia or Caryadiplosis, the abbreviation “C.” is used to mean Caryomyia or Caryadiplosis; in those discus- sions I use the common names of the hosts to avoid confusion. Both scientific and common names of hickories are listed in Table 1. Hickory identifications were made with the use of various keys and treatments, including those of Brown & Brown (1972), Elias (1980), Gleason (1963), Grauke & Pratt (1986), and Stone (1997). Keys do not all agree in the number of species. After gaining some experience I followed Elias (1980), Grauke & Pratt (1986), and Stone (1997) in which Carya ovalis is placed under C. glabra and C. septentrionalis un- der C. ovata. Hickories are highly prone to hy- bridization (Grauke 1988, 2003) and one notices considerable variation in this genus, but only rarely did I encounter a tree not readily attributa- ble to a species. As it happens, most cecidomyiids in this study do not appear to discern among hick- ories, particularly those feeding on the Eucarya section. Collections.—Most galls and their cecidomyi- ids used for this study were collected by me since 1993. Observations and collections were largely made at many sites within 20 miles of my resi- dence in Silver Spring, Maryland and at one site near Hedgesville, West Virginia. A major local area sampled was the Beltsville Agricultural Re- search Center in Beltsville, Maryland where trees remained undisturbed during this time period and where I was free to collect as an employee of the U.S. Department of Agriculture. I made three separate circuit trips across midwestern and southern United States to collect galls, two sepa- rate trips to Florida, several to New England and New York, including once an extension into On- tario, and one trip to Tamaulipas, Mexico with a stop on the way to southeastern Texas. Additional collections were made for me by collaborators in Connecticut, Georgia, and Florida. In addition, I studied types and other specimens from the Felt Collection, which is the property of the New York State Museum in Albany but currently on loan to the Systematic Entomology Laboratory in Wash- ington, DC, and the few miscellaneous specimens, including the existing Osten Sacken types, de- posited over the years in the National Museum of Natural History, Washington, DC. One needs to cover a lot of ground to find hick- ories with galls, especially the less common ones. The best collection sites were forest edges or open-grown trees where low-hanging branches made collecting convenient. Sampling while trav- eling is always haphazard, dependent on a place to park a car or obtaining permission from prop- erty owners. Old churchyards, cemeteries, and in- terstate highway rest stops, particularly the large midwestern ones, are productive sites. Paths through woods are not as good when time is pressing because finding hickories can be uncer- 6 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLAN DACEAE: CARYA) tain and branches in forests are often too high to reach. Some good collecting sites in central Mary- land were lost to development over the course of this work: one site is now a public ice skating rink with large adjacent parking lot, and another a shopping center. Collecting consisted of harvesting gall-bearing leaflets and placing them in a plastic bag with a locality and host label. I gathered galls at a site un- til |had a sufficient number of the available kinds. Because these galls are small, close examination was done subsequently, usually in the evening of the same day with the aid of a dissecting micro- scope. Representative galls were cut open and the larvae removed and saved in alcohol. Samples of whole and cross-sectioned galls were glued to a card and pinned together with a data label for permanent reference. Remaining galls that were excess but not numerous enough to attempt to rear were kept dry in labelled plastic bags for fu- ture comparison and extraction of larvae as needed. Larval samples were placed in alcohol and labelled, some eventually to be slide mounted or used for SEM photos. At the proper season, de- pending on the species, galls with full-grown lar- vae or, for genera other than Caryomyia, larvae that had escaped the galls, were kept through winter to rear adults. Some reared adults were pinned but most were kept in 70% ethyl alcohol for eventual slide mounting. Sweeping or otherwise trapping adults in flight is not practical. Associating both sexes of a species is usually not possible for most plant- feeding gall midges unless they are reared to- gether from similar galls. Any sweep of hickory branches can yield many species of gall midges, some just passing by and having nothing to do with hickory. While gall midges caught in flight may with practice be identifiable to generic level when viewed dried or in alcohol, they must be painstakingly mounted on microscope slides for further critical viewing, and even then most flight-caught phytophagous cecidomyiids cannot usually be determined beyond genus with confi- dence. Slide mounting is a time-consuming activ- ity better spent on reared samples. The best method for obtaining adults of phytophagous gall midges is to collect a sufficiency of galls before they begin dropping from the trees, usually in September and October for hickory galls, and keep them over the winter. Galls with full-grown larvae are placed in pots filled with dampened peat moss. If larvae normally exit the galls when full grown, they will crawl into the peat moss. In this way it is possible to obtain all stages of a species and learn something of their biology. Even so, I was unable to rear adults or obtain pupae of all of the species treated here. Many larvae die from parasitism or other causes before they are fully developed, so oné needs a large number of galls to successfully obtain gall midge pupae and adults. Galls of the less common species were never found in sufficiently large numbers to suc- cessfully rear adults. Adult rearing.—I have tried various methods to rear adults but that outlined in Gagné (1989, 1994) still works satisfactorily, with one change. I place galls, or larvae when these escape the galls to pupate in the soil, in pots filled with dampened peat moss and place them in an outdoor cold frame for the winter. Many galls in that situation undergo deterioration from the dampness and fungi, allowing entry by various arthropods, mainly mites and sciarids. This is probably no dif- ferent than what happens in nature, but it means that one has to have a large number of galls to in- sure success. At the end of February when I bring the pots indoors, I place each pot in a separate clear plastic bag. The bag is large enough to spread well away from the pot at the bottom so any emerging specimens are noticeable when the bag is held against the light. The top of the bag is closed by bringing together two ends of the open- ing in a simple knot. Condensation does not col- lect on the inside of the bag if the pot is not overly wet in the first place and the bag is kept out of di- rect sunlight. If moisture does condense on the in- side of the bag, a wet bag can be changed repeat- edly to a dry one until the pot loses the excess moisture. This seems to be a more efficient method than the one I formerly used, when I placed the pots individually in cardboard shoe- boxes, each with a vial fit tightly into an opening at the end. With that method I occasionally lost some specimens that could squeeze out through a cover that did not fit tightly or escape when I opened the cover for a peek. Specimens examined and their preparation.— Slide mounting of specimens follows the tech- nique in Gagné (1989, 1994) using Canada balsam. Most of the larvae and adults studied during this study can be traced to particular galls that are ei- ther on pinned cards or in alcohol. Holotypes of new species and remaining specimens used in this RAYMOND J. GAGNE 7 study, except those from the Felt Collection, are deposited in the insect collection of the National Museum of Natural History in Washington, DC (USNM). Descriptions.—In the species descriptions that follow, the gall is described first because it is the most obvious manifestation of a species. Each gall description lists elements of the mature gall in this order: frequency (common, occasional, or rare), distribution on hickories (Apocarya or Eucarya sec- tion or both); arrangement of galls on a leaflet (single or clustered), position on a leaflet (upper or lower leaf surface, between or on veins); height or length in mm and external shape; external tex- ture; connection to the leaflet; wall structure in longitudinal section; and shape and texture of the larval chamber. A concluding sentence or two usually follows to differentiate the gall from other galls with which it might be confused. Some species are named and described here on the basis of only the gall and larva because during the period of this study I did not succeed in rear- ing the adult. In a comprehensive study the ab- sence of adults of some species should not pre- clude describing species from larvae alone. Larvae have good distinguishing characters, their biology and hosts are known, and their character- istic galls can be found again, possibly to success- fully rear adults. Adult abdominal tergal setation is described from the third tergum posteriad because the pos- terior setal rows of the first and second tergites are always discontinuous mesally. Larval length of- fers a rough size estimate. In any instar a larva grows considerably, so the larger number of the range is the more meaningful one. Lists of collection and specimen localities.— Collection localities of galls of each cecidomyiid species in this revision are listed alphabetically by state or province and county or town. Voucher specimens for the gall records are deposited in the USNM, either pinned or in alcohol. Listed also, with citations, are galls from publications when accompanied by identifiable figures. A separate list of larvae, pupae, and adults of each species studied gives the collection and rearing (if any) data in alphabetical order by state or province, and town or other site, followed by collection date, collector if not me, and the associated life stage. All these specimens, including types of new species, but excepting the Felt Collection speci- mens, are deposited in the USNM. Abbeviations include: For. = Forest; Mt. = Mountain; Natl = Na- tional; MCZ = Museum of Comparative Zoology, Cambridge, Massachusetts; Ref. = Refuge; St. = State; USDA = U.S. Department of Agricuture; USNM = National Museum of Natural History, Smithsonian Institution, Washington, DC. Species concept within Caryomyia.—I consid- ered a distinct gall the phenotypic extension of a discrete species. In only one case might the gall distinction seem minor, viz. between C. leviglobus and C. spiniglobus. The first (Fig. 62) has a smooth exterior, the other (Fig. 64) is covered with stiff se- tae. Because the two kinds were never found inter- spersed, I consider the gall makers distinct, even though I cannot separate them morphologically. Two other species are noteworthy here because of their variability. Caryomyia thompsoni has two dis- tinct variations in gall surface, one kind more or less sparsely haired and the gall surface visible (Fig. 113), the other with thick hair obscuring the gall surface (Fig. 114). These can often be found in- terspersed on a leaflet (Fig. 112), and the morphol- ogy of Caryomyia specimens from each is indistin- guishable. Except for the type specimens under the species name, I made no further distinction in the list between the two kinds and considered them to be made by the same species. The other special case is C. caryae, in which the galls inter- grade. The extreme forms (cf. Figs. 55-56 with Figs. 57-58) each seen alone certainly appear dif- ferent and I might have treated them as separate species were it not for commonly found inter- grades. No differences in any stage of the gall midge from the two extremes of galls could be de- tected. GENUS CARYOMYIA FELT Caryomyia Felt 1909: 292. Type species, Cecidomyia tubi- cola Osten Sacken, by original designation. Felt 1911: 56 (review); Felt 1921: 94 (review); Gagné 2004: 100 (catalog). Diagnosis.—This genus may be differentiated from other genera of Cecidomyiidi by the follow- ing combination of characters: Adult without dor- sal protuberance on occiput; wing (Fig. 245) broad- est on basal half, C unbroken at juncture with Rs; legs considerably foreshortened (Fig. 183), shorter than wing length; abdominal sternites lacking an- terior pair of trichoid sensilla; gonostylus short, broad, with wide tooth; protrusible ovipositor ta- 8 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) pered abruptly beyond eighth segment, short, the ninth tergite dorsally with narrow, pigmented, stiff, dermal structure, and cerci partially fused dorsally at base (Figs. 184-185). Larva with only 1-4 lateral papillae; dorsal thoracic papillae re- duced to 4; terminal papillae all similar, flat or con- vex, asetose (Fig. 271). Description.—Adult: Head: Eyes connate; facets mostly hexagonoid and closely adjacent ex- cept slightly farther apart and circular at mid- height of head. Occiput without dorsal protuber- ance. Frons setose. Labrum with or without setae. Labella separate or coalesced, with or without se- tae. Palpus 1- to 4-segmented. Antenna with 12-20 flagellomeres, when 12, often with an additional bud at apex of twelfth counted as a thirteenth, but this one without circumfila; male flagellomeres (Figs. 235, 237, 239) generally foreshortened, bin- odal with foreshortened internodes and necks, or cylindrical (Fig. 240) and lacking internodes and necks; either with 3 circumfila, 1 on basal node and 2 on distal node, or the 3 evenly distributed along flagellomere length, loops of the circumfila sub- equal in length, short, not attaining bases of next distal circumfilum, rarely a loop connected be- tween two circumfila, or with 9-10 interconnected horizontal circumfila; female flagellomeres (Figs. 236, 238, 241) each progressively shorter, cylindri- cal with short necks, ringed with 2 appressed cir- cumfila connected by two longitudinal bands or several anastomozing bands. Thorax: Scutum with 4 longitudinal rows of setae. Scutellum with lateral group of setae on each side. Pleura with vestiture only on setose mesepimeron. Wing (Fig. 245) broadest on basal half, unmarked, width 0.4—0.5 wing length; C unbroken at juncture with Rs; Rs curved apically, joining C posterior to wing apex; Rs incomplete, closer to arculus than to apex of R;. Legs generally foreshortened, femora, tibiae and tarsi of approximately equal length or pro- gressively shorter from femora to tarsi. Tarsal claws (Fig. 246) untoothed, curved beyond mid- length; empodia as long as claws; pulvilli minus- cule, much shorter than claws. Male abdomen: First through seventh tergites entire, rectangular, about 5 times as wide as long; with setae in 1-4 rows along posterior margin or only posterolater- ally and with anterior pair of trichoid sensilla but no other vestiture except for microsetulae entirely covering sclerites; eighth tergite may be more weakly sclerotized than preceding tergites and have fewer setae; tenth tergum with narrow, elon- gate, stiff, pigmented dermal structure along its length. Sternites rectangular, 2-3 times as wide as long, without trichoid sensilla; second through sixth sternites with horizontal row or rows of pos- terior setae and variable horizontal rows of mostly shorter setae near midlength; seventh and eighth sternites with more numerous setae and both rows adjacent. Genitalia (Figs. 186, 247-257): cerci short, broad, with setae along margin; hypoproct paral- lel-sided to broadly widened posteriorly, with scattered short setae along posterior and lateral edges, covered with setulae except, in most species, dorsum bare from base to posterior third or fourth; aedeagus tapering from base to apex or broad, cylindrical, rounded or notched apically; gonocoxite cylindrical, short, broad; gonostylus short, broad, ovate, with dense setae and entirely setulose. Female abdomen (Figs. 183-185): First through eighth tergites entire, rectangular, about 6 times as wide as long; vestiture as in male. Sec- ond through seventh sternites rectangular, about 2.5 times as wide as long, vestiture as in male. Ovipositor slightly protrusible, soft to rigid, grad- ually tapering beyond eighth segment; ninth ter- gite dorsally with narrow, pigmented, stiff dermal sclerite, laterally and ventrally setose; cerci ovoid or triangular, partially fused anterodorsally, cov- ered with setae but without distal peglike setae; hypoproct short, broad, with 2 posterior setae. Pupa (Figs. 187-214, 267-268): Integument un- pigmented. Head: Vertex convex, on each side with anterolateral pair of papillae, one of each pair, occasionally both, with conspicuous seta. Anten- nal bases forming anteriorly directed horns of var- ious length and shape. Anteriormost angles of face between antennal bases and flagella often convex. Face anterior to midlength smooth or indented with an extruded posterior lip on each side. Frons and clypeus convex or conspicuously expanded, base of frons with pair of setose papillae. Labella triangular, convex. Thorax: Prothoracic spiracles elongate, pointed. Thoracic papillae short-setose, situated on each side as follows: 1 on prothorax an- terior and medial to spiracles, 3 along anterior edge of notum, 1 anterior to wing base, and one at posterior end of notum. Notum with row of tiny spicules on each side of medial suture. Wing pads extending just beyond second abdominal seg- ment, legs extending approximately to end of third abdominal segment. Abdomen: Spiracles sessile. First abdominal tergum of all species covered with short spicules and additionally with horizontal RAYMOND J. GAGNE 9 row of 6 short-setose papillae barely longer than spicules; intersegmental tergal areas smooth, bare; second through eighth tergites as for first or with several horizontal rows of large, closely set, spine- like spicules on anterior third of segment. Pleura smooth anteriorly and posteriorly but elsewhere covered with small spicules, a pair of setose papil- lae present similar to those on terga. Sterna wrin- kled, rough, with minute pointed verrucae anteri- orly and laterally, papillae not detected. Larva: Third instar (Figs. 215-231, 269-271): White, cylindrical to ovoid, anterior and posterior ends rounded or tapered. Head directed anteri- orly or ventrally. Antenna about twice as long as wide. Spatula (Figs. 274-327) present, variously shaped. Papillae with or without setae, their bases prominent or not. Lateral papillae 1-4. Ventral papilla on each side of spatula closely adjacent to lateral papillae. Dorsal papillae of thorax 4, of first through seventh abdominal segments 4—6. First through eighth abdominal segments always with verrucae laterally, dorsum and venter smooth or lined and/or variously covered with rounded to pointed verrucae and ventrally with rounded ver- rucae or pointed spicules. Terminal segment of same length to more than twice as long as eighth abdominal segment, with 8 similar papillae with- out setae, 4 on each side, slightly to conspicuously convex. Anal region with 0-2 papillae. Second in- star (272-273): White, ovoid. Head directed ante- riorly or ventrally. Antenna about twice as long as wide. Spatula present in all species except C. albi- pilosa and C. striolata, with apically pointed tooth and elliptical to quadrate shaft. When spatula present, collar segment on each side with patch of enlarged spicules. Papillae with or without setae, their bases prominently raised or not. Lateral papillae 1-4. Dorsal papillae of thorax 4, of first through seventh abdominal segments 4-6. First through eighth abdominal segments variously covered dorsally with rounded and or pointed verrucae. First instar (Fig. 35): Translucent to white, fusiform at first, becoming flattened and discoid. Spiracles present on first thoracic and eighth abdominal segments only. Antenna about three times as long as wide. Egg (Fig. 34): Elongate-ovoid, red. AFFINITIES Caryomyia belongs to the supertribe Ce- cidomyiidi because of the separate many-looped circumfila of the male flagellomeres (Gagné 1989, 1994). Below that level the genus fits into the large tribe Cecidomyiini that contains most of the Hol- arctic plant-feeding Cecidomyiidi, mostly genera that lack a dorsal occipital protuberance, have usually simple tarsal claws that are curved be- yond midlength, and larvae with a particular makeup of the terminal papillae (Gagné 1989). While Cecidomyiini typically have four pairs of terminal papillae, three of them short-setose and the other corniform, enlarged, and recurved dor- sally (as in Fig. 334), all eight terminal papillae of Caryomyia are without setae and similar in ap- pearance (Fig. 271). The relationship of the genus to other Cecidomyiini is nonetheless clear due to the fortunate discovery during this study of the new genus Caryadiplosis that is described below. While the two genera are evidently related due to shared, derived adult characters, Caryadiplosis has retained the larval form commonly found in the tribe. In the key to genera of Cecidomyiidae in Gagné (1981), Caryomyia runs to couplet 135 with Contarinia, but that placement does not argue for any affinity between those two genera. Felt (1909) first described Caryomyia to include eight species and eventually added another seven (Felt 1921). Three were included only for conve- nience because they occurred on hickories. Felt (1909, 1921) wrote that Caryomyia bore some re- semblance to Planetella (as Hormomyia), but the only grounds he offered was that the two genera shared “somewhat the appearance,” a suggestion that does not bear scrutiny. Felt (1921) later wrote that Caryomyia might be a relative of Polystepha (as Cincticornia), a genus in the tribe Asphondyliini, because the male antennal circumfila of both gen- era are short and Carya and Quercus, the host of Polystepha, are somewhat related. Because of this, Caryomyia was listed in Felt (1958) with the tribe Asphondyliini, and that placement was contin- ued in Foote (1965). Male antennal flagellomeres and their circumfila are foreshortened in some species of Caryomyia that Felt knew (Fig. 237), but had he seen a male of the Caryomyia cilidolium group with its reticulated flagellomeres (Fig. 240) he would have had better but still superficial sup- port. The antennae of the C. cilidolium species group are generally similar to those found in sev- eral species of Polystepha spp. (cf. Fig. 30a, p. 154 of Felt 1916). We now know that the two genera are in separate tribes because Polystepha shares a number of apomorphies with the Asphondyliini 10 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) (Mohn 1961, Gagné 1994), the most important be- ing the enlarged female seventh sternite. That the antennae of two groups of species in two separate tribes are so similar is only one more example of adaptive homoplasy that can be found in Ce- cidomyiidae. Other examples of parallel modifi- cations are listed in the following discussion. These serve as a good argument for comprehen- sive revisions, in which context these happen- stances can be appreciated better. A relationship between Caryomyia and Cary- adiplosis is supported by three synapomorphies and one other character they have in common. The first synapomorphy is the presence of a nar- row, elongate, dorsal sclerite on the female ninth segment, the posterior, more rigid half of the ovi- positor (Figs. 258-264, 331). Both genera have pro- trusible, yet relatively short ovipositors on which the structure is presumably used as a stiffener. I know of a corresponding structure on only two other genera of Cecidomyiini, Macrodiplosis and Paradiplosis. Macrodiplosis, a large genus of leaf gall makers on oaks (Gagné 1989), has a relatively short ovipositor also, but the stiffener extends only along the posterior two-thirds of the seg- ment. In Paradiplosis, a genus of four species on fir and spruce, the ovipositor is extremely long (Os- good and Gagné 1978). The second synapomorphy is the unbroken costal vein at its junction with Rs. An unbroken costal wing vein is rare in Cecidomyiini and un- known in other tribes of the supertribe Ce- cidomyiidi. The trait occurs in two cecidomyiine genera that infest Pinaceae, Paradiplosis and The- codiplosis, neither of which seems especially closely related to one another or to Caryomyia and Caryadiplosis, so I suppose the closed costa evolved independently in this tribe at least three times. Macrodiplosis, mentioned in the previous paragraph, has a broken costa. The third synapomorphy is the apparent loss of a pair of peglike sensilla usually found near the apex of the female cerci. Almost all Cecidomyiidi have these two (sometimes more) distinctive sen- silla that have broad sockets and wide, blunt- tipped setae (as in Fig. 362). Paradiplosis and The- codiplosis both have the pair of sensilla. Another character Caryomyia and Caryadiplosis have in common is a completely setulose gonosty- lus, but I cannot say with confidence whether this is primitive or derived. The alternative state, the gonostylus setulose only on the basal half and longitudinally ridged beyond, is the one most commonly found in the subfamily Cecidomyi- inae, including Macrodiplosis and Thecodiplosis, but not Paradiplosis. A completely setulose gonostylus is definitely the less common state in Cecidomyi- ini. It will be noticed that Paradiplosis shares at least three characters with Caryomyia and Ca- ryadiplosis: a sclerite on the distal half of the ovipositor, the unbroken costal vein, and the com- pletely setulose gonostylus. This genus is dis- cussed further under Caryadiplosis. Caryomyia differs from Caryadiplosis in many ways, presumably all apomorphic, as follows: 1.Galls are complex, the result of a complete re- structuring of the host tissue, instead of simple swellings. 2. Larvae have only one to four lateral thoracic papillae on each side of the midline (Fig. 274) instead of six lateral thoracic papillae in two triplets on each side (Figs. 335). Such reduction is not uncommon in Cecidomyiinae that form galls, especially complex galls, but the full complement is the rule in non-phytophagous Cecidomyiinae. 3. Larvae have only four instead of six thoracic dorsal papillae. Except for Caryomyia, all other Cecidomyiidi known to me have six dorsal thoracic papillae. I know of a similar reduction in two genera, Asteromyia (Gagné 1968) and Se- mudobia (Roskam 1977), both members of the supertribe Lasiopteridi but in separate tribes. 4. The larval terminal papillae are all similar and asetose (Fig. 271), instead of dissimilar and ei- ther setose or corniform (Fig. 334). This reduc- tion is unique in Cecidomyiidi. 5.Pupation occurs in the gall rather than in the soil. 6. Adult tarsi of Caryomyia are foreshortened, no longer than the tibiae and 0.33 or shorter than the entire leg, while in Caryadiplosis and most other Cecidomyiini the tarsi are appreciably longer than the tibiae and 0.40 or more than the entire leg. 7. Adult abdominal tergites have neither a lateral group of setae (setae that lie just anterior to but separate from the posterior row of setae) nor scales (Fig. 258). In Caryadiplosis and most other Cecidomyiini, these setae and scales are usually present (seen in Fig. 330). RAYMOND J. GAGNE 11 8. Adult abdominal sternites lack an anterior pair of trichoid sensilla (Fig. 330). Their loss here appears to be unique in Cecidomyiini. 9. The male ninth tergum has a narrow, elongate sclerite along the midline (Figs. 248-249). In my experience, the presence in Caryomyia of this sclerite is unique. As to apomorphies of Caryadiplosis, there are possibly two, the presence of a midlength hori- zontal ridge on the male hypoproct (Fig. 333) and the short, wide, cylindrical aedeagus (Fig. 333). Caryomyia and other Cecidomyiini known to me have no horizontal ridge at midlength on the hy- poproct (Fig. 247), and I know of no other ce- cidomyiine with a similar aedeagus. INTRAGENERIC MORPHOLOGY AND RELATIONSHIPS Caryomyia species have diverged in many ways. Some of their differences, particularly lar- val shape, are evidently adaptations for life in galls as these changed with time. Reasons for other differences are not so clear, as, e.g., why the male flagellomeres of the C. cilidolium species group are unlike anything seen elsewhere among Cecidomyiini, or how the presence of setae on the sternal and hind ventral papillae of C. leviglobus and C. spiniglobus are again expressed after hav- ing been lost, possibly before the tribe’s genesis. The genus can be divided into four groups that I informally designate here as the striolata (2 spp.), cilidolium (4 spp.), caryae (9 spp.), and tubicola (14 spp.) groups. The first is ancestral to the remain- der, while the second is sister to the third and fourth. Other species are not referable to these groups because I have not to date succeeded in rearing their male sex. The striolata group consists of C. striolata and C. albipilosa whose galls develop quickly in early spring instead of slowly through late spring and summer, the habit of the remaining species. Two apomorphic characters support the monophyly of these two species, the foreshortened male and fe- male flagellomeres (Figs. 235-236) and an elon- gate ovipositor (as in Fig. 258). These characters evidently occurred twice in Caryomyia, as will be outlined in the following discussion, but this group lacks the unique developments found in the rest of the genus. The main characters supporting the mono- phyly of Caryomyia species exclusive of the strio- lata group are the presence in the second instar of a spatula and enlarged spicules on each side of the collar segment (Fig. 273). A second instar spatula is a rare event in Cecidomylidae, known else- where only in five other genera, all gall makers: Paradiplosis, a cecidomyiine genus mentioned ear- lier (Osgood and Gagné 1978); in at least two species of Lophodiplosis, a cecidomyiine genus from Melaleuca in Australia (Gagné, Balciunas and Burrows 1997); in two of the 11 known species of Acacidiplosis, also cecidomyiine, responsible for complex galls on Acacia in Africa (Gagné and Marohasy 1993); in Celticecis of the supertribe La- siopteridi responsible for complex galls on Celtis (Moser 1965), and, again in Lasiopteridi, in at least one species of Rabdophaga (pers. obs.) responsible for galls on Salix. The second instar spatula ap- pears de novo in each of these genera where, pre- sumably, it acts as a scraper. Where this second in- star spatula does occur in Caryomyia, a group of enlarged spicules is present on each side of the collar segment of that instar (Fig. 273). These en- larged spicules are presumably used either for scraping the internal gall surface or gaining pur- chase for the head. Analogous enlarged anterior spicules in the second instar are present in two other cecidomyiids known to me, Mikiola fagi Har- tig (Dziurzynski 1961) and Lopesia armata Gagné (Gagné & Marohasy 1993), but instead of on the collar segment, the enlarged spicules appear on the first thoracic segment. Neither of these species has a second instar spatula. The cilidolium species group, comprising C. cilidolium, C. hirtidolium, C. tuberidolium, and C. viscidolium, is distinguished by several apomor- phies as follows: 1.Galls are distinguished by an apical, circular, weakened area through which pupae exit in spring (Fig. 45). All other Caryomyia galls have no such defined exit. The striolata group pupae force their way through the gall apex but the galls do not have a defined, breakaway apex, and all other Caryomyia species force their way through the gall side or base through a tunnel dug by the larvae the previous autumn. 2. The larval head points ventrally (Fig. 231), pre- sumably an adaptation for the mouthparts to anchor to the feeding surface ina large spherical 12 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) larval chamber. This character is found also in some species of the caryae and tubicola species groups that also have very large larval cham- bers, larger than the larvae can fill when full grown. The character is presumably separately derived where it occurs. The head capsule of gall midges usually points anteriorly (Fig. 217). 3. The spatula has a single, pointed anterior tooth (Figs. 277-280). Both the caryae and tubicola species groups also have species with a single, triangular apical tooth but those are evidently separately modified. 4.The pupal face has a pronounced convexity just posteriad of the antennal bases and a verti- cal median furrow (Figs. 193-196). The striolata species group has a smooth face without these characteristics (Figs. 267-268). 5.The pupal antennal bases are conspicuously elongated and dorsoventrally flattened (Figs. 193-194), unique to this group within the genus and presumably a modification for pushing open the apical gall exit. 6. Circumfila of the male flagellomeres are reticu- late and appressed (Figs. 240-241) instead of being made up of three separate circumfila with regular loops (Figs. 235-239). This is a unique development in Cecidomyiini. It is known elsewhere in Polystepha and Eocincticor- nia of the tribe Asphondyliini. Female circum- fila also are reticulate (Fig. 241) instead of be- ing disposed in the more usual two horizontal rings with two vertical connectives (Fig. 238). I should point out here that in the cilidolium group the circumfilar strands are more numer- ous on the venter of the flagellomeres, just as both loops and setae are usually more numer- ous on the venter of other cecidomyiinae. I as- sume that the significance of augmented cir- cumfilar strands or setae on the ventral surface is for optimal sensory perception. When a gall midge is in flight, its antennae are normally held aloft with the venter the leading edge. 7. The adult labella are entirely fused and asetose (Fig. 243), while in other Caryomyia they are partly or entirely discrete and long-setose (Figs. 242, 244). The caryae and tubicola species groups share at least four synapomorphies, as follows: 1. The pupal face has prominent anterolateral in- dentations and a conspicuously produced frons (Figs. 188-189), presumably used during emergence from the gall in spring. In the other species groups the face is generally smooth, without lateral indentations or bumps. Non- gall making cecidomyiines have a primitively convex, smooth face. Many gall makers that pupate in their galls have various facial ridges and projections, but the particular makeup of these in Caryomyia is unique in Cecidomyiinae. 2.The pupal abdomen has conspicuous spines on the anterior half of the second through eighth abdominal terga (Fig. 192). These are used in exiting galls. Abdominal spines are of- ten found in plant-feeding Cecidomyiidae that pupate in their galls, but gall makers do not universally have them. 3. The ovipositor is longer (Fig. 258) than in the cilidolium group (Fig. 261), as measured by the length of the cerci that are 1/4 or less the length of the ninth segment. I consider a short ovipositor to be ancestral to a longer one; the reverse hypothesis would involve many si- multaneous reversals of a complex organ, its tubes, muscles, and integument. 4. The male hypoproct is setulose only on its pos- terior third (Fig. 253), an obvious reduction from the usually completely setulose ce- cidomyiid hypoproct. The tubicola species group, which includes C. tubicola, C. conoidea, C. deflexipili, C. flaticrustum, C. inanis, C. inclinata, C. inflata, C. leviglobus, C. mar- ginata, C. recurvata, C. sanguinolenta, C. spherica, C. spiniglobus, and C. turbinata, is separated from the caryae group by two synapomorphies. The aedea- gus is notched at the apex (Fig. 247), unlike all other Caryomyia and other cecidomyiines known to me that have a rounded or blunt apex (Fig. 333), and the male flagellomeres are foreshortened (Figs. 237), without the distinct internodes and necks found only in the caryae species group (Fig. 239) and almost universally in Cecidomyiini. As- suming that foreshortened flagellomeres are de- rived, they evidently appeared at least twice in the genus: foreshortening appears in the basalmost striolata species group and again in the tubicola group, but here it is correlated with an apically notched aedeagus. The caryae species group has, unfortunately, no synapomorphy that I can determine. The C. caryae group includes, besides C. caryae, at least C. aggregata, C. hirtiglobus, C. persicoides, C. pur- RAYMOND J. GAGNE 13 purea, C. shmoo, C. thompsont, C. tuberculata, and C. turbanella. Other noteworthy characters of Caryomyia os- tensibly appeared more than once and are not useful for grouping species or are found in single species: 1. Gall structure can be useful for defining groups, especially when supported by characters of the gall midges themselves. Some structures evi- dently appeared only once, as for instance the apical pupal exit in galls of the C. cilidolium species group (Fig. 45). Other changes, such as extraneous tissue having nothing to do with the larval chamber, appears to have come about more than once. Galls of C. sanguinolenta (Figs. 134-135) and C. persicoides (Figs. 126-129) re- semble one another in general shape: both are conical, the extensive tissue surrounding the larval chamber is large-celled and soft, and the larval chamber itself is lined with a distinct pel- licle that eventually hardens and presumably serves to protect the larva through the winter as the surrounding soft tissue breaks down and sloughs off. Nonetheless the galls have distinct differences: the gall of C. sanguinolenta has a shield-like base (Fig. 136) through which the pupa eventually emerges; that of C. persicoides lacks the differently textured base and its pupa emerges through the side of the gall. These dif- ferences take on more weight with the realiza- tion that adults from the two galls belong to two different species groups. 2. Larval shape reflects the space available in the larval chamber. Larvae constrained in ovoid or cylindrical chambers are ovoid or cylindrical (Figs. 215-216) and ultimately fill the available space. Larvae of some species in chambers with more space than they can fill may have a ventrally disposed head, a wide, flat body or convex segments, an elongate terminal seg- ment, and papillae that may be situated on lobes (Figs. 226-229). 3. Verrucae and spicules that evidently serve for traction may cover all or only parts of a larva. Larvae free to crawl within a large space gen- erally have more verrucae than larvae that are more constrained. 4.The third instar spatula is variously shaped and not predictable by gall form. One of its uses for most species of the caryae and tubicola species groups is for boring a hole through the gall wall. A one-toothed spatula (Fig. 276) is presumably derived from a two-toothed one, the ancestral form (Fig. 285), but a secondarily notched tooth (Fig. 286) may be derived from a one-toothed spatula, and a notched spatula may possibly revert to a simple, triangular tooth. Several species responsible for generally similar galls show a gradual narrowing of the two-toothed apex, and one of them has a single tooth (see under C. flaticrustum and cf. Figs. 287-290). The two widely spaced spatula teeth of C. caryae (Fig. 282) and relatives is evidently a derived state because it is unique to this genus. 5.Setation of larval papillae is uncommon in Caryomyia and occurs in seemingly unrelated groups of species, in either or both second and third instars. In two species, C. leviglobus and C. spiniglobus, all papillae except the termi- nals have setae, a unique development in Ce- cidomyiidae. 6. Dorsal papillae of the first through seventh lar- val abdominal segments are reduced to four in about half the species from the basic six found elsewhere in the genus and customarily in Ce- cidomyiidi. This loss does not appear to follow natural groupings in the genus, so probably happened more than once. 7. Adult antennal flagellomeres usually number 12 in the supertribe Cecidomyiidi, but may be augmented to more than the basic number, most often by an incomplete thirteenth flagel- lomere in one or both sexes. In C. deflexipili the number is augmented to 18-21. Augmentation in number of flagellomeres beyond a short thirteenth segment is rare in Cecidomyiidi, but occurs elsewhere in some species of Planetella (Gagné 1981). 8. Adult palpi usually have four segments (Fig. 342). Only occasionally has a specimen only three segments on one side, but this is not a specific difference. One species, C. spherica, which belongs to the tubicola species group, is unique in the genus for having only one or two palpal segments (Fig. 244). Noteworthy here is the corresponding short palpal sheath on the pupa of this species (Figs. 205-206). Interest- ingly, in other Cecidomyiidae that have a short palpus, e.g., in Rhopalomyia (Jones et al. 1983), the labella are also reduced in size and seta- tion, but in C. spherica the labella are large, sep- arate, and have exceptionally strong setae. 14 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) 9. The ovipositor of Caryomyia ranges from gen- erally short and pliable with short setae (Fig. 261) to somewhat longer, rigid in some species, and with a small number of long setae (262). Large, mostly separate cerci become variably reduced in size and can be almost completely joined dorsally. These modifications seem to have arisen independently within Caryomyia: C. inanis and C. inclinata have generally simi- lar, rigid, completely pigmented ovipositors, but otherwise show no particular evidence of close affinity. BIOLOGY General life cycle.—All Caryomyia species are univoltine. In central Maryland adults generally emerge in late March through April from galls fallen to the ground the previous summer and fall. Later emergence evidently does not occur: adults almost never emerged from my collections after April, and second flushes of hickory leaves that replace occasional early-season damage by Lepidoptera never have Caryomyia galls. Within a few minutes after eclosion, adult wings and legs expand to full length and harden. Females espe- cially are heavy-bodied and do not fly readily. In traps where they have been observed, they gen- erally walk rather than fly when disturbed, so are easy to pick up with a dampened artist’s brush. Mating in nature presumably occurs close to the emergence site. Females then fly to burgeoning hickory buds or rapidly developing leaves where they deposit eggs on the bud or leaf surface (Fig. 34). The newly hatched larva is at first elongate- ovoid, the same shape as the egg itself. Whether the larva crawls initially, it soon settles and begins to feed, whereupon its salivary secretions induce the leaf tissue to encompass it in a gall distinctive to its species. As the gall develops, this first instar becomes flat and discoid (Fig. 35). Galls of Cary- omyia species are essentially full-grown by the time the larva molts to the second instar. Follow- ing the molt to the second instar and an additional period of larval growth, the larva then molts to the third and last instar. Once the third instar is fully grown, in late spring for a few species or summer for the remainder, the gall either dehisces or hangs on until the leaves drop from the tree in autumn. The larva diapauses inside the gall through winter. In early spring, the larva molts to the pupal stage and the adult develops quickly. The fully-developed adult still inside the pupal skin then forces its way through the gall epider- mis, for most species through a cylindrical tunnel burrowed in the gall wall the previous autumn by the full-grown last instar. When the pupa is halfway out of the gall, the greater width of its ab- domen blocks further progress through the aper- ture. Major sutures of the pupal head and thorax then split open, allowing the adult to emerge. Within that generalized life cycle, Caryomyia species show a remarkable variety of oviposition and host selection, seasonal development, gall structure, and larval and pupal adaptations. These aspects are taken up here in turn. Oviposition and host selection.—The female selects the host and the site where the galls will subsequently develop, whether on the lamina or veins. Eggs are laid singly, in clusters, or in rows, presumably a species-specific feature. In one closely related group of four species (see under C. cilidolium), one species occurs only on bitternut of the Apocarya section, while those of the remaining three occur on the Eucarya section. Two of the three on the Eucarya section occur on the lower surface of the leaf, while one occurs exclusively on the upper surface. The top leaf surface is not a fa- vored site for Caryomyia galls, with only two of the 56 species found exclusively on the top sur- face and a third that occurs equally regularly on either surface. Galls of about a third of the species are always found attached to veins and some of them are situated so closely together that they sometimes partly coalesce. These site selections by the female are evidently also species specific. Seasonal development.—Galls once initiated develop quickly, but their triggering is staggered in most species. Galls of two species, C. albipilosa and C. striolata, are exceptional in that their galls appear as soon as or even before the leaves have fully unfurled in early spring. Their larvae, from first through third instar, develop within about two weeks. In central Maryland their galls nor- mally dehisce by late May and early June. Larval development of another, more variable species, C. urnula, is not quite so rapid, but the galls appear soon after the two earlier species. Galls of C. ur- nula vary in time of appearance, and some galls dehisce as early as mid-June, while others remain on the leaves into September or until leaf fall. The rapid development in spring of C. albipilosa and C. striolata galls may possibly be the ancient mode RAYMOND J. GAGNE 15 for the genus because those species share some primitive larval and adult character states of Cary- omyia. A possible advantage of early development is that the galls are fully developed and dehisce before the hickory leaves can be affected by heavy Phylloxera populations that may cause leaf drop or before the leaves can be eaten by caterpillars, events I have observed for leaves bearing the more slowly developing Caryomyia species. Alter- natively, there may be disadvantages to a gall ly- ing on the ground through the summer. It hap- pens that the spring galls are among the most weak-walled of Caryomyia galls. The appearance of galls of the remaining species is variable. Depending on a species but also within a species, galls may become apparent only in June, July, or August. Once galls are fully grown and first instars have molted, larval devel- opment progresses fairly slowly through most of the summer. Galls may dehisce from the leaves be- ginning in August but galls of some species remain on the leaves until frost or even after leaf fall. Gall structure —All Caryomyia species induce complex galls on hickory leaflets. A complex gall is the result of reorganized tissue that is different from any structure normally found on the host plant. Most of the great diversity of galls caused by Caryomyia spp. (Figs. 36-164), including color, shape, relative hairiness, and in some a sticky coat- ing and/or false chamber, occurs outside the bounds of the larval chamber and has no apparent effect on the larva, at least during feeding. Exam- ples of benign structures would be the knob at the apex of the C. leviglobus gall (Fig. 62) or the re- curved apex of the C. recurvata gall (Fig. 90). I ob- served no sign of any insect, including eggs of par- asitoids, in any partial or complete false chamber, so a false chamber does not appear to be a protec- tive adaptation in the same manner as it is for galls of Cynipidae (Cso6ka et al. 2005). Whether relative stickiness, hairiness, or hardness deter some para- sitoids from laying their eggs and have some sig- nificance for gall diversity is a determination that must await further study, but one explanation for gall diversity might be that the differences are only fortuitous and trivial accidents due to genetic drift during repeated isolation events over time. Larval adaptations.—Larvae of Caryomyia are less homogeneous than adults. Adults need to steer their way out of the galls while still encom- passed by the pupal skin, emerge from the pupal skin, then harden, and then mate, and the females have to lay their eggs at a particular site on a suit- able substrate. In contrast, the varied needs of the larvae, in part determined by the shape of the lar- val chamber in the gall, gall intrusions, and rela- tive thickness and hardness of the wall through which larvae need to burrow, have led to some re- markable anatomical adaptations variously dis- tributed among Caryomyia spp. Depending on the species, larvae are either sessile or active while still growing. Larvae of C. tubicola and C. sanguinolenta, e.g., fill the width or entire larval chamber, at least by the time they are full-grown. Their larvae are evenly cylindrical or ovoid, respectively, and have a modicum of horizontal rows of raised verrucae or denticles to effect crawling or other movement. Larvae of species that live in large, roomy cham- bers, e.g., C. cilidolium and C. caryae, are more or less dorsoventrally flattened, have expanded lat- eral and terminal extremities, and are variously haired or lobed for better traction. In addition, their bodies are completely covered with a rough integument and the head capsules of some of those species are oriented ventrally instead of anteriorly. In a freshly cut gall with a spherical larval cham- ber, one can see that the ventral side of the larva is closely appressed and curved along the interior surface of the large chamber. The third instar spatula shows some variety of size and shape, which seems partly associated with gall form. Before entering diapause in sum- mer or fall the third instar uses its spatula to cut a cylindrical tunnel on the side or near the bottom of the galls, up to but not including the outer epider- mis. The pupal-encased adult will emerge through this tunnel or weakened area the following spring. Conspicuous wear of the spatula itself through the effects of such abrasion can be seen on third instar exuviae sloughed off after pupation. A particularly striking adaptation within Cary- omyia is the development of a spatula on all Cary- omyia second instars, except in C. albipilosa and C. striolata, the two spring gallmakers discussed ear- lier. A spatula is always present in third instar ce- cidomyiids except in the few species or genera that have secondarily lost it. Its appearance in sec- ond instars is rare, found only in gall makers, and presumably separately derived in the few genera where it occurs (see under Intrageneric Morphol- ogy and Relationships). This structure is presum- ably employed as a scraper to encourage growth of food-producing cells in the galls. A trait associ- ated with this development in Caryomyia is the 16 THE GALL MIDGES (DIPTERA: CECIDOMYITIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) presence of an enlarged group of spicules on each side of the collar segment behind the head (Fig. 273). The enlarged spicules presumably also aid as a scraper or anchor for the head capsule against the internal gall wall surface. Pupal adaptations.—Adaptive structures in- clude the different shapes and disposition of an- tennal bases, facial protuberances, and rows of ab- dominal spines on most species. These structures are presumably used in exiting from the harder galls because they are undeveloped in C. albipilosa and C. striolata, the two species that make softer, thin-walled galls, and in C. urnula and the C. cili- dolium group that escape their galls through an apical trap door. Host-SPECIFICITY The host range of individual Caryomyia species is mostly hickory section specific rather than species specific, and some species may occur on both sections of hickories (Table 2). Fifty-three of the 56 species feed on Eucarya, and there is no evi- dence to show that they discriminate among the species of the section. Hickories themselves com- monly hybridize in nature (Grauke 1988, 2003), which may indicate a lack of barriers for Cary- omyia spp. Further, species of other genera on hickories, Caryadiplosis spp., Contarinia cucumata, and Gliaspilota glutinosa (q.v. this paper), are regu- larly found on various species of Eucarya as well as of Apocarya. Of the 53 Caryomyia species that feed on Eucarya, one is commonly, two occasion- ally, and three rarely found also on Apocarya. Galls of Caryomyia tubicola, the species most commonly found on both sections, appear on different tis- sues from one section to another. On Eucarya galls are most often found on the leaf lamina, but on bitternut they are usually situated adjacent to leaf veins. Possibly the open lamina of bitternut leaves cannot always sustain the galls because its leaves seem generally not as thick and stiff as those of the Eucarya group. Only three species of Caryomyia, C. ansericol- lum, C. cilidolium, and C. spinulosa, are restricted to bitternut of the Apocarya section. They possibly occur also on that section’s other species, but those trees were not sampled as often during this study as was bitternut. Nevertheless, I only once found a Caryomyia gall on either of two other species of the Apocarya section, pecan and water hickory, and neither one of the collections was of the species restricted to bitternut. Interestingly, the three species found only on bitternut each ap- pear more closely related to species on Eucarya than to one another, so colonization from one to the other section evidently happened at least three times. DISTRIBUTION Many genera of gall midges restricted to par- ticular host genera occur naturally across the en- tire breadth of the Holarctic Region. Examples include Semudobia on birches (Roskam 1977), Ce- cidomyia on pines (Gagné 1978), and Macrodiplosis (spring galls) and Polystepha (summer galls) on oaks (Gagné 1989, q.v. for other examples). These associations presumably antedate the separation between Europe and North America in the mid- Eocene. As mentioned earlier, hickories, while now much more restricted in distribution than are birches, pines, or oaks, once ranged across the Holarctic before being extinguished in Europe and elsewhere during the Pleistocene (Manches- ter 1987). It is probable that Caryomyia, too, once also ranged across the Holarctic so that whatever influences shaped the distribution of hickories during geological time presumably also affected Caryomyia. Most Caryomyia species occur throughout the range of hickories in eastern North America. Some, such as C. caryae, C. thompsoni, and C. tubi- cola, can be anticipated on almost any tree of the Eucarya section, while others, e.g., C. cynipsea and C. glebosa, although geographically widespread, are rare. A separate phenomenon is individual tree susceptibility. Trees of the Eucarya section may annually have large populations of galls of many species while adjacent trees of the same species may have almost none. Over a period of several years 26 species of Caryomyia were found on one open-grown pignut in the National Ar- boretum in Washington, DC. while its near neigh- bor, a pignut of similar age, size, and situation, never bore but a few individuals of only one species. A mockernut at another site in the Na- tional Arboretum sustains each year a large popu- lation of C. tubicola galls so heavy that the leaflets droop, but bears few other kinds. Even some of the more common galls can have a patchy distribution. A case in point is the great difference only 75 miles apart at two of the sites where I collected hickory galls for over 10 years. RAYMOND J. GAGNE 17 Table 2. List of gall midges (excl. of Lestodiplosis spp. and Dasineura sp.) on hickories, their scientific and common names. Presence on Eucarya and/or Apocarya sections is indicated by an asterisk. Scientific Name Caryadiplosis biconvexa Gagné Caryadiplosis venicola Gagné Caryomyia aggregata Gagné Caryomyia albipilosa Gagné Caryomyia ansericollum Gagné Caryomyia arcuata Gagné Caryomyia asteris Gagné Caryomyia antennata Felt Caryomyia biretta Gagné Caryomyia caminata Gagné Caryomyia caryae (O.S.) Caryomyia caryaecola (O.S.) Caryomyia cilidolium Gagné Caryomyia conoidea Gagné Caryomyia cucurbitata Gagné Caryomyjia cynipsea (O.S.) Caryomyia deflexipili Gagné Caryomyjia echinata Gagné Caryomyia eumaris Gagné Caryomyia flaticrustum Gagné Caryomyia glauciglobus Gagné Caryomyia glebosa Gagné Caryomyia guttata Gagné Caryomyjia hirtidolium Gagné Caryomyia hirtiglobus Gagné Caryomyia holotricha (O.S.) Caryomyia inanis Felt Caryomyia inclinata Gagné Caryomyia inflata Gagné Caryomyia lenta Gagné Caryomyia levicrustum Gagné Caryomyia leviglobus Gagné Caryomyia marginata Gagné Caryomyia melicrustum Gagné Caryomyia ovalis Gagné Caryomyia persicoides (O.S.) Caryomyia procumbens Gagné Caryomyia purpurea Gagné Caryomyjia recurvata Gagné Caryomyia sanguinolenta (O.S.) Caryomyia shmoo Gagné Caryomyia spherica Gagné Caryomyia spiniglobus Gagné Caryomyia spinulosa Gagné Caryomyia stellata Gagné Caryomyia striolacrustum Gagné Caryomyia striolata Gagné Caryomyia subulata Gagné Caryomyia supina Gagné Caryomyia thompsoni Felt Caryomyia tuberculata Gagné Common Name of Gall hickory convex leaf swelling hickory midrib swelling hickory aggregate gall hickory white-haired gall hickory gooseneck gall hickory sombrero gall hickory starburst gall hickory spindle gall hickory beret gall hickory chimney gall hickory sticky globe gall hickory onion gall hickory fuzzy ginger jar gall hickory fuzzy gumdrop gall hickory pumpkin gall hickory smooth woody gall hickory reflexed hair gall hickory spiny gall hickory Persian shoe gall hickory puff tart gall hickory blue-powder gall hickory knobby cone gall hickory teardrop gall hickory hairy ginger jar gall hickory woody globe gall hickory hairy gumdrop gall hickory papery gall hickory inclined gall hickory rubbery gall hickory popover gall -hickory plain tart gall hickory smooth ball gall hickory marginate gall hickory honey tart gall hickory ovoid gall hickory peach-haired gall hickory hairy squash gall hickory purple gumdrop gall hickory smooth squash gall hickory smooth gumdrop gall hickory shmoo gall hickory spherical woody gall hickory spiny ball gall hickory thorn gall hickory starry-base gall hickory striate tart gall hickory fluted cone gall hickory awl-shaped gall hickory supine squash gall hickory placenta gall hickory bumpy woody gall on Eucarya * * * + Che OSES: EF ON AE Cee a ECS OEE OS SO EN SG ES NS SH CS RES. e's eh Oe Ee EN ES 8 CM ea Ch ES Ey £3 PEP ER Ee on Apocarya + * (continued) 18 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Table 2. List of gall midges (excl. of Lestodiplosis spp. and Dasineura sp.) on hickories, their scientific and common names. Presence on Eucarya and/or Apocarya sections is indicated by an asterisk. (continued) Scientific Name Caryomyia tuberidolium Gagné Caryomyia tubicola (O.S.) hickory bullet gall Caryomyia tumida Gagné hickory frosted gall Caryomyia turbanella Gagné hickory turban gall Caryomyia turbinata Gagné Caryomyia urnula Gagné Caryomyia viscidolium Gagné hickory urn gall Contarinia bulliformis Gagné hickory leaf bubble Contarinia cucumata Gagné hickory cucumber gall Gliaspilota glutinosa (O.S.) hickory sticky leaf spot Harmandiola nucicola (O.S.) hickory husk swelling Parallelodiplosis caryae Felt an inquiline The two sites are the National Arboretum in Washington, DC and a private property on Cherry Run, 6 mi NW of Hedgesville, WV, both sites close to the Potomac River. These sites share 18 species. The Arboretum has 11 species not known from the Cherry Run property, while the latter site has 13 not found at the Arboretum. Even two sites sepa- rated by only 10 miles, the Arboretum and the Beltsville Agricultural Research Center show some differences. These two sites have 26 species in common, but the Arboretum has one not found at Beltsville, while Beltsville has five not found at the Arboretum. Besides local differences in occurence, 11 species are so far unknown east of the Appalachi- ans in the mid-Atlantic Region. These are: C. as- teris, C. biretta, C. deflexipili, C. echinata, C. inanis, C. inclinata, C. inflata, C. lenta, C. marginata, C. procum- bens, and C. subulata. One other species, C. arcuata, has a similar distribution except that it has also been found near the South Carolina coast. This dis- tribution is fairly definite (Map 2); if these species occurred along the Piedmont of Virginia and Maryland, where I collected widely and often over 13 years, I should have collected them. This pat- tern is surprising inasmuch as only fairly low mountains separate the Piedmont from eastern West Virginia where those “western” Caryomyia species can be found. Interesting also is that most of these 12 species occur to the north into New En- gland. Possibly they occurred long ago on the Piedmont but were extirpated as the forests of the Piedmont were largely cut for farmland early in this country’s history. Still, with regrowth of Common Name of Gall hickory bumpy ginger jar gall hickory top-shaped gall hickory sticky ginger jar gall on Eucarya on Apocarya * % F FF F F * % ¥® forests, prevailing winds should by now have wafted females in an easterly direction, or the species should have spread via watercourses that cut through the mountains and run through the mid-Atlantic region. Nor do these Caryomyia species appear to be particularly affected by alti- tude or temperature, inasmuch as several occur from Mississippi and Arkansas to New England. In addition to that continental distribution pat- tern are two examples of local species paucity. The first is on scrub hickory, a low-growing tree of sand dunes and sandy ridges of central Florida and the sole species of Eucarya growing there. It is host to only four Caryomyia species, three of them widespread, but the fourth, C. guttata, an en- demic, is the only Caryomyia so limited in distri- bution. Galls from scrub hickory were collected over several seasons by K. Hibbard and one year at several sites also by me. As a comparison, only about 150 miles north of those collections on scrub hickory, in Gainesville, Florida, where two species of the Eucarya section, mockernut and pignut, grow together, I found 15 species of Caryomyia ina two-hour search. The other example of a locally restricted num- ber of species is on Mexican hickory in eastern Mexico. On one isolated population of this tree, only about 12 sq. mi. in extent in a cloud forest of the Sierra Madre Oriental southwest of Ciudad Victoria, Tamaulipas, I observed galls of only two species, C. caryae and C. inclinata, in about equal numbers during several days’ search. Galls of C. caryae there belong to one of the two regular forms, the more robust type of that species; those of C. in- RAYMOND J. GAGNE 19 clinata, a highly variable species even in the United States, mostly show a greatly expanded base for that species. This population of a Eucarya hickory is separated by about 550 miles from the next clos- est United States hickories, north of Houston, Texas. Near New Waverly, Texas, about 35 miles north of Houston, I collected from black and mock- ernut hickories 30 species of Caryomyia, including C. caryae and C. inclinata in about three hours. As- suming that the Mexican population of hickories was once conterminous with the Texas population, possibly as long ago as before the Pleistocene (Gra- ham 1999), galls of the two species of Caryomyia in Mexico would seem not to have changed at all dur- ing that long period of time. The two Caryomyia species that occur on Mexican hickory are possibly all that remain of a more diverse assemblage of species that became extinct over time. Those two examples of a reduced number of species at isolated sites have in common the avail- ability of only one host in a restricted area. Possi- bly two host species growing together preserve more variability in Caryomyia spp. than only one restricted host. This seems to hold true for other in- sects on other hosts: the more widely distributed willow species tend to support more gall-making sawflies (Hymenoptera: Tenthredinidae) (Roini- nen et al. 2005) and the numerous species of oaks in the cloud forests of Costa Rica support many species of gall wasps (Hymenoptera: Cynipidae) while the one oak species in the lowland forest supports only two (Hanson & Gomez-Laurito 2005). Geographical restriction over. long periods of time must also be a factor in the loss of species. The complete lack of gall midges on beeches in North America may be an example of this. Unlike birches, pines, and oaks mentioned earlier that to- gether with their particular cecidomyiids are dis- tributed across the Holarctic Region, American beeches are devoid of cecidomyiids in contrast to European and Asian beeches that are hosts to three cecidomyiid genera comprising many species (Buhr 1964 & 1965, Tsuda 1982, Sato & Yukawa 2001, Yukawa & Masuda 1996). Gall midges of beech possibly also once occurred in the Nearctic but became extinct through a bottleneck effect as the range of beeches became severely constricted during successive Pleistocene glaciations. In Caryomyia, speciation seems best explained by geographic isolation. Glacial phases comprised about 90% of the Pleistocene age, time when de- ciduous tree species survived in small popula- tions (Davis 1983). The probability of extinction for individual species was affected by factors such as climate severity, extent of geographical dis- placement of particular species, the size of popu- lations, and the community composition of the forest in refuge areas (Davis 1983). While most ex- tant species of Caryomyia are widely distributed, some are more common than others, and some species are able to live on trees in both sections of Carya. A species of Caryomyia, e.g., C. tubicola, that presently has a widespread distribution (Map 1) and does not discriminate between hickory sec- tions, would appear to have a better chance to survive an ice age than would rarer species. Con- versely, as the range of hickories became con- stricted and fragmented, those rare or more local species of Caryomyia now known from only two or three localities might be more likely extinguished. In addition, if these rare species did not already exist in what was to become a refuge during an ice age, their chances for survival would be further diminished. In sum, survival during Pleistocene glaciations would favor those species already in the areas that became refugia, which would likely be the commonest and most widely disseminated species, and the rarer species would be the most likely to be lost. Once isolated during a glacial pe- riod, various aspects of a species’s variability might be selected and the process of speciation be- gun. Subsequent gradual dispersal north and re- stored sympatry of all the species throughout eastern North America during the interglacials might then serve to discontinue the process of speciation until the next glacial period. MortTALITY FACTORS Caryomyia populations are greatly affected by parasitoids and predators, but also by less definite causes. Galls cut open in spring after adults should have emerged often revealed first instars, sometimes also second instars, that died the pre- vious year. The galls were otherwise clean, with no sign of fungus or parasitoids. Possibly death in these cases was due to weather or plant resistance, and two examples I know of indicate that weather is a factor. At a regular collecting site in West Vir- ginia one year, a late frost on May 10 killed the early flush of leaves of many hickories on which galls were already evident. On another occasion, in late summer in Sallisaw, Oklahoma, when lar- vae should have been in the third instar, I col- 20 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) lected from a grove of trees hundreds of stunted C. caryaecola galls, all only half-grown and each containing a dead first instar. These were evi- dently killed at the same time and by the same cause earlier in the season. Interestingly, galls of this species of Caryomyia were the only stunted ones I found at that site. Dissection of galls showing no pupal or para- sitic hymenopteran exit holes in spring often showed that larvae were killed or eaten by fungi or mites over the winter. My method of rearing, with 100 or so galls in a small pot, may attract a greater concentration of predators than do galls in a more natural situation. From my rearing pots, I occasionally reared Bradysia coprophila (Lintner) (Diptera: Sciaridae) in large numbers. I observed several times that sciarid larvae, possibly of that species, had broken through the epidermis at the end of exit tunnels and were actively feeding on larvae or pupae inside. In one case three sciarid larvae were inside a gall, wound around and in- side a Caryomyia pupa that they had almost com- pletely consumed. In another, a sciarid had bro- ken through the base of a gall and was feeding on a larva. Predaceous mites could occasionally be seen in the pots, and I once observed a male Cary- omyia being attacked by mites as it was emerging. Hymenopterous endo- and_ ectoparasitoids were often seen when freshly collected galls were cut open to extract Caryomyia larvae or determine their progress. Many parasitoids emerged from galls in spring, and representatives are now kept in the Hymenoptera section of the National Collec- tion of Insects. Most common were Platygastridae, some not yet identified that were reared from C. caryaecola, C. flaticrustum, C. inclinata, C. leviglobus, C. melicrustum, C. persicoides, C. procumbens, C. pur- purea, C. thompsoni, C. tuberculata, and C. tubicola, and others belonging to Eritrissomerus n. sp. near cecidomyiae Ashmead that were reared from C. ansericollum, C. aggregata, C. cilidolium, C. deflexipili, C. eumaris, C. glauciglobus, C. hirtidolium, C. persi- coides, C. sanguinolenta, C. spinulosa, C. thompsoni, and C. tuberculata. Other less commonly reared parasitoids were: an unidentified species of Gas- trancistus (Pteromalidae) from: C. ansericollum, C. flaticrustum, C. guttata, C. holotricha, C. inclinata, C. leviglobus, C. persicoides, C. purpurea, C. sanguino- lenta, C. thompsoni, C. tuberculata, and C. tubicola; a pirenine pteromalid from C. aggregata, C. ansericol- lum, C. caryae, C. holotricha, C. purpurea, and C. tu- berculata; and a possible new species of Torymoides (Torymidae) from C. aggregata, C. flaticrustum, and C. glauciglobus. Additionally, three species of para- sitoids from Caryomyia are known from the litera- ture (Krombein et al. 1979): a torymid, Torymus du- rus (Osten Sacken) reared from C. caryae, and two platygastrids, Platygaster caryae Ashmead, report- edly reared from C. holotricha, and Platygaster hyalinipennis (Ashmead), reportedly reared from C. tubicola. As mentioned earlier, predaceous cecidomyi- ids, Lestodiplosis spp., were occasionally found in galls or reared from galls in spring. In addition, an inquiline gall midge, Parallelodiplosis caryae (Felt), treated separately in this paper, was occasionally found in or around galls. Its impact and precise ef- fect in galls of Caryomyia spp. is not known. SPECIES TREATMENTS Caryomyia aggregata Gagné, new species Description.—Gall (Figs. 21a-b, 156-157): Com- mon, on most Eucarya hickories; on lower leaf sur- face on vein, usually clustered in a row along ma- jor vein and often in such close aggregation as to partially coalesce; 3.0-5.0 mm in height, spheroid to bilaterally compressed, sometimes pointed at apex, when coalesced bearing two or more larval chambers in an apparently single gall; covered with thick, mostly brown hair obscuring gall sur- face; base with small, irregular, shallow, central to off-center excavation; wall woody, thick, espe- cially above, larval chamber irregularly ovoid, glabrous, green to brown. The gall of this species can be confused with several other spheroid- conical, densely hairy galls, but its position along the major veins and usual crowding sets it apart from galls of the other species, e.g., C. holotricha and C. purpurea. Galls were described in Osten Sacken (1862) as a variant of typical galls of C. holotricha, but galls of the latter always have an api- cal intrusion of hairs in the larval chamber and oc- cur between instead of on the veins. Adult: Head: Antenna with 12 flagellomeres; male flagellomeres binodal with definite inter- node and neck, circumfila with short loops, their bases widely separated (as in Fig. 239); female fla- gellomeres with 2 horizontal circumfila connected by 2 vertical strands. Labella (as in Fig. 242) sepa- rated from one another anteriorly, hemispherical, short-setose, short-setulose. Palpus 4 segmented. Thorax: Wing length 3.1-3.6 mm in male (n=5), 3.9 mm in female (n=5). Male abdomen: Third through eighth tergites each with continuous sin- RAYMOND J. GAGNE 21 gle row of posterior setae, those of eighth as strong as on preceding tergite. Genitalia (Fig. 186): hypoproct nearly parallel sided, concave pos- teromesally, the margin partly setose, but not in concavity, dorsum setulose only on distal third; apex of aedeagus convex; gonostylus broad at apex. Female abdomen: Third through eighth ter- gites each with mostly single, continuous row of posterior setae, the eighth with double continu- ous or mesally broken row of setae weaker than those of preceding tergite. Ninth segment pliable except for slender dorsal sclerite, setae long but sparse. Cerci elongate-ovoid, pointed apically, pigmented, fused dorsally on basal third. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, moderately separated, acutely pointed apicolaterally. Face with anterolateral pair of hori- zontal incisions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with several anterior rows of enlarged spinules. Larva: Third instar: Length 2.2-3.2 mm (n=10). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 323) with single tooth tapering from wide base to pointed apex and elongate, wide shaft. Papillae without setae. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdominal segments with dorsum almost completely covered with ver- rucae, and venter spiculose anterior to sternal papillae and on posterior third. Terminal segment shorter than eighth. Second instar: Length 1.0-1.3 mm (n=4). Collar segment with patch of enlarged spicules on each side. Spatula with acutely trian- gular anterior tooth and longer, wider, concave- sided shaft. Bases of dorsal and pleural papillae slightly raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material—Holotype: Third instar, left- most of 5 larvae on slide mount, from gall on C. fo- mentosa, Beltsville, Maryland, IX-28-1999, RJ. Gagné, deposited in USNM. Etymology.—The name aggregata is a Latin ad- jective meaning clustered, with reference to the typically closely juxtaposed galls of this species. Affinities.—The closest relative of this species among the C. caryae species group is not apparent. Galls of this species and those of C. guttata are the only ones known to grow so closely together, coa- lescing at times. Biological notes.——Small, tan, white haired galls in tight or loose rows along midrib were found with first instars in central Maryland on May 26. In mid-June galls were full-sized with second instars. In mid-July, third instars could be found but until mid-August most galls contained second instars. By late August all galls contained third instars. Collections of galls——Atasama: Ashville (to- mentosa). ARKANSAS: 2 mi S DeQueen (tomentosa); Osage (texana); Piney (texana). CONNECTICUT: Am- ston (ovata); Danbury (ovata); Litchfield (ovata); Phoenixville (ovata); Plainfield (ovata). FLORIDA: Gainesville (tomentosa); Lake City (tomentosa). Georolia: Forsyth (glabra, ovata, tomentosa). INDI- ANA: Spencer Co. (ovata). KENTucKy: Brownsville (tomentosa); Grayson Lake St. Pk, Carter Co. (glabra, tomentosa); Winchester (laciniosa). ILLINOIs: Marshall (tomentosa). INDIANA: Spencer Co. (tomen- tosa). MARYLAND: Beltsville, USDA Agricultural Research Center (tomentosa); Calverton (tomen- tosa); Clarksburg (tomentosa); College Park (tomen- tosa); Gaithersburg (tomentosa); Lusby (glabra, tomentosa); Potomac (tomentosa); Ridgely (tomen- tosa); Silver Spring (tomentosa); Spencerville (to- mentosa); Wheaton (glabra, tomentosa). MASSACHU- SETTS: Leominster (ovata); Merrimac (ovata); Petersham (ovata); vic. Worcester (Carya sp., pho- tographed in Thompson 1915: 92, Fig. 242); Wren- tham (ovata). Mississippi: Natchez Trace Pkwy, Mile 193, Choctaw Co. (tomentosa). Missouri: Crowley Ridge Roadside Pk, Stoddard Co. (glabra, tomen- tosa); Holly Ridge State For., Stoddard Co. (glabra, tomentosa); Rolla (texana); Wildwood (tomentosa). New Hampsuire: Greenland (ovata); Kingston (ovata). New York: Herkimer (ovata); Hyde Park (ovata); Ithaca (ovata); Ravena (ovata); Shokan (to- mentosa). NORTH CAROLINA: Alberta (glabra); Dortches (glabra, tomentosa); Providence (tomen- tosa). oH10: Hocking St. For., Hocking Co. (ovata); Hocking Co. (Carya spp., Wells 1916: Figs. 21a—b). PENNSYLVANIA: Pavia (ovata); Schellsburg (ovata). TENNESSEE: Dandridge (tomentosa). VIRGINIA: Car- son (tomentosa); Huntley Meadows Pk, Fairfax Co. (tomentosa); Ladysmith (tomentosa). WASHINGTON, DC: National Arboretum (glabra, tomentosa); Wal- ter Reed Army Medical Center (tomentosa). WEST VIRGINIA: 6 mi NW Hedgesville (ovata). Specimens of C. aggregata examined.— ARKANSAS: Osage, IX-9-2004 (larvae). CONNECTI- cut: Amston, IX-17-1999 (larvae). INDIANA: Spencer Co., IX-20-1995 (larvae). KENTUCKY: Brownsville, IX-21-1995 (larvae); Winchester, IX- 22-1995 (larvae). MARYLAND: Beltsville, USDA Agricultural Research Center, X-13-1999, IX- 22 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) 28-1999 & VI-22—2004 (larvae), X-24-2005 (larvae; 285; lo) pupae,wem: “HIE30stomlV-7—2006); Gaithersburg, X-4-1994 (larvae); Ridgely, IX-2003 (4 5,2 2, pupae, em. IV-2004); Silver Spring, X- 24-1999 (larvae). MassacuusetTtTs: Leominster, IX- 21-1994 (larvae); Petersham, IX-6-2000 (¢, em. TV-2001); vic. Worcester, M.T. Thompson (2 4, 2 2); Wrentham, IX-20-1994 (larvae). Missouri: Wildwood, IX-8-2004 (larvae). Onto: Hocking St. For., Hocking Co., IX-19-1995 (larvae). WASHING- TON, DC: National Arboretum, IX-30-1999 (lar- vae; 3 5,3¢, pupae, em. IV-2000), X-3-1994 & VII- 13-2000 (larvae); Walter Reed Army Medical Center, X-1999 (larvae; 2, em. IV-2000) and X-2004 (3 2, em. IV-1-2005). West VirGINiA: 6 mi NW Hedgesville, VIII-27—2000 (larvae). Caryomyia albipilosa Gagné, new species Description—Gall (Figs. 39-40): Common, a spring gall, on both Apocarya and Eucarya hicko- ries; found singly or in groups on lower leaf surface between veins; 3.5-5.0 mm in height, elongate-con- ical, narrowing gradually from base to pointed apex; surface whitish green, eventually turning brown, obscured except at apex by white hairs longer than gall diameter; gall base shallowly con- cave; wall uniformly thin, becoming brittle; larval chamber rough with weak longitudinal ridges. This gall differs from that of C. striolata only in hav- ing a thick covering of long, white hair. Adult: Head: Antenna with 12 flagellomeres and usually a short 13th without circumfila in male; each of the regular 12 male flagellomeres (Fig. 235) scarcely binodal, basal flagellomeres only slightly narrowed beyond basal circumfilum, less constricted on more distal flagellomeres, loops of circumfila short, subequal in length; female fla- gellomeres (Fig. 236) with 2 horizontal circumfila connected by 2 vertical strands. Thorax: Wing length 1.7—2.0 mm in male (n=5); 1.9-2.2 mm in fe- male (n=4). Male abdomen: Third through sixth tergites each with posterior setae continuous across width, in mostly single row; seventh tergite with only a few posterolateral setae; eighth tergite more weakly sclerotized, with anterior pair of tri- choid sensilla the only vestiture. Genitalia (Figs. 249-250): hypoproct broadest posteriorly, weakly concave posteromedially, posterior margin setose, dorsal surface extensively setulose; aedeagus nar- rowing from base to apex, rounded at apex. Fe- male abdomen: Third through seventh tergites each with posterior setae continuous across width, in mostly single row; eighth tergite more weakly sclerotized than preceding tergite, with sparse posterior row of setae, the setae not as long as on seventh tergite. Ninth segment (Fig. 260) pliable except for slender dorsal sclerite, setae shorter than cerci, in two groups, one dorsolateral, one ventral. Cerci elongate-ovoid in lateral view, fused dorsally on basal third. Pupa (Figs. 267-268): Antennal bases parallel in ventral view, carinate anteroventrally, the mesal corner elongated, terminating anteriorly in short, obtuse projection. Face without incisions, frons and clypeus not protruding. Abdominal tergal spinules all similarly minute. Larva: Third instar: Length 1.8—2.1 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 275) with 2 separate, equilateral teeth an- teriorly, the intervening space with 1+ denticles. Two, occasionally 3, lateral papillae on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Papillae without setae, their bases not raised. Abdominal segments dorsally lined, sub- verrucose, with sparse, pointed spicules mainly surrounding the 2 lateral pairs of dorsal papillae, and ventrally- with spicules anterior to sternal papillae. Terminal segment as long as eighth. Sec- ond instar: Length 0.7-1.1 mm (n=10). Collar seg- ment without enlarged spicules laterally. Without spatula. Bases of dorsal and pleural papillae slightly raised, asetose. Six dorsal papillae on first 7 abdominal segments. Type material—Holotype: Third instar, from gall on C. tomentosa, Entomology Road, Beltsville, Maryland, V-29-1994, R.J. Gagne. Etymology.—The name albipilosa is an adjec- tive from the Latin meaning white haired and refers to the gall’s thick covering of white hair. Affinities —Except for the presence of white hair covering the gall, I can detect no difference between this species and that of C. striolata. These are the only strictly spring-occurring galls in this genus and differ in several other important ways from their congeners: the second instar lacks both a sternal spatula and a group of enlarged spicules on each side of the collar segment; the third instar has a clove-like spatula most like that of the ma- jority of Cecidomyiini. The absence in the second instar of a spatula and of enlarged prothoracic papillae place this species and C. striolata as the sister group of the remaining Caryomyia. That the RAYMOND J. GAGNE 23 galls of these two species appear in spring and de- hisce soon after further separates them from re- maining Caryomyia spp. Biological notes.—In central Maryland this species is one of two spring galls with larvae de- veloping between late April and early June. A se- ries of adults emerged from caged previous year’s galls on 16-18 April when hickory leaves were un- furling. A specimen of Lestodiplosis, a predaceous cecidomyiid, was reared with this series of adults. Very small, unidentifiable galls were evident on May 17, 2005. Circled with a marker, these traces were full-grown galls of C. albipilosa with full- grown larvae on May 24. Galls still on the leaves after mid-June are parasitized or contain dead lar- vae. See under C. striolata for more details on tim- ing of galls. Collections of galls.—Galls of this species are presumably more common than this short list indi- cates, but most of my collections outside of the Washington area were made in summer, when galls with live, fully developed larvae would al- ready have fallen from the leaves. CONNECTICUT: Meriden (tomentosa); Southbury (glabra). Mary- LAND: Beltsville, USDA Agricultural Research Cen- ter (tomentosa); Gaithersburg (tomentosa); Kemp Mill (glabra); Silver Spring (tomentosa); Wheaton (tomentosa). NEw York: Addison (glabra). VIRGINIA: Carson (tomentosa); Leesburg (tomentosa). WasH- INGTON, DC: National Arboretum (glabra). West VirGINIA: Falling Waters (cordiformis) Specimens of C. albipilosa examined.—Mary- LAND: Beltsville, USDA Agricultural Research Center, VI-2-1993 (larvae; 7 5,4 2, pupae, em. IV- 17 to 19-1994), V-30-1993, V-29-1994, V-4-1997, and V-16 to 19-2000 (larvae); Gaithersburg, V- 23-2000 (larvae); Silver Spring, V-21-1994 (lar- vae); Wheaton, V-21-1994 (larvae). Caryomyia ansericollum Gagné, new name Caryomyia caryaecola Felt 1921: 114. Junior secondary homonym of Cecidomyia caryaecola Osten Sacken; Gagné 2004: 100, as new junior synonym of Cary- omyia caryaecola (Osten Sacken) to avoid coining new name for C. caryaecola (Felt). Description.—Gall (Figs. 147-148): Infrequent, known only from bitternut of the Apocarya section; usually in closely set series, attached to vein on lower leaf surface, occasionally so recurved as to break through leaf surface to upper surface; 4.1-9.0 mm in height, upright, leaning, or re- curved, base spheroidal, abruptly narrowed to elongate, narrow, curved or sinuous neck, the apex digitate; with short to long, crinkled, white to light brown hair not obscuring gall surface; green, turn- ing brown, not sticky; base with irregular, shallow, central depression; larval chamber basal, ovoid, white, surrounded by thick, green tissue turning brown and woody; a bundle of fibers present be- tween larval chamber and gall apex. This is the only gall on bitternut with thick, woody tissue; the others are thin-walled and brittle. Adult (female only): Head: Antenna with 12 flagellomeres, flagellomeres with 2 horizontal cir- cumfila connected by 2 vertical strands. Labella separated from one another anteriorly, hemi- spherical, short-setose, short-setulose. Palpus 4 segmented. Thorax: Wing length 3.3-3.5 mm (n=3). Abdomen: Third through seventh tergites each with continuous mostly double posterior row of setae, thinning to single row mesally, eighth tergite with single row of posterior setae. Ninth segment (Fig. 263) pliable except for slen- der dorsal sclerite, setae long but sparse. Cerci elongate-ovoid, pointed apically, pigmented, fused dorsally on basal third. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 2.2-3.2 mm (n=10). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 319) with tapering, acutely-pointed tooth and elongate shaft, wide anteriorly, narrow- ing beyond. Papillae without setae, their bases not raised. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae present on first 7 abdomi- nal segments. Abdominal segments with verrucae extensively covering dorsum and rows of small, pointed spicules covering venter except at mid- length. Terminal segment shorter than eighth. Second instar: Length 1.0-1.1 mm (n=3). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider shaft, narrowing gradu- ally posteriorly. Bases of dorsal and pleural papil- lae slightly raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. 24 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Type material—tLectotype of Caryomyjia caryaecola, here designated, third instar, a1786, ex Carya cordiformis, Hamburg, New York, X- 17-1907, in Felt Collection, New York State Mu- seum. Paralectotypes as follows: galls of C. anseri- collum, Felt 1915d, Pl. 4, Fig. 9 (colored drawing by L.H. Joutel); galls of C. spinulosa on Carya cordi- formis, Felt 1921, Pl. 10, Fig. 2 (photo); galls of Carya stellata on Carya sp., Felt 1921, Pl. 10, Fig. 3 (photo), (ex Stebbins 1910: Fig. 13). No paralecto- type galls remain in the Felt Collection; they exist only as published drawings and photographs. Felt (1921) redescribed C. caryaecola Osten Sacken as a new species because he did not con- sider Osten Sacken’s name to be valid, Ostern Sacken having described it on the basis of the gall only. Unfortunately, Felt misidentified Osten Sacken’s species, so Felt’s homonym must be re- named here. Felt (1921) had a broad concept of the gall of this species that included three kinds of conical galls shown here to yield three separate Caryomyia spp. According to Felt’s unpublished notebook, the lectotype larva, #a1786, from Ham- burg, NY, came from bitternut. This narrows the conical type gall, now lost, to either C. ansericollum or C. spinulosa, but the type larva with its triangu- lar spatula tooth, fits larvae taken from C. anseri- collum galls. In fact, Felt’s (1921) gall description matches the gall of C. ansericollum, and the gall is the same as Felt (1915, Pl. 4, Fig. 9) referred to in his 1921 synonymy, not the two photos of Felt (1921, Pl. 10, Figs. 2-3), that he refers to in the 1921 description. The gall was also correctly figured in Felt (1906, Pl. 1; Fig: 21). Etymology.—The name ansericollum is a noun in apposition coined from the Latin anser (goose) and neck (collum) with reference to the long, usu- ally sinuous neck of these galls. Affinities —This species is one of three found only on bitternut; each one appears more similar to species on the Eucarya section than to one an- other. Females (male unknown), pupae, and lar- vae of C. ansericollum are generally similar to those of C. inclinata, q.v. for further discussion of the group of species that have conical, woody, api- cally digitate galls. Collections of galls (all from Carya cordi- formis).—INpIANA: Plainville. MARYLAND: Ellicott City; Kensington; Ridgely; Wheaton. New JERSEY: Worthington. New York: Hamburg; Ithaca; Ravena. PENNSYLVANIA: Pavia. VERMONT: Guilford. ViRGINIA: Rt. 81, mi. 13.9, Washington Co. WEsT VIRGINIA: Falling Waters. Specimens of C. ansericollum examined.—InbI- ANA: Plainville, IX-7—2004 (larvae). MARYLAND: El- licott City, IX-2-2000 (larva); Kensington, VIII-11 and IX-28-2000 (larvae); Ridgely, IX-23-1999 & X- 6-2005 (larvae); Wheaton, [X-1-1994 (larvae). NEw Jersey: Worthington, IX-8-2000 (larvae; 2 °, 3 pu- pae, em. IV-2001). New York: Hamburg, IX- 17-1907, E.P. Felt (larva, lectotype); Ithaca, IX- 3-2003 (larvae). VERMONT: Guilford, IX-7—2005 (larvae). West VirGciniA: Falling Waters, IX- 27-2003 (larvae), VI-21-2004 (larvae), IX-2005 (2 ?, pupae, em. IV-19 to 25, 2006). Caryomyia antennata Felt Caryomyia antennata Felt 1909: 292. Description.—Gall (Figs. 158-159): Occasional, on Eucarya hickories; on or closely adjacent to veins of lower leaf surface; 3-6 mm in height, spheroidal, usually tapered to conical apex, cov- ered with moderate to thick, white to tan hair not quite obscuring gall surface; base with central hemispherical pedicel fitting into corresponding leaf depression; some short exfoliation of leaf sur- rounding connection; wall woody, uniformly thick, larval chamber narrowed at both ends, with longitudinal ridges. This gall could be mistaken for one of the less hairy C. holotricha or C. aggregata galls, but galls of those species have a basal exca- vation rather than the prominent hemispherical connection to the leaf of the present species. The larval chamber of this gall is unique in that it ex- tends into this connection. Adult: Head: Antenna with 12 regular flagel- lomeres; male flagellomeres binodal with definite internode and neck, circumfila with short loops, their bases widely separated; first few female fla- gellomeres with basal circumfilum weakly anas- tomozing on venter. Labella separated from one another anteriorly, hemispherical, short-setose, short-setulose. Palpus 4 segmented. Thorax: Wing length 3.1-3.3 mm in female (n=10). Male ab- domen: Third through eighth tergites each with posterior setae separated into two lateral groups, in mostly double row; eighth tergite not more weakly sclerotized than seventh. Genitalia: hy- poproct widest on posterior third, slightly con- cave posteromesally, margin setose except in con- RAYMOND J. GAGNE 25 cavity, setulose on dorsum only on posterior third; apex of aedeagus slightly rounded. Female abdomen: Third through eighth tergites each with posterior setae in three rows, diminishing to two rows mesally, continuous across sclerite except in- terrupted mesally on eighth. Ninth segment pli- able except fairly wide dorsal sclerite, setae long but sparse. Cerci elongate-ovoid, pointed apically, pigmented, fused dorsally on basal third. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 2.0-2.9 mm (n=10). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 324) with acutely triangular anterior tooth, shaft wide at both ends and narrowing to- wards midlength. Papillae without setae, their bases not raised. Lateral papillae numbering 3 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Abdominal segments with sparse verrucae at midlength dorsally and with rows of tiny, horizontal rows of pointed spicules antero- and posteroventrally. Terminal segment shorter than eighth. Second instar: Length 2.4-3.5 mm (n=8). Collar segment with patch of enlarged spicules on each side. Spatula with elongate, acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae slightly raised, papillae asetose. Six dorsal papillae on first 7 abdominal segments. Type material—Lectotype male, here desig- nated, leftmost specimen of three specimens on slide #a1944, emerged 1909 from Carya sp. galls, Nassau, New York, deposited in New York State Museum, Albany. Paralectotypes: male, on same slide as lectotype (another specimen on same slide is a female Neolasioptera, not part of type se- ries); 2 females, each on separate slide, same data as lectotype except with mounting date of “2 May ‘09”; one female with same data as lectotype with mounting date of “3 May ‘09,” belongs to a different, undetermined species of Caryomyia. Two cards labelled with the type number, a1944, in the Felt collection bear galls, all of one kind, from which this species was reared. They appear at first glance to have no hair covering but they have been wetted and worn by handling and weather. Etymology.—The name antennata is a Latin ad- jective that refers to the antennae, but further con- text is not manifest. Affinities —The gall of this species is unique for the larval chamber extending into the conical basal connection with the leaf. Its closest relative among the other woody gall makers with long an- tennal flagellomeres and an apically rounded aedeagus is not apparent. Biological notes.—First noticed as full-size, hard galls in eastern West Virginia in early July, all were inhabited by second instars. By early August galls contained either a second or third instar. Full-grown larvae fully fill the larval chamber. Collections of galls—Arkansas: Palestine (ovata). CONNECTICUT: Amston (ovata); Danbury (ovata); Hartland (ovata); Harwinton (ovata); Litch- field (ovata) Meriden (ovata); Phoenixville (ovata); Plainfield (ovata); Westchester (ovata). GEORGIA: Forsyth (ovata); Lizella (ovata). ILtiNots: Highland (laciniosa). INDIANA: Spencer Co. (ovata). KEN- Tucky: Brownsville (ovata); Winchester (laciniosa). MaryLAND: Lusby (glabra, tomentosa); Piney Grove (ovata); Ridgely (glabra). Massacuusetts: Boston (tomentosa); Great Barrington (ovata); Holyoke (ovata); Merrimac (ovata); Petersham (ovata). NEw Hampsuire: Greenland (ovata); Seabrook (ovata). New York: Herkimer (ovata); Kinderhook (Carya sp.); Nassau (Carya sp.); New York (Carya sp.). TENNESSEE: Mohawk (tomentosa). WEST VIRGINIA: 6 mi NW Hedgesville (ovata). Specimens of C. antennata examined.—Ar- KANSAS: Palestine, IX-10—2004 (larva). CONNECTI- cut: Amston, IX-17-1999 (larvae); Danbury, IX- 6-2005 (larvae; pupa, excised from gall V-2-2006); Hartland, IX-7—2005 (larva); Meriden, IX-16-1992 (larva). Georeta: Lizella, X-12-1993, J. & R. Payne (larva). INDIANA: Spencer Co., IX-19-1995 (larva). Kentucky: Brownsville, IX-21-1995 (larvae); Win- chester, IX-22-1995 (larvae). MARYLAND: Lusby, IX-16-2003 (larvae); Ridgely, IX-11-2003 (glabra). New York: Herkimer, [X-4—2003 (larvae); Nassau, E.P. Felt (type series of 23,3 2, em. 1909; also pu- pae, larva). Caryomyia arcuata Gagné, new species Description.—Gall (Figs. 141-142): Rare, on Eu- carya hickories; found singly on lower leaf surface 26 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) between veins; 2.5—2.7 mm high, base lunulate, its sides flared toward apex and surrounding base of central, apically pointed cone; hairless, not sticky, brown to purple; base with shallow, central exca- vation; larval chamber located at base of gall, ovoid, lined with yellowish membrane, contrast- ing with brown to purple, large-celled, spongy, vis- cous tissue surrounding larval chamber, a bundle of longitudinal fibers present between larval chamber and gall apex. This gall shows some re- semblance to that of C. sanguinolenta from which it differs in the sides of the gall being conspicuously flared upwards around the apical cone. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.5-1.8 mm (n=8). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 304) with anterior tooth elongate- triangular but notched apically, the shaft widened anteriorly and tapered behind. Papillae without setae, their bases not raised. Lateral papillae 2-3 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdominal segments covered on dorsum at midlength with verrucae and on ventral anterior and posterior thirds with horizontal rows of spicules. Terminal segment shorter than eighth. Second instar: Length 0.9-1.0 mm (n=5). Collar segment with patch of enlarged spicules on each side. Spatula with acutely trian- gular anterior tooth and longer, wider, concave- sided shaft. Bases of dorsal and pleural papillae not raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. tomentosa, St. Clair, Missouri, IX-8-2004, R.J. Gagné, deposited in USNM. Etymology.—The name arcuata is a Latin adjec- tive meaning curved, with reference to the re- curved base of the gall. Affinities—See under C. sanguinolenta, from which this species differs in the shape of the gall. Collections of galls—AtaBaMa: Auburn (to- mentosa). Greenville (tomentosa). FLORIDA: Gainesville (tomentosa). ILLINOIS: Rock Island (to- mentosa, B.D. Walsh). LouIs!1ANA: Lake Bistineau St. Pk, Webster Parish (tomentosa). Missouri: Con- way (texana); St. Clair (tomentosa). SouTH Car- OLINA: Georgetown (tomentosa). Specimens of C. arcuata examined.—ALABAMA: Auburn, IX-19-2000 (larva); Greenville, IX- 20-2000 (larvae). FLoripa: Gainesville, IX-21—2000 (larvae). Missouri: Conway IX-8-2004 (larva); St. Clair, IX-8-2004 (larva). SourH CAROLINA: George- town X-1—-2005, D. & B. Gagné (larva; pupa, ex- cised from gall III-28-2006). Caryomyjia asteris Gagné, new species Description.—Gall (Figs. 32, 100-101): Rare, on Eucarya hickories, not found east of the Ap- palachian Mountains or Ohio, except for one spec- imen from near the South Carolina Coast; on lower leaf surface, between veins; 2.0-2.5 mm in height, short-obconic, the apex almost flat, expanded lat- erally into flat, circular, horizontal disk with raggedly serrate edge, with small, central umbo at apex usually still bearing thin, circular, disk (the ephemeral cut-out leaf lamina beneath which gall issued; easily detachable and often no longer pres- ent) equal to more than 1/3 expanded gall diame- ter; surface covered with fine, short, brown hairs not obscuring gall surface; base tapering to small, conical pedicel, leaf with some irregular exfolia- tion surrounding connection with gall and show- ing discolored, slight convexity on reverse side of leaf; wall thin, becoming brittle, larval chamber glabrous, longitudinally striate. The architecture of this gall is generally similar to that of C. inanis except that the extraneous tissue surmounting the larval chamber is disposed horizontally rather than vertically and is more papery in C. inanis. Adult and pupa: Unknown. Larva: Third instar: Length 2.1-2.5 mm (n=7). Body elongate-fusiform, anterior end broadly rounded, posterior end tapering to weak point. Head directed anteroventrally. Spatula (Fig. 300) with parallel-sided, apically-notched tooth and narrow shaft. Papillae without setae, raised, espe- cially those of dorsal, pleural, and terminal papil- lae. Lateral papillae 3 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Ab- dominal segments completely covered with ver- rucae. Terminal segment as long as eighth and nearly pointed. Second instar: Not seen. Type material_—Holotype: Third instar, (right- most of 3 larvae on slide), from gall on C. ovata, I 64 rest stop, 2 mi E US 231, Spencer Co., Indiana, IX-20-1995, R.J. Gagné, deposited in USNM. RAYMOND J. GAGNE 27 Etymology.—The name asteris is a Latin adjec- tive meaning starlike with reference to the radiate, flared rim of the gall. Affinities.—See under C. inanis. Collections of galls—ILuinots: Marshall (to- mentosa). INDIANA: Richmond (tomentosa); Spencer Co. (ovata). Mississipri: Noxubee Natl Wildlife Ref., Oktibbeha Co. (tomentosa). On1o: Gypsum (ovata, Wells 1916: Fig. 32); Huron Co. (ovata, Sears 1914: Fig. 13). Specimens of C. asteris examined.—ILLINoIs: Marshall, IX-7—2004 (larva). INDIANA: Richmond, IX-7-2004 (larvae); Spencer Co., IX-20-1995 (lar- vae). Caryomyia biretta Gagné, new species Description.—Gall (Figs. 16, 144): Rare, on Eu- carya hickories, but not E of Appalachian Mts.; on lower leaf surface between veins; 2.5-3.0 mm high, depressed-spheroid, broad apical surface flat to slightly concave with central elongate nip- ple; hairless, not sticky, green to purple; base with central, shallow, central excavation, the leaf slightly convex and discolored on reverse side; larval chamber basal, depressed-ovoid, lined with yellowish membrane, contrasting with purple, woody tissue surrounding larval chamber, espe- cially laterally. Exteriorly this gall simulates that of C. inflata except that the latter does not have the tassel at the apex. The internal shape of these two galls is very different with the larval chamber of C. biretta being surrounded by thick walls and that of C. inflata being thin walled and surmounted by a false chamber. Adult and pupa: Unknown. Larva: Third instar: Length 1.7-2.6 mm (n=5). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 302) with tooth nearly parallel-sided, the apex slightly flaring and weakly notched, the shaft widened anteriorly and tapered behind. Papillae without setae, their bases not raised. Lat- eral papillae 2-3 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Ab- dominal segments covered on dorsal midlength with verrucae and on ventral anterior and poste- rior thirds with horizontal rows of spicules. Ter- minal segment shorter than eighth. Second instar: Length 0.8 mm (n=1). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular tooth and longer, wider, con- cave-sided shaft. Bases of dorsal and pleural papillae not raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material—Holotype: Third instar (right- most specimen of three larvae on slide), from gall on C. ovata, Petersham, Massachusetts, IX-6—2000, R.J. Gagné, deposited in USNM. Etymology.—The name Diretta is a Latin noun in apposition meaning beret, simulated by the outline of this gall. Affinities—See under C. sanguinolenta, from which this species differs in the shape of the gall. Collections of galls——Connecticut: Hopeville (ovata); Meriden (ovata); Phoenixville (ovata). Georcia: Lizella (ovata). ILLinots: Dexter (laciniosa). MAssacuuseTts: Petersham (ovata). New Hampsuire: Barrington (ovata). On10: Hock- ing Co. (Carya sp., Wells 1916: Fig. 16. Wells wrote that his gall was also illustrated in Thompson 1915: Fig. 228, but Thompson’s gall belongs in- stead to C. inflata); St. Clairsville (tomentosa). Specimens of C. biretta examined.—Massa- CHUSETTS: Petersham, IX-6-2000 (larvae). NEw Hampsuire: Barrington, IX-22-1994 (larva). OHIO: St. Clairsville, 6-IX-2004 (larvae). Caryomyia caminata Gagné, new species Description.—Gall (Figs. 7, 108-109): Rare, on Eucarya hickories; on lower leaf surface, between veins; 5 mm in height, columnar, widest near base, tapering to apex, a small aperture present at apex; matte, hairless, green, turning brown; base of gall truncate, leaf with slight exfoliation around connection forming short, tubular socket, a corre- sponding convexity present on opposite leaf sur- face; two-chambered, the basal larval chamber about 1/4 length of gall, ovoid, smooth, longitu- dinally ridged, the apical false chamber also ridged and open or closed apically, the partition between chambers with minute, central perfora- tion. This gall is known from only two specimens, one a figure in Wells (1916). Adult and pupa: Unknown. Larva: Third instar: Length 2.2 mm (n=1). Body elongate-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 297) with anterior tooth gradually narrowing to erose apex, the shaft broad anteri- orly, tapering beyond. Papillae without setae, their bases not raised. Lateral papillae 2 on each side of spatula. Six dorsal papillae on first 7 ab- 28 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) dominal segments. Abdominal segments dorso- laterally with verrucae along line of dorsal papil- lae and with horizontal rows of pointed spicules anteroventrally but not posteroventrally. Terminal segment shorter than eighth. Second instar: Not seen. Type material.—Holotype: Third instar, from gall on C. ovata, I 64 rest stop, 2 mi E U.S. 231, Spencer Co., Indiana, IX-20-1995, R.J. Gagné, de- posited in USNM. - Etymology.—The name caminata is a Latin ad- jective for chimney-like, referring to the general shape of the gall with the apical aperture. Affinities—The gall vaguely resembles an- other bicameral gall, that of C. ovalis, but the wall of the latter is thicker and shiny, not brittle, folia- ceous, and matte as in the present species. The gall of C. tubicola is also elongate-cylindrical and set into a leaf socket, but it does not have a false chamber; also the larva of C. tubicola has only 4 dorsal abdominal papillae, while the present species has 6. Collections of galls——INpIANA: Spencer Co. (ovata). Onto: Hocking Co. (glabra, Wells 1916: Fig. 7). Specimens of C. caminata examined.—INDIANA: Spencer Co., IX-20-1995 (larva, holotype). Caryomyia caryae (Osten Sacken) Diplosis caryae Osten Sacken 1862: 191; Felt 1909:293 (Caryomyia). Hormomyia arcuaria Felt 1908: 388; Felt 1918: 46 (Cary- omyia). New synonym. Description.—Gall (Figs. 18, 20-20a, 55-58): Common, found on Eucarya hickories; single or clustered, on lower leaf surface, usually between veins but may occur on vein; 2.5-3.5 mm in height, spheroidal, sometimes bulging out to one side at midlength, apex with short umbo, less of- ten gall subcylindrical and constricted near basal third; green or yellow, becoming tan to red or brown, hairless, more or less sticky, variably cov- ered with small resin glands; base with weak to prominent conical pedicel in center of circular in- dentation, leaf in most specimens slightly exfoli- ate surrounding base of gall, in subcylindrical form more conspicuously so; wall firm, brittle, of uniform thickness, usually thin, larval chamber shaped as for gall, yellow, brown, or purple, glabrous, with longitudinal ridges. This gall is generally similar to that of C. shmoo, but the gall of the latter is conspicuously narrow at its apical third, while the present gall is more evenly spher- oidal. Adult: Head: Antenna with 12 flagellomeres, in male usually with additional short 13th flagellom- ere without circumfila and in the two New Wa- verly Texas, females, 13-14 flagellomeres; each of the regular 12 male flagellomeres (Fig. 239) bin- odal with definite internode and neck, circumfila loops short, their bases widely separated; female flagellomeres with 2 horizontal circumfila con- nected by 2 vertical strands. Labella separated from one another anteriorly, hemispherical, short- setose, short-setulose. Thorax: Wing length 3.44.0 mm in male (n=10), 2.9-3.9 mm in female (n=10). Male abdomen: Third through eighth ter- gites each with posterior setae separated into two lateral groups, in 1 to 3 horizontal rows; eighth tergite not more weakly sclerotized than seventh. Genitalia: hypoproct nearly parallel sided, weakly concave posteromesally, the margin partly setose; apex of aedeagus convex. Female abdomen: Third through seventh tergites each with poste- rior setae as for male; eighth tergite more weakly sclerotized than preceding tergite, with posterior setae only laterally and not as long as on preced- ing tergite. Ninth segment pliable except for dor- sal sclerite that widens from base to apex, setae sparse laterally and ventrally, many longer than cerci. Cerci ovoid in lateral view, fused dorsally for about 2/3 length. Pupa (Figs. 197-198): Antennal bases oblique in ventral view, converging and narrowing anteri- orly, their rounded apices closely adjacent. Face with anterolateral pair of horizontal incisions, their posterior lip protruding to form concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with several anterior rows of en- larged spinules. Larva: Third instar (Figs. 218-219): Length 2.4-3.6 mm (n=10). Body dorsoventrally flattened, elongate-ovoid, rounded at ends, segments con- vex. Head directed antero-ventrally. Spatula (Fig. 282) with 2 widely separated anterior teeth, shaft widest anteriorly, narrowing abruptly to posterior half. Papillae mostly on short lobes, dorsals and pleurals except for prothorax with setae, also ven- trals with setae on first through seventh abdomi- nal segments. Lateral papillae usually 2, occasion- RAYMOND J. GAGNE 29 ally 3 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Body entirely cov- ered with verrucae. Terminal segment as long as eighth. Second instar: Length 1.0-1.5 mm (n=10). Collar segment with patch of enlarged spicules on each side. Spatula with apically pointed tooth and elliptical or triangular shaft. Bases of dorsal and pleural papillae raised and all short-setose except asetose on prothorax. Six dorsal papillae on first 7 abdominal segments. Type material.—Diplosis caryae Osten Sacken: Lectotype, here designated, third instar (excised from galls of the more common, less robust, sub- spherical gall form), from gall on Carya sp., Wash- ington, DC., collected by Osten Sacken, deposited in USNM. Paralectotypes, same data as lectotype, a second instar excised from a separate gall, and one remaining gall, in USNM; also male(s) and fe- male(s), mentioned by Osten Sacken (1862) but now lost. For unstated reasons, Felt (1921) wrote that the adult of Osten Sacken’s type series “could hardly be a Caryomyia” and was “probably an in- quiline.” Nothing about Osten Sacken’s descrip- tion of the adult would disallow it as a Caryomyia. Perhaps Felt objected to the April date of emer- gence that Osten Sacken (1862) reported. Felt (1921) thought that the adult emerged “probably early in June,” possibly because that is when galls of this species become apparent, but my rearings show that adults of this species and other Cary- omyia species emerge in March and April in the Washington, DC area. Hormomyia arcuaria Felt: Holotype.d, vicinity of Worcester, Massachusetts, reared from Carya sp., Thompson # 254-255, Felt #C1104z, deposited in Felt Collection. This specimen came from a smooth, subglobular, nippled gall (Felt 1908). Felt (1921) later added that this male was separated from a series of adults reared from an apparently typical lot of C. caryae galls. The holotype is a ten- eral specimen of C. caryae with the flagellomeres not yet fully expanded (cf. Felt 1921: 99, fig. 9), which may be the reason Felt differentiated the specimens from the remainder of the series. Etymology.—The name caryae means “of Carya,” and arcuaria means arcuate. The latter name was evidently coined for the arched anten- nal circumfila as drawn in Felt (1921: 99, fig. 9). Affinities—As mentioned in the gall descrip- tion, two intergrading kinds of galls are formed by C. caryae as defined here (compare Figs. 55-56 with 57-58). This species is apparently related to C. hirtiglobus and C. shmoo because of the general conformation of the gall with its acute-conical connection to the leaf and similarity of the larvae, most conspicuously in the widely spaced teeth of the third instar spatula. Males of all three species have comparably shaped flagellomeres and simi- lar genitalia and abdominal tergite setation. Third instars of this group of three species have setae on the dorsal and pleural papillae except that C. caryae lacks them on the prothorax; C. caryae and C. shmoo carry the same pattern along on the sec- ond instar but C. hirtiglobus lacks any papillar se- tae on that instar. Pupae of C. caryae have antennal bases slightly longer than those of C. hirtiglobus and C. shmoo, and differ also in that the apices touch one another. Biological notes.—In central Maryland, galls were first noticed in late May when they con- tained first and second instars, but the galls were immature, soft and very sticky. Small, barely no- ticeable adjacent galls with first instars were rec- ognizable but smaller and stickier. Through the first week of August most galls are brittle and con- tain second instars, but during this period occa- sional galls may contain a third instar. After early August galls usually contain third instars. Adults were reared in early April the following year. Collections of galls.—Collections of the sub- cylindrical form of this gall are marked with an as- terisk in this list. In addition to the following galls from Canada and United States, I have photo- graphs of some of the subcylindrical gall type from Tamaulipas, Mexico. ALABAMA: Ashville (fomen- tosa); Auburn (glabra, ovata, pallida, tomentosa*); Birmingham (pallida); 2 mi S DeQueen (texana, to- mentosa). Greenville (tomentosa); Oak Mt. St. Pk, Shelby Co. (ovata*, pallida*). ARKANSAS: Beulah (to- mentosa); Chester (texana); 3 mi S Harrison (tomen- tosa*); London (tomentosa); Magazine Mt., Logan Co. (ovata*, texana*, tomentosa*); Osage (texana); Palestine (ovata); Piney (texana). CONNECTICUT: Amston (ovata); Danbury (ovata); Hartland (ovata); Harwinton (ovata); Hopeville (ovata); Litchfield (ovata); Meriden (ovata); New Haven (glabra); Oakville (glabra, ovata); Phoenixville (ovata); Plain- field (ovata); Southbury (glabra*, tomentosa*); Southington (ovata); Westchester (ovata). FLORIDA: Avon Park (floridana); Chattahoochee (glabra, to- mentosa); Gainesville (glabra, tomentosa); Lake City (tomentosa); GEorGIA: Carnesville (glabra); Forsyth 30 THE GALL MIDGES (DIPTERA: CECIDOMYTIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) (glabra, ovata); 8 km E Fort Valley (pallida); Lake Park (glabra); Lizella (ovata, tomentosa). ILLINOIS: Dexter (laciniosa, tomentosa); Highland (laciniosa); Marshall (tomentosa*). INDIANA: Richmond (tomen- tosa); Spencer Co. (ovata). KENTUCKY: Brownsville (tomentosa); Grayson Lake St. Pk, Carter Co. (to- mentosa); Morehead (tomentosa); My Old Kentucky Home St. Pk, Nelson Co. (ovata*); Pennyrile St. Pk, Christian Co. (ovata); Winchester (laciniosa, ovata). LouIsiIANa: Lake Bistineau St. Pk, Webster Parish (texana, tomentosa). MARYLAND: Beaver Dam (to- mentosa); Beltsville, USDA Agricultural Research Center (glabra, tomentosa); Brighton (tomentosa*); Calverton (tomentosa); Clarksburg (glabra, tomen- tosa); College Park (tomentosa); Ellicott City (tomen- tosa); Gaithersburg (tomentosa*); Lusby (glabra, to- mentosa); Piney Grove (ovata); Potomac (tomentosa); Ridgely (glabra, tomentosa); St. Leonard (tomentosa*); Silver Spring (tomentosa); Snow Hill (tomentosa); Spencerville (tomentosa); Wheaton (glabra, tomentosa). MassacuusetTts: Bolton (ovata); Great Barrington (ovata); Holyoke (ovata); Leomin- ster (ovata); Merrimac (ovata); Petersham (ovata); Worcester (tomentosa); Wrentham (ovata). MIssIs- sippi: Natchez Trace Pkwy, Mile 193, Choctaw Co. (glabra, pallida, tomentosa); Noxubee Natl Wildlfe Ref., Oktibbeha Co. (tomentosa); 7 mi E Starkville (ovata, pallida, tomentosa). Mussourt: Conway (ovata, texana); Crowley Ridge Roadside Pk, Stod- dard Co. (glabra, tomentosa); Holly Ridge State For., Stoddard Co. (glabra*, tomentosa*); Rolla (texana); Wildwood (tomentosa). NEw Hampsuire: Barring- ton (ovata); Dover (ovata); Exeter (ovata); Seabrook (ovata). New York: Albany (Carya sp.); E. Schodack (Carya sp.); Herkimer (ovata); Ithaca (ovata); Nas- sau (Carya sp.); New York (Carya sp.); Ravena (glabra, ovata); Saratoga Springs (ovata); Shokan (glabra). NorTH CAROLINA: Alamance (tomentosa); Alberta (glabra); Dortches (glabra, tomentosa); Prov- idence (glabra, tomentosa); Rowland (tomentosa); Wise (tomentosa). Onto: Hocking St. For., Hocking Co. (glabra, ovata); Hocking Co. (ovata, tomentosa, Wells 1916: Figs. 18, 20—20a); St. Clairsville (tomen- tosa); Summerford (tomentosa); Yellow Springs (glabra). OKLAHOMA: Sallisaw (tomentosa). ON- TARIO: Dundas (ovata). PENNSYLVANIA: Kirby (ovata); Julian (glabra); Millsboro (glabra); Schellsburg (glabra, ovata). SouTH CAROLINA: Dillon (tomentosa); Georgetown (tomentosa). TENNESSEE: Crossville (pallida); Dandridge (tomentosa*); Dickson (tomen- tosa). TAMAULIPAS: SW of Ciudad Victoria (ovata, var. mexicana; photos only). Texas: Daingerfield St. Pk, Morris Co. (tomentosa); New Waverly (tomen- tosa). VIRGINIA: Warren Co., Blue Ridge Pkwy Mile 4.6 (ovata); Carson (glabra); Dinwiddie (glabra, to- mentosa); Huntley Meadows Pk, Fairfax Co. (to- mentosa); Ladysmith (tomentosa); Radford (tomen- tosa); Skippers (tomentosa). WASHINGTON, DC: National Arboretum (glabra, tomentosa); Walter Reed Army Medical Center (tomentosa). WEsT ViR- GINIA: 6 mi. NW Hedgesville (ovata, tomentosa). Specimens of C. caryae examined.— ALABAMA: Auburn, IX-19-2000 (larvae). ARKANSAS: Beulah, IX-10-2004 (larva); Chester, IX-9-2004 (larvae); 3 mi S Harrison, VII-22-1988 (larva); Osage, IX- 9-2004 (larva). CONNECTICUT: Amston, IX-19-1994 & IX-17-1999 (larvae); Phoenixville, IX-20-1994 (larvae). FLoripA: Avon Park, XI-7—13-2002, K. Hi- bbard (larva). GeorGiA: 8 km E Fort Valley, IX- 11-1993, J. & R. Payne (larvae; 2 °, pupae, em. IV- 1994). Ittinors: Marshall, IX-7—2004 (larvae). INDIANA: Spencer Co., IX-20-1995 (larva). KEN- Tucky: Brownsville, IX-21-1995 (larvae); My Old Kentucky Home St. Pk, Nelson Co., VII-1988 (larva); Winchester, IX-22-1995 (larva). Mary- LAND: Beltsville, USDA Agricultural Research Center, V-5-1998, VII-1-1998, VII-5-1998 & VII- 1-2000 (larvae); Brighton, X-10-1993 & X-8-1994 (larvae; 5 3,5 ¢, pupae, em. IV-1995); Clarksburg, X-4-2005 (pupae, 19 d, em. IV-5 to 17-2006); Gaithersburg, X-4-1994, VII-20-2000 (larvae); Lusby, IX-14-2005 (pupae, 2, em. IV-2-2006); Ridgely, X-1999 (larvae; 6, 2, pupae, em. IV- 2000); Silver Spring, IX-17-1993 & X-24-1993 (lar- vae); Spencerville, IX-18—2004 (larvae); Wheaton, IX-19-1993 (larvae). MASSACHUSETTS: Leominster, IX-21-1994 (larvae); vic. Worcester, M.T. Thomp- son (larva; d (holotype of C. arcuaria), °); Wren- tham, IX-20-1994 (larvae). Missourr: Conway, IX- 8-2004 (larva); Holly Ridge State For., Stoddard Co., VII-1988 (larva). New HAmpsuire: Barrington, IX-22-1994 (larvae; 5, 2, pupae, em. IV-1995); Dover, IX-21-1994 (larva); Exeter, [X-21-1994 (larva). NEw York: E. Schodack, E.P. Felt (larva); Nassau, E.P. Felt (16 6, 13 2); New York, E.P. Felt (larva; 3°); Saratoga Springs, IX-25-1994 (larvae). Onto: Hocking St. For., Hocking Co., IX-19-1995 (larva), Yellow Springs, IX-7—2004. OKLAHOMA: Sallisaw, IX-9-2004 (larvae). Ontario: Dundas, VII-17-1995 (larvae). TENNESSEE: Dandridge, IX- 12-2004 (larva). Texas: New Waverly, X-6—2001 (3 5,10 2, pupae, em. IV-2002). WASHINGTON, DC: type series of C. caryae, locality not specified fur- ther, C. glabra); National Arboretum, X-13-1997 RAYMOND J. GAGNE 31 (larvae); Walter Reed Army Medical Center, IX- 2000 (larvae). West VIRGINIA: 6 mi. NW Hedgesville, IX-12 & 19-1999 (larvae). Caryomyjia caryaecola (Osten Sacken) Cecidomyia caryaecola Osten Sacken 1862: 192; Felt 1915: 222 (Caryomyia). Description.—Gall (Figs. 12-13, 70-71).—Com- mon, on Eucarya hickories; single or in groups, at- tached to vein on lower leaf surface; 4.8-8.0 mm in height, onion-shaped, upright to leaning, tapered from rounded base to pointed apex; hairless, glabrous, green or yellow, becoming tan to brown; base broadly rounded with large, central conical pedicel in deep, circular indentation; wall hard, of uniform thickness, larval chamber shaped as for gall, glabrous, with longitudinal ridges. This gall vaguely resembles those of C. caryae and C. shmoo in its connection to the leaf and its hard, hairless surface, but it is definitely onion-shaped and, un- like the other two, it is always attached to veins instead of the lamina. Adult (female only): Head: Antenna with 12 flagellomeres, flagellomeres with 2 or, on more basal flagellomeres, a third horizontal circum- filum connected by vertical strands. Labella sepa- rated from one another anteriorly, hemispherical, short-setose, short-setulose. Palpus 4 segmented. Thorax: Wing length 2.5 mm (n=1). Abdomen: Third through seventh tergites each with poste- rior setae continuous across width,.in 2-3 rows laterally, tapering to 1-2 rows medially; eighth tergite with 3-4 weak lateral posterior setae. Ninth segment (generally as for Fig. 260, but not well displayed on slide mount) pliable except for slender dorsal sclerite, setae shorter than cerci, in two groups, one dorsolateral, one ventral. Cerci elongate-ovoid in lateral view, fused dorsally on basal third. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 2.6-4.4 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 286) with short tooth, barely narrowing from base to apex, nearly parallel-sided, shal- lowly notched apically, on some specimens merely erose, and with elongate, narrow shaft be- coming broader anteriorly. Papillae without setae, bases of dorsals and pleurals somewhat raised. Lateral papillae usually 4, occasionally 3, on each side of spatula. Six dorsal papillae on first 7 ab- dominal segments. Abdominal segments exten- sively covered with verrucae. Terminal segment shorter than eighth. Second instar: Length 0.7-1.1 mm (n=5). Collar segment with patch of enlarged spicules on each side. Spatula with acutely trian- gular anterior tooth and longer, wider, concave- sided shaft. Bases of dorsal and pleural papillae raised, papillae with long setae. Six dorsal papil- lae on first 7 abdominal segments. Type material.—Syntypes: Four galls from C. tomentosa (as C. alba), Washington, DC, C. Osten Sacken, deposited in USNM. Etymology.—The name caryaecola is a noun meaning hickory inhabitant. Affinities—This species may be related to C. caryae (q.v.) and its two close relatives. As in those species, C. caryaecola has setose papillae in the sec- ond instar but, unlike them, the third instar papil- lae are asetose. The pupae of C. caryaecola have an- tennal bases with apicolateral points unlike those of C. caryae and relatives in which the pointed apices are mesally situated. The gall of the present species shares with the other three a similar re- cessed, conical base and thin, but hard wall. Biological notes.—In central Maryland, galls appear by May 23. At first they are light green and soft, but grow very quickly so that by June 9 and until June 15, galls are mostly full-size, green, and brittle, and still contain first instars. By June 17, most galls are hardened and contain second in- stars. Until the second week of August most galls still contain second instars, but an occasional gall contains a third instar. Adults emerge in spring from a subapical hole near the thinnest part of the gall. In Sallisaw, Oklahoma, I saw hundreds of par- tially developed galls that contained dead, dried first instars. All apparently died of the same cause and gall growth was terminated. Those partially developed galls correspond to the gall drawn in Wells (1916, Fig. 13), a stunted example of the one of the same species he drew in his Fig. 12. Collections of galls—AtaBaMa: Auburn (glabra, ovata, tomentosa); Oak Mt. St. Pk, Shelby Co. (ovata). ARKANSAS: Beulah (tomentosa); Chester (texana); London (tomentosa); Magazine Mt., Logan 32 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Co. (texana); Piney (texana). CONNECTICUT: Amston (ovata); Meriden (ovata); New Haven (glabra); Phoenixville (ovata); Plainfield (ovata); Westchester (ovata). GrorGia: Forsyth (glabra, ovata); Lizella (ovata, tomentosa. ILLiNois: Highland (laciniosa); Marshall (tomentosa). INDIANA: New Castle (glabra); Spencer Co. (ovata). KENTUCKY: Pennyrile St. Pk, Christian Co. (glabra); Winchester (laciniosa). MAINE: Eliot (tomentosa). MARYLAND: Beltsville, USDA Agricultural Research Center (glabra, tomen- tosa); Clarksburg (tomentosa); College Park (glabra); Ellicott City (tomentosa); Gaithersburg (tomentosa); Kemp Mill (glabra); Lusby (glabra, tomentosa); Ridgely (tomentosa); St. Leonard (tomentosa); Silver Spring (glabra); Spencerville (tomentosa); Wheaton (glabra). Massacuusetts: vic. Worcester (Carya sp., Thompson 1915: 89, Fig. 226); Wrentham (ovata). Mississippi: Natchez Trace Pkwy, Mile 123, Rankin Co. (tomentosa); nr. Sessums, Oktibbeha Co. (ovata). Missouri: Conway (texana); Holly Ridge State For., Stoddard Co. (glabra, tomentosa); Wildwood (to- mentosa). NEW HAMPSHIRE: Kingston (ovata). NEw York: Nassau (Carya sp.); New York (Carya sp, Beutenmiiller 1904: 27, fig. 57). NORTH CAROLINA: Alberta (glabra), Providence (glabra, tomentosa). Onto: Hocking Co. (Carya spp., Wells 1916: Figs. 12, 13); Huron Co. (C. glabra, Sears 1914: Fig. 17). OKLAHOMA: Sallisaw (tomentosa). TENNESSEE: Greene Co., 181, Mile 38.5 (tomentosa); Jackson (to- mentosa); Lebanon (glabra); Mohawk (tomentosa). VirGiniA: Warren Co., Blue Ridge Pkwy Mile 4.6 (ovata); Huntley Meadows Pk, Fairfax Co. (tomen- tosa); Ladysmith (tomentosa); 181, Mile 13.9, Wash- ington Co. (ovata). WASHINGTON, DC: Walter Reed Army Medical Center (tomentosa); unspecified (Carya sp. (syntypes)). West VircInia: Beech Fork St. Pk, Wayne Co. (tomentosa); Cacapon (ovata); Cooper’s Rock St. Pk, Preston Co. (glabra); 6 mi NW Hedgesville (ovata). Specimens of C. caryaecola examined.—Con- NEcTICUT: Amston, IX-19-1994 (larvae). KENTUCKY: Winchester, IX-22-1995 (larvae). MARYLAND: Beltsville, USDA Agricultural Research Center, IX-28-1999, VII-1-2000, & VII-25-2000 (larvae); Clarksburg, V-23—2000 (larvae) & X-4—2005 (pupal exuviae, IV-1—2006, & pupa, taken from gall IV- 12-2006); Gaithersburg, X-4-1994 (larva); Kemp Mill, IX-6-2003 (larvae); Lusby, IX-16-2003 (¢, pupae, em. IV-10—2004); Spencerville, IX-7—2004 (larvae); Wheaton, X-12-—1997 (larvae). MASSACHU- sETTS: Wrentham, IX-20-—1994 (larvae). OKLAHOMA: Sallisaw, IX-9-2004 (larvae). TENNESSEE: Toma- hawk, IX-12-2004 (larvae). Vircinia: Huntley Meadows Pk, Fairfax Co., VIH-30-2000 (larvae). WEST VIRGINIA: 6 mi NW Hedgesville, VIII- 27-2000 (larvae). Caryomyia cilidolium Gagné, new species Description.—Gall (Figs. 51-52).—Occasional, known only from C. cordiformis of the Apocarya section; found singly, usually on lower leaf sur- face, rarely on upper, usually between but occa- sionally on veins; 3.1-3.9 mm in height, spher- oidal, slightly longer than wide, base broadly rounded to truncate in profile, apex with small central nipple surrounded by extensive areola that with pressure can separate from gall; surface not sticky, occasionally slightly bumpy, tan to brown, with few to many short, fine, white hairs not obscuring surface, areola usually paler than surrounding surface; base with deep, wide, circu- lar indentation; wall firm, brittle, almost uni- formly thin, larval chamber glabrous, with longi- tudinal ridges. For notes on galls of related species, see below under affinities. Adult: Head: Antenna with 12 flagellomeres. Each of the 12 male flagellomeres (as in Fig. 240) evenly cylindrical, girdled by 8-9 more or less par- allel, occasionally interconnected, appressed cir- cumfila interspersed with setae and covered with setulae. Female flagellomeres with anastomozing circumfila on venter (as in Fig. 241). Labella fused, the pair semicircular, setulose but asetose. Thorax: Wing length 2.1-2.2 mm (n=2) in male, 3.1 mm (n=1) in female. Male abdomen: Third through seventh tergites each with posterior setae continu- ous across width in mostly double, partly triple row; eighth tergite weakly sclerotized, with short posterior setae. Genitalia (Fig. 251): hypoproct broadest posteriorly and expanded posteroven- trally to partly cover sides of aedeagus, each of the 2 lobes with 2-3 apical setae, dorsal surface com- pletely covered with setulae; aedeagus tapering gradually from wide base to narrow apex; gonos- tylus dorsoventrally flattened, the tooth large, dorsoventrally compressed. Female abdomen: Third through seventh tergites each with posterior setae in 3 horizontal rows laterally, thinning to 2, then one, with slight medial interruption in setae; eighth tergite more weakly sclerotized than pre- ceding tergite, with weak posterior setae numer- ous laterally, continuing mesally in single, sparse row. Ninth segment (as in Fig. 261) pliable except for slender dorsal sclerite, with numerous apico- lateral and ventral setae, none longer than the RAYMOND J. GAGNE 33 cerci, the two groups separated laterally. Cerci elongate-ovoid in lateral view, pointed apically, fused dorsally for about 1/2 length. Pupa (as in Figs. 193-194): Antennal bases elon- gate, in ventral view parallel, closely adjacent, rounded anteriorly, the apices dorsoventrally flat- tened. Face with longitudinal impression running along length, a slight anteromesal convexity pos- terior to each antennal base, and without antero- lateral pair of horizontal incisions. Frons and clypeus slightly convex. Abdominal tergal spin- ules all similarly small. Larva: Third instar (Figs. 230-231): Length 2.0—2.9 mm (n=10). Body dorsoventrally flattened, elongate-ovoid, broadly rounded anteriorly, spindleform posteriorly. Head directed ventrally, surmounted dorsally by collar segment. Spatula (Fig. 277) with pointed anterior tooth and broad shaft. Papillae prominently raised above sur- rounding surface, without setae. Lateral papillae 2 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Body covered with verru- cae. Terminal segment as long as eighth, rounded. Second instar: Length 1.0 1.4 mm (n=10). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Six dorsal papillae on first 7 abdominal segments. Bases of dorsal and pleural papillae raised, papil- lae asetose. Type material—Holotype: Third instar, from gall on C. cordiformis, Ellicott City, Maryland, IX- 2-2000, R.J. Gagné, deposited in USNM. Etymology.—The name cilidolium is a noun combining the Latin ciliatus (fuzzy) and dolium (jar), with reference to the fuzzy, barrel-like gall. Affinities —Caryomyia cilidolium, C. hirtidolium, C. tuberidolium, and C. viscidolium form similarly shaped, spheroidal, thin walled galls (Figs. 43-52). At their apex is a large, circular areola surrounding a small central nipple. With a little pressure the cir- cular apex can be broken off as a unit from the rest of the gall and forms the exit from which pupae emerge in spring. Galls of C. cilidolium occur only on bitternut of the Apocarya section of Carya, usu- ally on the lower surface, and are covered with sparse, fine, short hairs. The other three species of this group occur on the Eucarya section. Galls of C. hirtidolium are hairy, much more so than those of C. cilidolium, so that the long hair obscures the gall surface. Galls of C. tuberidolium and C. viscidolium are both sticky, the former very much so and also bumpy and more spherical, the latter merely slightly resinous and smooth. A major difference between these two last species is that C. tuberi- dolium occurs on the underside of the leaf (except for one example known to me) while C. viscidolium occurs on the upperside. This difference indicates a different egg-laying strategy by the female. The pupal stages are known for all except C. hirtidolium, both sexes for C. cilidolium and C. visci- dolium, and the female for C. tuberidolium. The known pupal and adult stages are each very simi- lar among species, as are the larvae. Females of this group have the shortest ovipositors for the genus, and the pupae and male genitalia are dis- tinctive. Pupae of this group lack spiny setulae on the pupal abdomen, as do C. albipilosa and C. strio- lata, but the second instars, unlike those two species, have a spatula. One of the most striking characteristics of this group is the male antenna that, instead of having three separate circumfila as do all other Cary- omyia, is instead girdled by 8-10 occasionally in- terconnected, unlooped circumfila reminiscent of those found in Polystepha of the tribe Asphondyli- ini. Females also have extra anastomozing strands of circumfila on the venter of the flagellomeres. Labella are completely joined mesally and lack se- tae, unlike all other Caryomyia species. Pupae in this group of species are distinct from all other Caryomyia in the shape of the antennal bases and the lack of horizontal facial incisions with pro- truding lower lips, but they do have a characteris- tic anteromesal facial prominence and a longitudi- nal facial furrow. The pupae show some similarity to that of C. urnula, except that the antennal horns are differently shaped. The third instars of this group are active, large, dorsoventrally flattened and have a spatula with a single anterior tooth. The head is situated ventrally with the dorsal part of the neck segment projecting over it. The dorsal and pleural papillae are on prominent lobes. The posterior end of the larva is generally rounded and as long as the previous segment. The larval spatulas of the four species in this group are each distinctive. That of C. cilidolium has the anterior corners of the shaft rounded, that of C. viscidolium has a very long tooth compared to the shaft, and the shaft of C. tuberidolium is greatly widened anteriorly and posteriorly. Biological notes.—In mid-June in the eastern panhandle of West Virginia fresh galls were green and brittle and with either second or third instars. In London, Arkansas, galls collected on Sept. 14 still contained second instars. 34 THE GALL MIDGES (DIPTERA: CECIDOMYITDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Collections of galls (all from Carya cordi- formis).—ARKANSAS: London. CONNECTICUT: Meri- den, Thomaston. ILLINots: Highland. INDIANA: Plainville. MaryYLAND: Cabin John; Ellicott City; Kensington; Long Beach; Potomac; Ridgely; Wheaton. Missourt: Holly Ridge State For., Stod- dard Co.; Oregon Co., junc. Rts. 11 & 142. New Jersey: Columbia. New York: Ithaca; Ravena. PENNSYLVANIA: Pavia. TENNESSEE: Gatlinburg. VER- MONT: Guilford. WesT VIRGINIA: Falling Waters. Specimens of C. cilidolium examined.— ARKANSAS: London, IX-10-2004 (larvae). Con- NEcTicuT: Meriden, [X-16—1992 (larvae). ILLINOIs: Highland, IX-8-2004 (larva). INDIANA: Plainville, IX-7-2004 (larva). MARYLAND: Ellicott City, IX- 2-2000 (larvae); Kensington, IX-28-2000 (larva); Ridgely, IX-23-1999 (larvae); Wheaton, IX-1-1994 & VIII-11-2000 (larvae). New York: Ithaca, IX- 3-2003 (larvae). VERMONT: Guilford, IX-7—2005 (larvae; 1 d, 1 ¢, pupae, em. IV-15 to 28-2006). WesT VirGINIA: Falling Waters, VI-21—2004 (lar- vae), X-2005 (larva; 2 d, pupae, em. IV-12—2006). Caryomyjia conoidea Gagné, new species Description.—Gall (Figs. 137-138): Occasional, on Eucarya hickories; single or in groups, on lower leaf surface between veins; height 3.8-5.0 mm, generally conical, edge at base rounded, base ad- jacent to leaf uniformly hard, conical and often with cylindrical extension at apex, sides develop- ing longitudinal ridges and furrows upon matu- rity; green to red or brown surface smooth, with sparse, short, erect, white to tan hairs not obscur- ing surface; base with central, circular, shallow depression, leaf with short, irregular exfoliation surrounding basal connection to gall; larval cham- ber basal, depressed-ovoid, lined with yellowish membrane, contrasting with brown to purple, large-celled, spongy, viscous tissue surrounding larval chamber that shrivels with age and tra- versed by bundle of fibers between larval cham- ber and gall apex. This gall is most similar to that of C. sanguinolenta, q.v. for remarks. Adult (Fig. 183-185): Head: Antenna with 12 flagellomeres; male flagellomeres scarcely bi- nodal (as in Fig. 237), only slightly narrowed be- yond basal circumfilum, circumfila loops short, subequal in length; female flagellomeres with 2 horizontal circumfila connected by 2. vertical strands. Labella separated from one another ante- riorly, hemispherical, short-setose, short-setulose. Palpus 3 to 4 segmented. Thorax: Wing length 2.2-2.7 mm in male (n=5); 3.1-3.5 mm in female (n=5). Male abdomen: Third through seventh ter- gites each with posterior setae continuous across width, in mostly single row; eighth tergite with several posterolateral setae. Genitalia (as in Figs. 247-248): hypoproct broadest near apex, slightly concave posteromedially, margin setose except in concavity, dorsal setulae only on distal third; aedeagus furrowed apically and ventrally. Gonos- tylus broad at apex. Female abdomen (Figs. 183-185): Third through seventh tergites each with posterior setae continuous across width, in mostly single row, usually double at lateral limits; eighth tergite not less sclerotized than seventh and posterior row of setae in mostly single row but double laterally. Ninth segment pliable except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ven- trally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.8-2.8 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 312) with long, nearly parallel sided, apically notched tooth and long shaft. Papillae without setae, their bases not raised. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdomi- nal segments with rounded verrucae extensively covering dorsum and with horizontal rows of spicules on venter anterior to sternal papillae and posterior to ventral papillae. Terminal seg- ment shorter than eighth. Second instar: Length 1.0-1.4 mm (n=5). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and_ longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae slightly raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal seg- ments. Type material—Holotype: Third instar, left specimen of two on slide, from gall on C. glabra, Wheaton Park, Montgomery Co., Maryland, X- 12-1997, R.J. Gagné, deposited in USNM. Etymology—The name conoidea is a Latin ad- jective referring to the conical shape of the gall. RAYMOND J. GAGNE 35 Affinities—See under C. sanguinolenta, from which this species differs in the shape of the gall. Biological note.—In spring the pupa emerges from the base of the gall (as in Fig. 136). Collections of galls—Connecticut: Amston (ovata); Danbury (ovata); Hartland (ovata); Harwin- ton (ovata); Litchfield (ovata); Meriden (glabra); New Haven (glabra); Oakville (ovata); Phoenixville (ovata); Plainfield (ovata); Thomaston (ovata); Westchester (ovata). KeENTucKy: Grayson Lake St. Pk, Carter Co. (tomentosa); Winchester (laciniosa). Maine: Eliot (tomentosa); North Berwick (ovata). MaryLanb: Ellicott City (tomentosa); Gaithersburg (tomentosa); Lusby (tomentosa); Wheaton (glabra). MassacuuseTts: Holyoke (ovata); Leominster (ovata); Merrimac (ovata); Petersham (ovata); Wren- tham (ovata). New Hampsuire: Barrington (ovata); Dover (ovata); Exeter (ovata). New York: E. Schodack (Carya sp.); Herkimer (ovata); Ithaca (ovata); Ravena (glabra, ovata). NORTH CAROLINA: Kannapolis (glabra). Onto: Hocking St. For., Hock- ing Co. (ovata). ONTARIO: Dundas (ovata). PENNSYL- VANIA: Julian (glabra); Pavia (ovata); Schellsburg (ovata). WASHINGTON, DC: National Arboretum (glabra). WEST VIRGINIA: Cooper’s Rock St. Pk, Pres- ton Co. (glabra); 6 mi. NW Hedgesville (tomentosa). Specimens of C. conoidea examined.—Con- NEcTiIcuT: Amston, [X-19-1994 (larvae); Hartland, IX-7-2005 (larvae; 5 3,3 2, pupae, em. IV-15 to 28-2006); Harwinton, IX-6—-2005 (larva); New Haven, IX-19-1994 (larvae); Phoenixville, IX- 20-1994 (larvae); Plainfield, VIII-27—1988 (larvae). Kentucky: Winchester, IX-22-1995 (larvae). Maine: North Berwick, IX-21-1994 (larva). Mary- LAND: Ellicott City, IX-20-2000 (larva); Gaithers- burg, X-4-1994 & VIII-20-2000 (larvae); Wheaton, X-12-1997 (larvae). MAssAcHusETTs: Petersham, IX-6-2000 (2 5, 2, em. IV-2001); Wrentham, IX- 20-1994 (larvae). New Hampsuire: Barrington, IX- 22-1994 (larvae; 10 5,9 2, pupae, em. IV-1995); Exeter, IX-21-1994 (larvae). NorTH CAROLINA: Kannapolis, IX-18-2000 (larvae). ONTARIO: Dun- das, VIII-17-1994 (larvae). WASHINGTON, DC: Na- tional Arboretum, X-28-2002 (larvae; 4 3,5 2, pu- pae, em. IV-2003). West VircINiIA: 6 mi. NW Hedgesville, X-1-1995 & VIII-8-2000 (larvae). Caryomyia cucurbitata Gagné, new species Description.—Gall (Figs. 149-151): Infrequent, on Eucarya hickories; on lower leaf surface, single or in row along major vein; length 3.0-5.0 mm, spheroidal at base, abruptly narrowed to slender neck of variable length, digitate at apex; yellow with spherical, red, resinous dots and sparse long or short white hair not obscuring surface, the neck brown or black, with white hairs; base with circu- lar, shallow, central excavation; larval chamber basal, ovoid, white, surrounded by green tissue turning brown and woody; a bundle of fibers present between larval chamber and gall apex. The gall shows some similarity with that of C. ansericollum but the latter is not covered with resinous dots and occurs only on bitternut. Also generally close in shape is the gall of C. echinata but the latter is covered with long, stiff, straight hairs and occurs on the leaf lamina rather than the vein. Adult: unknown. Pupa (Figs. as for 213-214): Antennal bases py- ramidal, moderately separated, acutely pointed apicolaterally. Face with anterolateral pair of hori- zontal incisions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with several anterior rows of enlarged spinules. Larva: Third instar: Length 1.7—2.5 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 308) with long, nearly parallel-sided, apically-notched tooth and long shaft, widest an- teriorly, tapered beyond. Papillae without setae, bases not raised. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 ab- dominal segments. Abdominal segments with rounded verrucae covering dorsum at midlength and horizontal rows of spicules on venter ante- rior to sternal papillae and posterior to ventral papillae. Terminal segment shorter than eighth. Second instar: Length 0.9-1.1 mm (n=7). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae not raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. tomentosa, Lusby, Maryland, IX-16—2003, R.J. Gagné, deposited in USNM. Etymology.—The name cucurbitata, a Latin ad- jective meaning pumpkin-like, refers to the gen- eral resemblance of the gall to a pumpkin. Affinities.—See discussion under C. inclinata. Collections of galls—ALaBama: Auburn (to- mentosa); Greenville (tomentosa). ARKANSAS: Chester (fexana). KENTUCKY: Brownsville (fomen- 36 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) tosa); My Old Kentucky Home St. Pk, Nelson Co. (ovata, tomentosa). MARYLAND: Beltsville, USDA Agricultural Research Center (tomentosa); Clarks- burg (tomentosa); Ellicott City (tomentosa); Gaith- ersburg (tomentosa); Lusby (tomentosa); Ridgely (to- mentosa). Missouri: St. Clair (tomentosa). NORTH CAROLINA: Alamance (tomentosa). Onto: Hocking St. For., Hocking Co. (tomentosa). TENNESSEE: Mohawk (tomentosa). WEST VIRGINIA: 6 mi NW Hedgesville (ovata). Specimens examined.—ALABAMA: Auburn, IX- 19-2000 (larvae); Greenville, IX-20—2000 (larvae). ARKANSAS: Chester, IX-9—2004 (larvae). KENTUCKY: Brownsville, IX-21-1995 (larvae). MARYLAND: Beltsville, USDA Agricultural Research Center, V- 5-1994 & VIII-31-2004 (larvae); Clarksburg, X- 42005 (larvae; pupa excised from gall III- 28-2006); Gaithersburg, X-4—1994 (larvae); Lusby, IX-16-2003 (larvae); Ridgely, IX-23-1999 (larvae). Missouri: St. Clair, IX-8—2004 (larva). NorTH CAR- OLINA: Alamance, IX-18-2000 (larvae). Onto: Hocking St. For., Hocking Co., IX-19-1995 (lar- vae). TENNESSEE: Mohawk, IX-12—2004 (larvae). WEsT VIRGINIA: 6 mi NW Hedgesville, VIII- 27-2000 (larvae). Caryomyia cynipsea (Osten Sacken) Cecidomyia cynipsea Osten Sacken 1862: 193; Felt 1918: 47 (Caryomyia); Gagné 2004: 276, as unplaced sp. of Ce- cidomyiinae. Description.—Gall (Figs. 162-164): Rare, on Eucarya hickories; on lower leaf surface, along midrib; 2.6-5.0 mm in height, irregularly sphe- roid, smooth, hairless, yellow turning brown; base with irregular, shallow, central excavation; woody, polythalamous, with numerous, discrete, irregularly ovoid, smooth walled, larval cham- bers. This is the only polythalamous gall formed by a Caryomyia. Some other species, e.g., C. aggre- gata, may appear to be so because their galls will occasionally be found so crowded as to be par- tially fused, but below each larval chamber will be a basal connection to the leaf. The gall of C. cynipsea has a single connection to the vein. Adult (female only): Head: Antenna with 12 flagellomeres; female flagellomeres mostly with 2 horizontal circumfila connected by 2 vertical strands. the basal circumfilum on the 2 basal fla- gellomeres doubled. Labella separated from one another, rounded anteriorly, hemispherical, short- setulose, without setae. Palpus 4 segmented. Tho- rax: Wing length 3.6 mm in female (n=1). Female abdomen: Third through eighth tergites usually with posterior setae in 2-3 lateral rows, some- times almost joined mesally, especially on poste- rior segments, setae not as long or strong on eighth as on previous tergite. Ninth segment (Fig. 264) broad, pliable except for slender dorsal scle- rite, setae shorter than cerci, in separate dorsolat- eral and ventral groups. Cerci elongate-ovoid in lateral view, fused dorsally on basal third. Pupa (as for Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventraliy. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 2.2-3.7 mm (n=7). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 311) with elongate, parallel-sided, apically notched tooth and strong shaft. Papillae without setae, their bases not raised. Lateral papillae 3 on each side of spatula. Six dorsal papil- lae on first 7 abdominal segments. Abdominal segments extensively covered with verrucae mesodorsally and antero- and posteroventrally. Terminal segment shorter than eighth. Second in- star: Not seen. Type material —Syntypes: galls and larva, from Carya sp., vic. Washington, DC, July, Osten Sacken, evidently lost. No examples exist in either the MCZ or the USNM. Neotype designated here: third instar (middle of 5 specimens on slide), from gall on C. glabra, Wheaton Park, Wheaton, Maryland, VII-7-1985, R.J. Gagné, deposited in USNM. Affinities.—This species forms the only poly- thalamous gall in the genus. The female is distinc- tive for its short, deep ovipositor with short setae. Biological notes.—In Spencerville, MD, on July 7, 2004, two separate young galls were found, each containing many first instars in discrete chambers, showing that the gall is truly polythala- mous (Fig. 162). Third instars were found in galls from the same site the same year in mid-Septem- ber. The following spring 3 females were reared, all emerging from separate exit holes from one of the galls. Collections of galls.—GeorcIa: Forsyth (glabra, ovata). MARYLAND: Wheaton (glabra); Gaithersburg (tomentosa); Spencerville (tomentosa). Specimens of C. cynipsea examined.—GEORGIA: Forsyth, IX-22-2000 (larvae). MARYLAND: Gaith- RAYMOND J. GAGNE 37 ersburg, X-4-1994 (larvae); Spencerville, IX- 20-2004 (larvae; 3 2, pupae, em. IV-2-2004); Wheaton, VIII-7-1985 (larvae). Caryomyia deflexipili Gagné, new species Description.—Gall (Figs. 66-67): Occasional, on Eucarya hickories, but not east of the Appalachi- ans, except for New England and on the Georgia Piedmont; single or dispersed, on lower leaf sur- face between veins; 2.8-3.7 mm in height, spheri- cal with small umbo at apex; green, becoming tan to brown, surface matte, non-sticky, with short, sparse, white, decumbent hairs not obscuring sur- face; base with small conical pedicel, leaf irregu- larly exfoliate surrounding base of gall; wall firm, brittle, uniformly thin, enclosing glabrous, longi- tudinally ridged larval chamber of same shape as gall. This gall is most similar to that of C. glau- ciglobus (Fig. 68), which, instead of having a sparse covering of deflexed hairs, is naked and covered with a bluish bloom. Adult: Head: Antenna with 18-21 flagellom- eres in male, 19-20 in female; each of the male fla- gellomeres (as in Fig. 237) scarcely binodal, area between nodes marked mainly by lack of setulae and slight compression between basal and middle circumfila, circumfila loops short, subequal in length; female flagellomeres with 2 horizontal cir- cumfila connected by 2 vertical strands. Labella separated from one another anteriorly, hemi- spherical, short-setose, short-setulose. Palpus 4- segmented. Thorax: Wing length 2.3-2.4 mm in male (n=2); 2.8-3.3 mm in female (n=2). Male ab- domen: Third through seventh tergites each with posterior setae in 3-4 rows laterally, narrowing to 1-2 mesally; eighth tergite without posterior se- tae. Genitalia: hypoproct broadest before poste- rior convex margin, margin setose, dorsally with setulae only on apical fourth; aedeagus evenly cylindrical, wide, notched apically. Female ab- domen: Third through seventh tergites as for male; eighth tergite not so strongly sclerotized as preceding tergite, with few posterior setae, the se- tae not as long as on seventh tergite. Ninth seg- ment pliable except for slender dorsal sclerite that widens apically to invest part of cerci, setae sparse laterally, mostly as long as cerci, setae more nu- merous ventrally, mostly shorter than lateral se- tae. Cerci elongate-ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (as in Figs. 203-204): Antennal bases oblique in ventral view, converging and narrow- ing anteriorly, their narrowly rounded apices closely adjacent. Face with anterolateral pair of horizontal incisions, their posterior lip protruding to form concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar (Figs. 226-228): Length 1.7-3.4 mm (n=6). Body elongate-fusiform, broad and knobby anteriorly, strongly tapered posteri- orly. Head directed ventrally. Spatula (Fig. 298) with parallel-sided, deeply notched anterior tooth and parallel-sided shaft not appreciably wider than tooth. Papillae without setae, their bases prominently raised, especially those of dorsal, pleural, and terminal papillae. Lateral papillae 3 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Abdominal segments com- pletely covered by verrucae. Terminal segment ta- pered to narrow terminus, prolonged beyond the ventrally disposed terminal papillae, appreciably longer than eighth segment. Second instar: Length 0.9-1.4 mm (n=8). Collar segment with patch of enlarged spicules on each side. Spatula with apically pointed tooth and longer, wider shaft. Bases of dorsal and pleural papillae raised but asetose. Six dorsal papillae on first 7 abdomi- nal segments. Collar segment with enlarged group of spicules anterolaterally. Spatula with rectangu- lar shaft and large pointed tooth. Dorsal and pleu- ral papillae with short setae, their bases raised. Type material.—Holotype: Third instar (middle of 3 larvae on slide), from gall on C. ovata, Amston, Connecticut, IX-17-1999, R.J. Gagné, deposited in USNM. Etymology—The name deflexipili is a noun formed from the Latin deflexus (decumbent) and pilus (fine hair) with reference to the fine decum- bent hair on the gall surface. Affinities——This species is remarkable and unique in the genus for having an augmented number, 18 or more, of antennal flagellomeres. In many ways it is most like C. glauciglobus, but adults have not yet been reared for the latter species. Both have similarly-shaped galls but with distinctively different surfaces, that of the present species with short, decumbent hairs, that of C. glauciglobus without hairs and glaucous. Larvae of both are generally similar, particularly for their ventrally-directed heads, fusiform shape, and very long terminal segment. The two differ in the shape of the spatula. Biological notes.—Pupae exit from the side of the gall in April and May after winter diapause. 38 THE GALL MIDGES (DIPTERA: CECIDOMYITDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Collections of galls—— Arkansas: Chester (tex- ana); Palestine (ovata); Winslow (ovata). CONNECTI- cut: Amston (ovata); Danbury (ovata); Hartland (ovata); Harwinton (ovata); Hopeville (ovata); Litch- field (ovata); Meriden (ovata); Phoenixville (ovata); Plainfield (ovata); Thomaston (ovata). GEORGIA: 8 km E Fort Valley (pallida); Lizella (ovata). ILLINOIs: Dexter (laciniosa); Highland (laciniosa); Marshall (tomentosa). INDIANA: Richmond (tomentosa); Spencer Co. (ovata). Kentucky: Winchester (laci- niosa). MAINE: Eliot (tomentosa); North Berwick (ovata). MassAcuusETTs: Great Barrington (ovata); Holyoke (ovata). Mussissippi: Noxubee Natl Wildlife Ref. (ovata). Missouri: Rolla (texana); Wildwood (tomentosa). NEw HAmpsHIRE: Barring- ton (ovata); Exeter (ovata); Greenland (ovata); Sea- brook (ovata). NEw Jersey: Columbia (ovata). NEw York: Herkimer (ovata); Ithaca (ovata); Ravena (glabra, ovata); Saratoga Springs (ovata). OHIO: Hocking St. For., Hocking Co. (ovata); St. Clairs- ville ((tomentosa); Summerford (tomentosa). PENN- SYLVANIA: Kirby (ovata); Mooresville (tomentosa); Schellsburg (ovata). TENNESSEE: Gatlinburg (ovata); Jackson (tomentosa). West VIRGINIA: Cacapon (ovata); 6 mi NW Hedgesville (ovata). Specimens of C. deflexipili examined.—Con- NECTICUT: Amston, IX-17-1999 (larvae); Hartland, IX-7-2005 (larvae; pupal exuviae, 1 d,7 2, em. IV- 15 to V-2—2006); Phoenixville, IX-20-1994 (larvae). Georeia: Lizella, X-12-1993, J. & R. Payne (lar- vae). INDIANA: Spencer Co., IX-20-1995 (larvae). Kentucky: Winchester, IX-22-1995 _ (larvae). Maine: North Berwick, [X-22-1994 (larvae). NEw Hampsuire: Barrington, IX-22-1994 (larvae); Ex- eter, IX-21-1994 (larvae). New York: Saratoga Springs, IX-25-1994 (larvae). Onto: Hocking St. For., Hocking Co., IX-19-1995 (larvae). WesT VirR- GINIA: 6 mi NW Hedgesville, IX-4-19- to X-17- 1999 & VII- 26-2000 (larvae), & X-2000 (larvae; 2 3,2 ¢,em. IV-2001). Caryomyia echinata Gagné, new species Gall (Figs. 14, 152-153): Rare, on Eucarya hicko- ries west of the Appalachian Mts; on lower leaf surface, between veins; length 2.4-5.0 mm, spher- oidal at base, tapering abruptly to long, narrow neck, 2-4 parted apically; covered with long, stiff, brown hairs mixed with short, soft brown hairs, surface obscured; base with circular, shallow, cen- tral depression; larval chamber basal, ovoid, white to violet, remainder of gall made up of green tissue turning brown and woody; a bundle of fibers present between larval chamber and gall apex. See under C. inclinata for similar species. This gall is the only woody Caryomyia gall that is covered with very long, stiff hairs and occurs be- tween instead of on veins. Adult and pupa: Unknown. Larva: Third instar: Length 1.7—2.2 mm (n=6). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 315) with long, nearly parallel-sided, api- cally-notched tooth and long shaft. Papillae with- out setae, their bases not raised. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdominal segments with rounded and pointed verrucae covering dor- sum across midlength and horizontal rows of pointed spicules across venter anterior to sternal papillae and posterior to ventral papillae. Termi- nal segment shorter than eighth. Second instar: Not seen. Type material.—Holotype: Third instar, from gall on C. texana, Rolla, Missouri, IX-8—2004, R.J. Gagné, deposited in USNM. Etymology—The name echinata is a Latin ad- jective meaning spiny, with reference to the long, stiff hairs covering the gall. Affinities.—See discussion under C. inclinata. Collections of galls—Arkansas: Beulah (to- mentosa); 2 mi S DeQueen (tomentosa); Rt.1 at Red River, Little River Co. (tomentosa). ILLINoIs: Mar- shall (tomentosa). Mississirpr: Natchez Trace Pkwy, Mile 193, Choctaw Co. (pallida, tomentosa); Nox- ubee Natl Wildlife Ref., Oktibbeha Co. (ovata); 7 mi E Starkville (pallida). Missouri: Conway (ovata); Rolla (texana). Onto: Gypsum (ovata, Wells 1916: Fig. 14); Summerford (tomentosa). ONTARIO: Dundas (ovata). Texas: Daingerfield St. Pk, Morris Co. (tomentosa). Specimens of C. echinata examined.—ARKAN- sas: Beulah, IX-10—2004 (larvae). Missourt: Rolla, IX-8-2004 (larva). On1o: Summerford, IX-6—2004 (larvae). Caryomyia eumaris Gagné, new species Description.—Gall (Figs. 9a, 88-89): Common, found on Eucarya hickories; usually single or scat- tered, attached to small veins on lower leaf sur- face; 2.2-2.7 mm in length, recumbent, subovoid except for apical, curved, tapered extension; rough textured, white, green, to tan, with short, RAYMOND J. GAGNE 39 sparse, white hairs not obscuring surface; base of gall with short, central conical projection sur- rounded by shallow, circular indentation; wall firm, brittle, uniformly thin, larval chamber glabrous with longitudinal ridges following gall axis from leaf vein connection to recurved apex. This gall is most similar to that of C. recurvata but has a rough surface with sparse hair. The gall of C. recurvata has a hairless, smooth, glaucous surface. Adult (female only): Head: Antenna with 12 fla- gellomeres, flagellomeres with 2 horizontal cir- cumfila connected by 2 vertical strands (as in Fig. 238). Labella separated from one another anteri- orly, hemispherical, short-setose, short-setulose. Palpus 4 segmented. Thorax: Wing length 1.8-1.9 mm (n=2). Abdomen: Third through seventh ter- gites each with single posterior row of setae con- tinuous across width, eighth tergite without setae. Ninth segment pliable except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ventrally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.2-1.7 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 291) with tooth nearly parallel-sided, usually narrowed toward apex, notched at apex, the shaft wide anteriorly, narrowed beyond. Papillae without setae, their bases not raised. Lat- eral papillae 2-3 on each side of spatula. Only 4 dorsal papillae present on first 7 abdominal seg- ments. Abdomen with small field of verrucae at midlength of dorsum, usually only around the 2 lateralmost pairs of dorsal setae, but occasionally more extensive, and with horizontal rows of spicules present on venter anterior to sternal papillae and a few rows of spicules along poste- rior margin. Terminal segment shorter than eighth. Second instar: Length 0.7-0.8 (2). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae not raised, papillae asetose. Only 4 dorsal papillae present on first 7 abdominal segments. Type material—Holotype: Third instar, from gall on C. tomentosa, Entomology Road, BARC, Beltsville, Maryland, IX-26-1993, R.J. Gagné, de- posited in USNM. Etymology——The name eumaris is a Greek noun for Persian shoe, with reference to the re- cumbent gall’s upturned apex. Affinities.—Galls of several other species, C. procumbens, C. recurvata, C. spinulosa, and C. su- pina, have similar attachments to leaf veins, grow horizontally, are thin-walled but brittle, and have longitudinally ridged larval chambers. Larvae of all are generally similar with narrow but two- toothed spatulas and only four dorsal papillae on each abdominal segment. The adult stage has been reared for only two species of this group, the female for C. eumaris and the male for C. recurvata. Biological notes—In mid-June in central Maryland, galls are small to full-grown, green and soft, and contain first instars. By end of June through August, most galls examined contained second instars but a few still had first instars. By end of August galls mostly contained third instars and many galls were turning or had turned red. Collections of galls——Atapama: Ashville (to- mentosa); Auburn (ovata, tomentosa); Greenville (to- mentosa). ARKANSAS: Beulah (tomentosa); 2 miS De- Queen (tomentosa); 3 mi S Harrison (tomentosa); Osage (tomentosa). CONNECTICUT: Oakville (ovata); Phoenixville (ovata); Plainfield (ovata); Southbury (tomentosa); Westchester (ovata). FLORIDA: Chatta- hoochee (tomentosa); Gainesville (tomentosa); Ponce de Leon (glabra). Georata: Lizella (ovata, to- mentosa). ILLINOIS: Dexter (laciniosa, tomentosa). Kentucky: Brownsville (tomentosa); Grayson Lake St. Pk, Carter Co. (ovata); Winchester (glabra). LouIsIANa: Lake Bistineau St. Pk, Webster Parish (tomentosa). MARYLAND: Beltsville, USDA Agricul- tural Research Center (tomentosa); Brighton (to- mentosa); Calverton (tomentosa); Clarksburg (glabra, tomentosa); College Park (glabra); Ellicott City (tomentosa); Gaithersburg (tomentosa); Kemp Mill (glabra); Lusby (tomentosa); Potomac (tomen- tosa); Ridgely (glabra, tomentosa); Silver Spring (to- mentosa); Spencerville (tomentosa); Wheaton (to- mentosa). Massacuusetts: Merrimac (ovata). MississiPPi: Noxubee Natl Wildlife Ref., Oktib- beha Co. (tomentosa); 7 mi E Starkville (tomentosa). Missouri: Holly Ridge State For., Stoddard Co. (tomentosa); Conway (ovata). New York: Ravena (ovata). NorTH CAROLINA: Alamance (tomentosa); 40 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Fayetteville (tomentosa); Providence (tomentosa); Rowland (tomentosa). Onto: Hocking Co. (ovata, Wells 1916: Fig. 9a). OKLAHOMA: Sallisaw (tomen- tosa). SOUTH CAROLINA: Georgetown (tomentosa). TENNESSEE: Crossville (pallida, tomentosa); Dan- dridge (tomentosa); Greene Co., I 81, Mile 38.5 (to- mentosa); Mohawk (tomentosa). Texas: Dainger- field St. Pk, Morris Co. (tomentosa). VIRGINIA: Carson (tomentosa); Dinwiddie (tomentosa); Hunt- ley Meadows Pk,-Fairfax Co. (tomentosa); Lady- smith (tomentosa); Radford (tomentosa); Skippers (tomentosa). WASHINGTON, DC: National Arbore- tum. (glabra, tomentosa); Walter Reed Army Med- ical Center (tomentosa). Specimens of C. eumaris examined.—ARKAN- sas: Osage, IX-9-2004 (larva). CONNECTICUT: Phoenixville, IX-20-1994 (larvae). KENTUCKY: Brownsville, X-21-1995 (larvae); Grayson Lake St. Pk, Carter Co., IX-22-1995 (larvae); Winchester, IX-22-1995 (larvae). MARYLAND: Beltsville, USDA Agricultural Research Center, IX-26—1993 (larvae), IX-23-2005 (pupal exuviae, 3 2, em. IV-10—2006); Ridgely, X-1999 (larvae; 2 2, pupa, em. IV-2000), X-6-2005 (larvae); Silver Spring, IX-17-1993 (lar- vae). SOUTH CAROLINA: Georgetown, X-1—2005, D. & B. Gagné (larvae). Caryomyia flaticrustum Gagné, new species Description.—Gall (Figs. 16a, 78-81): Com- mon, on Eucarya hickories; on lower leaf surface between veins, usually clustered; 2-4 mm in height, usually longer than wide, conical to cylin- drical, deeply concave apically, walls encircling concavity flared outwards, cylindrical, or in- grown toward apex as to form nearly complete false chamber, base of concavity with short central umbo; green at first, drying to tan or brown, walls surrounding concavity soft, eventually turning mucilaginous and contracting to form circular brown crust atop basal portion of gall; leaf with only slight exfoliation surrounding conical base of gall; lateral wall of larval chamber firm, brittle, thin apical wall usually thicker, larval chamber glabrous, with longitudinal ridges. When dried this gall simulates that of C. melicrustum but the gall of the latter is surrounded basally with an ex- tensive, thick leaf exfoliation. Adult: Head: Antenna with 12 flagellomeres; male flagellomeres (as in Fig. 237) scarcely bin- odal, only slightly narrowed beyond basal cir- cumfilum, circumfila loops short, subequal in length; female flagellomeres with 2 horizontal cir- cumfila connected by 2 vertical strands. Labella separated from one another anteriorly, hemi- spherical, short-setose or asetose, setulose. Palpus 4 segmented. Thorax: Wing length 1.6-2.0 mm in male (n=5); 1.4-2.2 mm in female (n=5). Male ab- domen: Third through seventh tergites each with posterior setae in single row, continuous across width; eighth tergite without posterior setae. Gen- italia (as in Figs. 247-248): gonostylus tooth nearly as wide as gonostylus; hypoproct broadest near apex, concave posteromedially, margin setose ex- cept in concavity; setulose dorsally only on distal third; aedeagus furrowed apically and ventrally. Female abdomen: Third through seventh tergites each with single row of posterior setae, continu- ous across width; eighth tergite without posterior setae. Ninth segment pliable except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ventrally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.8—2.2 mm (n=6). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 289) with anterior tooth narrowing from base to slightly notched or erose apex, shaft wide anteriorly, tapering abruptly and linear beyond. Papillae without setae, their bases not raised. Lat- eral papillae 2 on each side of spatula. Only 4 dor- sal papillae on first 7 abdominal segments. Ab- dominal segments essentially smooth dorsally, occasionally with some verrucae around lateral pairs of dorsal papillae, with fine, pointed spicules on venter anterior to sternal papillae and without spicules posteriorly. Terminal segment shorter than eighth. Second instar: Length 0.6-0.9 mm (n=5). Collar segment with patch of enlarged spicules on each side. Spatula with acutely trian- gular anterior tooth and longer, wider, concave- sided shaft. Bases of dorsal and pleural papillae not raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Specimen on left of two on slide, third instar, from gall on C. tomen- RAYMOND J. GAGNE 41 tosa, BARC, Beltsville, Maryland, IX-30-2003, R.J. Gagné, deposited in USNM. Etymology.—The name flaticrustum, a noun, is formed from the Latin flatus for puff of air and crustum for tart, with reference to the ephemeral apical funnel-like ring of tissue that shrinks atop the tart-like base of the gall. Affinities.—Caryomyia flaticrustum, C. levicrus- tum, C. melicrustum, and C. striolacrustum are four common, widely distributed species on the Eu- carya section with diminutive, more or less dis- coid galls with conical bases (Figs. 78-87). The main differences among the four species are in the shape and surface texture of the galls. That of C. flaticrustum, has a soft, sticky, concave apex that eventually contracts to form a thick crust: These mature galls simulate somewhat galls of C. meli- crustum, except that galls of the latter are sub- tended by a definite exfoliation of the leaf. Larvae of all four species are generally similar with a fairly smooth integument and cruciform spatula. The spatula tooth is barely notched in three species, but acutely pointed in C. levicrustum. Collections of galls—AtaBama: Ashville (to- mentosa); Auburn (ovata, tomentosa); Greenville (to- mentosa); Oak Mt. St. Pk, Shelby Co. (ovata, pallida). ARKANSAS: Beulah (tomentosa); Chester (texana); 2 mi S DeQueen (texana, tomentosa); 3 mi S Harrison (tomentosa); London (tomentosa); Magazine Mt., Logan Co. (ovata, texana, tomentosa); Osage (texana, tomentosa); Palestine (ovata, tomentosa); Piney (tex- ana); Rt.1 at Red River, Little River Co. (tomentosa). CoNNEcTICUT: Amston (ovata); Meriden (glabra, ovata); New Haven (glabra); Phoenixville (ovata); Plainfield (ovata); Southbury (glabra, tomentosa); Westchester (ovata). FLorRIDA: Ponce de Leon (glabra). Georcta: Carnesville (glabra); 8 km E Fort Valley (pallida); Forsyth (glabra, ovata); Lizella (ovata, tomentosa). ILLINOIS: Dexter (laciniosa, tomen- tosa); Marshall (tomentosa). INDIANA: Spencer Co. (ovata). Kentucky: Brownsville (tomentosa); Grayson Lake St. Pk, Carter Co. (tomentosa); More- head (tomentosa); My Old Kentucky Home St. Pk, Nelson Co. (ovata); Pennyrile St. Pk, Christian Co. (glabra, ovata); Winchester (lacinosa). MAINE: North Berwick (ovata). MARYLAND: Beaver Dam (tomen- tosa); Beltsville, USDA Agricultural Research Cen- ter (glabra, tomentosa); Brighton (tomentosa); Calverton (tomentosa); Clarksburg (glabra, tomen- tosa); College Park (glabra); Ellicott City (tomen- tosa); Gaithersburg (tomentosa); Lusby (glabra, tomentosa); Piney Grove (ovata); Potomac (tomen- tosa); Ridgely (glabra, tomentosa); Silver Spring (glabra, tomentosa); Snow Hill (tomentosa); Spen- cerville (tomentosa); Wheaton (glabra, tomentosa). MassacuusttTts: Great Barrington (ovata); Holy- oke (ovata); Merrimac (ovata); Petersham (ovata); Wrentham (ovata). Missouri: Conway (ovata, tex- ana); Crowley Ridge Roadside Pk, Stoddard Co. (glabra, tomentosa); Holly Ridge State For., Stod- dard Co. (glabra, tomentosa); Rolla (texana); Wild- wood (tomentosa). Mississippi: Natchez ‘Trace Pkwy, Mile 193, Choctaw Co. (glabra, pallida, to- mentosa); Noxubee Natl Wildlife Ref., Oktibbeha Co. (ovata, tomentosa); nr. Sessums, Oktibbeha Co. (ovata); 7 mi E Starkville (pallida). Missourt: St. Clair (tomentosa). New Hampsuire: Barrington (ovata); Dover (ovata); Kingston (ovata). New JER- sEY: Columbia (ovata); Nyack (tomentosa); Colum- bia (tomentosa). NEw York: Ithaca (ovata); Saratoga Springs (ovata). NORTH CAROLINA: Alamance (to- mentosa); Alberta (glabra); Dortches (tomentosa); Fayetteville (tomentosa); Gastonia (tomentosa); Providence (glabra, tomentosa); Rowland (tomen- tosa); Wise (tomentosa). Ounto: Hocking St. For., Hocking Co. (tomentosa); Hocking Co. (Carya sp., Wells 1916: Fig. 16a); St. Clairsville (tomentosa). OKLAHOMA: Sallisaw (tomentosa). PENNSYLVANIA: Pavia (ovata); Schellsburg (glabra). SouTH Car- OLINA: Georgetown (tomentosa); Richland Co. TEN- NESSEE: Crossville (pallida); Dandridge (tomentosa); Dickson (tomentosa); Gatlinburg (glabra, tomen- tosa); Greene Co., 1 81, Mile 38.5 (ovata, tomentosa); Jackson (tomentosa); Lebanon (glabra); Mohawk (tomentosa). TExAs: Daingerfield St. Pk, Morris Co. (tomentosa). VirGINIA: Warren Co., Blue Ridge Pkwy Mile 4.6 (ovata); Carson (tomentosa); Dinwid- die (glabra, tomentosa); Huntley Meadows Pk, Fair- fax Co. (tomentosa); Ladysmith (tomentosa); Rt. 81, mi. 13.9, Washington Co. (ovata). WASHINGTON, DC: National Arboretum (glabra, tomentosa); Wal- ter Reed Army Medical Center (tomentosa). WEST VirGINIA: Beech Fork St. Pk, Wayne Co. (tomen- tosa); Cooper’s Rock St. Pk, Preston Co. (glabra); 6 mi. NW Hedgesville (tomentosa). Specimens of C. flaticrustum examined.—ILLI- Nols: Dexter, X-8—2004 (larvae; 3 2, em. IV-11- 2005). MARYLAND: Beltsville, USDA Agricultural Research Center, IX-8—2002 (larvae; 5 5,2 @, pu- pae, em IV-2003) & IX-30-2003 (larvae; 3 5,3 2, pupae, em IV-11—2004); College Park, IX-23-2003 (larvae; 2 5,2 2, pupae, em. IV-11-2004); Ridgely, IX-2-2000 (larvae; 3 6, 2 2, em. IV-2001), IX- 11-2003 (larvae). NORTH CAROLINA: Providence, 42 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) IX-18-2000 (larvae). Texas: New Waverly, X-6- 2001 (larvae). Caryomyia glauciglobus Gagné, new species Description—Gall (Figs. 68-69): Occasional, on Eucarya hickories; single or dispersed on lower leaf surface, between veins; 3.6-4.2 mm in height, spherical with small umbo at apex, green becom- ing tan to brown, smooth, not sticky, hairless, cov- ered with blue-gray powdery bloom; base with small conical pedicel, leaf with irregular exfolia- tion around base of gall; wall firm, brittle, uni- formly thin, larval chamber glabrous with longi- tudinal ridges. This gall appears superficially similar to those of C. deflexipili and C. caryae be- cause of their spherical shape without conspicu- ous hairs. Galls of C. deflexipili have deflexed, fine hair that is detectable with a hand lens, while galls of C. caryae are sticky to the touch and not glaucous. Adult and pupa: Unknown. Larva: Third instar: Length 3.2-4.5 mm (n=6). Body elongate-fusiform, broad and knobby ante- riorly, strongly tapered posteriorly. Head directed ventrally. Spatula (Fig. 299) with parallel-sided, deeply notched anterior tooth and parallel-sided shaft not appreciably wider than tooth. Papillae without setae, their bases prominently raised, es- pecially those of dorsal, pleural, and terminal papillae. Lateral papillae 3 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Abdominal segments completely covered by ver- rucae. Terminal segment tapered to narrow termi- nus, prolonged beyond the ventrally disposed ter- minal papillae, appreciably longer than eighth segment. Second instar: Not seen. Type material.—Holotype: Third instar, from gall on C. ovata, Amston, Connecticut, IX-7-1999, R.J. Gagné, deposited in USNM. Etymology.—The name glauciglobus is a noun formed from the Latin glaucus (bluish gray) and globus (globe), referring to the blue-powdery, spherical gall. Affinities—See under C. deflexipili. Collections of galls—Connecticut: Amston (ovata); Danbury (ovata); Litchfield (ovata); Meri- den (ovata); Phoenixville (ovata). GkorGia: Forsyth (glabra); Lizella (tomentosa). INDIANA: Spencer Co. (ovata). KENTUCKY: Winchester (laciniosa). MAINE: North Berwick (ovata). MARYLAND: Beltsville, USDA Agricultural Research Center (tomentosa). Massacuusetts: Magnolia (Carya sp.); Petersham (ovata); Wrentham (ovata). Mississippi: nr. Ses- sums, Oktibbeha Co. (ovata). NEw York: Herkimer (ovata); Ithaca (ovata). New York: Albany (Carya sp.); Orchard Park (Carya sp.); Ravena (ovata). Onto: Hocking St. For., Hocking Co. (tomentosa); St. Clairsville (tomentosa). TENNESSEE: Jackson (to- mentosa). West VIRGINIA: Beech Fork St. Pk, Wayne Co. (ovata); Cacapon (ovata); 6 mi NW Hedgesville (ovata). Specimens of C. glauciglobus examined.—Con- NEcTIcuT: Amston, IX-17-1999 (larva); Danbury, IX-6-2005 (larvae). KeNTUcKy: Winchester, IX- 22-1995 (larva). Maine: North Berwick, IX-21- 1995 (larvae). Massacuusetts: Petersham, IX- 7-2000 (larvae). New York: Ithaca, IX-3-2003 (larva). Onto: Hocking St. For., Hocking Co., IX- 19-1995 (larvae). TENNESSEE: Jackson, IX-11—2004 (larva). WEST VIRGINIA: 6 mi NW Hedgesville, VI- 25-2004 (larvae). Caryomyia glebosa Gagné, new species Description.—Gall (Figs. 130-131): Rare but widespread, on Eucarya hickories; on lower leaf surface on vein; 3.0-3.2 mm in height, conical, hairless, brown to blood-red, bumpy, the bumps sometimes yellow; base with narrow, circular, shallow, central excavation; larval chamber basal, ovoid, lined with yellowish pellicle, remainder of gall tissue large-celled, blood-red, viscous, tra- versed between larval chamber and apex of gall by bundle of long fibers. The gall of this species is the only soft, conical Caryomyia gall that is hairless and covered with bumps. Adult and pupa: Unknown. Larva: Third instar (Figs. 223-225): Length 1.7— 2.2 mm (n=6). Body cylindrical, anterior and pos- terior ends broadly rounded. Head directed ante- riorly. Spatula (Fig. 307) with long, nearly paral- lel-sided, apically-notched tooth and long shaft widest just posterior to tooth, parallel-sided be- yond. Papillae without setae, their bases not raised. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal seg- ments. Abdominal segments with rounded verru- cae covering dorsum at midlength and horizontal rows of spicules on venter anterior to sternal papillae and posterior to ventral papillae. Termi- nal segment shorter than eighth. Second instar: Length 0.8-1.5 mm (n=6). Collar segment with patch of enlarged spicules on each side. Spatula RAYMOND J. GAGNE 43 with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae slightly raised, asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material—Holotype: Third instar, from gall on C. tomentosa, Wildwood, Missouri, IX- 8—2004, R.J. Gagné, deposited in USNM. Etymology.—The name glebosa is an adjective from the Latin meaning bumpy and refers to the characteristic surface on the gall of this species. Affinities—Galls of this species resemble somewhat those of C. turbanella because of the conical shape and at least partly bumpy surface. Larvae of the two species are similar, but adults of C. glebosa are not yet known. Collections of galls——Arkansas: 3 mi S Harri- son (tomentosa). CONNECTICUT: Meriden (ovata); Phoenixville (ovata). ILLinoIs: Marshall (tomen- tosa). MARYLAND: Kemp Mill (glabra); Gaithersburg (tomentosa); Silver Spring (glabra); Spencerville (to- mentosa). Missouri: Wildwood (tomentosa). NORTH CAROLINA: Wise (tomentosa). TENNESSEE: Dandridge (tomentosa); Dickson (tomentosa). WEST VIRGINIA: Cacapon (ovata). Specimens of C. glebosa examined.—ILunots: Marshall, IX-7—2004 (larvae). MARYLAND: Gai- thersburg, X-4-1994 (larvae); Silver Spring, IX- 22-1993 (larvae); Spencerville IX-21—2003 (larvae). Missouri: Wildwood, IX-8—2004 (larvae). Caryomyia guttata Gagné, new species Description.—Gall (Figs. 154-155): Found commonly but only on the central Florida en- demic, scrub hickory, of the Eucarya section; on lower leaf surface on vein, occasionally on leaf rachis, usually clustered in a row along major vein and often coalescing in series; 3-6 mm in height, spheroid basally, usually teardrop shaped, the apex tapering to a point; surface rugose, covered with tiny, red-brown and/or gold, resinous dots (as is the tree) and sparse setae; base with circular to irregular, shallow, central excavation; wall woody, thick, larval chamber basal, irregularly ovoid, glabrous, green to brown, a bundle of fibers present between apex of larval chamber and gall apex. This is the only hickory gall with such a local distribution and the only one known to occur sometimes on the leaf rachis. Galls resem- ble most those of C. aggregata and C. holotricha in shape but lack a dense covering of long hair and have no intrusion of hairs into the larval chamber. As are galls of C. aggregata, galls of C. guttata are found in close series along leaf veins. Adult and pupa: Unknown. Larva: Third instar: Length 2.4-3.5 mm (n=6). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 321) with single tooth tapering from wide base to pointed apex and with shaft broad anteriorly and tapering beyond. Papillae without setae. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal seg- ments. Abdominal segments with dorsum exten- sively covered with verrucae and venter spiculose anterior to sternal papillae and on posterior third. Terminal segment shorter than eighth. Second in- star: Not seen. Type material.—Holotype: Third instar, right- most of 3 specimens on slide mount, from gall on C. floridana, St. Lucie, Florida, XI-7 to 12-2002, K. Hibbard, deposited in USNM. Etymology.—The name guttata is a Latin adjec- tive meaning speckled, with reference to the resin- flecked surface of the gall. Affinities.—See discussion under C. inclinata. Collections of galls—Ftoripa (all on Carya floridana): Avon Park; Ft. Pierce; St. Lucie; Vero Beach; White City. Specimens of C. guttata examined.—FLoRIDA: Avon Park, XI-7 to 12-2002 (larvae); St. Lucie, XI-7 to 12-2002 (larvae), White City, XI-7 to 12-2002 (larvae). Caryomyia hirtidolium Gagné, new species Description.—Gall (Figs. 49-50): Occasional, on Eucarya hickories; on lower leaf surface, between veins; 3.0-3.9 mm in height, spheroidal, slightly longer than wide, base broadly rounded to trun- cate in profile, apex with small central nipple sur- rounded by extensive areola that with pressure can separate from gall; covered with long, silvery brown to brown hair obscuring surface, the hair denser and longer, rosette-like, on areola; base with wide, deep, circular indentation; wall firm, brittle, almost uniformly thin, larval chamber glabrous, with longitudinal ridges. For notes on galls of related species, see under C. cilidolium. Adult and pupa: Unknown. Larva: Third instar: Length 2.8-3.8 mm (n=10). Body dorsoventrally flattened, elongate-ovoid, broadly rounded anteriorly, spindleform posteri- orly. Head directed ventrally, surmounted dor- 44 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) sally by collar segment. Spatula (Fig. 278) with pointed anterior tooth and long, broad shaft. Papillae without setae, their bases raised. Lateral papillae 2 on each side of spatula. Six dorsal papil- lae on first 7 abdominal segments. Body covered with verrucae. Terminal segment as long as eighth, rounded posteriorly. Second instar: Length 1.2 mm (n=1). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular: anterior tooth and_ longer, wider, concave-sided shaft. Six dorsal papillae on first 7 abdominal segments. Bases of dorsal and pleural papillae elongate, asetose. Type material.—Holotype: Third instar, from gall on C. tomentosa, Lusby, Maryland, IX-16—2003, R.J. Gagné, deposited in USNM. Etymology.—The name hirtidolium, a noun, combines the Latin words hirtus (hairy) and dolium (jar) and refers to the hairy, barrel-like gall. Affinities —See under C. cilidolium. Collections of galls—ALaBamMa: Auburn (ovata); Greenville (tomentosa); Oak Mt. St. Pk, Shelby Co. (ovata). ARKANSAS: Beulah (tomentosa). Connecticut: Phoenixville (ovata). GEORGIA: Forsyth (glabra); Lizella (tomentosa). MARYLAND: Beltsville, USDA Agricultural Research Center (to- mentosa); Brighton (tomentosa); Clarksburg (tomen- tosa); Ellicott City (tomentosa); Gaithersburg (to- mentosa); Kemp Mill (glabra); Lusby (tomentosa); St. Leonard (tomentosa); Snow Hill (tomentosa). OKLA- HOMA: Sallisaw (tomentosa). SOUTH CAROLINA: Georgetown (tomentosa). VirGis1a: Radford (to- mentosa). WASHINGTON, DC: National Arboretum (tomentosa); Walter Reed Army Medical Center (to- mentosa). Specimens of C. hirtidolium examined.—ALa- BAMA: Greenville, IX-20—2000 (larva). ARKANSAS: Beulah, IX-10-2004 (larva). MARYLAND: Beltsville, USDA Agricultural Research Center, VII-25—2000 (larvae); Lusby, IX-16-2003 (larva); Silver Spring, IX-6—2003 (larvae). Caryomyia hirtiglobus Gagné, new species Description.—Gall (Figs. 53-54): Occasional, found on Eucarya hickories; single or clustered, on lower leaf surface between veins; 2.5-3.0 mm in height, depressed-spherical, occasionally weakly conical apically; covered with short, thick, brown hair obscuring gall surface; base of gall at center with short, conical pedicel at center of circular in- dentation, the corresponding leaf connection sur- rounded by inconspicuous exfoliation of the leaf; wall firm, brittle, uniformly thin except slightly thicker at apex; larval chamber of same shape as gall, without conspicuous extension into basal pedicel, glabrous, green to purple, with longitudi- nal ridges. In outward appearance the gall of this species is generally similar to those of other spherical, hairy galls, especially those made by C. purpurea and C. holotricha, but when cut longitudi- nally, the gall’s uniformly thin wall and its conical connection to the leaf will serve to distinguish it from otherwise superficially similar galls. Adult: Head: Antenna with 12 flagellomeres, in male a short 13th flagellomere without circumfila present; each of regular 12 male flagellomeres (as in Fig. 239) binodal with definite internode and neck, circumfila with short loops, their bases widely separated; female flagellomeres with 2 horizontal circumfila connected by 2 vertical strands. Labella separated from one another ante- riorly, hemispherical, short-setose, short-setulose. Palpus 4-segmented. Thorax: Wing length 3.3-3.4 mm in male (n=2), 3.1-3.4 mm in female (n=4). Male abdomen: Third through eighth tergites each with posterior setae separated into 2 lateral groups, in single to double rows; eighth tergite not more weakly sclerotized than seventh. Geni- talia: hypoproct parallel-sided, slightly concave posteromesally, margin setose, dorsum setulose only apically; apex of aedeagus convex. Female abdomen: Third through seventh tergites each with posterior setae in 2 lateral, mostly double rows; eighth tergite more weakly sclerotized than preceding tergite, with posterior setae only later- ally and not as long as on preceding tergite. Ninth segment pliable except for dorsal sclerite that widens from base to apex, setae sparse laterally and ventrally, many longer than cerci. Cerci ovoid in lateral view, fused dorsally for about 2/3 length. Pupa (Figs. 199-200): Antennal bases oblique in ventral view, converging and narrowing anteri- orly, their rounded apices closely adjacent. Face with anterolateral pair of horizontal incisions, their posterior lip protruding to form concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with several anterior rows of en- larged spinules. Larva: Third instar: Length 2.0-2.6 mm (n=10). Body dorsoventrally flattened, elongate-ovoid, rounded at ends, segments convex. Head directed antero-ventrally. Spatula (Fig. 283) with 2 widely RAYMOND J. GAGNE 45 separated anterior teeth, shaft widest anteriorly, narrowing abruptly to posterior half. Papillae mostly on short lobes, dorsals and pleurals with setae, also ventrals with setae on first through sev- enth abdominal segments. Lateral papillae usu- ally 2, occasionally 3 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Body entirely covered with verrucae. Terminal segment as long as eighth. Second instar: Length 0.6-1.2 mm (n=9). Collar segment with patch of enlarged spicules on each side. Spatula present, with acutely triangular anterior tooth and longer, elliptical shaft. Dorsal and pleural papillae raised, about as long as wide, without setae. Six dorsal papillae on first 7 abdominal segments. — Type material—Holotype: Third instar, from gall on C. tomentosa, Silver Spring, Maryland, IX- 17-1993, R.J. Gagné, deposited in USNM. Etymology.—The name hirtiglobus is a noun in apposition formed from the Latin hirtus (hairy) and globus (globe) with reference to the hairy, spherical gall. Affinities.—This species forms a natural group with C. caryae and C. shmoo because of the general shape of the gall and the similarity of the larvae, particularly the 2 widely separated anterior teeth of the third instar spatula. A difference between this species and the other two is the lack of setae on the dorsal and pleural papillae of the second instar. See under C. caryae for further discussion. Biological notes.—Galls of various sizes were first noticed on June 16 in central Maryland, with first instars in the smaller galls, second instars in the larger. The hair covering the gall is at first white, turning to brown by the time the gall is full size. Galls cut open on July 27 contained either sec- ond or third instars, both instars very active when touched. Their ventrally situated heads are closely appressed to the gall tissue so that the leading edge of the larva as viewed dorsally is the prono- tum. Small specimens of third instars seen on Au- gust 19 were closely appressed to the curved gall interior. By mid-September fully-developed third instars are opaque white and stout with expanded segments, and are no longer so closely appressed to the larval chamber surface. Collections of galls—A.aBama: Ashville (fo- mentosa); Auburn (tomentosa); Greenville (tomen- tosa). ARKANSAS: Beulah (tomentosa); 2 mi S De- Queen (tomentosa); London (tomentosa); Palestine (ovata); Piney (texana). CONNECTICUT: Amston (ovata); Meriden (glabra); Plainfield (ovata); Westchester (ovata). FLoripa: Gainesville (glabra, tomentosa). GEorGIA: 8 km E Fort Valley (pallida); Lizella (tomentosa). KeENTucKy: Grayson Lake St. Pk, Carter Co. (ovata). MARYLAND: Beltsville, USDA Agricultural Research Center (glabra, to- mentosa); Gaithersburg (tomentosa); Piney Grove (ovata); Ridgely (tomentosa); Silver Spring (tomen- tosa). MassacuuseTts: Holyoke (ovata). Missis- sippi: Noxubee Natl Wildlife Ref., Oktibbeha Co. (tomentosa); 7 mi E Starkville (pallida, tomentosa). Missouri: Holly Ridge St. For., Stoddard Co. (glabra, tomentosa). New York: Hyde Park (ovata); Ithaca (ovata). NortTH CAROLINA: Gastonia (fomen- tosa). OKLAHOMA: Sallisaw (tomentosa). TENNESSEE: Mohawk (tomentosa). VirGintA: Huntley Mead- ows Pk, Fairfax Co. (tomentosa); Ladysmith (to- mentosa). WASHINGTON, DC: National Arboretum (tomentosa). West VirRGINIA: Beech Fork St. Pk, Wayne Co. (tomentosa). Specimens of C. hirtiglobus examined.— ARKANSAS: Piney, X-10—2004 (larvae). CONNECTI- cut: Amston, IX-19-1994 (larva). GEoraGia: 8 km E Fort Valley, X-11-1993, J. & R. Payne (larvae). KEN- Tucky: Grayson Lake St. Pk, Carter Co., IX- 22-1995 (larvae). MARYLAND: Beltsville, USDA Agricultural Research Center, X-5-1994 (larvae) & X-13-1999 (larvae; 3 2, pupae, emerged IV-2000); Gaithersburg, X-4-1994 (larvae); Ridgely, IX- 23-1999 (larvae); Silver Spring, IX-20-1988, IX- 17-1993, VII-27-1994, & VIII-1—2000 (larvae). NEw York: Hyde Park, IX-26—2004 (larva); Ithaca, IX- 3-2003 (larva). OKLAHOMA: Sallisaw, IX-19-2004 (larvae). VirGINIA: Ladysmith, IX-23-2000 (larvae; 23,1 2, pupae, em. IV-2001). WasHINGTON, DC: National Arboretum, V-24—1994 (larvae). Caryomyia holotricha (Osten Sacken) Cecidomyia holotricha Osten Sacken 1862: 193; Felt 1909: 293 (Caryomyia). Dirhiza caryae Felt 1907: 47; Felt 1921: 101 (Caryomyia, as synonym of C. holotricha); Gagné 2004: 100, as junior synonym of C. caryae (Osten Sacken). Description.—Gall (Figs. 116-117): Common, on Eucarya hickories; usually clustered, on lower leaf surface between veins; 2.5-4.5 mm in height, from depressed-spherical with slight peak to onion shaped; with long, thick, brown hair ob- scuring gall surface; base with circular, deep, cen- tral excavation, leaf without exfoliation surround- ing connection; larval chamber basal, ovoid, floccose, green to brown, occasionally with slight 46 THE GALL MIDGES (DIPTERA: CECIDOMYITDAE) OF HICKORIES (JUGLANDACEAE: CARYA) violet hue, a tuft of brown hairs issuing from apex of larval chamber at end of fibrous bundle that terminates at gall apex; gall wall thick, woody. Galls are superficially similar to those of C. aggre- gata and C. purpurea because of their dense hair covering. Osten Sacken (1862) described two kinds of galls for this species. One kind is limited here as C. holotricha by the designation of the lec- totype that shows a characteristic intrusion of hairs from the gall apex into the larval cavity. The other kind presumably belongs to C. aggregata but none of Osten Sacken’s galls of that sort remain. Unlike galls of C. holotricha that are usually dis- crete and grow between veins, those of C. aggre- gata appear in closely packed rows along the midrib. It is not always possible to differentiate the more onion-shaped galls of C. holotricha from those of C. purpurea without opening them to see whether there is an apical intrusion of hairs that C. purpurea galls do not have. In addition, the inside surface of the C. purpurea galls is almost always purple instead of green to brown. Adult: Head: Antenna with 12 flagellomeres; male flagellomeres (as in Fig. 239) binodal with definite internode and neck, circumfila loops short, their bases widely separated; female flagel- lomeres with 2 horizontal circumfila connected by 2 vertical strands. Labella separated from one an- other anteriorly, hemispherical, short-setose, short-setulose. Palpus 4 segmented. Thorax: Wing length 2.9-3.7 mm in male (n=5), 2.3-3.9 mm in fe- male (n=5). Male abdomen: Third through eighth tergites each with posterior setae separated into two extensive lateral groups, almost joining mesally in posterior tergites, in single to double rows, sometimes partially triple rows on posterior tergites. Genitalia: hypoproct nearly parallel- sided, slightly concave posteromesally, margin se- tose except along concavity, setulose dorsally only on apical third; apex of aedeagus rounded; gonos- tylus not tapering, with wide tooth. Female ab- domen: Third through eighth tergites usually with posterior setae in 2-3 lateral rows, some- times almost joined mesally, especially on poste- rior segments, setae not as long or strong on eighth as on previous tergite. Ninth segment pli- able except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more nu- merous ventrally. Cerci elongate-ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (Figs. 207-208): Antennal bases pyrami- dal, moderately separated, acutely pointed apico- laterally. Face with anterolateral pair of horizontal incisions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 2.5-3.3 mm (n=10). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 327) with anterior tooth tapering from wide base to pointed apex, shaft widened at both ends, attenuated along midlength. Papillae without setae, their bases not raised. Lateral papillae 3 on each side of spatula, occasionally 2 on one side. Six dorsal papillae on first 7 abdomi- nal segments. Abdominal segments completely covered with verrucae, those anteriorly and pos- teriorly on venter spiculose. Terminal segment shorter than eighth. Second instar: Length 2.5-3.3 mm (n=10). Collar segment with patch of en- larged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, con- cave-sided shaft. Bases of dorsal and pleural papillae somewhat raised, asetose. Six dorsal papillae on first 7 abdominal segments. Type material—Caryomyia holotricha (O.S.): Lec- totype, here designated, gall from Carya sp., Wash- ington, DC, Osten Sacken, deposited in USNM. Paralectotypes, 14 galls on one leaf, marked “typi- cal form,” same data as lectotype. Osten Sacken (1862) also evidently described for this species galls of C. aggregata as a second form, but none of these is present in the Osten Sacken collection. Os- ten Sacken wrote that if there were two species in- volved, he would retain the name for the first form, which is done here. Caryomyia caryae (Felt): Holotype 2°, cecid. 58, taken on hickory, V-20-1906, Albany, New York, deposited in the New York State Museum in Al- bany. Felt (1921) listed his C. caryae as a junior syn- onym of both C. caryae (Osten Sacken) and C. holotricha. Felt’s species definitely fits within the parameters of C. holotricha. The holotype is even labeled C. holotricha in Felt’s handwriting, so the placement under C. caryae (O.S.) in Felt (1921) was evidently a lapsus. Etymology.—Osten Sacken (1862) coined the name holotricha with reference to the hair that completely covers the gall. The name caryae means “of Carya.” Affinities—Galls of this species and of C. thompsoni are the only ones with hairs intruding into the larval chamber from the apex of the gall. RAYMOND J. GAGNE 47 The two species are alike in most particulars of adults, pupae, and larvae, but galls of C. holotricha lack the basal intrusion into the larval chamber present in C. thompsoni galls and are almost al- ways conical rather than spheroidal. Biological notes.— In central Maryland, soft, whitish-green galls were first noticed on June 9 and contained first instars. On June 15, galls were nearing full size and contained first instars. By June 17, some galls were hard and contained sec- ond instars, as well as young hymenopterous par- asitoids. By June 22 and through July 16, all galls were at full size and hard. On August 3, galls con- tained large second instars. By August 31, more than 2/3 of the galls contained third instars. The full-grown larva is opaque white and practically fills the larval chamber. The exit tunnel is made to one side of the gall. Collections of galls—Atasama: Ashville (to- mentosa); Auburn (glabra, pallida, tomentosa); Greenville (tomentosa); Oak Mt. St. Pk, Shelby Co. (pallida). ARKANSAS: Beulah (tomentosa); Chester (texana); 2 mi S DeQueen (tomentosa); 3 mi S Harri- son (tomentosa); London (tomentosa); Magazine Mt., Logan Co. (texana, tomentosa); Piney (texana). CONNECTICUT: Amston (ovata); Danbury (ovata); Litchfield (ovata); Hartland (ovata); Meriden (glabra, ovata); New Haven (glabra); Oakville (ovata); Phoenixville (ovata); Southbury (glabra, to- mentosa); Southington (ovata); Thomaston (ovata). FLtoripA: Chattahoochee (glabra, tomentosa); Gainesville (glabra, tomentosa); Lake City (glabra, tomentosa). GEORGIA: Carnesville (glabra); 8 km E Fort Valley (pallida). ItuiNots: Dexter (laciniosa); Marshall (tomentosa). INDIANA: Spencer Co. (ovata). Kentucky: Grayson Lake St. Pk, Carter Co. (glabra, ovata); Morehead (tomentosa); My Old Kentucky Home St. Pk, Nelson Co. (ovata); Pennyrile St. Pk, Christian Co. (ovata); Winchester (laciniosa). LouIsiANa: Lake Bistineau St. Pk, Webster Parish (texana, tomentosa). MARYLAND: Beaver Dam (to- mentosa); Beltsville, USDA Agricultural Research Center (tomentosa); Brighton (tomentosa); Clarks- burg (glabra, tomentosa); Ellicott City (tomentosa); Gaithersburg (tomentosa); Lusby (glabra, tomen- tosa); Potomac (tomentosa); Ridgely (glabra, tomen- tosa); Silver Spring (tomentosa); Snow Hill (tomen- tosa); Wheaton (glabra). MASSACHUSETTS: Great Barrington (ovata); Leominster (ovata); Merrimac (ovata); Petersham (ovata); Wrentham (ovata). MIs- sissipPi: Natchez Trace Pkwy, Mile 123, Rankin Co. (pallida, tomentosa); Noxubee Natl Wildlife Ref., Oktibbeha Co. (tomentosa). Missourt: Conway (texana); Holly Ridge State For., Stoddard Co. (glabra); St. Clair (texana, tomentosa). NEw Hamp- SHIRE: Kingston (ovata). New York: Albany (Carya sp.); Hyde Park (ovata); Nassau (Carya sp.); New York (Carya sp.); Ravena (glabra); Shokan (glabra); West Point (Carya sp.). NortH CAROLINA: Ala- mance (tomentosa); Fayetteville (tomentosa); Provi- dence (tomentosa); Rowland (tomentosa); Wise (to- mentosa). Onto: Hocking St. For., Hocking Co. (ovata); St. Clairsville ((fomentosa); Yellow Springs (glabra). OKLAHOMA: Sallisaw (tomentosa). PENN- SYLVANIA: Julian (ovata); Millsboro (glabra); Schells- burg (glabra, tomentosa). SouTH CAROLINA: Dillon (tomentosa); Georgetown (tomentosa). TENNESSEE: Dandridge (tomentosa); Mohawk _ (tomentosa). Texas: Daingerfield St. Pk, Morris Co. (tomentosa); New Waverly (tomentosa). ViRGINIA: Warren Co., Blue Ridge Pkwy Mile 4.6 (ovata); Dinwiddie (to- mentosa); Huntley Meadows Pk, Fairfax Co. (to- mentosa); Ladysmith (tomentosa). WASHINGTON, DC: unspecified (Carya sp.); National Arboretum (tomentosa); Walter Reed Army Medical Center (tomentosa). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (ovata, tomentosa); 6 mi NW Hedges- ville (tomentosa). Specimens of C. holotricha examined.—Con- NECTICUT: Amston, IX-19-1994 (larvae); New Haven, IX-19-1994 (larvae); Phoenixville, IX- 20-1994 (larvae). Georcia: 8 km E Fort Valley, X- 11-1993, J. & R. Payne (larvae). ILLINOIS: Marshall, IX-7-2004 (larvae). KeNTUuCcKy: Winchester, IX- 22-1995 (larvae). MARYLAND: Beltsville, USDA Agricultural Research Center, X-5-1994 & VII- 12-2000 (larvae), X-13-1999 (larvae; 3 3,2 ¢, pu- pae, em. IV-2000), X-11-2005 (larvae; 5 6, 84 @, pupae, em. IV-2006); Brighton, X-8-1994 (2 6,5 @, pupae, em. IV-1995); Gaithersburg, X-3-1999 (lar- vae); Lusby, IX-14-2005 (larvae; pupa excised from gall IV-7—2006); Ridgely, X-2-2000 (larvae; 9 3, 7 2, pupae, em. IV-2001); Silver Spring, X- 3-1993 (larvae; 2 5, 3 2, pupae, em. IV-1994); Wheaton, X-13-1999 (larvae). MASSACHUSETTS: Beverly, 190?, C.H. Clarke (d, 2 2, larva); Boston, 190?, C.H. Clarke (3d, 2, larva); Leominster, IX- 21-1994 (larva). New Hampsuire: Kingston, IX- 21-1994 (larvae; 3 ¢,4 2,em. IV-1995). New York: Albany, found on hickory, V-20-1906 (¢, holotype of C. caryae (Felt)); E. Schodack, 190?, E.P. Felt (2 3,2 2); New York City, X-15-1908, E.P. Felt (larva; 3, 2, pupae, em. IV-22-1909). NorTH CAROLINA: Asheville, E.P. Felt, IX-16-2006 (larva). Onto: 48 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Hocking St. For., Hocking Co., IX-19-1995 (lar- vae); Yellow Springs, IX-7-2004 (larvae). TEN- NESSEE: Mohawk, IX-12—2004 (larvae). Texas: New Waverly, X-2001 (3 ¢, 4 2, pupae, em. IV-2002). WASHINGTON, DC: unspecified (type series of C. holotricha); National Arboretum, IX-27—2005 (lar- vae; 3 5,12 &, pupae, em. IV-2006); Walter Reed Army Medical Center, X-1999 (larvae; 2 2, pupae, em. IV-2000) and X-2004 (4 2, em. IV-1—2005). WEST VIRGINIA: 6 mai NW Hedgesville, IX-19-1999 (larvae). Caryomyia inanis Felt Caryomyia inanis Felt 1909: 292. Description.—Gall (Figs. 8, 98-99): Occasional, on Eucarya hickories, not found east of the Ap- palachian Mts (Map 2); found singly or in groups on lower leaf surface, between veins; length 2.5-4.0 mm, spheroidal at base with short to long, foliate, papery, lateral extension forming partial false chamber, the distal opening broadly agape to nearly closed; apex of closed part of gall slightly convex with central nipple often bearing thin, horizontal, flat disk (the leaf surface beneath which gall issued) equal to more than 1/3 gall di- ameter; green to brown with sparse, long, white or brown hairs not obscuring gall surface; base with prominent conical projection in center of cir- cular indentation, leaf with thin exfoliation sur- rounding gall base; larval chamber ovoid, wall firm, brittle, uniformly thin, smooth with longitu- dinal ridges. This gall has a similar structure to that of C. asteris, although that is not at first appar- ent. The foliate extension of the gall of C. inanis is vertical and tubular, while that of C. asteris is hori- zontally disposed. Adult: Head: Antenna with 12 flagellomeres; each of male flagellomeres (as in Fig. 237) scarcely binodal, distance between nodes marked mainly by disruption of setulae and slight compression between basal and middle circumfila, circumfila loops short, subequal in length; female flagello- meres with 2 horizontal circumfila connected by 2 vertical strands. Labella separated from one an- other anteriorly, hemispherical, short-setose, short-setulose. Palpus 4 segmented. Thorax: Wing length 2.1—2.2 mm in male (n=2); 2.5—2.8 mm in fe- male n=(5). Male abdomen: Third through sev- enth tergites each with posterior setae in 2-3 rows laterally, narrowing to 1-2 mesally; eighth tergite without posterior setae. Genitalia: hypoproct broadest before posterior convex margin, margin setose, dorsal surface setulose only on apical fourth; aedeagus evenly cylindrical, wide, barely notched apically. Female abdomen: Third through seventh tergites as for male; eighth ter- gite not so strongly sclerotized as preceding ter- gite, with 0-2 posterior weak posterior setae each side. Ninth segment (Fig. 266) rigid, pigmented, with sparse, long setae. Cerci pointed, almost fully fused dorsally. Pupa (as for Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.5-1.8 mm (n=8). Body elongate-fusiform, anterior end broadly rounded, posterior end tapering to weak point. Head directed anteroventrally. Spatula (Fig. 301) with parallel-sided, apically notched tooth and narrow shaft. Papillae without setae, their bases raised, especially those of dorsal, pleural, and ter- minal papillae. Lateral papillae 3 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Abdominal segments completely cov- ered with verrucae. Terminal segment as long as eighth and nearly pointed. Second instar: Length 1.0-1.3 mm (n=4). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, posteriorly widened shaft. Bases of dorsal and pleural papillae raised, papillae asetose. Six dorsal papillae on first 7 abdominal segments. Type material.—Lectotype here designated, 2, reared from Carya sp., IV-29-1909, Nassau, New York, Ca1950, deposited in New York State Mu- seum, Albany. Paralectotype °, same data as lecto- type. Etymology.— The name inanis is a Latin adjec- tive for “empty,” probably coined with reference to the false chamber above the larval chamber. Affinities.—This species and C. asteris are simi- lar for their fusiform larvae with raised, but ase- tose papillae. An important difference between the two is the presence in only C. asteris of setae on the dorsal and pleural papillae of the second in- star. Galls of the two also are generally similar, the main difference being that the exfoliate part of the gall beyond the larval chamber is vertical and RAYMOND J. GAGNE 49 continuous in C. inanis but horizontal and starlike in C. asteris. The present species is striking for its completely sclerotized ovipositor with its nearly completely joined cerci. The adult stage of C. as- teris has not yet been reared. Biological notes.—In mid-June in eastern West Virginia, galls were almost full grown and light green with the exfoliate part still easily pliable and contained second instars. In early July, galls were still green but with third instars that were very active when the galls were opened. In Au- gust the galls had turned brown. Collections of galls—Connecticut: Amston (ovata); Danbury (ovata); Harwinton (ovata); Hopeville (ovata); Litchfield (ovata); Meriden (ovata); Phoenixville (ovata); Plainfield (ovata); Southington (ovata); Thomaston (ovata). KEN- Tucky: My Old Kentucky Home St. Pk, Nelson Co. (ovata); Winchester (laciniosa). MAINE: Eliot (tomen- tosa); North Berwick (ovata). MASSACHUSETTS: Great Barrington (ovata); Holyoke (ovata); Mag- nolia (Carya sp.); Merrimac (ovata); Petersham (ovata). NEw HAmpsuire: Exeter (ovata); Greenland (ovata). New Jersey: Worthington (ovata). New York: Herkimer (ovata); Ithaca (ovata); Nassau (Carya sp.); New York (Carya sp.); Ravena (ovata); Shokan (tomentosa). Ou1o: Athens Co. (tomentosa, Wells 1916: Fig. 8); Summerford (tomentosa). PENN- SYLVANIA: Kirby (ovata); Pavia (ovata). TENNESSEE: Greene Co., I 81, Mile 38.5 (tomentosa). WEST VIR- GINIA: 6 mi NW Hedgesville (ovata, tomentosa). Specimens of C. inanis examined.—COoNNECTI- cut: Amston, IX-19-1994 & IX-17-1999 (larvae); Litchfield, IX-7-2005 (7 2, pupal exuviae, em. IV- 24 to 25-2006); Phoenixville, IX-20-1994 (larvae). Kentucky: Winchester, IX-22—1995 (larva). MAINE: North Berwick, IX-21-1994 (larva). NEw Hamp- SHIRE: Exeter, IX-21-1994 (larvae; 3d, 2 2, pupae, em. IV-1995). New Jersey: Worthington, IX-8-2000 (larva). New York: Nassau, E.P. Felt (2 2, type se- ries, em. [V-29-1909). Onto: Summerford, IX- 7—2004 (larvae). WEST VirGINIA: 6 mi. NW Hedges- ville, X-2000 (larvae; 5d, 2 2, em. IV-2001), X-1- 1995, IX-19 & X-17-1999, & VII-4—2000 (larvae). Caryomyia inclinata Gagné, new species Description.—Gall (Figs. 145-146): Occasional, locally common but not east of Appalachian Mts, on Eucarya hickories; on lower leaf surface, on veins; length from center of base to tip 3.3-7.0 mm, leaning, irregularly conical, base prolonged and irregularly lobed opposite direction of in- cline, the apex with short or long taper, digitate when long; sparsely covered with short, white hair not obscuring green, red, to brown surface; base at center flat or tapered to short-conical pedi- cel; larval chamber basal, green, with longitudinal ridges, remaining gall tissue woody. Galls are most like those of C. ansericollum, but in that species the base is not conspicuously flattened and outspread to one side opposite the direction of the cone. Adult: Head: Antenna with 12 flagellomeres; male flagellomeres scarcely binodal, only slightly narrowed beyond basal circumfilum, circumfila loops short, subequal in length; female flagellom- eres with 2 horizontal circumfila connected by 2 vertical strands. Labella separated from one an- other anteriorly, hemispherical, short-setose, short-setulose (as in Fig. 242). Palpus 4 seg- mented. Thorax: Wing length 1.9-2.1 mm in male (n=3); 2.5-2.7 mm in female (n=3). Male ab- domen: Third through seventh tergites each with posterior setae continuous across width, in mostly single row; eighth tergite more weakly sclero- tized, with 0-2 posterior setae laterally. Genitalia: hypoproct broadest just anterior to apex, poste- rior margin sinuous, setose except along mesal concavity, setulose dorsally on distal fourth; apex and venter of aedeagus furrowed. Female ab- domen: Third through seventh tergites each with posterior setae continuous across width, in mostly double row laterally, tapering to single row mesad; eighth tergite with a group of weak setae laterally. Ninth segment (Fig. 262) pliable except for fairly wide dorsal sclerite, setae long but sparse. Cerci elongate-ovoid, pointed apically, pigmented, fused dorsally on basal third. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva : Third instar: Length 1.4—-2.3 mm (n=10). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 320) with acutely pointed tooth and elongate, wide shaft. Papillae without setae, not conspicuously raised. Lateral papillae 2-3 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdominal segments with 50 THE GALL MIDGES (DIPTERA: CECIDOMYTIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) verrucae present at midlength on dorsum and with spicules present anterior to sternal papillae and on posterior third of venter. Terminal seg- ment shorter than eighth. Second instar: Length 0.7-0.9 mm (n=6). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae not raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. tomentosa, 6 mi NW Hedgesville, West Virginia, IX-19-1999, RJ. Gagné, deposited in USNM. Etymology.—The name inclinata is a Latin ad- jective with reference to the gall’s leaning habit. Affinities.—Galls of a group of five species are conical, woody, and have more or less digitate apices, but are still distinct in shape and vestiture. Both sexes of the present species and the female of C. ansericollum are known; the ovipositor of both species is similarly long and sparsely setose. The other three species are C. guttata, C. cucurbitata, and C. echinata, known only from their larvae. Larvae of all five species have only four papillae on the first through seventh abdominal segments, but the spatula of the last two named species are notched anteriorly and the other three have a tri- angular tooth. Biological notes.— Between June and begin- ning August in eastern West Virginia, young and full-grown galls can be found, containing first and second instars, respectively. By the first week of August galls contain either second or third in- stars. Pupae emerge through the side of the gall near the base. Collections of galls—Arkansas: Palestine (ovata); Winslow (ovata). CONNECTICUT: Amston (ovata); Danbury (ovata); Hartland (ovata); Har- winton (ovata); Hopeville (ovata); Meriden (tomen- tosa); Phoenixville (ovata); Plainfield (ovata); Thomaston (ovata). ILLINoIs: Dexter (laciniosa); Marshall (tomentosa). INDIANA: Spencer Co. (ovata). Kentucky: Grayson Lake St. Pk, Carter Co. (ovata); My Old Kentucky Home St. Pk, Nelson Co. (ovata). Winchester (laciniosa). MAINE: Eliot (fomen- tosa); North Berwick (ovata). MASSACHUSETTS: Great Barrington (ovata); Holyoke (ovata); Leominster (ovata); Petersham (ovata). NEw HAmp- SHIRE: Barrington (ovata); Greenland (ovata). NEw Jersey: Columbia (ovata, tomentosa). NEW YORK: Herkimer (ovata); Ithaca (ovata); Saratoga Springs (ovata). NORTH CAROLINA: Kannapolis (tomentosa). Ouro: St. Clairsville (tomentosa); Summerford (to- mentosa). ONTARIO: Dundas (ovata). TAMAULIPAS: SW of Ciudad Victoria (ovata, var. mexicana; pho- tos only). West VirGINiA: Cacapon (ovata); 6 mi NW Hedgesville (ovata). Specimens of C. inclinata examined.—Con- NEcTicuT: Amston, IX-19-1994 & IX-17-1999 (lar- vae); Phoenixville, IX-20-1994 (larva). KENTUCKY: Grayson Lake St. Pk, Carter Co., IX-22-1995 (larva). MassacuuseTTs: Leominster, IX-21-1994 (larva). New York: Herkimer , IX-4—2003 ( larvae); Saratoga Springs, IX-25-1994 (larva). OnIo: Sum- merford, IX-7¢2004 (larva). ONTARIO: Dundas, VIII-17-1994 (larvae). West VIRGINIA: 6 mi NW Hedgesville, X-1999 (larvae; 5, 2, pupae, em. TV-2000), X-2000 (larvae; 6, 2 @, em. IV-12- 2001), X-3-2003 (5 2, em. IV-12-2004), IX-19-1999 & VII-23-2000 (larvae), IX-30-2005 (2, em. IV- 12-2006). Caryomyia inflata Gagné, new species Description.—Gall (Figs. 5a, 102-103): Occa- sional, on Eucarya hickories, but not found on At- lantic side of Appalachians NE of central Georgia (Map 2); usually clustered, on lower leaf surface between veins; 2.2-4.0 mm in height, obconic, the apical cap convex early, becoming flat or slightly depressed at maturity, with central umbo; green, occasionally red, turning gray or brown with age, smooth, hairless, not sticky; base tapered to coni- cal pedicel, leaf with slight exfoliation around connection forming short, tubular socket and on reverse side showing discolored convexity; inside are a basal ovoid larval chamber and apical, equal-sized to larger and much broader, closed, false chamber, the basal larval chamber separated from apical chamber by thin wall with central umbo, walls thin, elastic at first, becoming brittle, both chambers longitudinally striate. This gall is most similar to that of C. tumida. The two are sim- ilarly shaped except that the gall of C. tumida is completely covered with short, decumbent hair. Adult: Head: Antenna with 12 flagellomeres; male flagellomeres (as in Fig. 237) scarcely bin- odal, distance between nodes marked mainly by disruption of setulae and slight compression be- tween basal and middle circumfila, circumfila loops short, subequal in length; female flagellom- eres with 2 horizontal circumfila connected by 2 vertical strands. Labella (Fig. 242) separated from one another anteriorly, hemispherical, short-se- tose, short-setulose. Palpus 4-segmented. Thorax: RAYMOND J. GAGNE 51 Wing length 2.0-2.1 mm in male (n=2); 2.2-2.5 mm in female (n=3). Male abdomen: Third through seventh tergites each with posterior setae in mesally interrupted, mostly single row; eighth tergite with 2-3 posterior setae laterally. Genitalia (as in Figs. 247-248): hypoproct broadest before posterior concave margin, margin setose except in concavity, dorsum setulose only on apical fourth; aedeagus deeply furrowed apically. Female ab- domen: Third through seventh tergites as for male; eighth tergite with 0-3 posterior setae later- ally. Ninth segment pliable except for slender dor- sal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ventrally. Cerci elon- gate-ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (Figs. 209-210): Antennal bases separate, acutely pointed apicolaterally. Face with antero- lateral pair of horizontal incisions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with several anterior rows of en- larged spinules. Larva: Third instar: Length 2.4-2.8 mm (n=6). Body cylindrical, anterior and posterior ends rounded. Head directed anteroventrally. Spatula (Fig. 313) with nearly parallel-sided anterior tooth only about as long as wide, obtusely notched at apex, the shaft parallel-sided, approximately as wide as tooth. Two lateral papillae each side of spatula. Papillae without setae, bases not raised. Six dorsal papillae on first 7 abdominal segments. Abdominal segments with sparse verrucae dor- sally at midlength surrounding dorsal papillae, and with horizontal rows of sharp, pointed spicules ventrally, anterior to sternal papillae, but not on posterior margin of segments. Terminal segment shorter than eighth. Second instar: Length 0.9-1.2 mm (n=10). Collar segment with patch of enlarged spicules on each side. Spatula with short right-triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae slightly raised, short-setose. Usually 6 dorsal papillae on first 6 abdominal seg- ments, but outer pair not always expressed; with only 4 on seventh segment. Type material.—Holotype: Third instar (right- most specimen of 4 larvae on slide), from gall on C. tomentosa, 6 mi NW Hedgesville, West Virginia, IX-26-1999, R.J. Gagné, deposited in USNM. Etymology.—The name inflata is a Latin adjec- tive referring to the inflated false chamber of this gall. Affinities.—This species and C. tumida come from similarly shaped, two-chambered galls, but that of C. inflata is smooth, the other thickly cov- ered with short, decumbent hairs. Beyond that, the galls have certain similarities to that of C. as- teris. Although the latter gall is not two-cham- bered, second instars of all three species have se- tose dorsal and pleural papillae. The rays of the C. asteris gall may indicate some midpoint develop- ment of a strictly two-chambered gall. Biological note-—Pupae emerge from a hole at the base of the gall next to the leaf connection. Collections of galls—ALABAMA: Oak Mt. St. Pk, Shelby Co. (pallida). CoNnNecticut: Amston (ovata); Danbury (ovata); Litchfield (ovata); Hart- land (ovata); Harwinton (ovata); Meriden (ovata); Phoenixville (ovata); Plainfield (ovata); West- chester (ovata). Georcia: Forsyth (glabra, ovata); Lizella (ovata). ILuiNots: Highland (laciniosa). INDI- ANA: New Castle (glabra). KeNTrucky: Grayson Lake St. Pk, Carter Co. (glabra, tomentosa). MAINE: Eliot (tomentosa); North Berwick (ovata); Seabrook (ovata). MassacuusetTts: Boston (ovata); Holyoke (ovata); Leominster (ovata); Merrimac (ovata); Pe- tersham (ovata); vic. Worcester (Carya sp., Thomp- son 1915: 93, Fig. 228). Mississippi: Noxubee Natl Wildlife Ref., Oktibbeha Co. (tomentosa). NEw Jersey: Columbia (ovata). New York: Ithaca (ovata). Onto: Hocking St. For., Hocking Co. (ovata); unspecified (Carya sp., Wells 1916: Fig. 5a); Huron Co. (C. glabra, Sears 1914: Fig. 18). TEN- NESSEE: Greene Co., I 81, Mile 38.5 (ovata). WEST ViRGINIA: Beech Fork St. Pk, Wayne Co. (ftomen- tosa); 6 mi NW Hedgesville (ovata, tomentosa). Specimens of C. inflata examined.—CONNECTI- cut: Harwinton, IX-6—2005 (pupae, 6 ¢, em. IV-25 to V-7-2006). Georeia: Lizella, X-12-1993, J. & R. Payne (larva). Kentucky: Grayson Lake St. Pk, Carter Co., IX-22-1995 (larva). Maine: North Berwick, IX-21-1994 (larvae; 2 6,3 ?, pupae, em. IV-1995). Massacuusetts: Leominster, [IX-21-1994 (larvae). WesT VIRGINIA: Beech Fork St. Pk, Wayne Co., VII-18-1988 (larvae); 6 mi NW Hedgesville, IX-4 & 26-1999 (larvae; 5 2, pupae, em. IV-2000), X-1-1995, VII-8—2000 & VIII-27—2000 (larvae). Caryomyia lenta Gagné, new species Description.—Gall (Figs. 5, 106-107): Occa- sional, on Eucarya hickories, not found east of the Appalachian Mts (Map 2); on lower or upper (one example) leaf surface between veins; height 3.3— 5.0 mm, depressed-spheroidal, sometimes 52 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) conical, with small nipple or tiny opening at cen- ter apex; rough textured, sticky, hairless, light green turning brown; base with central conical pedicel, leaf with raised, wide, mucilaginous, brown ring surrounding connection with gall and on reverse side showing discolored, hardened convexity; wall thin, rubbery, larval chamber sur- rounded laterally and apically by much larger, false chamber, the chambers separated by hemi- spherical, thin, sticky wall, both chambers longi- tudinally striate. The one instance of a gall on the upper leaf surface (Amston, CT) occurred on a leaf that also had a gall on the lower surface. This gall is superfically similar to that of C. inflata ex- cept that the present gall is extremely sticky to the touch and its false chamber nearly completely surrounds the top and sides rather than sits atop the larval chamber. Adult and pupa: Unknown. Larva : Third instar: Length 2.0-2.6 mm (n=10). Body cylindrical, anterior and posterior ends rounded. Head directed anteroventrally. Spatula (Fig. 318) tooth with single, acute point, the shaft wider, elongate. Lateral papillae 2 on each side of spatula. Usually 6 papillae on first 7 abdominal segments but outer pair on 6th and 7th segments not expressed on some specimens. Papillae with- out setae, bases not conspicuously raised. Ab- dominal segments dorsally with verrucae only at midlength close to dorsal papillae and ventrally with horizontal rows of sharp spicules anterior to sternal papillae and at posterior fourth of seg- ments. Terminal segment shorter than eighth. Sec- ond instar: Length 1.1-1.2 mm (n=4). Collar seg- ment with patch of enlarged spicules on each side. Spatula with equilateral anterior tooth and longer, wider, rectangular shaft. Bases of dorsal and pleu- ral papillae slightly raised, papillae asetose. Six dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. tomentosa, 6 mi NW Hedgesville, West Virginia, X-17—-1999, R.J. Gagné, deposited in USNM. Etymology.—The name lenta is a Latin adjec- tive meaning sticky or pliant, both qualities of the gall. Affinities—The two-chambered gall has cer- tain aspects in common with those of C. inflata and C. tumida, all of them with a false chamber, but the surface of the gall of the present species is sticky, the false chamber is much larger to cover the sides as well as the top of the larval chamber, and its base is surrounded by a large leaf socket. The third instar spatula of the present species is acutely toothed but those of the other two species are notched. Collections of galls—Connecticut: Amston (ovata); Phoenixville (ovata); Plainfield (ovata). ILtinots: Marshall (tomentosa); Rock Island (tomen- tosa, B.D. Walsh). INDIANA: Spencer Co. (ovata). Kentucky: Winchester (laciniosa). Maine: North Berwick (ovata). MassacuusettTs: Magnolia (Carya sp.); Petersham (ovata). PENNSYLVANIA: Pavia (ovata). New York: Herkimer (ovata). Onto: Athens & Hocking Counties (tomentosa, Wells 1916: Fig. 5); Summerford (tomentosa). WEST VIR- GINIA: Beech Fork St. Pk, Wayne Co. (tomentosa); 6 mi NW Hedgesville (ovata, tomentosa). Specimens of C. lenta examined.—ConneECTI- cut: Amston, IX-19-1994 (larvae). ILLINOIS: Mar- shall, IX-7—2004 (larva); Rock Island, B.D. Walsh (larva). Maine: North Berwick, IX-21-1994 (lar- vae). Onto: Summerford, IX-7—2004 (larvae). WEsT VIRGINIA: 6 mi. NW Hedgesville, IX-3 to 19-1999, VITI-16-2000 & IX-30-2005 (larvae). Caryomyia levicrustum Gagné, new species Description—Gall (Figs. 86-87): Common, on Eucarya hickories; on lower leaf surface between or on veins, usually clustered; height 2.2-3.0 mm, conical at base, cylindrical or flaring beyond, about as wide as long, the apex flat or shallowly to deeply convex, the walls surrounding any apical concavity variously flared outwards, evenly cylindrical, or almost closed to form partial false chamber, the center of apex with short umbo; sur- face glabrous, not sticky, green to tan; base coni- cal, area of contact with leaf surrounded by thin leaf exfoliation; wall firm, brittle, thin basally, usually slightly thicker apically, larval chamber shiny, smooth, with longitudinal ridges. A variety of gall shapes is included under this heading, but the variation involves only the structure above the larval chamber and the galls are otherwise similar in color and lack of sticky exudate. Adult (female only): Head: Antenna with 12 flagellomeres; flagellomeres with 2 horizontal cir- cumfila connected by 2 vertical strands. Labella separated from one another anteriorly, hemi- spherical, short-setose, and setulose. Palpus 4 seg- mented. Thorax: Wing length 1.8—2.1 mm (n=3). Female abdomen: Third through seventh tergites each with single row posterior setae, sparse mesally; eighth tergite with 0-2 posterior setae RAYMOND J. GAGNE 53 laterally. Ninth segment pliable except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ventrally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa: Unknown. Larva: Third instar: Length 1.2-1.6 mm (n=10). Body elongate-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 290) with anterior tooth an equilat- eral triangle, the shaft wide anteriorly, narrowed and linear beyond. Papillae without setae, bases not raised. Lateral papillae 2 on each side of spat- ula. Only 4 dorsal papillae on first 7 abdominal segments. Abdominal segments with no verrucae on dorsum or sparse verrucae only near bases of lateral pairs of dorsal papillae and with spicules on venter anterior to sternal papillae. Terminal seg- ment shorter than eighth. Second instar: Not seen. Type material—Holotype: Third instar, from gall on C. tomentosa, Entomology Road, BARC, Beltsville, Maryland, IX-26-1993, R.J. Gagné, de- posited in USNM. Etymology.—The name levicrustum, a noun, combines the Latin levis (smooth) and crustum (tart) and refers to the glabrous surface of the tart- like gall. Affinities—Galls of this species appear most like those of C. flaticrustum but instead of being soft and resinous, are hard and glabrous. For fur- ther remarks, see under C. flaticrustum. Biological notes.—In central Maryland galls were noticed as early as mid-June. Galls with sec- ond and third instars could be found from then through August. In late March and early April, pupae break out from the side of the galls near the base. Collections of galls—Atasama: Ashville (to- mentosa); Auburn (ovata, tomentosa); Greenville (to- mentosa); Oak Mt. St. Pk, Shelby Co. (ovata, pall- ida). ARKANSAS: 3 mi S Harrison (tomentosa); London (tomentosa); Magazine Mt., Logan Co. (ovata, texana, tomentosa); 2 mi S DeQueen (texana, tomentosa); Rt.1 at Red River, Little River Co. (to- mentosa). CONNECTICUT: Amston (ovata); Meriden (glabra, ovata); New Haven (glabra); Phoenixville (ovata); Plainfield (ovata); Southbury (glabra, to- mentosa). FLORIDA: Ponce de Leon (glabra). GEor- cia: Carnesville (glabra); 8 km E Fort Valley (pall- ida); Forsyth (glabra, ovata); Lizella (ovata, tomentosa). INDIANA: Spencer Co. (ovata). KEN- Tucky: Brownsville (tomentosa); Grayson Lake St. Pk, Carter Co. (tomentosa); Morehead (tomentosa); My Old Kentucky Home St. Pk, Nelson Co. (ovata); Pennyrile St. Pk, Christian Co. (glabra, ovata); Winchester (lacinosa). Maine: North Berwick (ovata). MaAryYLAND: Beltsville, USDA Agricultural Research Center (glabra, tomentosa); Brighton (tomentosa); Clarksburg (tomentosa); Elli- cott City (tomentosa); Gaithersburg (tomentosa); Kemp Mill (glabra); Lusby (glabra, tomentosa); Po- tomac (tomentosa); Ridgely (tomentosa); St. Leonard (tomentosa); Silver Spring (glabra, tomen- tosa); Spencerville (tomentosa); Wheaton (glabra, to- mentosa). MaAssaAcHusETTs: Great Barrington (ovata); Merrimac (ovata); Petersham (ovata); Wrentham (ovata). Mississippi: Natchez Trace Pkwy, Mile 123, Rankin Co. (tomentosa); Natchez Trace Pkwy, Mile 193, Choctaw Co. (glabra, pallida, tomentosa); Noxubee Natl Wildlife Ref., Oktibbeha Co. (ovata, tomentosa); nr. Sessums, Oktibbeha Co. (ovata); 7 mi E Starkville (pallida). Missouri: Con- way (ovata); Crowley Ridge Roadside Pk, Stod- dard Co. (glabra, tomentosa); Holly Ridge State For., Stoddard Co. (glabra, tomentosa). NEw HAmp- SHIRE: Barrington (ovata); Dover (ovata); Kingston (ovata). New Jersey: Columbia (ovata); Nyack (to- mentosa); Columbia (tomentosa). NEw YORK: Saratoga Springs (ovata). NORTH CAROLINA: Ala- mance (tomentosa); Alberta (glabra); Fayetteville (tomentosa); Gastonia (tomentosa); Providence (glabra, tomentosa); Rowland (tomentosa); Wise (to- mentosa). Onto: Hocking St. For., Hocking Co. (to- mentosa). OKLAHOMA: Sallisaw (tomentosa). SOUTH CAROLINA: Richland Co. TENNESSEE: Crossville (pallida, tomentosa); Gatlinburg (glabra, tomentosa); Greene Co., I 81, Mile 38.5 (ovata, tomentosa). Texas: Daingerfield St. Pk, Morris Co. (tomentosa). VirciniA: Warren Co., Blue Ridge Pkwy Mile 4.6 (ovata); Carson (tomentosa); Dinwiddie (glabra, to- mentosa); Huntley Meadows Pk, Fairfax Co. (to- mentosa); Ladysmith (tomentosa); I 81, Mile 13.9, Washington Co. (ovata). WASHINGTON, DC: Na- tional Arboretum (glabra, tomentosa); Walter Reed Army Medical Center (tomentosa). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (tomentosa); Cooper’s Rock St. Pk, Preston Co. (glabra). 6 mi NW Hedgesville (ovata, tomentosa). Specimens of C. levicrustum examined.—ALa- BAMA: Auburn, IX-19-2000 (larvae); Greenville, IX-20-2000 (larvae; 2 2, em. IV-2001). CONNECTI- cut: Phoenixville, [IX-20-1994 (larvae); Plainfield, VIII-27-1988 (larvae). MARYLAND: Beltsville, USDA Agricultural Research Center, IX-26-1993 54 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) (larvae) & X-28-1994 (larvae; 2, em. IV-1995); Lusby, IX-16-2003 (larvae). OKLAHOMA: Sallisaw, IX-9-2004 (larvae). Vircinia: Huntley Meadows Pk, Fairfax Co., VIII-30—2000 (larvae). Caryomyia leviglobus Gagné, new species Description.—Gall (Figs. 19, 62-63): Common, on Eucarya hickories; single or dispersed on lower leaf surface between veins; 2.8-3.5 mm in height, spherical with a conspicuous apical terminus made up of a nipple atop an abruptly raised are- ola; hairless, not sticky, glabrous, light green, be- coming yellow, green, or brown; base with small conical pedicel, leaf without exfoliation; wall firm, brittle, uniformly thin, larval chamber glabrous with longitudinal ridges. Wells (1916) wrote that the gall resembled a small bomb. This gall differs from that of C. spiniglobus in that it lacks hair. Both galls are distinctive for their conspicuous apical knob. Adult: Head: Antenna with 12 flagellomeres; each of the 12 male flagellomeres (as for Fig. 237) scarcely binodal, only slightly narrowed beyond basal circumfilum, circumfila loops short, sub- equal in length; female flagellomeres with 2 hori- zontal circumfila connected by 2 vertical strands. Labella (as in Fig. 242) separated from one an- other anteriorly, hemispherical, short-setose, short-setulose. Palpus 4-segmented. Thorax: Wing length 2.0-2.1 mm in male (n=3); 2.2-2.5 mm in female (n=5). Male abdomen: Third through seventh tergites each with mostly single row of posterior setae nearly continuous across width, interrupted for a short space mesally; eighth tergite without posterior setae. Genitalia: hypoproct broadest before convex posterior mar- gin, margin setose; aedeagus evenly cylindrical, notched apically. Female abdomen: Third through seventh tergites each with single row of posterior setae nearly continuous across width except interrupted mesally; eighth tergite as scle- rotized as preceding tergite, with a few posterior setae laterally, the setae as long as on seventh ter- gite. Ninth segment pliable except for slender dorsal sclerite that widens apically to invest part of cerci, setae sparse laterally, mostly as long as cerci, setae more numerous ventrally, mostly shorter than lateral setae. Cerci elongate-ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (Figs. 203-204): Antennal bases oblique in ventral view, converging and narrowing anteri- orly, their narrowly rounded apices closely adja- cent. Face with anterolateral pair of horizontal in- cisions, their posterior lip protruding to form con- cavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with several anterior rows of enlarged spinules. Larva: Third instar: Length 2.5-3.5 mm (n=10). Body spindleform, segments convex, head end tapered, posterior end rounded. Head ventral. Spatula (Fig. 281) with large, simple anterior tooth nearly as long as broad shaft. All papillae except terminals setose, papillar bases not con- spicuously raised. Lateral papillae numbering 2 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Abdominal segments completely covered with raised, pointed verru- cae. Terminal segment as long as eighth. Second instar: Length 0.9-1.5 mm (n=10). Collar segment with patch of enlarged spicules on each side. Spatula with apically pointed tooth and rectan- gular or triangular shaft. Bases of dorsal and pleural papillae raised and all short-setose in- cluding on prothorax. Six dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Larva, from gall on C. tomentosa, vic. Ridgely, Maryland, IX-23-1999, R.J. Gagné, deposited in USNM. Etymology.—The name leviglobus is a noun in apposition formed from the Latin levis (smooth) and globus (globe) with reference to the hairless, spherical gall. Affinities —This species does not differ from C. spiniglobus except in the exterior of the gall that is smooth in C. leviglobus and stiff-haired in C. spiniglobus. I found galls of both species at the same site, but not in proximity on the same leaflet, so consider them separate species. While C. leviglobus is more common than C. spiniglobus, both are generally widespread. Together these species are unique in Cecidomyiini for their setose ventral larval papillae of the eighth abdominal segment. This character state is a defining one for the tribe Lasiopteridi; as far as I know these two species are the only Cecidomyiidi with setose ventral papillae, which is evidently a reversal here. The larval sternal papillae are also setose, unlike all Cecidomyiinae with the exception of the subtribe Asphondyliina. As with other species (viz. the C. caryae and C. cilidolium groups) that live unconstrained in a rather large, spherical lar- val chamber, the integument is completely cov- ered with spiny verrucae that evidently allow RAYMOND J. GAGNE 55 these species purchase in crawling across the curved surface of the larval chamber. Biological notes.—Tiny, soft, light green galls with first instars were first noted in central Mary- land on June 10. After mid-June galls in various stages of hardening yielded all instars including thirds. Adults emerged from a slight bulge in the side of the gall that was evidently weakened by the larva the previous autumn. Collections of galls—AtasBama: Auburn (pall- ida, tomentosa); Greenville (tomentosa); Oak Mt. St. Pk, Shelby Co. (ovata, pallida, tomentosa). ARKAN- sas: Beulah (tomentosa); 2 mi S DeQueen (texana, tomentosa); Magazine Mt., Logan Co. (ovata); Os- age (texana); Piney (texana). CONNECTICUT: Amston (ovata); Meriden (glabra, ovata); Natchaug St. For. (ovata); New Haven (glabra); Oakville (ovata); Westchester (ovata). FLoriDA: Gainesville (glabra). Georcia: Carnesville (glabra); Forsyth (glabra, ovata); 8 km E Fort Valley (pallida); Lake Park (glabra); Lizella (ovata, tomentosa). KENTUCKY: Brownsville (tomentosa); Grayson L. St. Pk, Carter Co. (glabra); Pennyrile St. Pk, Christian Co. (ovata). LOUISIANA: Lake Bistineau St. Pk, Webster Parish (texana, tomentosa). MARYLAND: Beltsville, USDA Agricultural Research Center (glabra, tomentosa); Calverton (tomentosa); Clarksburg (glabra, ovata, tomentosa); College Park (glabra); Gaithersburg (to- mentosa); Kemp Mill (ovata, tomentosa); Lusby (glabra, tomentosa); Potomac (tomentosa); Ridgely (tomentosa); St. Leonard (tomentosa); Silver Spring (glabra); Spencerville (tomentosa); Wheaton (glabra). MAssacuusetTtTs: Wrentham (ovata). Mis- sissippi: Noxubee Natl Wildlife Refuge (ovata); Ok- tibbeha Co., nr. Sessums (ovata). Missouri: Con- way (ovata, texana); Holly Ridge St. For., Stoddard Co. (tomentosa); Rolla (texana); St. Clair (tomen- tosa); Wildwood (tomentosa). NEw HAMPSHIRE: Kingston (ovata). New York: Shokan (glabra). NortH Caro.iina: Alamance (tomentosa); Alberta (glabra); Gastonia (tomentosa); Kannapolis (tomen- tosa); Rowland (tomentosa); Wise (tomentosa). Onto: Hocking St. For., Hocking Co. (ovata); Hock- ing Co. (ovata, Wells 1916: Fig. 19); Yellow Springs (glabra). OKLAHOMA: Sallisaw (tomentosa). TEN- NESSEE: Crossville (pallida); Dandridge (tomentosa); Dickson (tomentosa); Greene Co., I 81, Mile 38.5 (tomentosa); Jackson (tomentosa); Lebanon (glabra); Mohawk (tomentosa). Texas: Daingerfield St Pk, Morris Co. (tomentosa); New Waverly (tomentosa). ViRGINIA: Carson (tomentosa); Dinwiddie (glabra); Huntley Meadows, Fairfax Co. (tomentosa); War- ren Co., Blue Ridge Pkwy Mile 4.6 (ovata). WAsH- INGTON, DC: National Arboretum (glabra, tomen- tosa); Walter Reed Army Medical Center (tomen- tosa). WEST VIRGINIA: 6 mi NW _ Hedgesville (ovata). Specimens of C. leviglobus examined.— Con- NEcTICUT: Amston, IX-19-1994 (larva); New Haven, IX-19-1994 (larva). Georaia: 8 km E Fort Valley, IX-11-1993, J. & R. Payne (larvae). KEN- TucKy: Brownsville, IX-29-1995 (larvae). Mary- LAND: Beltsville, USDA Agricultural Research Center, [X-28-1999 & VII-22-2000 (larvae); Gai- thersburg, VII-20—2000 (larvae); Lusby, IX-2003 (2 3,4 2, em. IV-11-2004); vic. Ridgely, IX-23-1999 (larva); Wheaton, IX-19-1993 (larvae). MASSACHU- sETTS: Wrentham, [X-20-1994 (larvae; 2 2, em IV- 1995). New Hampsuire: Kingston, IX-21-1994 (larva). Onto: Hocking St. For., Hocking Co., IX- 19-1995 (larva). Texas: New Waverly, X-6—2001, (d,2 2, pupae, em. IV-2002). Vircinia: Huntley Meadows, Fairfax Co., VII-30-2000 (larvae). WASHINGTON, DC: National Arboretum, IX- 22-1988 (larvae). Caryomyia marginata Gagné, new species Description.—Gall (Figs. 139-140): Infrequent, on Eucarya hickories; on lower leaf surface, be- tween or on veins; length 3.2-5.0 mm, conical, the base flat against leaf surface, circular, with flat- tened lip surrounding cone that gradually and evenly narrows to pointed apex; green to tan, smooth, covered evenly with long, sparse hairs not obscuring surface; base with narrow, shallow, circular, central excavation, the leaf without exfo- liation surrounding connection; larval chamber ovoid, located at base of gall, surrounded by woody tissue. This conical, sparsely haired gall is most similar to that of C. conoidea but the sides of the cone of the C. marginata gall are concave and the basal angle is acute and marginate. This gall is entirely woody unlike that of C. conoidea, which is partially filled with soft tissue that eventually col- lapses. Adult: Head: Antenna with 12 flagellomeres; male flagellomeres scarcely binodal, only slightly narrowed beyond basal circumfilum, circumfila loops short, subequal in length; female flagellom- eres with 2 horizontal circumfila connected by 2 vertical strands. Labella separated from one an- other anteriorly, hemispherical, short-setose, short-setulose (as in Fig. 242). Palpus 4-seg- 56 THE GALL MIDGES (DIPTERA: CECIDOMYTIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) mented. Thorax: Wing length 2.0-2.2 mm in male (n=5); 2.6-3.0 mm in female (n=4). Male ab- domen: Third through seventh tergites each with single row of posterior setae continuous across width; eighth tergite with 0-2 setae laterally. Gen- italia: hypoproct broadest near apex, slightly con- cave posteromedially, margin setose except in concavity, dorsal setulae only on distal third; aedeagus furrowed. apically and ventrally; gonos- tylus narrowed towards apex. Female abdomen: Third through eighth tergites each with single row of posterior setae continuous across width, those on eighth tergite sparse and not as long. Ninth segment pliable except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ventrally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (Figs. 211-212): Antennal bases pyrami- dal, separate, acutely pointed apically. Face with anterolateral pair of horizontal incisions, their posterior lip protruding to form a craterlike con- cavity. Frons and clypeus strongly protruded ven- trally. Abdominal terga with several anterior rows of enlarged spinules. Larva: Third instar: Length 1.3-2.1 (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 309) with long, nearly parallel sided, api- cally notched tooth and long shaft. Papillae with- out setae, their bases not raised. Lateral papillae 2-3 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdominal seg- ments with rounded verrucae dorsum at mi- dlength and horizontal rows of spicules on venter anterior to sternal papillae and posterior to ven- tral papillae. Terminal segment shorter than eighth. Second instar: Length 0.8—-0.9 (n=3). Col- lar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, elliptical shaft. Bases of dorsal and pleural papillae slightly raised, papil- lae asetose. Only 4 dorsal papillae on first 7 ab- dominal segments. Type material.—Holotype: Third instar, from gall on C. ovata, Leominster, Massachusetts, IX- 21-1994, R.J. Gagné, deposited in USNM. Etymology.—The name marginata is a Latin ad- jective that refers to the circular, thin lateral mar- gin of the gall. Affinities.—See under C. sanguinolenta, from which this species differs in the shape of the gall. Collections of galls —ALABAMa: Greenville (to- mentosa). ARKANSAS: Chester (texana). CONNECTI- cut: Amston (ovata); Danbury (ovata); Litchfield (ovata). Kentucky: My Old Kentucky Home St. Pk, Nelson Co. (tomentosa). MAINE: East Lebanon (ovata); Eliot (tomentosa); North Berwick (ovata). MAssAcHuseETTs: Great Barrington (ovata); Leo- minster (ovata). Mississippi: Noxubee Natl Wildlife Ref., Oktibbeha Co. (tomentosa). Missouri: Holly Ridge State For., Stoddard Co. (tomentosa). NEw Hampsuire: Barrington (ovata); Dover (ovata); Greenland (ovata). New York: Herkimer (ovata); Ravena (ovata). Onto: St. Clairsville (tomentosa); Summerford (tomentosa). ONTARIO: Dundas (ovata). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (tomentosa); 6 mi NW Hedgesville (ovata). Specimens of C. marginata examined.—ARKAN- sas: Chester, IX-9-2004 (larva; 3 °, pupae, em. IV- 6/9-2005). MassacuusetTts: Great Barrington, IX- 6-2000 (larva); Leominster, IX-21-1994 (larvae; 5 3,4 2, pupae, em. IV-1995). New York: Herkimer, IX-4—2003 (larva). Onto: St. Clairsville, IX-6—2004 (larvae); Summerford, IX-9-2004 (larvae). ON- TARIO: Dundas, VIII-17—1994 (larva). West ViR- GINIA: 6 mi NW Hedgesville, VII-23-2000 & VII- 8-2004 (larvae). Caryomyia melicrustum Gagné, new species Description.—Gall (Figs. 30, 82-83): Common, on Eucarya hickories; on lower leaf surface be- tween veins, usually clustered; 2.0-3.0 mm in height, cylindrical, usually slightly wider than long, conical at base, circular, flat to concave at apex; lateral surface tan to brown, shiny, viscid, the apex orange to brown, crusty, thickly mucil- laginous; subtending leaf tissue with conspicu- ous, extensive, irregular exfoliation surrounding base of gall, corresponding top surface of leaf usu- ally discolored, convex; lateral wall of larval chamber firm, brittle, thin laterally, apical wall usually thicker, larval chamber glabrous, with longitudinal ridges. These galls can sometimes be confused with those of mature, dried C. flaticrus- tum galls, but galls of the latter lack the conspicu- ous leaf exfoliation surrounding the base of galls of C. melicrustum. Adult (female only): Head: Antenna with 12 flagellomeres; flagellomeres with 2 horizontal cir- cumfila connected by 2 vertical strands. Labella separated from one another anteriorly, hemi- spherical, short-setose, short-setulose. Palpus 4 RAYMOND J. GAGNE 57 segmented. Thorax: Wing length 2.1-2.5 mm (n=3). Female abdomen: Third through seventh tergites each with posterior setae in a single row, continuous across width; eighth tergite without posterior setae. Ninth segment pliable except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ven- trally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.4-2.4 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 287) with anterior tooth narrowing from base to slightly notched apex, shaft wide anteri- orly, tapering abruptly and linear beyond. Papil- lae without setae, not raised. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdominal segments essentially smooth dorsally, occasionally with some verrucae around lateral pairs of dorsal papillae, with fine, pointed spicules on venter an- terior to sternal papillae and none posteriorly. Ter- minal segment shorter than eighth. Second instar: Length 0.7-0.9 mm (n=4). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae slightly raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal seg- ments. Type material—Holotype: Third instar, from gall on C. glabra, Wheaton Park, Maryland, IX- 19-1993, R.J. Gagné, deposited in USNM. Etymology.—The name melicrustum, a noun, is formed from the Latin mel for honey and crustum for tart, with reference to the resinous covering of the tart-like gall. Affinities—Galls of this species, short with a conical base set into an encircling leaf socket, are most like those of C. striolacrustum except that they are unlined laterally and sticky to the touch. For further remarks, see under C. flaticrustum. Biological notes.—In late May in Central Maryland, one can find young galls not yet com- pletely broken out of the leaf covering and simu- lating young galls of C. tubicola (Fig. 74) and con- taining first instars. In mid- to late June, galls are full size but still soft and contain first and second instars. In August one can still see green, softer galls with second instars, but also harder, brown galls with third instars. Pupae emerge from a hole in the thinner walls adjacent to the conical base. Collections of galls—AtLaBama: Ashville (to- mentosa); Auburn (ovata, tomentosa); Greenville (tomentosa); Oak Mt. St. Pk, Shelby Co. (ovata, pal- lida). ARKANSAS: 3 mi S Harrison (tomentosa); Magazine Mt., Logan Co. (ovata, texana, tomen- tosa); 2 mi S DeQueen (texana, tomentosa); Rt.1 at Red River, Little River Co. (tomentosa). CONNECTI- cut: Amston (ovata); Meriden (glabra, ovata); New Haven (glabra); Phoenixville (ovata); Plainfield (ovata); Southbury (glabra, tomentosa); Westchester (ovata). FLORIDA: Ponce de Leon (glabra). GEORGIA: Carnesville (glabra); 8 km E Fort Valley (pallida); Forsyth (glabra, ovata); Lizella (ovata, tomen- tosa). INDIANA: Spencer Co. (ovata). KENTUCKY: Brownsville (tomentosa); Grayson Lake St. Pk, Carter Co. (tomentosa); Morehead (tomentosa); My Old Kentucky Home St. Pk, Nelson Co. (ovata); Pennyrile St. Pk, Christian Co. (glabra, ovata); Winchester (lacinosa). Maine: North Berwick (ovata). MARYLAND: Beltsville, USDA Agricultural Research Center (glabra, tomentosa); Brighton (to- mentosa); Clarksburg (tomentosa); College Park (glabra); Ellicott City (tomentosa); Gaithersburg (tomentosa); Kemp Mill (glabra); Lusby (glabra); Potomac (tomentosa); Ridgely (tomentosa); St. Leonard (tomentosa); Silver Spring (glabra, tomen- tosa); Snow Hill (tomentosa); Spencerville (tomen- tosa); Wheaton (glabra, tomentosa). MassAcHu- seTTs: Great Barrington (ovata); Merrimac (ovata); Petersham (ovata); Wrentham (ovata). MISSISSIPPI: Natchez Trace Pkwy, Mile 193, Choctaw Co. (glabra, pallida, tomentosa); Noxubee Natl Wildlife Ref., Oktibbeha Co. (ovata, tomentosa); nr. Ses- sums, Oktibbeha Co. (ovata); 7 mi E Starkville (pallida). Missouri: Crowley Ridge Roadside Pk, Stoddard Co. (glabra, tomentosa); Holly Ridge State For., Stoddard Co. (glabra, tomentosa). NEw HampsuHire: Barrington (ovata); Dover (ovata); Kingston (ovata). New Jersey: Columbia (ovata); Nyack (tomentosa); Columbia (tomentosa). NEw York: Saratoga Springs (ovata). NORTH CAROLINA: Alamance (tomentosa); Alberta (glabra); Dortches (glabra); Fayetteville (tomentosa); Gastonia (tomen- tosa); Providence (glabra, tomentosa); Rowland (to- mentosa); Wise (tomentosa). Onto: Hocking St. For., 58 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Hocking Co. (tomentosa); Hocking Co. (ovata, Wells 1916: Fig. 30). Ontario: Dundas (ovata). SouTH CAROLINA: Georgetown (tomentosa); Rich- land Co. (Carya sp.). TENNESSEE: Gatlinburg (glabra, tomentosa); Greene Co., I 81, Mile 38.5 (ovata, tomentosa). TExAs: Daingerfield St. Pk, Morris Co. (tomentosa). ViRGINIA: Warren Co., Blue Ridge Pkwy Mile 4.6 (ovata); Carson (tomen- tosa); Dinwiddie (glabra, tomentosa); Huntley Meadows Pk, Fairfax Co. (tomentosa); Ladysmith (tomentosa); Rt. 81, mi. 13.9, Washington Co. (ovata). WASHINGTON, DC: National Arboretum (glabra, tomentosa); Walter Reed Army Medical Center (tomentosa). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (tomentosa); Cooper’s Rock St. Pk, Preston Co. (glabra); 6 mi. NW Hedgesville (to- mentosa). Specimens of C. melicrustum examined.—ALa- BAMA: Greenville, X-20—2000 (larva). CONNECTI- cuT: Plainfield, VII]I-27-1988 (larvae). INDIANA: Spencer Co., IX-20-1995 (larva). MARYLAND: Beltsville, USDA Agricultural Research Center, IX-26-1993 & VIII-22-2000 (larvae), & College Park, IX-23-2003 (larvae; 2 2, pupae, em. IV- 10-2000); ‘Kemp: Mill, X-3-1993 (2, em. IV- 26-1994); Lusby, IX-16—2003 (larvae); Ridgely, IX- 23-1999 & X-2-2000 (larvae); Silver Spring, IX-17-1999 & IX-12-1999 (larvae); Wheaton, IX- 19-1993 (larvae). NORTH CAROLINA: Providence, X- 18-2000 (larva). ONTARIO: Dundas, VIII-17-1995 (larva). VircintA: Huntley Meadows Pk, Fairfax Co., VII-30—2000 (larvae). WASHINGTON, DC: Na- tional Arboretum, V-24—1994 (larvae). WEsT Vir- GINIA: 6 mi NW Hedgesville, IX-19-1999 (larvae). Caryomyia ovalis Gagné, new species Description.—Gall (Figs. 110-111): Rare, on Eu- carya hickories; on lower leaf surface, between veins; 2.7-3.5 mm in height, ovoid to elongate ovoid, usually longer than wide, slightly bilater- ally compressed; smooth, shining, hairless, not sticky, green, turning brown; base of gall abruptly tapered to conical pedicel, leaf with short, thick exfoliation at base of gall forming short, tubular socket, with corresponding convexity on opposite leaf surface; walls hard, enclosing two chambers, the basal larval chamber about 1/3 length of gall, ovoid, smooth, longitudinally ridged, the apical false chamber elongate, thicker walled, gradually narrowed to apex. This gall is two-chambered as is that of C. caminata, but the walls of the present galls are thick and shiny, unlike those of C. cami- nata that are brittle, foliaceous, and matte. Adult and pupa: Unknown. Larva: Third instar: Length 1.7 mm (n=1). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 325) with anterior tooth equilateral, the shaft broadest anteriorly, parallel-sided beyond. Papillae without setae, their bases not raised. Lat- eral papillae 2 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Abdomi- nal segments with verrucae dorsally along mid- length and with horizontal rows of pointed spicules anteroventrally but not posteroventrally. Terminal segment shorter than eighth. Second in- star: Length 1.1-1.2 mm (n=3). Collar segment with patch of enlarged spicules on each side. Spat- ula with acutely triangular anterior tooth and longer, slightly wider shaft. Bases of dorsal and pleural papillae not raised, papillae asetose. Six dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. glabra, Kemp Mill, Maryland, X-1—2005, R.J. Gagné, deposited in USNM. Etymology.—The name ovalis, a Latin adjective meaning egg-shaped, refers to the general shape of the gall. Affinities—No obvious relative of this rare species presents itself. The distinctive gall is most similar to that of C. inflata, but taller and much thicker and woodier. Collections of galls: MaryLaANp: Kemp Mill (glabra); Silver Spring (glabra); Spencerville (to- mentosa). Specimens of C. ovalis examined: MARYLAND: Kemp Mill, X-1-2005 (larva); Spencerville, VII- 8—2004 (larvae). Caryomyia persicoides (Osten Sacken) Cecidomyia persicoides Osten Sacken 1862: 193; Felt 1918: 46 (Caryomyia). Description—Gall (Figs. 17, 126-129): Com- mon, found on Eucarya hickories; single or clus- tered, on veins of lower leaf surface; 5.1—-9.0 mm in height, spheroidal or conical, base flat against leaf surface, clasping leaf vein, edge around base acute and lobed to obtusely rounded, tissue on apical third eventually collapsing somewhat and becoming furrowed when mature; covered with short, dense, velvety, green, yellow, tan, or red RAYMOND J. GAGNE 59 hair obscuring surface; connection to leaf circular and shallowly concave or flush with leaf; larval chamber located basally, ovoid, lined with yel- lowish pellicle different in texture from surround- ing soft, spongy, green to purple, viscous tissue that partially collapses with age; a bundle of fibers present between apex of larval chamber and gall apex. Galls may be gum-drop shaped to nearly spherical. The uniform covering of dense, short hair is only slightly longer in galls of C. spherica, but the latter galls are always spherical, hard and woody, and the larval chamber is located at the very center of the gall. Adult: Head: Antenna with 12 binodal flagel- lomeres, male flagellomeres (as in Fig. 239) each with definite internode and neck, the circumfila with short loops, their bases widely separated; fe- male flagellomeres with 2 horizontal circumfila connected by 2 vertical strands. Labella (as in Fig. 242) separated from one another anteriorly, hemi- spherical, short-setose, short-setulose. Palpus 4- segmented. Thorax: Wing length 2.9-4.0 mm in male (n=6), 2.9-4.1 mm in female (n=8). Male ab- domen: Third to eighth tergites each with poste- rior setae continuous across width in mostly dou- ble row laterally, tapering to single row mesad, eighth tergite with 2-3 setal rows laterally. Geni- talia (Figs. 256-257): hypoproct widened laterally to curve around sides of aedeagus, posterior mar- gin setose, dorsal surface setulose only on poste- rior fourth; apex of aedeagus convex; gonostylus broad apically. Female abdomen: Third through eighth tergites as for male, setae of eighth tergite may be interrupted mesally but almost as long as on preceding tergite. Ninth segment (Fig. 265) pli- able except for wide dorsal sclerite that partly in- vests cerci, setae sparse laterally and ventrally, mostly longer than cerci. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 2.0-3.1 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 314) with long, nearly parallel-sided, api- cally notched tooth and long shaft. Papillae with- out setae, bases not raised. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdominal segments with rounded verrucae covering dorsum at mid- length and horizontal rows of spicules present on venter anterior to sternal papillae and posterior to ventral papillae. Terminal segment shorter than eighth. Second instar: Length 1.0-1.4 mm (n=5). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae not raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Syntypes: Three young galls, from Carya sp., vic. Washington, DC. Osten Sacken, deposited in USNM. Etymology.—Osten Sacken (1862) named this species persicoides, Latin for peachlike, for the dense short hair covering the gall. Affinities.—Antennal and male genitalic char- acters of this species are similar to those of C. tur- banella. The male antennal flagellomeres are long with definite nodes and internodes. The aedeagus is rounded apically and the hypoproct is ex- panded laterally to cover the sides of the aedea- gus. The ovipositor of the present species (Fig. 265), however, is elongate and the cerci are re- duced in size, while in C. turbanella the ovipositor is similar to that in Fig. 259, the more common form found in the genus. Galls somewhat resem- ble those of the C. sanguinolenta species group be- cause of their extensive soft tissue and the pres- ence of a pellicle lining the larval chamber. A large difference between their galls is that only the top section of C. persicoides and C. turbanella galls breaks off over the winter as a unit, while all the soft tissue on the top and sides of the galls of the C. sanguinolenta group eventually separates from the brittle pellicle. Also, pupae of C. persicoides and C. turbanella break out of the side of the gall, while those of the C. sanguinolenta group break out of the bottom of the larval chamber. In addi- tion to these differences, adults of the two groups are different enough that it appears soft-tissued galls evolved at least twice in Caryomyia. Biological notes.—Galls of this species seem to be more commonly found in late summer. In cen- tral Maryland I occasionally found small galls with first instars beginning in mid-June but the galls seemed more numerous beginning in early August, when I found galls with larvae in first or second instars. Third instars were apparent in late 60 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) August and September. Pupae break through the side of the gall. [once noticed a sciarid pupal exu- viae sticking out of a hole near the gall’s apex; possibly some other organism made the hole. Collections of galls—AtaBamMa: Auburn (glabra, tomentosa); Greenville (tomentosa). ARKAN- sas: Beulah (tomentosa); Chester (texana); Osage (texana); Palestine (ovata). CONNECTICUT: Amston (ovata); Danbury (ovata); Harwinton (ovata); Meri- den (ovata); New Haven (glabra); Oakville (ovata); Phoenixville (ovata); Southbury (tomentosa). FLoR- IDA: Chattahoochee (tomentosa); Gainesville (gla- bra, tomentosa). Georcia: Carnesville (glabra); Forsyth (glabra); 8 km E Fort Valley (pallida); Lake Park (glabra); Lizella (ovata, tomentosa). ILLINOISs: Dexter (fomentosa); Marshall (tomentosa); Rock Is- land (tomentosa). INDIANA: New Castle (glabra). Kentucky: Brownsville (tomentosa). MARYLAND: Beaver Dam (tomentosa); Beltsville, USDA Agricul- tural Research Center (glabra, tomentosa); Brighton (tomentosa); Clarksburg (glabra, tomentosa); Colum- bia (glabra); Ellicott City (tomentosa); Gaithersburg (tomentosa); Kemp Mill (glabra); Lusby (tomentosa); Piney Grove (ovata); Potomac (tomentosa); Ridgely (glabra, tomentosa); Silver Spring (glabra, tomentosa); Snow Hill (tomentosa); Spencerville (tomentosa); Wheaton (glabra, tomentosa). MAssACHUSETTS: Wrentham (ovata). Missourt: Conway (ovata, tex- ana); St. Clair (tomentosa); Wildwood (tomentosa). New Hampsuire: Greenland (ovata); Kingston (ovata). New York: Addison (glabra); East Hamp- ton (Carya sp.); Herkimer (ovata); Ithaca (ovata); Ravena (glabra, ovata); Shokan (glabra). NortH CaroLina: Alamance (tomentosa); Alberta (glabra); Dortches (glabra, tomentosa); Fayetteville (tomen- tosa); Providence (glabra, tomentosa); Wise (tomen- tosa). Onto: Hocking St. For., Hocking Co. (ovata); Hocking Co. (glabra, ovata, tomentosa, Wells 1916: Fig. 17). OKLAHOMA: Sallisaw (tomentosa). Ontario: Dundas (ovata). PENNSYLVANIA: Kirby (ovata); Schellsburg (glabra). SourH CAROLINA: Dil- lon (tomentosa); Georgetown (tomentosa). TEN- NESSEE: Crossville (pallida, tomentosa); Dandridge (tomentosa); Dickson (tomentosa). Texas: New Wa- verly (texana). VIRGINIA: Carson (tomentosa); Din- widdie (glabra, tomentosa); Huntley Meadows Pk, Fairfax Co. (tomentosa); Ladysmith (tomentosa); Radford (tomentosa); Skippers (tomentosa). WasH- INGTON, DC: National Arboretum (glabra, tomen- tosa); Walter Reed Army Medical Center (ftomen- tosa); unspecified (types of C. persicoides). WEST VIRGINIA: 6 mi NW Hedgesville (ovata). Specimens of C. persicoides examined.—Con- NEcTicuT: Amston, VIII-27-1997 (larvae). FLORIDA: Gainesville, IX-21—2000 (larvae). GEorGIA: 8 km E Fort Valley, X-11-1993, J. & R. Payne (larvae); Lizella, X-11-1993, J. & R. Payne (larvae). KEN- Tucky: Brownsville, IX-21-1995 (larvae). MAry- LAND: Brighton, X-VIII-1994 (larvae); Gaithers- burg, X-4-1994 (larvae); Beltsville, USDA Agricultural Research Center, [X-11-1994 (larvae; 3 ¢, pupae, em. IV-1995), IX-26-1993, VIII- 25-2000 & IX-30-2003 (larvae); Gaithersburg, X- 4-1994 (larvae); Lusby, IX-16-2005 (d, 2, pupae, em. IV-2 to 3-2006); Ridgely, IX-23-1999 (larvae); Silver Spring, IX-22-1993 & X-24-1999 (larvae). Missouri: Wildwood, IX-8-2004 (larva). NEw Hampsuire: Kingston, IX-21-1994 (larvae). New York: Herkimer, [X-4—2003 (larvae). Onto: Hock- ing St. For., Hocking Co., IX-19-1995 (larvae). ON- TARIO: Dundas, VIII-17—1995 (larvae). TExas: New Waverly, X-6-2001 (3, 2, em. IV-2002). WASHING- TON, DC: National Arboretum, IX-3-1994 (larvae; 2 6,2 2, pupae, em. IV-1995), X-3-1995 (3 6,5 2, pupae, em. IV-1996), X-13-1997, IX-22-1998, and VII-13—2000 (larvae); Walter Reed Army Medical Center, IX-25-2003 (larvae); X-13-2004 (d, 2 9, em. IV-3-2005). West VIRGINIA: 6 mi NW Hedgesville,- IX-19-1999, X-17-1999 & VIII- 20-2000 (larvae). Caryomyia procumbens Gagné, new species Description —Gall (Figs. 94-95): Occasional, found on Eucarya hickories, but not east of the Ap- palachians, except in New England; usually found singly, attached to vein on lower leaf surface; 4—6 mm in length, horizontal, elongate, straight, nar- rowing gradually from rounded base to pointed apex; white, green, to tan, with stiff, sparse white hairs not obscuring surface; base of gall with cen- tral conical pedicel in shallow, circular indenta- tion; wall firm, brittle, uniformly thin, larval cham- ber glabrous with longitudinal ridges following gall axis from leaf vein connection to apex. The gall is generally similar to that of C. spinulosa, q.v. for further remarks about similar galls. Adult and pupa: Unknown. Larva: Third instar: Length 1.8—2.1 (10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 293) with single, almost parallel-sided tooth with shallow notch anteriorly, shaft wide anteriorly, narrowed beyond. Papillae without setae, their RAYMOND J. GAGNE 61 bases not raised. Lateral papillae 2-3 on each side of spatula. Only 4 dorsal papillae on first 7 ab- dominal segments. Abdomen with extensive ver- rucae on dorsum, particularly at midlength, and with horizontal rows of spicules on venter ante- rior to sternal papillae and covering posterior third. Terminal segment shorter than eighth. Sec- ond instar: Length: 0.9 mm (n=1). Collar segment with patch of enlarged spicules on each side. Spat- ula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae not appreciably raised, papil- lae asetose. Only 4 dorsal papillae on first 7 ab- dominal segments. Type material—Holotype: Third instar, from gall on C. ovata, Morgan Co., West Virginia, X- 12000, R.J. Gagné, deposited in USNM. Etymology.—The name procumbens is a Latin adjective and refers to the procumbent nature of the gall. Affinities —See under C. eumaris. Collections of galls—Connecticut: Danbury (ovata); Phoenixville (ovata); Plainfield (ovata); Thomaston (ovata). ILtinois: Dexter (laciniosa); Marshall (tomentosa). INDIANA: Spencer Co. (ovata). Kentucky: My Old Kentucky Home St. Pk, Nelson Co. (ovata); Pennyrile St. Pk, Christian Co. (ovata); Winchester (laciniosa). Mississippi: Natchez Trace Pkwy, Mile 193, Choctaw Co. (fomentosa); Noxubee Natl Wildlfe Refuge, Oktibbeha Co. (tomentosa). Missouri: Wildwood (tomentosa). OHIO: Summer- ford (tomentosa). PENNSYLVANIA: Mooresville (to- mentosa). TENNESSEE: Greene Co., 1 81, Mile 38.5 (to- mentosa). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (tomentosa); Cacapon (tomentosa); 6 mi NW Hedgesville (glabra, ovata). Specimens of C. procumbens examined.—ILLI- Nots: Marshall, IX-7-2004 (larva). INDIANA: Spencer Co., IX-20-1995 (larvae). KENTucky: Win- chester, IX-22-1995 (larvae). On1o: Summerford, IX-7-2004 (larvae). WEST VIRGINIA: 6 mi NW Hedgesville, IX-12-1999, VII-8-2000, & X-3-2003 (larvae); Morgan Co., X-1—2000 (larva). Caryomyia purpurea Gagné, new species Description.—Gall (Figs. 15, 120-122): Com- mon, found on Eucarya hickories; on lower leaf surface, usually found in groups between veins but may occur close to veins; 2.2-3.0 mm in height, spheroidal, occasionally with slight, coni- cal prominence at apex, surface covered with long, thick, mostly dark brown hair obscuring gall surface; base with deep, circular, central excava- tion, the leaf without exfoliation surrounding con- nection; wall woody, of even thickness but thicker apically if peak present, green to brown; larval chamber at base of gall ovoid, violet to purple. Ex- teriorly, the more conical galls are similar to the more short-conical galls of C. holotricha; such galls must be cut open to see whether hairs intrude into the larval chamber from the apex as they do in galls of C. holotricha. Adult: Head: Antenna with 12 flagellomeres; each of male flagellomeres (as in Fig. 239) binodal with definite internode and neck, circumfila with short loops, their bases widely separated; female flagellomeres with 2 horizontal circumfila con- nected by 2 vertical strands. Labella separated from one another anteriorly, hemispherical, short- setose, short-setulose. Thorax: Wing length 3.1-3.4 mm in male (n=5), 2.5-3.5 mm in female (n=5). Male abdomen: Third through eighth ter- gites each with posterior setae continuous across width in mostly single row, occasionally in partly double row, the line broken for short distance mesally; eighth tergite not more weakly sclero- tized than seventh, the setae as long. Genitalia (Figs. 254-255): hypoproct nearly parallel sided, deeply concave posteromesally, posterior margin setose except in concavity; apex of aedeagus con- vex. Female abdomen: Third through eighth ter- gites each with posterior setae continuous across width in mostly single row, occasionally in partly double row, the line broken for short space mesally; eighth tergite not more weakly sclero- tized than seventh, the setae as long. Ninth seg- ment pliable except for fairly wide dorsal sclerite, setae long but sparse. Cerci elongate-ovoid, pointed apically, pigmented, fused dorsally on basal third. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.8—2.2 mm (n=10). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 306) with parallel-sided apically- notched apical tooth, the space in notch erose in some specimens, and elongate, narrow shaft. 62 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Papillae without setae, their bases not raised. Lat- eral papillae numbering 2-3 on each side of spat- ula. Only 4 dorsal papillae present on first 7 ab- dominal segments. Abdominal segments with verrucae along midlength of dorsum and horizon- tal rows of spicules anterior to sternal papillae on venter and also posterior to ventral papillae. Ter- minal segment shorter than eighth. Second instar: Length 0.9-1.3 mm (n=4). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae not raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. glabra, Beltsville, Maryland, IX-28-1999, R.J. Gagné, deposited in USNM. Etymology.—The name purpurea is a Latin ad- jective meaning purple, with reference to the color of the larval chamber walls. Affinities—This species differs from others, such as C. tuberculata and C. holotricha, which also have long male flagellomeres, an apically- rounded aedeagus, and a deep, circular, basal ex- cavation of the gall, but the larva of the present species has a notched spatula and the first though seventh abdominal segments each have only four dorsal papillae. The other two species have a tri- angular spatula tooth and six dorsal abdominal papillae. Collections of galls —ALaBaAmMa: Auburn (ovata, tomentosa); Greenville (tomentosa). ARKANSAS: Piney (texana). Connecticut: Danbury (ovata); Oakville (ovata); Phoenixville (ovata); Plainfield (ovata); Southbury (tomentosa); Thomaston (ovata); Westchester (ovata). GEORGIA: Forsyth (glabra); 8 km E Fort Valley (pallida). Lake Park (glabra). TLu1- NoIs: Dexter (laciniosa, tomentosa). MARYLAND: Beltsville, USDA Agricultural Research Center (fo- mentosa); Brighton (tomentosa); College Park (glabra); Ellicott City (tomentosa); Gaithersburg (to- mentosa); Lusby (glabra, tomentosa); Ridgely (glabra, tomentosa); Silver Spring (tomentosa); Spencerville (tomentosa); Wheaton (glabra, tomentosa). Massa- CHUSETTS: Great Barrington (ovata). Missourt: Con- way (texana); Rolla (texana); St. Clair (tomentosa). New Hampsuire: Kingston (ovata). NEw JERSEY: Co- lumbia (glabra, tomentosa). New York: E. Schodack (Carya sp.); Hyde Park (ovata); New York (Carya sp.). NortH Carona: Alamance (glabra, tomen- tosa); Fayetteville (tomentosa); Providence (glabra, tomentosa); Wise (tomentosa). Onto: Hocking Co. (tomentosa, Wells 1916: Fig. 15); Yellow Springs (glabra). ONTARIO: Dundas (ovata). SouTH CarR- OLINA: Georgetown (tomentosa). ‘TENNESSEE: Crossville (pallida); Dickson (tomentosa). VIRGINIA: Dinwiddie (glabra, tomentosa); Huntley Meadows Pk, Fairfax Co. (tomentosa).; Ladysmith (tomentosa). WasHINGTON, DC: National Arboretum (glabra, to- mentosa); Rock Creek Park (ovata); Walter Reed Army Medical Center (tomentosa). Specimens of C. purpurea examined.—Con- NEcTICcUT: Phoenixville, IX-20-1994 (larvae). Georaia: 8 km E Fort Valley, IX-11-1993, J. & R. Payne (larvae). MARYLAND: Beltsville, USDA Agri- cultural Research Center, X-5-1994 & IX-28-1999 (larvae); Brighton, X-8-1994 (larvae; 3 6,4 2, em. IV-1995); Ridgely, IX-23-1999 (larvae), X-2000 (3 3,em. IV-2001); Gaithersburg, X-4-1994 (larvae; 2 ?, pupae, em. IV-1995); Silver Spring, X-3-1993 (larvae; 10 5,9 2, pupae, em. IV-26-1994); Spen- cerville, IX-22—2003 (larvae; 2, em. IV-14—2004); Wheaton, IX-19-1993 & X-13-1999 (larvae). Mis- souRI: Conway, IX-8—2004 (larvae). NEw Hamp- SHIRE: Kingston, IX-21—1994 (larvae). Onto: Yellow Springs, IX-7-2004 (larvae). Ontario: Dundas, VIH-17-1995 (larvae). WASHINGTON, DC: National Arboretum, X-3-1994 (larvae); Walter Reed Army Medical Center, X-7-1993 (larvae; 3 5,39, pupae, em. IV-23—26-1994), X-2005 (larvae; pupae, 5 d, 26 2, em. III-28 to IV-15—2006). Caryomyia recurvata Gagné, new species Description. —Gall (Figs. 9, 90-91): Occasional, found on Eucarya hickories; usually in groups, on lower leaf surface on vein; 2.5—-4.0 mm in length, recumbent, truncate near base, abruptly nar- rowed and recurved beyond basal third; white, green, to tan, hairless, glaucous; base of gall with central, conical pedicel surrounded by shallow, circular indentation; wall firm, brittle, uniformly thin except for slightly thicker blunt end, larval chamber glabrous, with longitudinal ridges fol- lowing gall axis from leaf vein connection to re- curved apex. This gall is most similar to that of C. eumaris but is hairless and has a smooth glaucous surface and has a more truncate base. The gall of C. eumaris has a rough and sparsely hairy surface. Adult (male only): Head: Antenna with 12 fla- gellomeres; male flagellomeres (as in Fig. 237) scarcely binodal, only slightly narrowed beyond basal circumfilum, circumfila loops short, sube- qual in length. Labella (as in Fig. 242) separated RAYMOND J. GAGNE 63 from one another anteriorly, hemispherical, short- setose, short-setulose. Palpus 4-segmented. Tho- rax: Wing length 1.5 mm (n=1). Male abdomen: Third through seventh tergites each with poste- rior setae continuous across width, in single row; eighth tergite weakly sclerotized, with 3 posterior setae laterally. Genitalia: gonostylus gradually narrowed from base to apex, the tooth relatively narrow; hypoproct broadest near apex with slight concavity posteromesally, apical margin setose except in concavity, dorsal setulae present only on distal third; aedeagus furrowed apically and ventrally. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.6—2.2 mm (n=7). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 294) with nearly parallel-sided, shallowly notched anterior tooth, the shaft widest anteriorly, tapering beyond. Papillae without setae, their bases not raised. Lateral papillae 2-3 on each side of spatula. Only 4 dorsal papillae on first 7 abdom- inal segments. Abdomen with extensive verrucae on dorsum, particularly at midlength, and with horizontal rows of spicules on venter anterior to sternal papillae and covering posterior third. Ter- minal segment shorter than eighth. Second instar: Length 0.8 mm (n=1). Collar segment with patch of enlarged spicules on each side. Spatula with large, apically pointed tooth and longer, wider, concave- sided shaft. Bases of dorsal and pleural papillae not appreciably raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material—Holotype: Third instar, from gall on C. glabra, National Arboretum, Washing- ton, DC, IX-23-1988, R.J. Gagné, deposited in USNM. Etymology.—The name recurvata is a Latin ad- jective refering to the recurved neck of the gall. Affinities.—See under C. eumaris. Collections of galls—AtaBama: Auburn (gla- bra, pallida). ARKANSAS: Magazine Mt., Logan Co. (tomentosa); 2 mi S DeQueen (tomentosa); Rt.1 at Red River, Little River Co. (tomentosa). CONNECTI- cut: Phoenixville (ovata); Westchester (ovata). TLLI- NoIs: Dexter (tomentosa); Marshall (tomentosa). Kentucky: Grayson Lake St. Pk, Carter Co. (to- mentosa); Pennyrile St. Pk, Christian Co. (glabra, ovata). LOUISIANA: Lake Bistineau St. Pk, Webster Parish (tomentosa). MARYLAND: Beltsville, USDA Agricultural Research Center (tomentosa); Ellicott City (tomentosa); Gaithersburg (tomentosa); Ridgely (tomentosa); Snow Hill (tomentosa). Mis- sissippi: Noxubee Natl Wildlife Ref., Oktibbeha Co. (ovata, tomentosa). Onto: Hocking Co. (ovata, Wells 1916: Fig. 9). Missourt: Conway (ovata, tex- ana); Holly Ridge State For., Stoddard Co. (tomen- tosa); St. Clair (tomentosa); Wildwood (tomentosa). TENNESSEE: Dickson (tomentosa). WASHINGTON, DC: National Arboretum (glabra). Specimens of C. recurvata examined.—Mary- LAND: Beltsville, USDA Agricultural Research Center, X-5-1994 & VII-1-2000 (larvae); Ridgely, IX-23-1999 (larvae). Missourt: Conway, IX-8-2004 (3, pupae, em. IV-9-2005); Wildwood, IX-8-2004 (larvae). WASHINGTON, DC: National Arboretum, IX-22-1988 (larva). Caryomyia sanguinolenta (Osten Sacken) Cecidomyia sanguinolenta Osten Sacken 1862: 192; Felt 1909: 293 (Caryomyia). Description —Gall (Figs. 10, 134-136): Com- mon, found on Eucarya hickories; single or clus- tered, on lower leaf surface between veins; height 1.9-4.1 mm, conical, basal edge flared, acute or rounded, the base adjacent to leaf either flat or convex, the central part of the base permanently hard and differing in texture from softer outer part (Fig. 136), lateral sides of cone in profile con- vex at first, becoming concave, then furrowed upon further drying, the apex obtuse to acute; hairless, green, turning brown, crimson, or pur- ple, not sticky; base with shallow, circular, central excavation, the leaf without exfoliation around connection; larval chamber ovoid, located at base of gall, surrounded by yellowish pellicle different in texture from the soft, large-celled usually pur- ple tissue beyond that partly collapses with age; a bundle of fibers present between apex of larval chamber and gall apex. Galls of C. sanguinolenta and C. conoidea are most similar to one another in shape, but that of C conoidea bears short, rigid, sparse hairs. Adult: Head: Antenna with 12 flagellomeres; male flagellomeres (as in Fig. 237) scarcely bin- odal, only slightly narrowed beyond basal circum- 64 THE GALL MIDGES (DIPTERA: CECIDOMYITIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) filum, circumfila loops short, subequal in length; female flagellomeres with 2 horizontal circumfila connected by 2 vertical strands. Labella (as in Fig. 242) separated from one another anteriorly, hemi- spherical, short-setose, short-setulose. Palpus 4- segmented. Thorax: Wing length 1.7-2.1 mm in male (n=5); 2.1—-2.7 mm in female (n=4). Male ab- domen: Third through seventh tergites each with posterior setae continuous across width, in single row; eighth tergite weakly sclerotized, without posterior setae. Genitalia (as for Figs. 247-248): gonostylus gradually narrowed from base to apex, the tooth relatively narrow; hypoproct broadest near apex with slight concavity posteromesally, margin setose except in concavity, dorsal setulae present only on distal third; aedeagus furrowed apically. Female abdomen: Third through eighth tergites each with posterior setae continuous across width, in mostly single row, usually double at lateral limits; eighth tergite not less sclerotized than seventh, setae nearly as long. Ninth segment pliable except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ventrally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (Figs. 213-214): Antennal bases pyrami- dal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal incisions, their posterior lip protruding to form craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with several anterior rows of enlarged spinules. Larva (Figs. 220-222): Third instar: Length 18-25 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 303) with long, nearly parallel sided, apically notched tooth and long shaft. Papillae without setae, their bases not raised. Lateral papillae 2 on each side of spatula. Four dorsal papillae on first 7 abdominal seg- ments. Abdominal segments with rounded and pointed verrucae covering dorsum extensively across midlength and with horizontal rows of pointed spicules across venter anterior to sternal papillae and posterior to ventral papillae. Termi- nal segment shorter than eighth. Second instar: Length 0.8-1.2 mm (n=8). Collar segment with patch of enlarged spicules on each side. Spatula present, with acutely triangular anterior tooth and elliptical shaft. Dorsal and pleural papillae not appreciably raised, about as long as wide, without setae. Four dorsal papillae on first 7 ab- dominal segments. Type material.—Neotype: The original Osten Sacken syntypes, larva(e) and galls collected from Carya sp. in the vicinity of Washington, DC, are presumed lost. They were not found in either the MCZ or the USNM. A neotype is designated here, a third instar (right hand specimen of two on a slide) from galls fitting Osten Sacken’s original description, Carya tomentosa, Walter Reed Army Medical Center Annex, Silver Spring, Maryland, X-24-1999, R.J. Gagné, deposited in the USNM. The galls from which the neotype and the other associated larva were taken are also preserved in the USNM. Etymology.—The name sanguinolenta is an ad- jective from the Latin meaning blood red, coined by Osten Sacken with reference to the crimson to purple external color of most mature galls of this species. Affinities.—Galls of this species resemble closely enough those of several other species, C. biretta, C. conoidea, C. lunata, C. marginata, and C. stellata, to indicate some relationship among them, given other similarities of adult and larval characters. These species differ only in the shape of their galls, which are conical with their exten- sive soft, large-celled tissue eventually collapsing. Between that tissue and the larval cell is a pellicle that is at first soft but eventually hardens into a brittle covering. The soft tissue mostly deterio- rates during the winter. The base of the galls, through which pupae eventually escape, is shield- like and smooth, even on otherwise hairy galls. The conical galls only superficially resemble those of C. persicoides and C. turbanella. Galls of these last two deteriorate differently and pupae break out of the side of the gall rather than the base. Also, the male antennae and genitalia of these two species place them in another group. Biological notes.—Tiny, green, occasionally red galls with first instars were found as early as mid- May in the Washington, DC area. By mid-June, both small and full-size galls were present with first and second instars, respectively. Small galls with first instars could still be found as late as early August, although the preponderance of galls found after mid-July contained second in- stars. Third instars were found beginning in early August and by late August most galls contained third instars. Galls remain on leaves until leaf drop in October. Many galls are red or purple by autumn but some remain green. Pupae escape through a hole made in the base of the gall, to the side of the central, harder part (Fig. 136). RAYMOND J. GAGNE 65 Collections of galls—ALaBama: Ashville (to- mentosa); Auburn (glabra, ovata, pallida); Oak Mt. St. Pk, Shelby Co. (pallida). ARKANSAS: 3 miS Harrison (tomentosa); Magazine Mt., Logan Co. (ovata, to- mentosa); Piney (texana). CONNECTICUT: Amston (ovata); Meriden (glabra, ovata); New Haven (glabra); Pachaug St. Pk (ovata); Phoenixville (ovata); Southbury (tomentosa); Westchester (ovata). FLoripa: Chattahootchee (tomentosa); Gainesville (tomentosa). GeorGIA: Carnesville (glabra); Forsyth (glabra, ovata); Fort Valley (pallida); Lake Park (glabra); Lizella (ovata). ILLINOIS: Dexter (tomen- tosa). KENTUCKY: Brownsville (tomentosa); Grayson Lake St. Pk, Carter Co. (tomentosa). LOUISIANA: Lake Bistineau St. Pk, Webster Parish (texana). MaryYLAND: Beaver Dam (tomentosa); Beltsville, USDA Agricultural Research Center (glabra, tomen- tosa); Brighton (tomentosa); Calverton (tomentosa); Clarksburg (glabra, tomentosa); College Park (glabra); Ellicott City (tomentosa); Gaithersburg (to- mentosa); Kemp Mill (glabra); Lusby (glabra, tomen- tosa) Potomac (tomentosa); Ridgely (glabra, tomen- tosa); St. Leonard (tomentosa); Silver Spring (glabra, tomentosa); Spencerville (tomentosa); Wheaton (glabra, tomentosa). MAssAcuusETTs: Bolton (ovata); Merrimac (ovata); Wrentham (ovata). Missouri: Holly Ridge State For., Stoddard Co. (glabra). New Hampsuire: Kingston (ovata). New Jersey: Colum- bia (tomentosa). NEw York: Central Pk, New York (tomentosa); Ravena (glabra, ovata); Skokan (glabra). NortH CAROLINA: Alamance (tomentosa); Alberta (glabra); Fayetteville (tomentosa); Kannapolis (to- mentosa); Providence (glabra, tomentosa); Rowland (tomentosa); Wise (tomentosa). Onto: Gypsum (glabra, Wells 1916, Fig. 10). Ontario: Dundas (ovata). PENNSYLVANIA: Millsboro (glabra); Schells- burg (glabra, ovata). TENNESSEE: Dandridge (tomen- tosa); Lebanon (glabra). VirGINIA: Carson (tomen- tosa); Leesburg (tomentosa); Skippers (tomentosa). WASHINGTON, DC: National Arboretum (glabra, to- mentosa); Walter Reed Army Medical Center (to- mentosa). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (tomentosa); Cacapon (ovata); 6 mi NW Hedgesville (ovata). Specimens of C. sanguinolenta examined.— ARKANSAS: Piney, IX-10-2004 (larva). CONNECTI- cut: New Haven, IX-19-1994 (larva); Phoenixville, IX-20-1994 (larvae). Georaia: Lizella, X-12, 1993, J. & R. Payne (larva). ILLINoIs: Dexter, IX-8-2004 (lar- vae). KeNTucKy: Brownsville, IX-21-1995 (larva); Grayson Lake St. Pk, Carter Co., IX-22-1995 (larva). MARYLAND: Beltsville, USDA Agricultural Research Center, I-7—2000 (larvae); Brighton, X- 1OF199301(26 7278 pupae, em. slV-25—28-1993); Calverton, X-10-2005 (3,5 2, pupa & pupal exu- viae, em. III-30 to IV-15—2006); College Park, IX- 23-2003 (2 5,2 ¢, pupae, em. IV-10—2004); Ellicott City, IX-2-2000 (larvae); Gaithersburg, X-4-1994 (larvae); Lusby, IX-16-2003 (larvae); Ridgely, IX- 23-1999 (larvae); Silver Spring, IX-14-1993, IX- 17-1993, IX-22-1993, and X-24-1999 (larvae); Wheaton, IX-19-1999 (larvae). WASHINGTON, DC: National Arboretum, X-3-1994 (larva); Walter Reed Army Medical Center, IX-2000 (larvae; ¢, 3 2, pupae, em. IV-2001). WEsT VIRGINIA: 6 mi NW Hedgesville, X-17-1999 (larvae). Caryomyia shmoo Gagné, new species Description.—Gall (Figs. 59-61): Common, found on Eucarya hickories; single or clustered, on lower leaf surface between veins; 2.5-3.5 mm in height, bilaterally symmetrical, truncate-spher- oidal at base, apical third abruptly narrowed, set to one side, and tipped with an umbo; green to yel- low, turning tan to brown, smooth, hairless, not sticky; base with prominent conical pedicel set into circular indentation, leaf without conspicuous ex- foliation surrounding base of gall; wall firm, brit- tle, thin, larval chamber shaped as for gall, glabrous, with longitudinal ridges. This gall re- sembles most that of the related species, C. caryae. The gall of the latter is spheroidal but that of C. shmoo is conspicuously narrowed at its apical third. Adult: Head: Antenna with 12 flagellomeres, in male with additional short 13th flagellomere without circumfila; each of regular 12 male flagel- lomeres (as for Fig. 239) binodal with definite in- ternode and neck, circumfila with short loops, their bases widely separated; female flagellom- eres with 2 horizontal circumfila connected by 2 vertical strands. Labella separated from one an- other anteriorly, hemispherical, short-setose, short-setulose. Thorax: Wing length 2.7-3.6 mm in male (n=2), 3.0-3.1 mm in female (n=2). Male abdomen: Third through eighth tergites each with posterior setae separated into 2 lateral groups, in 1 to 3 horizontal rows; eighth tergite not more weakly sclerotized than seventh. Genitalia: hy- poproct nearly parallel sided, weakly concave posteromesally, the margin partly setose, dorsum setulose only on distal third; apex of aedeagus convex. Female abdomen: Third through seventh tergites each with posterior setae as for male; eighth tergite more weakly sclerotized than pre- ceding tergite, with posterior setae only laterally 66 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) and not as long as on preceding tergite. Ninth seg- ment pliable except for dorsal sclerite that widens from base to apex, setae sparse laterally and ven- trally, many longer than cerci. Cerci ovoid in lat- eral view, fused dorsally for about 2/3 length. Pupa (Figs. 201-202): Antennal bases oblique in ventral view, converging and narrowing anteri- orly, their rounded apices closely adjacent. Face with anterolateral pair of horizontal incisions, their posterior lip protruding to form concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with several anterior rows of en- larged spinules. Larva: Third instar: Length 1.8-2.4 mm (n=10). Body ovoid-cylindrical, anterior end more broadly rounded than posterior end. Head di- rected ventrally. Spatula (Fig. 284) with 2 widely separated anterior teeth, the space between con- cave, shaft widest anteriorly, narrowing abruptly before posterior half, parallel-sided beyond. Pap- illar bases moderately raised, dorsals and pleurals with setae, also ventrals with setae on first through seventh abdominal segments. Lateral papillae 3 on each side of spatula. Six dorsal papil- lae on first 7 abdominal segments. Body entirely covered with verrucae laterally and dorsally and with spicules covering venter. Terminal segment as long as eighth. Second instar: Length 0.8-1.2 mm (n=5). Collar segment with patch of enlarged spicules on each side. Spatula with acutely trian- gular anterior tooth and longer, wider concave- sided shaft. Bases of dorsal and pleural papillae raised and all short-setose including on protho- rax. Six dorsal papillae on first 7 abdominal seg- ments. Type material—Holotype: Third instar, left specimen on slide mount, from gall on C. glabra, Wheaton, Maryland, IX-19-1993, R.J. Gagné, de- posited in USNM. Etymology.—The name shmoo is a noun in ap- position. The shape of this gall is reminiscent of one of the benevolent and irrepressible organisms with that name created in 1948 by Al Capp for his comic strip, Li'l Abner. Affinities.—This species forms a natural group with C. caryae and C. hirtiglobus because of the general shape of the gall and the similarity of the larvae, particularly the two widely separated an- terior teeth of the third instar spatula. As with C. hirtiglobus but not C. caryae, the dorsal and pleural papillae of the larval prothorax are setose. C. shmoo differs from C. hirtiglobus in the apices of the pupal antennal horns being separated instead of closely adjacent and the larval spatula teeth separated from one another by a distance shorter than their length. See under C. caryae for further discussion. Biological notes.—Soft, whitish-green galls were first noted in central Maryland on June 9 and contained first instars. On June 15, full grown galls were brittle and still contained first instars. By July 8, galls contained second and third instars. When full-grown, the white larva fills the entire larval chamber and is very active when disturbed. Pupae exit the gall from one side through which the larva has already scraped away part of the wall. Collections of galls—AtaBama: Ashville (to- mentosa); Auburn (glabra, ovata, pallida, tomentosa); Greenville (tomentosa). ARKANSAS: Chester (texana); 2 miS DeQueen (texana, tomentosa); Piney (texana); Rt.1 at Red River, Little River Co. (tomentosa). CON- NEcTIcUT: Hartland (ovata); Meriden (ovata); Phoenixville (ovata); Southbury (glabra). FLORIDA: Gainesville (glabra, tomentosa); Chattahoochee (to- mentosa). GgEorciA: 8 km E Fort Valley (ovata). LouIsIANA: Lake Bistineau St. Pk, Webster Parish (texana). MARYLAND: Beltsville, USDA Agricultural Research Center (glabra, tomentosa); Clarksburg (to- mentosa); Gaithersburg (tomentosa); Lusby (tomen- tosa); Ridgely (tomentosa); Spencerville (tomen- tosa); Wheaton (glabra). Mississippi: Natchez Trace Pkwy, Mile 123, Rankin Co. (glabra, pallida, tomen- tosa); Natchez Trace Pkwy, Mile 193, Choctaw Co. (tomentosa); 7 mi E Starkville (tomentosa). MissOurI: Conway (texana); Mark Twain Natl For. (tomen- tosa); St. Clair (tomentosa). NEw HAMPSHIRE: Kingston (ovata). New York: Shokan (glabra); Saratoga Springs (ovata). NORTH CAROLINA: Fay- etteville (tomentosa). Onto: Yellow Springs (glabra). PENNSYLVANIA: Schellsburg (ovata). SourH CaR- OLINA: Georgetown (tomentosa). TExAs: New Wa- verly (tomentosa). VIRGINIA: Dinwiddie (tomentosa); Huntley Meadows Pk, Fairfax Co. (tomentosa); La- dysmith (fomentosa). WASHINGTON, DC: National Arboretum (tomentosa). WEST VIRGINIA: Cacapon (ovata); 6 mi NW Hedgesville (ovata). Specimens of C. shmoo examined.— ALABAMA: Greenville, IX-21-2000 (2 3, pupae, em. IV-2001). Georcia: 8 km E Fort Valley, X-11-1993, J. & R. Payne (larvae). MARYLAND: Beltsville, USDA Agri- cultural Research Center, IX-26-1993 & X-5-1994 (larvae); Gaithersburg, VII-20—2000 (larva); Lusby, IX-16-2003 (larvae); Ridgely, IX-23-1999 (larva); Spencerville, IX-4 & 21-2003 (larvae; 2, pupa, em. IV-2004); Wheaton, IX-19-1993 (larvae). NEw RAYMOND J. GAGNE 67 Hampsuire: Kingston, IX-2-1994 (larva). New York: Saratoga Springs, IX-25-1994 (larvae). Onto: Yellow Springs, IX-7—2004 (larvae). Texas: New Waverly, X-6-2001 (2, pupae, em. IV-2002). WASHINGTON, DC: National Arboretum, IX- 21-2003 (larvae). Caryomyia spherica Gagné, new species Description —Gall (Figs. 21, 160-161): Infre- quent, on Eucarya hickories; usually scattered along veins, on lower leaf surface; 5.0-8.0 mm in diameter, spherical, covered with thick, short, brown hair obscuring gall surface; base with nar- row, shallow, circular depression; larval chamber at center of sphere, ovoid, tan to brown, the sur- rounding thick walls large-celled, woody. Galls of this species are superficially similar to the rounder galls of C. persicoides, but the larval cham- ber of C. spherica is centrally located instead of at the base of the gall. The gall is also much harder and not subject to shrinkage as is the softer gall of C. persicoides. Adult: Head: Antenna with 12 flagellomeres and a short apical bud on the 12th; the regular 12 male flagellomeres almost evenly cylindrical, slightly narrowed from base to apex, circumfila loops short, subequal in length; female flagellom- eres with circumfila anastomozing on venter. La- bella (Fig. 244) discrete, ovoid, with many long se- tae and covered with setulae. Palpus (Fig. 244) 1-2 segmented. Labella separate with several robust setae. Thorax: Wing length 3.3-3.5 mm in male (n=2); 4.14.2 mm in female (n=2). Male ab- domen: Third through seventh tergites each with posterior setae continuous across width, 3-4 rows laterally, diminishing to 1-2 rows mesally; eighth tergite with 0-3 posterolateral setae. Genitalia: hy- poproct broadest near apex, slightly bilobed at apex, the lobes rounded; apex of aedeagus weakly furrowed. Female abdomen: Third through sev- enth tergites each with posterior setae continuous across width, in mostly 4 rows laterally, tapering to 2-3 rows mesally; eighth tergite with mostly double row of posterior setae laterally, diminish- ing to 0-1 rows mesally, and shorter than setae on previous tergite. Ninth segment pliable except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ven- trally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (Figs. 205-206): Antennal bases oblique in ventral view, converging anteriorly, each termi- nating in prominent, closely adjacent point. Face with anterolateral pair of horizontal incisions, their posterior lip slightly protruding. Frons and clypeus moderately protruded ventrally. Abdomi- nal terga with several anterior rows of enlarged spinules. Larva: Third instar: Length 2.3-4.2 mm (n=5). Body ovoid, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 317) with tapered, apically-pointed tooth and long shaft widened at both ends. Dorsal and pleu- ral papillae with short setae; bases of papillae not raised. Lateral papillae 2-3 on each side of spat- ula. Six dorsal papillae on first 7 abdominal seg- ments. Abdominal segments with verrucae cover- ing most of dorsum and with horizontal spicule rows anteriorly and posteriorly on venter. Termi- nal segment shorter than eighth. Second instar: Length 1.4-1.5 mm (n=10). Collar segment with patch of enlarged spicules on each side. Spatula with elongate, acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae raised, papillae with setae. Six dorsal papillae on first 7 abdominal seg- ments. Type material.—Holotype: Third instar, middle of 3 specimens on slide, from gall on C. tomentosa, Beltsville Agricultural Research Center, Beltsville, Maryland, IX-30-2003, R.J. Gagné, deposited in USNM. Etymology.—The name spherica is a Latin ad- jective, referring to the spherical gall. Affinities—Among the Caryomyia species with foreshortened male flagellomeres and fur- rowed aedeagus, this species stands alone for the one or two instead of four-segmented palpi and the strongly instead of weakly setose labella. Galls of C. spherica and the more spherical galls of C. persicoides can be superfically similar in gross as- pect. Larvae of C. spherica have a triangular apical tooth on the spatula and setose dorsal and pleural papillae and the adult has a 1-2 segmented pal- pus; in C. persicoides, the larva has a parallel-sided, notched tooth and asetose papillae and the adult has a 4-segmented palpus. Biological notes.—In autumn the full-grown larva cuts a cylindrical passage from its chamber in the center of the gall up to but not including the epidermis. In spring the pupa pushes through the epidermis. Adults were reared in early April. In my experience this gall appears in late summer. Collections of galls—AtaBAma: Auburn (glabra, pallida, tomentosa); Greenville (tomentosa). 68 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) ConnectTicuT: Meriden (ovata); Southbury (tomen- tosa). FLortDA: Chattahoochee (tomentosa). GEOR- GIA: Forsyth (ovata); Lizella (tomentosa). ILLINOIs: Highland (laciniosa); Rock Island (Carya sp., B.D. Walsh). MaryLANnb: Beltsville, USDA Agricultural Research Center (tomentosa); Ridgely (tomentosa). Mississippi: Starkville (tomentosa). NorTH CArR- OLINA: Alamance (tomentosa); Fayetteville (tomen- tosa); Gastonia (tomentosa); Providence (tomen- tosa). Ohio: Unspecified (ovata, Wells 1916: Fig. 21). TENNESSEE: Crossville (pallida); Jackson (tomen- tosa). Texas: New Waverly (tomentosa). VIRGINIA: Dinwiddie (tomentosa). Specimens of C. spherica examined.—GEorGIA: Lizella, X-12-1993, J. & R. Payne (larvae). ILLINOIs: Highland, IX-8-2004 (larvae); Rock Island, B.D. Walsh (larvae). MARYLAND: Beltsville, USDA Agri- cultural Research Center, IX-30—2003 (larvae; 2 3, pupae, em. IV-12—2004); Ridgely, IX-23-1999 (lar- vae; 2 2, em. IV-2000). NorTH CAROLINA: Gasto- nia, IX-18—2000 (larvae). TENNESSEE: Crossville, IX- 11-2004 (larva). Texas: New Waverly, X-6-2001 (larva; 2, em. IV-2002). Caryomyia spiniglobus Gagné, new species Description.—Gall (Fig. 64-65): Common, on Eucarya hickories; single or in groups on lower leaf surface between veins; 2.8-3.5 mm in height, spherical with a conspicuous apical terminus made up of a nipple atop an abruptly raised are- ola; green to yellow, becoming tan to brown with widely spaced, short, stiff light brown hairs not obscuring surface; base with small conical pedi- cel, leaf without exfoliation; wall firm, brittle, uni- formly thin, larval chamber glabrous with longi- tudinal ridges. This gall differs from that of C. leviglobus in that it has hair. Both galls are distinc- tive for the raised apical knob. Adult: Head: Antenna with 12 flagellomeres; each of the 12 male flagellomeres (as in Fig. 238) scarcely binodal, only slightly narrowed beyond basal circumfilum, circumfila loops short, sub- equal in length, basal node with almost no setulae, distal node evenly setulose; female flagellomeres with 2 horizontal circumfila connected by 2 vertical strands. Labella separated from one another ante- riorly, hemispherical, short-setose, short-setulose. Thorax: Wing length 1.7 mm in male (n=1); 2.0 mm in female (n=1). Male abdomen: Third through seventh tergites each with single row of posterior setae nearly continuous across width except inter- rupted mesally; eighth tergite without posterior se- tae. Genitalia: hypoproct broadest before posterior convex margin, margin setose; aedeagus evenly cylindrical, wide, notched apically. Female ab- domen: Third through seventh tergites each with single row of posterior setae nearly continuous across width except interrupted mesally; eighth tergite as sclerotized as preceding tergite, with sin- gle row of posterior setae only laterally, the setae not as long as on seventh tergite. Pupa : Unknown. Larva: Third instar: Length: 1.7-2.5 mm (n=10). Body spindleform, segments convex, head end tapered, posterior end rounded. Head ven- tral. Spatula (as in Fig. 281) with large, simple an- terior tooth nearly as long as broad shaft. All papillae except terminals setose, papillar bases not conspicuously raised. Lateral papillae 2 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Abdominal segments com- pletely covered with raised, pointed verrucae. Terminal segment as long as eighth. Second in- star: Not seen. Type material.—Holotype: Third instar, from gall on C. tomentosa, vic. Ridgely, Caroline Co., Maryland, R.J. Gagné, IX-23-1999, deposited in USNM. Etymology—The name spiniglobus is a noun formed from the Latin spina (spine) and globus (globe), with reference to the erect-haired spheri- cal gall. Affinities—See under C. leviglobus, from which the present species differs only in the hairy galls. Biological notes.—Tiny, soft, whitish galls with first instars were first noted in central Maryland on June 9. In mid-June to mid-July soft galls could be found mixed with hard, brittle galls, the former with first, the latter with second instars. By begin- ning of August galls were regularly found with third instars. Feeding larvae lie prone against the concave surface of the larval chamber. Collections of galls ——ALABAMA: Ashville (to- mentosa); Auburn (tomentosa); Greenville (tomen- tosa). ARKANSAS: 2 mi S DeQueen (tomentosa); Palestine (ovata). CONNECTICUT: Amston (ovata); Danbury (ovata); Hartland (ovata); Phoenixville (ovata). FLortpa: Chattahoochee (tomentosa); Gainesville (tomentosa); Lake City (tomentosa). Georcia: Lizella (tomentosa). Kentucky: My Old Kentucky Home St. Pk, Nelson Co. (ovata); More- head (tomentosa); Pennyrile St. Pk, Christian Co. RAYMOND J. GAGNE 69 (ovata); Winchester (laciniosa). Louisiana: Lake Bistineau St Pk, Webster Parish (tomentosa). Mary- LAND: Beltsville, USDA Agricultural Research Cen- ter (tomentosa); Calverton (tomentosa); Clarksburg (tomentosa); Ellicott City (tomentosa); Lusby (tomen- tosa); Ridgely (tomentosa). MASSACHUSETTS: Peter- sham (ovata). Mississippi: Natchez Trace Pkwy, Mile 193, Choctaw Co. (glabra, tomentosa); Nox- ubee Natl Wildlife Ref., Oktibbeha Co. (tomentosa); 7 mi E Starkville (tomentosa). NEw JERSEY: Colum- bia (ovata). NEw York: Ithaca (ovata). PENNSYLVA- NIA: Julian (glabra); Pavia (ovata); Schellsburg (ovata). SouTH CaroLina: Dillon (tomentosa). Georgetown (tomentosa). TENNESSEE: Dickson (to- mentosa); Greene Co., I 81, Mile 38.5 (tomentosa); Gatlinburg (tomentosa). VirGINIA: Huntley Mead- ows Pk, Fairfax Co. (fomentosa); Radford (tomen- tosa). WASHINGTON, DC: National Arboretum (to- mentosa). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (tomentosa); 6 mi NW Hedgesville (ovata). Specimens of C. spiniglobus examined.— ARKANSAS: Palestine, IX-10-2004 (larva). CONn- NeEcTicuT: Danbury, IX-6—2005 (larva). MARYLAND: Ridgely, IX-23-1999 (larvae) & IX-2-2000 (d, @, em. IV-2001). TENNESSEE: Dickson, IX-11—2004 (larvae). Caryomyia spinulosa Gagné, new species Description.—Gall (Figs. 92-93): Occasional, known from bitternut of the Apocarya section; found singly or in small groups, attached to leaf vein, usually on lower leaf surface, occasionally on upper leaf surface; 6.2-8.0 mm in length, re- cumbent, base bulbous, tapered beyond into elon- gate, usually abaxially bowed extension with pointed apex; green, yellow, to tan, covered with short, white, downy, crinkled hairs not obscuring surface; base of gall with central conical pedicel in shallow circular indentation; wall firm, brittle, uniformly thin, larval chamber glabrous with lon- gitudinal ridges following gall axis from leaf vein connection to apex. This gall is superficially simi- lar to those of C. procumbens and C. supina. It is known only from bitternut, while those of the other two occur on the Eucarya group, except for one known exception in C. supina. The gall of C. spinulosa has hair, as does that of C. procumbens, but the hair on the gall of C. spinulosa is downy and crinkly rather than stiff and straight and the gall base is much more bulbous. The gall of C. supina is hairless and not bulbous basally. Adult: Unknown. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.72.1 (8). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 295) with single tooth narrrowing slightly from base to apex, shallowly notched apically, shaft an- teriorly, narrowed beyond to widen again posteri- orly. Papillae without setae, their bases not raised. Lateral papillae 2-3 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdomen with extensive verrucae on dorsum, particularly at midlength, and with horizontal rows of spicules on venter anterior to sternal papillae and on posterior third. Terminal segment shorter than eighth. Second instar: Length 0.9 (1). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae not apprecia- bly raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material—Holotype: Third instar, from gall on C. cordiformis, Pavia, Pennsylvania, IX- 3-2003, R.J. Gagné, deposited in USNM. Etymology.—The name spinulosa is a Latin ad- jective and refers to the spinelike shape of the gall. Affinities.—Caryomyia spinulosa is one of the three Caryomyia spp. restricted to bitternut. This species and the other two, C. cilidolium and C. ansericollum, each appear most closely related to species found on the Eucarya group and not to one another. For further discussion of C. spinulosa, see under C. eumaris. Collections (all from Carya cordiformis).— ARKANSAS: Harrison; London. INp1ana: Plainville. MARYLAND: Ellicott City. MassacuuseTtTs: Barre. Missouri: Oregon Co., junc. Rts 11 & 142. PENN- SYLVANIA: Pavia. New York: Ithaca. VirGINIA: Rt. 81, mi. 13.9, Washington Co. WeEsT VIRGINIA: Falling Waters. Specimens of C. spinulosa examined.—ARKAN- sas: London, IX-10-2004 (larva). INDIANA: Plainville, IX-7—2004 (larva). MARYLAND: Ellicott City, IX-2-2000 (larvae). Massacuusetts: Barre, IX-1910, C.H. Clarke (larva). PENNSYLVANIA: Pavia, 70 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) IX-3-2003 (larva). New York: Ithaca, IX-3-2003 (larvae). West VirciniA: Falling Waters, IX- 10-2005 (larva), X-2005 (larva; pupa, excised from gall IV-28—2006). Caryomyia stellata Gagné, new species Description —Gall (Figs. 11a, 143): Occasional, on Eucarya hickories; single or clustered, on lower leaf surface betwéen veins; 5.0-9.0 mm in height, elongate-conical, base flared laterally, the flat- tened edge often stellate, sometimes merely irreg- ularly lobed, cone usually greatly attenuate and slightly curved to apex, furrowed; surface smooth, bare, green turning purple; base with shallow, circular, central excavation, leaf with no exfoliation surrounding connection; larval cham- ber ovoid, located at base of gall, surrounded by yellowish pellicle different in texture from soft, viscous, usually purple surrounding tissue that collapses with age, a bundle of fibers traversing between larval chamber and gall apex. Galls from Florida, Georgia, and Maryland are only tenta- tively placed with this species. They are the only ones found east of the Appalachians and are only shallowly lobed at the base and the lobes are rounded. West of the Appalachians the galls are more common, the bases more stellate, and the cones much longer (Fig. 143). Galls of this species have the same basic shape as those of C. sanguino- lenta except for a much higher crown and flared sides. Adult and pupa: Unknown. Larva: Third instar: Length 1.8—2.6 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 296) with long, nearly parallel sided, api- cally notched tooth and long shaft. Papillae with- out setae, their bases not raised. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdominal segments with rounded verrucae covering dorsum at mid- length and horizontal rows of spicules on venter anterior to sternal papillae and posterior to ven- tral papillae. Terminal segment shorter than eighth. Second instar: Length 0.8 mm (n=1). Col- lar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae not raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. tomentosa, Jackson, Tennessee, IX- 11-2004, R.J. Gagné, deposited in USNM. Etymology.—The name stellata, a Latin adjec- tive, refers to the stellate base of the typical form of the gall. Affinities —See under C. sanguinolenta, from which this species differs in the shape of the gall. Collections of galls —ALABAmMa: Oak Mt. St. Pk, Shelby Co. (ovata, pallida). Arkansas: Beulah (ovata); Magazine Mt., Logan Co. (ovata). COon- NECTICUT: Amston (ovata); Danbury (ovata); Litch- field (ovata); Plainfield (ovata); Westchester (ovata). Fioriwa: Avon Park (floridana); Gainesville (glabra, tomentosa). GEorGIA: Lake Park (glabra). INDIANA: Spencer Co. (ovata). KENtucky: Grayson Lake St. Pk, Carter Co. (tomentosa); Pennyrile St. Pk, Chris- tian Co. (ovata); Winchester (laciniosa). MAINE: North Berwick (ovata). MARYLAND: Gaithersburg (tomentosa); Silver Spring (glabra). MASSACHUSETTS: Great Barrington (ovata); Boston (tomentosa); Leominster (ovata); Petersham (ovata); Springfield (Carya sp., Stebbins 1910: Fig. 13). Mussissiprt: Natchez Trace Pkwy, Mile 123, Rankin Co. (pallida, tomentosa); Noxubee Natl Wildlife Ref., Oktibbeha Co. (ovata).; nr. Sessums, Oktibbeha Co. (ovata); 7 mi E Starkviile (pallida). Missouri: Wildwood (to- mentosa). NEw HAmpsuire: Barrington (ovata); Ex- eter (ovata); Seabrook (ovata). New York: Albany (Carya sp.); E. Schodack (Carya sp.). On10: Hocking Co. (ovata, Wells 1916: Figs. 11a); Huron Co. (Carya sp., Sears 1914: Fig. 14). (tomentosa); Yellow Springs (glabra). OKLAHOMA: Sallisaw (tomentosa). TEN- NESSEE: Jackson (tomentosa). WEST VIRGINIA: 6 mi NW Hedgesville (ovata). Specimens of C. stellata examined.—ARKANSAS: Beulah, IX-10—2004 (larvae). FLoripa: Gainesville, IX-21-2000 (larvae). INDIANA: Spencer Co., IX-20-— 1995 (larvae). KENTucky: Winchester, IX-22-1995 (larvae). MaAssAcuusetTtTs: Leominster, [X-21-1994 (larvae). New HAmpsuire: Exeter, [X-21-1994 (lar- vae). OKLAHOMA: Sallisaw, IX-9-2004 (larvae). TENNESSEE: Jackson, IX-11—2004 (larvae). West VIRGINIA: 6 mi NW Hedgesville, VIII-16—2000 (larvae). Caryomyia striolacrustum Gagné, new species Description.—Gall (Figs. 31, 84-85): Common, on Eucarya hickories; on lower leaf surface be- tween veins, usually clustered; height 2-3 mm, usually about as wide as long, basal half conical RAYMOND J. GAGNE 71 and deeply set into leaf swelling detectable on top surface of leaf as tan to black, conspicuous con- vexity, apical half of gall short-cylindrical with funnel-like, deep, narrow to wide concavity sur- rounded by correspondingly narrow to thick walls, center of funnel with short umbo; surface not sticky, yellow to brown, minutely longitudi- nally striated, the striations partly fibrous on dis- tal half; gall often surrounded and occasionally partially covered by conspicuous, brown, irregu- larly shaped leaf exfoliation; larval chamber in basal half of gall, wall firm, thin basally, thicker distally than laterally, glabrous, with longitudinal ridges. The shape of the gall most closely resem- bles that of C. melicrustum but the surface of the latter is mucilaginous and smooth. Adult: Head: Antenna with 12 flagellomeres. Male flagellomeres (as in Fig. 237) scarcely bin- odal, only slightly narrowed beyond basal circum- filum, circumfila loops short, subequal in length; female flagellomeres with 2 horizontal circumfila connected by 2 vertical strands. Labella (as in Fig. 242) separated from one another anteriorly, hemi- spherical, short-setose, short-setulose. Thorax: Wing length 1.4 mm in male (n=1); 2.1 mm in fe- male (n=1). Male abdomen: Third through sev- enth tergites each with posterior setae in single, sparse row continuous across width; eighth tergite without posterior setae. Genitalia (as in Figs. 247-248): hypoproct broadest near apex, concave posteromedially, margin setose, dorsal surface de- void of setulae on anterior 2/3; apex and venter of aedeagus furrowed. Female abdomen: Third through seventh tergites each with posterior setae continuous across width, in single row; eighth ter- gite with no posterior setae. Ninth segment pliable except for slender dorsal sclerite, setae sparse lat- erally, mostly as long as cerci, setae more numer- ous ventrally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.3-2.7 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 288) with pointed, almost equilateral an- terior tooth, occasionally with slight notch at apex, and anteriorly widened, elongate shaft. Papillae without setae, their bases not raised. Lat- eral papillae 2 on each side of spatula. Only 4 dor- sal papillae on first 7 abdominal segments. Ab- dominal segments with no verrucae on dorsum or sparse verrucae near bases of lateral pairs of dor- sal papillae and with spicules on venter anterior to sternal papillae. Terminal segment shorter than eighth. Second instar: Length 0.7-0.9 mm (n=4). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae slightly raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gallon C. glabra, National Arboretum, Washington, DC, IX-22-1988, R.J. Gagné, deposited in USNM. Etymology.—The name striolacrustum, a noun, is formed from the Latin striolatus for finely lined and crustum for tart, with reference to the stria- tions covering the tart-like gall. Affinities.—Galls of this species, short with a conical base set into an encircling leaf socket, are most like those of C. melicrustum except that they are finely striate and not resinous. For further re- marks, see under C. flaticrustum. Biological notes.—This gall appears to burst out of the leaf after it is mostly formed. In mid- June in central Maryland new galls could be found in various stages of emergence from the leaf epidermis. These contained first instars. By mid-July to mid-August, fully emerged galls con- tained second instars or third instars. Collections of galls——AtaBama: Ashville (to- mentosa); Auburn (ovata, tomentosa); Greenville (to- mentosa); Oak Mt. St. Pk, Shelby Co. (ovata, pall- ida). ARKANSAS: 3 mi S Harrison (tomentosa); Magazine Mt., Logan Co. (ovata, texana, tomen- tosa); 2 mi S DeQueen (texana, tomentosa); Rt.1 at Red River, Little River Co. (tomentosa). CONNECTI- cut: Amston (ovata); Meriden (glabra, ovata); New Haven (glabra); Phoenixville (ovata); Southbury (glabra, tomentosa); Westchester (ovata). FLORIDA: St. Lucie (floridana); Sebring (floridana); Vero Beach (floridana). Grorata: Carnesville (glabra); 8 km E Fort Valley (pallida); Forsyth (glabra, ovata); Lizella (ovata, tomentosa). INDIANA: Spencer Co. (ovata). Kentucky: Brownsville (tomentosa); Grayson Lake St. Pk, Carter Co. (tomentosa); Morehead (tomen- tosa); My Old Kentucky Home St. Pk, Nelson Co. (ovata); Pennyrile St. Pk, Christian Co. (glabra, 72 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) ovata); Winchester (lacinosa). MAINE: North Berwick (ovata).. MARYLAND: Beaver Dam (tomen- tosa); Beltsville, USDA Agricultural Research Cen- ter (glabra, tomentosa); Clarksburg (tomentosa); Elli- cott City (tomentosa); Gaithersburg (tomentosa); Lusby (glabra, tomentosa); Potomac (tomentosa); Ridgely (tomentosa); Silver Spring (glabra, tomen- tosa); Snow Hill (tomentosa); Spencerville (tomen- tosa); Wheaton (glabra, tomentosa). MASSACHU- sETTS: Great Barrington (ovata); Merrimac (ovata); Petersham (ovata); Wrentham (ovata). Mississippi: Natchez Trace Pkwy, Mile 193, Choctaw Co. (glabra, pallida, tomentosa); Noxubee Natl Wildlife Ref., Oktibbeha Co. (ovata, tomentosa); nr. Ses- sums, Oktibbeha Co. (ovata); 7 mi E Starkville (pal- lida). Missourr: Crowley Ridge Roadside Pk, Stoddard Co. (glabra, tomentosa); Holly Ridge State For., Stoddard Co. (glabra, tomentosa). NEw HAmMp- SHIRE: Barrington (ovata); Dover (ovata); Kingston (ovata). NEw Jersey: Columbia (ovata); Nyack (to- mentosa); Columbia (tomentosa). NEw York: Saratoga Springs (ovata). NORTH CAROLINA: Ala- mance (tomentosa); Alberta (glabra); Dortches (glabra, tomentosa); Fayetteville (tomentosa); Gasto- nia (tomentosa); Providence (glabra, tomentosa); Rowland (tomentosa); Wise (tomentosa). Oto: Hocking St. For., Hocking Co. (tomentosa); Hock- ing Co. (Carya sp., Wells 1916: Fig. 31). OKLAHOMA: Sallisaw (tomentosa). SOUTH CAROLINA: Dillon (to- mentosa); Richland Co. TENNESSEE: Gatlinburg (glabra, tomentosa); Greene Co., I 81, Mile 38.5 (ovata, tomentosa). Texas: Daingerfield St. Pk, Mor- ris Co. (tomentosa). VirGINIA: Blue Ridge Pkwy Mile 4.6, Warren Co. (ovata); Carson (tomentosa); Dinwiddie (glabra, tomentosa); Huntley Meadows Pk, Fairfax Co. (tomentosa); Ladysmith (tomentosa); Rt. 81, mi. 13.9, Washington Co. (ovata). WASHING- TON, DC: National Arboretum (glabra, tomentosa); Walter Reed Army Medical Center (tomentosa). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (to- mentosa); Cooper’s Rock St. Pk, Preston Co. (glabra); 6 mi. NW Hedgesville (tomentosa). Specimens of C. striolacrustum examined.—AL- ABAMA: Greenville, IX-20—2000 (larvae). CONNECTI- cut: Phoenixville, IX-20-1994 (larvae). FLORIDA: St. Lucie, XI-7 to 12-2002, K. Hibbard (larvae); Se- bring, XI-13—2002 (larvae); Vero Beach, IX-12-2002 (larvae). MARYLAND: Beltsville, USDA Agricul- tural Research Center, X-8—2002 (larvae; 3, 2, pu- pae, em IV-2003); Lusby, IX-16—2003 (larvae). NortH CAROLINA: Wise, IX-18—2000 (larvae). OKLAHOMA: Sallisaw, IX-9-2004 (larva). WASHING- TON, DC: National Arboretum, IX-22-1988 (larva). Caryomyia striolata Gagné, new species Description.—Gall (Figs. 36-38): Common, a spring gall, on both Eucarya and Apocarya hicko- ries; found singly or in groups on lower, rarely on upper, leaf surface, between or adjacent to veins; 3.5-5.0 mm in height, usually upright, sometimes leaning, base truncate, cylindrical, more or less fluted, tapering gradually to pointed apex; with sparse resin glands sometimes simulating hairs es- pecially on young galls, white to green, occasion- ally with some red, turning brown; base of gall shallowly concave; wall uniformly thin, larval chamber rough with weak, longitudinal ridges. This gall is similar in shape to that of C. albipilosa, but lacks a thick covering of long white hair. Adult (male only): Head: Antenna with 12 fla- gellomeres; each of the regular 12 male flagellom- eres (as in Fig. 235) scarcely binodal, basal flagel- lomeres only slightly narrowed beyond basal circumfilum, less constricted on more distal fla- gellomeres, loops of circumfila short, subequal in length. Thorax: Wing length 1.7 mm (n=1). Male abdomen: Third through sixth tergites each with posterior setae continuous across width, in mostly single row; seventh tergite with only a few pos- terolateral setae; eighth tergite more weakly scle- rotized, with anterior pair of trichoid sensilla the only vestiture. Genitalia (as in Figs. 249-250): hy- poproct broadest posteriorly, weakly concave posteromedially, posterior margin setose, dorsal surface extensively setulose; aedeagus narrowing from base to apex, rounded at apex. Pupa (as for Figs. 267-268): Antennal bases par- allel in ventral view, carinate anteroventrally, the mesal corner elongated, terminating anteriorly in short, obtuse projection. Face without incisions, frons and clypeus not protruding. Abdominal ter- gal spinules all similarly small. Larva: Third instar (Figs. 215-217, 269-271): Length 1.2—2.0 mm (n=10). Body cylindrical, ante- rior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 274) with 2 sepa- rate, equilateral teeth anteriorly, the intervening space with 1-4 denticles. Two, occasionally 3, lat- eral papillae each side of spatula. Six dorsal papil- lae on first 7 abdominal segments. Papillae with- out setae, bases not raised. Abdominal segments RAYMOND J. GAGNE 73 laterally with pointed spicules, dorsally lined, subverrucose, with sparse, pointed spicules mainly surrounding the 2 lateral pairs of dorsal papillae, and ventrally covered with spicules an- terior to ventral papillae. Terminal segment as long as eighth. Second instar: Length 0.7—1.1 mm (n=10). Collar segment without enlarged spicules laterally. Without spatula. Bases of dorsal and pleural papillae slightly raised, asetose. Six dorsal papillae on first 7 abdominal segments. First in- star (Fig. 35): Ovoid. Spicules present anteroven- trally on abdominal segments. Type material.—Holotype: Third instar (second specimen from right of four specimens on slide), from gall on C. tomentosa, Entomology Road, BARC-East, Beltsville, Maryland, V-16—2000, R.J. Gagné. Etymology—The name striolata is a Latin ad- jective meaning furrowed or fluted and refers to the gall’s texture. Affinities.—This species can be separated from C. albipilosa only by the lack of hair on the gall of this species and the thick coat of hair on the galls of C. albipilosa. See under the latter species for fur- ther discussion. Biological notes.—Galls of this species and C. albipilosa are the earliest Caryomyia galls to appear in spring and grow to full size within a few days. Young galls are at first mostly green and soon de- velop some red tinge. While the host leaves were still flaccid in early May, galls were found already fully formed, all with similar mature shape and size, with each containing a first, second, or third instar. In other years, galls found on May 16-19 contained second and third instars and by May 23 most galls were hanging loosely on the leaves or had already dehisced as indicated by abscission scars. On May 28, galls remaining on the leaves contained dead or parasitized larvae. These galls eventually turned brown and did not dehisce be- cause the gall ceased development upon the death of the larva and never completed an abscission layer. Any galls found on leaves after June 5 were brown and contained only dead or para- sitized larvae. Such galls could be found through September. Collections of galls —ALABAMA: Auburn (aquat- ica). CONNECTICUT: Meriden (tomentosa); Plainfield (ovata); Southbury (tomentosa). Georcta: Forsyth (glabra, ovata). Kentucky: Cave Run Lake, nr. Morehead. (Carya sp.). Mary.anp: Beltsville, USDA Agricultural Research Center (glabra, tomen- tosa); Calverton (glabra, tomentosa); Ellicott City (cordiformis, tomentosa); Kemp Mill (glabra); My- ersville (tomentosa); Spencerville (tomentosa); Wheaton (cordiformis, tomentosa). NEW JERSEY: Co- lumbia (tomentosa). WASHINGTON, DC: National Arboretum (glabra, tomentosa); Walter Reed Army Medical Center (tomentosa). WEST VIRGINIA: Falling Waters (cordiformis); Hedgesville (ovata). Specimens of C. striolata examined.—Con- NEcTICcUT: Meriden, VI-4-1994 (larvae). MARYLAND: Beltsville, USDA Agricultural Research Center, V- 22-1993, V-26 & VI-4-1997, V-14-1998, V-4 & 16-2000, & V-4—2004 (larvae); Calverton, V-4—2004 & V-15-2005 (larvae); Ellicott City, VI-8—2005 (lar- vae); Wheaton, V-30-1994 (larvae; 3, pupa, em. IV- 1995) & V-19-2000 (larvae). Caryomyia subulata Gagné, new species Description.—Gall (Figs. 27, 72-73): Occa- sional, on Eucarya hickories but not east of Ap- palachian Mts; usually in scattered groups, on lower leaf surface between veins; length 6.0-9.0 mm, elongate-cylindrical, erect, slightly bowed, tapered to pointed apex; shiny, brown, becoming black; base of gall deeply recessed, and set into socket formed from thick, scabrous, leaf growth with sharp apical edge; larval chamber occupying full length of gall, green, longitudinally striated, the wall firm, brittle, uniformly thin. This gall su- perficially resembles that of C. tubicola but the apex of the present gall is tapered rather than blunt and its base is deeply recessed rather than conical. Adult and pupa: Unknown. Larva: Third instar: Length 1.8—2.2 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 285) with 2 widely separated anterior teeth, the space between concave, shaft long, widest anteriorly. Papillae without setae, their bases not raised. Lateral papillae 2-3 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Abdominal segments completely cov- ered on dorsum with verrucae and on venter with spicules except at midlength around sternal papil- lae. Terminal segment as long as eighth. Second instar: Length 0.9-1.1 mm (n=3). Collar segment with patch of enlarged spicules on each side. Spat- ula with acutely triangular anterior tooth and 74 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae raised, papillae asetose. Six dorsal papillae on first 7 abdominal segments. Type material—Holotype: Third instar (speci- men farthest to right of three on slide), from gall on C. tomentosa, 6 mi W Hedgesville, Morgan Co., West Virginia, IX-19-1999, R.J. Gagné, deposited in USNM. Etymology.—The name subulata is a Latin ad- jective meaning awl-shaped with reference to the elongate-conical, tapered gall. Affinities.—The closest relative of this species is not apparent. The spatula with its two widely separated teeth but long, narrow shaft is unique for the genus. The elongate gall set into a deep socket only superficially resembles that of C. tubi- cola (Figs. 75-77). Notable differences are that the gall of C. subulata is spinelike, completely hollow within, and has a deeply excavated base, whereas the gall of C. tubicola is evenly cylindrical, has some woody tissue within, and has a conical base. Collections of galls—Arkansas: Piney (texana). Connecticut: Amston (ovata); Danbury (ovata); Hartland (ovata); Litchfield (ovata); Meriden (ovata). ILtiNoIs: Marshall (tomentosa). INDIANA: Spencer Co. (ovata). KENTUCKY: Grayson Lake St. Pk, Carter Co. (tomentosa); Winchester (laciniosa). Maine: North Berwick (ovata). MASSACHUSETTS: Pe- tersham (ovata). Mississippi: nr. Sessums (ovata). New Hampsuire: Exeter (ovata). NEw JERSEY: Co- lumbia (ovata). NEw York: Herkimer (ovata). OHIO: Hocking Co. (ovata, Wells 1916: Fig. 27); St. Clairsville (tomentosa); Summerford (tomentosa). Ontario: Dundas (ovata). PENNSYLVANIA: Kirby (ovata). TENNESSEE: Greene Co., I 81, Mile 38.5 (to- mentosa). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (ovata); 6 mi NW Hedgesville (ovata, tomentosa). Specimens of C. subulata examined.—Con- NEcTIcuT: Amston, [X-19-1994 (larva); Hartland, IX-7-2006 (larvae). INDIANA: Spencer Co., IX- 20-1995 (larvae). KEeNTUCKy: Winchester, IX- 22-1995 (larvae). MAtNe: North Berwick, IX- 21-1994 (larva). Onto: St. Clairsville, IX-6—-2004 (larvae). ONTARIO: Dundas, VIII-17-1994 (larvae). West VIRGINIA: 6 mi. NW Hedgesville, VIII- 20-1997, IX-19-1999, & VII-4—2000 (larvae). Caryomyia supina Gagné, new species Description.—Gall (Figs. 96-97): Occasional on Eucarya hickories and found once on bitternut of Apocarya section, occurs singly or in small groups; attached to side of vein on lower leaf surface; 6.0-8.0 mm in length, recumbent, base lobed across attachment to vein but not bulbous, gradu- ally narrowing along length to pointed apex; green, yellow, to tan, hairless; base of gall with central conical pedicel in shallow, circular inden- tation; wall firm, brittle, uniformly thin, larval chamber glabrous with longitudinal ridges fol- lowing gall axis from leaf vein connection to apex. See remarks about similar galls under C. spinulosa. Adult and pupa: Unknown. Larva: Third instar: Length 1.3-2.2 mm (n=7). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 292) with single tooth almost parallel- sided, deeply notched anteriorly, the tines slightly splayed, shaft wide anteriorly, narrowed beyond. Papillae without setae, their bases not raised. Lat- eral papillae 2-3 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Ab- domen with extensive verrucae on dorsum, par- ticularly at midlength, and with horizontal rows of spicules on venter anterior to sternal papillae and covering posterior third. Terminal segment shorter than eighth. Second instar: Length 0.8-1.2 mm (n=5). Collar segment with patch of enlarged spicules on each side. Spatula with acutely trian- gular anterior tooth and longer, wider, concave- sided shaft. Bases of dorsal and pleural papillae not appreciably raised, papillae asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. glabra, Wheaton, Maryland, IX-19-1993, R.J. Gagné, deposited in USNM. Etymology.—The name supina is a Latin adjec- tive that refers to the supine habit of the gall. Affinities —See under C. eumaris. Collections of galls:—ALABAMA: Auburn (glabra, ovata, tomentosa). ARKANSAS: Piney (tex- ana). CONNeEcTicuT: New Haven _ (glabra); Phoenixville (ovata); Southbury (glabra); Westch- ester (ovata). GEORGIA: Forsyth (ovata). KENTUCKY: Pennyrile St. Pk, Christian Co. (ovata). MARYLAND: Beltsville, USDA Agricultural Research Center (glabra, tomentosa); Brighton (tomentosa); Clarks- burg (tomentosa); Kemp Mill (glabra); Gaithersburg (glabra, tomentosa); Silver Spring (glabra); Spencerville (tomentosa); Wheaton (cordiformis, glabra). MassacuuseTts: Bolton (ovata). MIssourt: Mark Twain Natl For., Ripley Co. (tomentosa). Specimens of C. supina examined.—ARKANSAS: Piney, IX-10-2004 (larva). CONNecTicuT: New RAYMOND J. GAGNE 75 Haven, IX-19-1994 (larvae). MARYLAND: Beltsville, USDA Agricultural Research Center, IX-11-1994, VI-15-2000 & VII-1-2000 (larvae); Silver Spring, IX-26-1988 & IX-6—2003 (larvae); Spencerville, IX- 21-2003 (larvae); Wheaton, IX-19-1993 (larvae). Caryomyia thompsoni (Felt) Hormomyia thompsoni Felt 1908: 388; Felt 1918d: 47 (Caryomyia). Caryomyia consobrina Felt 1909: 292. New synonym. Caryomyia similis Felt 1909: 292. New synonym. Description.—Gall (Figs. 23, 112-115): One of the commonest Caryomyia galls, on Eucarya hicko- ries; usually in clusters, on lower leaf surface be- tween veins; 1.5-2.6 mm in height, depressed- spheroidal, often distended to one side, with deep dimple at center top; hair whitish gray, either sparse and not obscuring green to brown shiny surface or dense and obscuring gall surface; base of gall at center with deep, central, circular, shal- low depression, the leaf without exfoliation sur- rounding connection; wall of uniform thickness, firm, woody, the larval chamber invaded from both apex and base by vertical cylindrical intru- sions, the apical intrusion bearing crinkly hairs, both intrusions nearly abutting one another at center of larval chamber. Galls of this species vary considerably in hairiness, ranging from sparsely haired (Fig. 113) to so hairy as to obscure the gall surface (Fig. 115). Sparsely and densely haired ex- amples are commonly found intermixed and closely adjacent on a leaf (Fig. 112). The gall is unique for the vertical column through the center of the larval chamber made of the combined api- cal and basal intrusions of the gall wall. The only other Caryomyia gall with any intrusion is that of C. holotricha, but the intrusion in the latter is only of hair growing inward from the apex of the gall. Adult: Head: Antenna with 12 flagellomeres, in male a short, thirteenth flagellomere present in some specimens, but without circumfila; each of regular male flagellomeres (as in Fig. 239) bin- odal, internodes and necks both shorter than basal node; loops of circumfila short, subequal in length; female flagellomeres with 2 horizontal cir- cumfila connected by 2 vertical strands. Labella (as in Fig. 242) partially fused, blunt apically, short-setose, short-setulose. Palpus 4-segmented. Thorax: Wing length 3.5-4.2 mm in male (n=5); 3.0-3.5 mm in female (n=5). Male abdomen: Third through eighth tergites each with posterior setae in mostly double row, narrowing to single row mesally, sometimes discontinuous in middle. Genitalia (Figs. 252-253) cerci short, broadly rounded apically, setose dorsally and posteriorly; hypoproct broadest near apex, concave pos- teromesally, margin setose, setulose only posteri- orly on dorsum, completely ventrally; aedeagus broad, apex rounded, not furrowed; gonocoxite short, broad; gonostylus broad, with scattered se- tae, covered with setulae, the tooth nearly as wide as gonostylus. Female abdomen: Third through eighth segments with 2-3 rows of setae, tapering to 1 row and slightly discontinuous mesally. Ninth segment pliable except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ventrally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva : Third instar: Length 1.6-2.8 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 316) with simple, apically-pointed tooth, the shaft long, widest anteriorly and posteriorly, tapering from both ends to narrow midlength. Papillae without setae, their bases not raised above level of verrucae. Lateral papillae usually 3, occasionally 2 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Body completely covered by verrucae. Terminal seg- ment shorter than eighth. Second instar: Length 1.0-1.4 mm (n=10). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae raised but asetose. Six dorsal papillae on first 7 abdominal segments. Type material—Hormomyia thompsoni Felt: Lectotype here designated, photograph of gall in Thompson (1915: 90, Fig. 227) from Carya sp., lo- cality unspecified, presumably vicinity of Worces- ter, Massachusetts, M.T. Thompson. Paralectotype 3, 1116a, Thompson 227, ex Carya sp., locality un- specified, presumably Worcester, Massachusetts, M.T. Thompson, in Felt Collection, New York 76 THE GALL MIDGES (DIPTERA: CECIDOMYTIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) State Museum. The photo in Thompson (1915) is of the sparse-haired form of the gall. I choose to make the gall photo (actual specimen not seen) the lectotype because the species it belongs to is definite, whereas the single male described by Felt (now the paralectotype) cannot have come from this gall and cannot otherwise be placed with confidence. Antennal flagellomeres of this male are of the more foreshortened type found elsewhere in the genus and the aedeagus is fur- rowed. Felt’s (1918, 1921) subsequent figures and descriptions of C. thompsoni also were not of this species. In both papers Felt showed a photo of galls from New York City of specimens still in the Felt Collection and labelled with his unique num- ber “al908.” These galls are a mixture of C. holotricha and C. purpurea galls. Felt (1921) de- scribed the larva and adults of purported C. thompsoni with the note number “a1908.” The larva he described belongs to C. purpurea. The ac- companying adults in the numbered series are not C. purpurea and fit neither the paralectotype of C. thompsoni or adults that I have reared from galls similar to the lectotype. Caryomyia consobrina Felt: Lectotype, here desig- nated, ¢, al948, collected IX-8-1908, reared IV to V-1909 from depressed globular gall of Carya sp., Nassau, NY, deposited in the Felt Collection, New York State Museum. Paralectotypes, 3, 2, pupa, and larva, all with the same data as the lectotype. Galls typical of the dense-haired form of C. thomp- soni and labelled “a1948” are also in the Felt Col- lection. Caryomyia similis Felt: Lectotype, here desig- nated, larva, a1946, collected IX-16-1908 from gall of Carya sp., Nassau, NY, Felt Collection, New York State Museum. Paralectotypes: 3 5,5 2, same pertinent data as lectotype, but belonging to C. an- tennata on the basis of abdominal vestiture and the shape of the male flagellomeres. Two paralec- totype series of galls in the Felt collection, both la- belled “a1946” and “C. similis,” are of two kinds, one of the dense-haired form of C. thompsoni, from which the lectotype was apparently taken, the other of C. antennata, presumably the source of the paralectotype adults. Etymology of names in this taxon.—Felt named C. thompsoni in honor of Millett Taylor Thompson, a professor at Clark University in Worcester, Massachusetts, who had by the time of his death at 32 in 1907 partially prepared an illus- trated catalog on the gall makers of North Amer- ica. His book (Thompson 1915), edited by Felt, was posthumously published in accordance with the provisions of Thompson's father’s will. The name consobrina is Latin meaning cousin; Felt (1921) remarked that he found a general resem- blance between galls of this species and that of C. antennata. The name similis is a Latin adjective meaning similar; as Felt (1921) explained, he thought that the galls were easily mistaken for those of C. caryae. Affinities.—In adult, pupal, and larval charac- ters this species is similar to C. holotricha. The two species are the only congeners with an apical in- trusion of hairs into the larval chamber. The galls are otherwise distinct in that those of the present species have an additional intrusion into the base of the larval chamber and the apex of the gall is dimpled, not conical. Biological notes.—In central Maryland, very tiny to full-size galls were found on the same leaflet as early as May 23. Young galls were green- ish white turning light yellow with bright white hair. The larvae were all first instars. By June 9, galls appeared full-size and brittle but the larvae were still all first instars. In mid-June through mid-July galls were mostly hard, brittle, and brown to black and contained second instars, but during this time soft galls could still be found with first instars. By the beginning of August galls contained second or third instars and by mid- August more than half the larvae were third in- stars. Adults emerged in late March and early April from galls collected in New York, New Hampshire, Maryland, and Texas. Collections of galls ——ALaBaMa: Ashville (to- mentosa); Auburn (glabra, pallida, ovata); Greenville (tomentosa); Oak Mt. St. Pk, Shelby Co. (pallida). ARKANSAS: Beulah (tomentosa); Chester (texana); Magazine Mt., Logan Co. (ovata); Palestine (ovata). Connecticut: Amston (ovata); Danbury (ovata); Hartland (ovata); Harwinton (ovata); Litchfield (ovata); Meriden (glabra, ovata); New Haven (glabra); Oakille (ovata); Phoenixville (ovata); Plain- field (ovata); Southbury (glabra); Southington (ovata); Thomaston (ovata); Westchester (ovata). Fioripa: Avon Park (floridana); Gainesville (glabra, tomentosa). GEorGIA: Carnesville (glabra); Forsyth (glabra); 8 km E Fort Valley (pallida); Lizella (ovata); Macon (glabra). Ituinots: Highland (laciniosa). Kentucky: Brownsville (tomentosa). LOUISIANA: Lake Bistineau St. Pk, Webster Parish (texana). MAINE: Eliot (tomentosa). MARYLAND: Beaver Dam RAYMOND J. GAGNE 77 (tomentosa); Beltsville, USDA Agricultural Re- search Center (glabra, tomentosa); Calverton (to- mentosa); Clarksburg (tomentosa); College Park (glabra); Gaithersburg (tomentosa); Kemp Mill (glabra); Lusby (glabra, tomentosa); Piney Grove (ovata); Potomac (tomentosa); Ridgely (glabra); Snow Hill (tomentosa); Spencerville (tomentosa); Silver Spring (glabra, tomentosa); Wheaton (glabra, tomentosa). MassacuusETTs: Holyoke (ovata); Mer- rimac (ovata); vic. Worcester (Carya sp., Thompson 1915: 91, Fig. 227);. Wrentham (ovata). MIssIssipPt: Noxubee Natl Wildlife Ref., Oktibbeha Co. (ovata); nr. Sessums, Oktibbeha Co. (ovata). Missouri: Crowley Ridge Roadside Pk, Stoddard Co. (glabra, tomentosa); Holly Ridge State For., Stoddard Co. (glabra, tomentosa); Rolla (texana); St. Clair (tomen- tosa); Wildwood (tomentosa). NEw HAMPSHIRE: Dover (ovata); Kingston (ovata). New JERsEy: Co- lumbia (ovata, tomentosa). New York: Herkimer (ovata); Hyde Park (ovata); Ithaca (ovata); Nassau (Carya sp.); Ravena (ovata); Shokan (glabra, tomen- tosa). NORTH CAROLINA: Dortches (glabra, tomen- tosa); Providence (glabra, tomentosa); Wise (tomen- tosa). Onto: Hocking St. For., Hocking Co. (ovata); Hocking & Lake Cos. (ovata, Wells 1916: Fig. 23); St. Clairsville (tomentosa). OKLAHOMA: Sallisaw (to- mentosa). ONTARIO: Dundas (ovata). PENNSYLVANIA: Mooresville (tomentosa); Pavia (ovata); Schellsburg (glabra). SoutTH Carouina: Dillon (tomentosa); Georgetown (tomentosa). TENNESSEE: Crossville (pallida); Dandridge (tomentosa); Dickson (tomen- tosa); Greene Co., I 81, mi 38.5 (ovata); Lebanon (glabra); Mohawk (tomentosa). Vircinia: Dinwid- die (glabra, tomentosa); Warren Co., Blue Ridge Pkwy Mile 4.6 (ovata); Huntley Meadows Pk, Fair- fax Co. (tomentosa); Rt. 81, mi. 13.9, Washington Co. (ovata). WASHINGTON, DC: Rock Creek Park (ovata); National Arboretum (glabra); Walter Reed Army Medical Center (tomentosa). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (tomentosa); Ca- capon (ovata); Cooper’s Rock St. Pk, Preston Co. (glabra, ovata); 6 mi NW Hedgesville (glabra, ovata, tomentosa). Specimens of C. thompsoni examined.—Con- NEcTIcuT: Amston, IX-19-1994 (larvae); New Haven, IX-19-1994 (larvae); Phoenixville, IX- 20-1994 (larvae). FLoripa: Avon Park, IX-7 to 13-2002, K. Hibbard (larva); Gainesville, IX-21- 2000 (larvae; 2 6, 2,em. IV-2001). Georeia: 8 km E Fort Valley, IX-11-1993, J. & R. Payne (larvae); Lizella, X-12-1993, J. & R. Payne (larvae). KEN- Tucky: Brownsville, IX-21-1995 (larvae). MAry- LAND: Beltsville, USDA Agricultural Research Cen- ter, V-29-, VI-29-, and VII-1-1998 (larvae), IX- 23-2005 (larvae; 7 d, 23 2, pupae, em. IV-7 to 20-2006); Gaithersburg, V-23—2000 (larvae); Silver Spring, X-24-1999 (larvae); Wheaton, IX-19-1993 (larvae). New Hampsnire: Kingston, IX-21-1994 (larvae; 5, 2 2, pupae, em. IV-1995). New York: Herkimer, IX-4—2003 (larvae; pupa, emerged IV- 10-2004); Nassau, IX-16-1908, E.P. Felt (larvae, incl. lectotype of C. similis), IX-8-1908, E.P. Felt (3, ?, pupae, em. IV to V-1909, type series of C. conso- brina). Onto: Hocking St. For., Hocking Co., IX- 19-1995 (larvae). ONTARIO: Dundas, VII-17-1995 (larvae). Texas: New Waverly, X-6-2001 (2, em. TV-2002). WASHINGTON, DC: National Arboretum, VI-11 & X-13-1997 (larvae), IX-30-1999 (larvae; 2 3, 2, pupae, em. IV-2000). WEST VIRGINIA: 6 mi NW Hedgesville, IX-12 & 17-1999 (larvae). Caryomyjia tuberculata Gagné, new species Description —Gall (Figs. 118-119): Common, on Eucarya hickories; often in groups, on lower leaf surface between veins; 3.5—5.1 mm in height, depressed-spherical to spheroidal, occasionally with slightly pointed apex; surface hairless, bumpy, sticky, green, yellow, or brown to almost purple; base with large, central, deep, circular ex- cavation, leaf without exfoliation surrounding connection; wall uniformly thick, woody, larval chamber ovoid, floccose, white, green, to pur- plish. The gall is most like that of C. purpurea be- cause of its shape, the thick, woody wall, and the leaf connection but, unlike that of C. purpurea, is bumpy and hairless. Adult: Head: Antenna with 12 flagellomeres, usually a short 13th flagellomere without circum- fila present; each of regular 12 male flagellomeres (as in Fig. 239) binodal with definite internode and neck, the circumfila on each with short loops, their bases widely separated; female flagellom- eres with 2 horizontal circumfila connected by 2 vertical strands, occasionally with an extra spur at base of vertical strand. Labella (as in Fig. 242) sep- arated from one another anteriorly, hemispheri- cal, short-setose, short-setulose. Palpus 4-seg- mented. Thorax: Wing length 3.6—-4.5 mm in male (n=5), 3.4-4.8 mm in female (n=5). Male ab- domen: Third through eighth tergites each with posterior setae separated into two lateral groups, 2-3 rows deep laterally, narrowing to one mesally. Genitalia: hypoproct somewhat widened posteri- 78 THE GALL MIDGES (DIPTERA: CECIDOMYTIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) orly, concave posteromesally, margin setose ex- cept in concavity, setulose on dorsum only on pos- terior fourth; apex of aedeagus convex. Female abdomen: Third through eighth tergites each with posterior setae separated into two lateral groups, 2-3 rows deep laterally, narrowing to one mesad; eighth tergite with posterior setae in 1-3 rows lat- erally and not as long or strong as on preceding tergite. Pupa (as in Frgs. 201-202): Antennal bases oblique in ventral view, converging and narrow- ing anteriorly, their rounded apices closely adja- cent. Face with anterolateral pair of horizontal in- cisions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.9-3.5 mm (n=10). Body ovoid-cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spatula (Fig. 326) with tooth tapering from wide base to pointed apex and with elongate shaft. Papillae without setae, their bases not raised. Lat- eral papillae 3 on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Abdomi- nal segments on dorsum extensively covered with verrucae at midlength and on venter with hori- zontal rows of spicules anteriorly and posteriorly. Terminal segment shorter than eighth. Second in- star: Length 1.7 mm (n=1). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae slightly raised, asetose. Six dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. tomentosa, vic. Ridgely, Maryland, IX- 23-1999, R.J. Gagné, deposited in USNM. Etymology.—The name tuberculata is an adjec- tive from the Latin meaning bumpy and refers to the conspicuous bumps covering the gall surface of this species. Affinities.—The closest relative among species with long male flagellomeres and an apically con- vex aedeagus is not apparent. Unlike other such species, such as C. purpurea and C. holotricha with spheroidal galls that occur on the leaf lamina and have a deep, circular, basal recess, C. tuberculata has pupal antennal bases that are closely juxta- posed apically instead of being distinctly sepa- rate. Biological notes.—Tiny, yellow-green, soft galls containing first instars were first noticed in central Maryland on June 10. On June 22 new and full grown galls were found on the same leaf, with first and second instars, respectively. Galls found from late June through mid-August contained second instars, after which third instars were reg- ularly found. The full-grown larva fills the larval chamber. Adults emerge through a hole in the side of the gall. Collections of galls—ALaBAMA: Auburn (fo- mentosa); Greenville (tomentosa). ARKANSAS: Chester (texana); Magazine Mt., Logan Co. (tomen- tosa); Piney (texana). CONNECTICUT: Amston (ovata); Meriden (glabra, ovata); New Haven (glabra); Oakville (ovata); Phoenixville (ovata); Plainfield (ovata); Southbury (glabra, ovata, tomentosa); Southington (ovata). GrorGIA: Forsyth (ovata); Lizella (tomentosa). INDIANA: Spencer Co. (ovata). Kentucky: Grayson Lake St. Pk, Carter Co. (tomen- tosa). LOUISIANA: Lake Bistineau St. Pk, Webster Parish (tomentosa). MARYLAND: Beaver Dam (to- mentosa); Beltsville, USDA Agricultural Research Center (glabra, tomentosa); Brighton (tomentosa); Clarksburg (tomentosa); College Park (glabra); Gaithersburg (tomentosa); Lusby (glabra, tomen- tosa); Ridgely (tomentosa); Gaithersburg (tomentosa); Silver Spring (tomentosa); Snow Hill (tomentosa); Spencerville (tomentosa). MASSACHU- sETTS: Leominster (ovata); vic. Worcester (Carya sp., Thompson 1915: 91, Fig. 244); Wrentham (ovata). Mississippi: Noxubee Natl Wildlife Ref., Ok- tibbeha Co. (ovata). Missouri: Crowley Ridge Roadside Pk, Stoddard Co. (glabra); Holly Ridge State For., Stoddard Co. (tomentosa); St. Clair (to- mentosa). New HAampsHire: Kingston (ovata). NortTH Caro.ina: Alamance (tomentosa); Dortches (glabra); Fayetteville (tomentosa); Providence (to- mentosa). OKLAHOMA: Sallisaw (tomentosa). PENN- SYLVANIA: Schellsburg (ovata). SOUTH CAROLINA: Dillon (tomentosa). TENNESSEE: Crossville (pallida, tomentosa); Dandridge (tomentosa); Mohawk (to- mentosa). VIRGINIA: Carson (tomentosa); Dinwiddie (glabra, tomentosa); Huntley Meadows Pk, Fairfax Co. (tomentosa). (tomentosa); Ladysmith (tomen- tosa). WASHINGTON, DC: National Arboretum (glabra, tomentosa); Walter Reed Army Medical Center (tomentosa). WEST ViIRGINIA: Beech Fork St. Pk, Wayne Co. (tomentosa); 6 mi NW Hedgesville (tomentosa). Specimens of C. tuberculata examined.—Ar- KANSAS: Chester, IX-9-2004 (larvae); Piney, X- 10-2004 (larvae). CONNeEcTICUT: Amston, IX- 19-1994 (larva). Georcia: 8 km E Fort Valley, X-11-1993, J. & R. Payne (larvae). NEw HAMPSHIRE: RAYMOND J. GAGNE 79 Kingston, IX-21-1994 (larvae; 5 3,5 2, pupae, em. TV-1995). MARYLAND: Beltsville, USDA Agricul- tural Research Center, X-13-1999 (larvae; 5, pu- pae, em. IV-2000) & IX-11-1994 (larvae); Gaithers- burg, X-4-1994 (larvae); Lusby, IX-16—2003 (¢, pupa, em. IV-2004), IX-14-2005 (larvae; 7 °, pu- pae, em. IV-2 to 10-2006); Ridgely, IX-23-2000 (larvae); Silver Spring, IX-22-1993 (larvae). Mas- SACHUSETTS: Wrentham, IX-20-1994 (larva). WASH- INGTON, DC: National Arboretum, IX-22-1988 & X-13-1997 (larvae), IX-30-1999 (larvae; 5, 2, em. TV-2000); Walter Reed Army Medical Center, X- 7-1993 (d, 5 2, pupae, em. IV-23 to 24-1994). WEsT VIRGINIA: 6 mi NW Hedgesville, IX-19-1999 (larvae). Caryomyjia tuberidolium Gagné, new species Description.—Gall (Figs. 46-48): Common, on Eucarya hickories; often in groups, usually on lower, rarely on upper leaf surface, between veins; 3.2-4.0 mm in height, spheroidal, slightly longer than wide, base broadly rounded to truncate in profile, apex with small central nipple sur- rounded by extensive areola that with pressure easily separates from gall; surface covered with large bumps and resin glands, the sticky exudate in young galls sometimes extruded in hairlike strands, hairless, yellow, tan, or brown, areola paler than surrounding surface; base with wide, deep, circular indentation; wall firm, brittle, al- most uniformly thin, larval chamber glabrous, with longitudinal ridges. For notes on galls of re- lated species, see under C. cilidolium. Adult (female only): Head: Antenna with 12 flagellomeres with anastomozing circumfila on venter (as in Fig. 241). Labella united, convex api- cally, asetose, covered with setulae. Thorax: Wing length 3.1-3.3 mm in female (n=2). Abdomen: Third through seventh tergites each with poste- rior setae in 3 rows laterally, thinning to 2, then one, with slight medial interruption in setae; eighth tergite more weakly sclerotized than pre- ceding tergite, with weak posterior setae numer- ous laterally, continuing mesally in single, sparse row. Ninth segment (as in Fig. 261) pliable except for slender dorsal sclerite, with numerous apico- lateral and ventral setae, none longer than the cerci, the two groups separated laterally. Cerci elongate-ovoid in lateral view, pointed apically, fused dorsally for about 1/2 length. Pupa (Figs. 193-194): Antennal bases elongate, in ventral view parallel, closely adjacent, rounded anteriorly, the apices dorsoventrally flattened. Face with longitudinal impression running along length, a slight anteromesal convexity posterior to each antennal base, and without anterolateral pair of horizontal incisions. Frons and clypeus slightly convex. Abdominal tergal spinules all similarly small. Larva: Third instar: Length 2.6-4.8 mm (n=10). Body broadly rounded but knobby anteriorly, spindleform posteriorly. Head directed ventrally. Spatula (Fig. 280) with long, pointed anterior tooth and elongate shaft. Papillae prominently raised above surrounding surface, without setae. Lateral papillae numbering 2 on each side of spat- ula. Six dorsal papillae on first 7 abdominal seg- ments. Body entirely covered with verrucae. Ter- minal segment as long as eighth, rounded posteriorly. Second instar: Length 1.0-1.5 mm (n=2). Collar segment with patch of enlarged spicules on each side. Spatula with acutely trian- gular anterior tooth and longer, wider, concave- sided shaft. Six dorsal papillae on first 7 abdomi- nal segments. Bases of dorsal and pleural papillae elongate, asetose. Type material.—Holotype: Third instar, from gall on C. tomentosa, Gaithersburg, Maryland, VII- 20-2000, R.J. Gagné, deposited in USNM. Etymology.—The name tuberidolium, a noun, combines the Latin words tuberosus (bumpy) and dolium (jar), with reference to the bumpy, barrel- like gall. Affinities.—See under C. cilidolium. Biological notes.— In central Maryland, tiny to full-size galls were first noticed on May 24, the galls all with first instars. Second instars were present in some galls by June 25, but most galls still contained first instars through July 8. Third instars began appearing by mid-July and second instars could still be found through Aug. 19. Sec- ond and third instars are active when disturbed. Collections of galls—AtaBama: Auburn (gla- bra, ovata); Greenville (tomentosa); Oak Mt. St. Pk, Shelby Co. (pallida). ARKANSAS: Beulah (tomen- tosa); Osage (texana); Winslow (ovata). CONNECTI- cut: Amston (ovata); Danbury (ovata); Litchfield (ovata); Meriden (glabra, ovata); Phoenixville (ovata); Southbury (glabra); Southington (ovata); Westchester (ovata). Georcia: Lake Park (glabra); Forsyth (glabra); 8 km E Fort Valley (pallida). Ken- TuCKY: Brownsville (tomentosa); Grayson Lake St. Pk (glabra); Morehead (tomentosa); My Old Ken- tucky Home St. Pk, Nelson Co. (ovata). ILLINOIS: Marshall (tomentosa). MARYLAND: Beltsville, USDA 80 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Agricultural Research Center (tomentosa); Brighton (tomentosa); Calverton (tomentosa); Clarksburg (glabra, tomentosa); Gaithersburg (to- mentosa); Lusby (tomentosa); St. Leonard (tomen- tosa); Silver Spring (glabra); Wheaton (tomentosa); Kemp Mill (glabra). Massacuusetts: Bolton (ovata); Petersham (ovata). Mississirri: nr. Ses- sums, Oktibbeha Co. (ovata). NEw HAMPSHIRE: Seabrook (ovata). NEw Jersey: Columbia (tomen- tosa). New York: Herkimer (ovata). NORTH CAR- OLINA: Fayetteville (tomentosa); Providence (glabra, tomentosa); Wise (tomentosa). OKLAHOMA: Sallisaw (tomentosa). PENNSYLVANIA: Pavia (ovata). SOUTH CAROLINA: Georgetown (tomentosa). TENNESSEE: Dandridge (tomentosa); Dickson (tomentosa); Jack- son (tomentosa); Mohawk (tomentosa). VIRGINIA: Warren Co., Blue Ridge Pkwy Mile 4.6 (ovata); Dinwiddie (tomentosa); Huntley Meadows Pk, Fairfax Co. (tomentosa); Rt. 81, Mile 13.9, Washing- ton Co. (ovata). WASHINGTON, DC: National Ar- boretum (glabra, tomentosa); Walter Reed Army Medical Center (tomentosa). WEST VIRGINIA: Ca- capon (ovata); 6 mi NW Hedgesville (ovata). Specimens of C. tuberidolium examined.— ALABAMA: Auburn, IX-19—2000 (larvae). ARKANSAS: Beulah, IX-10-2004 (larva); Osage, IX-9-2004 (larva). MARYLAND: Beltsville, USDA Agricultural Research Center, IX-26-1993 & IX-11-1994 (larvae), IX-30-2003 (larvae; 2 2, pupae, em. IV-10—2004), IX-28-2005 (larva; pupal exuviae, 3 2, em. IV- 12-2006); Gaithersburg, VII-20-2000 (larva); Wheaton, V-24—1993 (larvae). WASHINGTON, DC: National Arboretum, IX-22-1988 (larvae). Caryomyia tubicola (Osten Sacken) Cecidomyia tubicola Osten Sacken 1862: 192; Felt 1909:293 (Caryomyia). Description.—Gall (Figs. 26, 74-77).—Com- mon and widespread (Map 1) on Eucarya hicko- ries and occasional on C. cordiformis of Apocarya section; found singly or in groups; 3-7 mm in height, on lower leaf, usually between veins, occa- sionally adjacent to a vein; elongate-cylindrical, usually erect, knurled at apex; shiny, with minute longitudinal ridges; green, yellow, brown, or black; base of tube abruptly conical, deeply set into, and tightly sheathed by thick, scabrous, leaf growth with sharp apical edge; larval chamber variable in length, occupying basal half to nearly entire length of tube, lateral walls thin, hard, firm, with longitudinal ridges; distal part of cylinder not hollow, traversed from larval chamber to apex by thin fibrous bundle. This gall superficially re- sembles that of C. swbulata but the shape of the base is different, that of C. subulata being deeply recessed rather than conical. The long shaft of both galls set them apart from other Caryomyia galls, but conical bases and leaf sheaths are also seen in galls of C. melicrustum and C. striolacrus- tum. Adult: Head: Antenna with 12 flagellomeres, occasionally a short 13th without circumfila in male. Each of the regular 12 male flagellomeres (Fig. 237) scarcely binodal, only slightly narrowed beyond basal circumfilum, circumfila loops short, subequal in length; female flagellomeres (Fig. 238) with 2 horizontal circumfila connected by 2 verti- cal strands. Labella (as in Fig. 242) separated from one another anteriorly, hemispherical, short-se- tose, short-setulose. Thorax: Wing (Fig. 245), length 1.3-2.0 mm in male (n=10); 1.8-2.6 mm in female (n=10). Male abdomen: Third through seventh tergites each with posterior setae continu- ous across width in mostly single row; eighth ter- gite weakly sclerotized, with or without a few posterior setae. Genitalia (Figs. 247-248): hy- poproct broadest posteriorly, divided into 2 rounded lobes, each with several apical setae, dor- sal surface devoid of setulae on anterior half; apex and venter of aedeagus furrowed. Female ab- domen (Figs. 258-259): Third through seventh ter- gites each with posterior setae continuous across width in mostly single row; eighth tergite more weakly sclerotized than preceding tergite, with posterior row of setae variable, from setae in con- tinuous single row or only 1 to a few laterally, the setae not as long as on seventh tergite. Ninth seg- ment pliable except for slender dorsal sclerite, se- tae sparse laterally, mostly as long as cerci, setae more numerous ventrally. Cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (Figs. 187-192): Antennal bases oblique in ventral view, converging anteriorly, each termi- nating in short, obtuse point. Face with anterolat- eral pair of horizontal incisions, their posterior lip slightly protruding. Frons and clypeus moder- ately protruded ventrally. Abdominal terga with several anterior rows of enlarged spinules. Larva: Third instar: Length 1.2-1.8 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 322) with acutely triangular anterior RAYMOND J. GAGNE 81 tooth and elongate, linear shaft, usually broad- ened anteriorly. Papillae without setae, their bases not raised. Lateral papillae usually 1, occasionally 2, on each side of spatula. Only 4 dorsal papillae present on first 7 abdominal segments. Abdomi- nal segments with verrucae present dorsally at midlength, occasionally more extensively but not prominent or only sparsely around lateral pairs of dorsal papillae, and with sharp spicules ventrally anterior to sternal papillae and at posterior end of segment. Terminal segment shorter than eighth. Second instar (Figs. 272-273): Length 1.7-1.2 mm (n=10). Collar segment with patch of enlarged spicules on each side. Spatula with acutely trian- gular anterior tooth and elliptical shaft. Bases of dorsal and pleural papillae slightly raised, dorsal papillae with short setae, remainder asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Syntypes: Larva(e) and galls from Carya sp., vic. Washington, DC, Osten Sacken. The larvae are presumed lost but several galls remain on each of three leaflets that are de- posited in the USNM. Etymology.—The name tubicola is a Latin noun meaning “tube dweller.” Affinities.—The closest relative of this species is not apparent, but galls of C. melicrustum and C. striolacrustum also have conical bases that are set into a socket made by the surrounding leaf. An- tennal bases of C. tubicola pupae are, however, un- like those of the other two species. Larvae of C. tubicola usually have the lateral papillae reduced to only one per side, the greatest known reduction in Cecidomyiidae. Biological notes.—This may be the most com- mon species of Caryomyia. While tubes of mature galls are usually about four times as long as wide, many are much shorter. Larval success does not appear to depend on gall length. In the Washing- ton, DC area, adults emerge beginning the first week of April from galls collected the previous autumn. Pupae emerge from a weak area at the base of the tube that was scraped away by the larva in the previous season (Fig. 77). New galls are visible by mid-May as a thickening visible on both surfaces of the leaflets, but they are still green and the tube has not yet begun to elongate (Fig. 74). Additional, apparently new galls can be found through early August. First instars are translucent and elliptical and lie flat against what will become the base of the tube. As the gall grows, the exterior is quite hard but the base, hid- den in the enveloping leaf collar, is initially soft until it grows to enclose the first instar. The larva then becomes more evenly cylindrical and mi- grates up the lengthening tube. By late May some tubes have begun to emerge from the basal boot. By mid-June some of the tubes have achieved their full length and the larvae have mostly molted to the more cylindrical second instar. By mid-July some galls contain third instars. The tubes mostly dehisce from the leaves before leaf drop. This species is one of the few that occurs on both the Eucarya and Apocarya sections, although to date I have found it only on C. cordiformis of the Apocarya group. Galls on C. cordiformis are never so common, dense, or robust as those on the Eu- carya group, but more weak-walled galls may be a function of the thinner leaves of C. cordiformis. On C. cordiformis the galls are usually found along a vein. Collections of galls——AtaBama: Ashville (to- mentosa); Auburn (ovata, pallida, tomentosa); Greenville (tomentosa); Oak Mt. St. Pk, Shelby Co. (ovata, pallida). Arkansas: Beulah (tomentosa); Chester (texana); 2 mi S DeQueen (texana, tomen- tosa); 3 mi S Harrison (tomentosa); Osage (texana); Rt.1 at Red River, Little River Co. (tomentosa); Magazine Mt., Logan Co. (ovata, texana, tomen- tosa); Palestine (ovata); Piney (texana); Winslow (ovata). CONNECTICUT: Amston (ovata); Danbury (ovata); Harwinton (ovata); Litchfield (ovata); Meri- den (cordiformis, glabra, ovata); New Haven (glabra); Oakville (glabra, ovata); Southbury (cordi- formis, glabra, tomentosa); Phoenixville (ovata); Plainfield (ovata); Southington (ovata); Thomaston (ovata); Westchester (ovata). FLormpA: Chatta- hoochee (glabra, tomentosa); Gainesville (glabra, to- mentosa); Lake City (tomentosa); Ponce de Leon (glabra). Grorcia: Carnesville (glabra); Forsyth (glabra, ovata); 8 km E Fort Valley Lake Park (glabra, pallida); Lizella (ovata, tomentosa). ILLINOIS: Dexter (laciniosa, tomentosa); Highland (laciniosa); Marshall (tomentosa). INDIANA: New Castle (glabra); Plainville (cordiformis, tomentosa); Rich- mond (tomentosa); Spencer Co. (ovata). KENTUCKY: Brownsville (tomentosa); Grayson Lake St. Pk, Carter Co. (tomentosa); Morehead (tomentosa); My Old Kentucky Home St. Pk, Nelson Co. (ovata); Pennyrile St. Pk, Christian Co. (glabra, ovata); Win- chester (laciniosa). LOUISIANA: Lake Bistineau St. Pk, Webster Parish (texana). MAINE: Eliot (tomen- tosa); North Berwick (ovata). MARYLAND: Beaver Dam (tomentosa); Beltsville, USDA Agricultural 82 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Research Center (glabra, tomentosa); Bethesda (cordiformis); Brighton (tomentosa); Cabin John (cordiformis); Calverton (tomentosa); Clarksburg (tomentosa); College Park (glabra); Ellicott City (to- mentosa); Gaithersburg (tomentosa); Kemp Mill (glabra); Kensington (cordiformis); Long Beach (cordiformis); Lusby (glabra, tomentosa); Potomac (cordiformis, tomentosa); Ridgely (cordiformis, glabra, tomentosa); St. Leonard (tomentosa); Silver Spring (glabra, tontentosa); Snow Hill (tomentosa); Spencerville (tomentosa); Wheaton (cordiformis, glabra, tomentosa). MASSACHUSETTS: Great Barring- ton (ovata); Holyoke (ovata); Merrimac (ovata); Pe- tersham (ovata); Wrentham (ovata). Mississippi: Natchez Trace Pkwy, Mile 193, Choctaw Co. (pall- ida, tomentosa); Noxubee Natl Wildlife Ref., Ok- tibbeha Co. (ovata, tomentosa); nr. Sessums, Ok- tibbeha Co. (ovata); 7 mi E Starkville (pallida); Vicksburg (cordiformis). Missourt: Conway (ovata, texana); Crowley Ridge Roadside Pk, Stoddard Co. (glabra, tomentosa); Holly Ridge State For., Stoddard Co. (glabra, tomentosa); Oregon Co., junc. Rts. 11 & 142 (cordiformis); Rolla (texana); St. Clair (texana, tomentosa); Wildwood (tomentosa). NEw HampsHireE: Barrington (ovata); Dover (ovata); Kingston (ovata). New Jersey: Columbia (cordi- formis, ovata, tomentosa). New York: Albany (Carya sp.); East Schodack (Carya sp.) Herkimer (ovata); Hyde Park (ovata); Ithaca (cordiformis, ovata); Nas- sau (Carya sp.); New York (Carya sp., Beuten- miller 1904: 27, fig. 58); Nyack (tomentosa); Ravena (ovata); Saratoga Springs (ovata); Shokan (glabra). NortH Caro.ina: Alamance (tomentosa); Alberta (glabra); Dortches (tomentosa); Fayetteville (tomentosa); Gastonia (tomentosa); Providence (glabra) Rowland (tomentosa); Wise (tomentosa). Ouro: Hocking St. For., Hocking Co. (tomentosa); Hocking Co. (Carya spp., Wells 1916: Fig. 26); Huron Co. (Carya sp., Sears 1914: Fig. 19); St. Clairsville (tomentosa); Summerford (tomentosa); Yellow Springs (glabra). OKLAHOMA: Sallisaw (to- mentosa). ONTARIO: Dundas (ovata). PENNSYLVANIA: Kirby (ovata); Julian (glabra); Millsboro (glabra); Mooresville (tomentosa); Pavia (ovata); Schellsburg (glabra, ovata). SouTH CAROLINA: Dillon (tomen- tosa); Georgetown (tomentosa); Richland Co. (Carya sp.). TENNESSEE: Crossville (pallida, tomen- tosa); Dandridge (tomentosa); Dickson (tomentosa); Greene Co., I 81, Mile 38.5 (ovata); Jackson (tomen- tosa); Lebanon (glabra); Mohawk (tomentosa). Texas: Daingerfield St. Pk, Morris Co. (tomentosa); New Waverly (tomentosa). Vircinia: Dinwiddie (glabra, tomentosa); Huntley Meadows Pk, Fairfax Co. (tomentosa); Ladysmith (tomentosa); Leesburg (tomentosa); Skippers (tomentosa); Warren Co., Blue Ridge Pkwy Mile 4.6 (ovata); Rt. 81, mi. 13.9, Washington Co. (ovata). WASHINGTON, DC: Na- tional Arboretum (glabra, tomentosa); Walter Reed Army Medical Center (tomentosa). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co. (ovata, tomentosa); Cacapon (ovata); Cooper’s Rock St. Pk, Preston Co. (glabra); Falling Waters (cordiformis); 6 mi NW Hedgesville (glabra, ovata, tomentosa). Specimens of C. tubicola examined.—ConNneECTI- cut: Meriden, IX-30-1997, (larvae), IX-18-1998 (larvae; 2 5, em. V-1999); New Haven, IX-19-1994 (larvae); Phoenixville, IX-26-1994 (larva); Plain- field, VIII-27-1988 (larvae). Georcia: Fort Valley, IX-11-1993, J. & R. Payne (larva). KENTUCKY: Brownsville, IX-21-1995 (larvae). MAINE: North Berwick, [IX-21—1994 (larva). MARYLAND: Beltsville, USDA Agricultural Research Center, IV-10-1997, V-26-1997, V-14-1998, VI-17-1998, VII-1-1998, VII- 15-1998 & VIII-5-1998 (larvae), & IX-5-1991 (lar- vae;4 5,3 2,pupae, em. IV-25-1992); Bethesda, IX- 15-1997 (larvae); Brighton, X-10-1993 (4 5,5 9, pupae, em. IV-21 to 28-1994); Gaithersburg, V- 23-2000 (larvae); Kensington, VII-27—2003 (lar- vae); Ridgely, X-2-2000 (2, pupa, em. IV-2001), IX- 11-2003 (2 6, 3 2, pupae, em. IV-10-2004), X-6-2005 (pupae, 2 ¢,6 2, em. IV-10 to 21-2006); Silver Spring, IX-17-1993 (larvae); Wheaton, IX- 28-1991, VI-18-1993 & IX-19-1993 (larvae). Mas- SACHUSETTS: vic. Worcester, M.T. Thompson (larva, 2 2). New York: East Schodack, E.P. Felt (larvae; 3, ?,em. V-19-30-1907); Nassau, E.P. Felt (larvae; 3, ?,em. IV 21-1908); Saratoga Springs, IX-25-1994 (larva). ONTARIO: Dundas, VIII-17-1995 (larvae). Texas: New Waverly, X-6—2001 (3d, 2 2, em. IV- 2002). WesT VirGINIA: 6 mi NW Hedgesville, IX- 29-2002 (larvae). WASHINGTON, DC: National Ar- boretum, VIII-16-1988, IX-16-1988, IX-13-1991, IX-26-1991, V-28-1993, IX-22-1993, & IV-11-1997 (larvae), IX-24-1980 (2, em. IV-1981), IX-24-1988 (3 3,2 2, pupae, em. IV-1989), X-8-1991 (3,3 @, em. IV-25-1992), IX-30-1999 (2 3,3 2, pupae, em. IV-2000), X-2001 (4 6, 1 2, em. IV-2002), & X- 29-2002 (larvae; 8 3,8 2, pupae, em. IV-2003). Caryomyia tumida Gagné, new species Description—Gall (Figs. 104-105): Rare, known from only 2 collections, on Eucarya hicko- ries; on lower leaf surface, between veins; 3 mm in RAYMOND J. GAGNE 83 height, depressed-obconic, with small umbo at apex, made up of basal ovoid larval chamber and apical, much broader, false chamber; tan, rubbery, covered with short, decumbent hairs obscuring gall surface, not sticky; base of gall tapered to con- ical pedicel, leaf with slight exfoliation around gall connection forming short, tubular socket and on reverse side showing discolored convexity; wall thin, basal larval chamber separated from larger, apical false chamber by thin wall with umbo at center, both chambers longitudinally stri- ate. The gall is shaped as for that of C. inflata but is covered with decumbent hairs in contrast to the completely hairless, glabrous gall of C. inflata. Adult and pupa: Unknown. Larva: Third instar: Length 1.7—2.2 mm (n=3). Body cylindrical, anterior and posterior ends rounded. Head directed anteroventrally. Spatula (as in Fig. 313) with anterior tooth about as wide as long, tapering from base to apex, almost trian- gular, obtusely notched at apex, the shaft long and narrow. Two lateral papillae present on each side of spatula. Six dorsal papillae on first 7 abdominal segments. Papillae without setae, not raised. Ab- dominal segments with sparse pointed verrucae dorsally at midlength surrounding dorsal papil- lae and horizontal rows of sharp, pointed spicules ventrally, anterior to sternal papillae, but none posteroventrally. Terminal segment shorter than eighth. Second instar: Not seen, but, because of similarity between the gall of this species and that of C. inflata, may be expected to have short-haired dorsal and pleural papillae. Type material—Holotype: Third instar (right- most specimen of 3 larvae on slide), from gall on Carya ovata, Schellsburg, Pennsylvania, IX-2-2003, R.J. Gagné, deposited in USNM. Etymology.—The name tumida is a Latin adjec- tive meaning swollen, referring to the inflated false chamber of the gall. Affinities —See under C. inflata. Collections of galls.—ILLiNots: Rock Island (tomentosa; as alba, B.D. Walsh). PENNSYLVANIA: Schellsburg (ovata). Specimens of C. tumida examined.—PENNSYL- VANIA: Schellsburg, IX-2-2003 (larvae). Caryomyia turbanella Gagné, new species Description.—Gall (Figs. 132-133): Rare but widely distributed, on Eucarya hickories; on veins of upper leaf surface; length 4.0-5.0 mm, spher- oidal, abruptly narrowed to conical apex; tan to purple, base with long, sparse, white hairs not ob- scuring surface, apical cone all or partly covered with small bumps and mostly hairless; base with circular, shallow, central excavation, leaf not exfo- liate surrounding connection; larval chamber near base of gall, depressed-ovoid, lined with yellow- ish membrane, contrasting with brown to purple, large-celled, spongy, viscous tissue surrounding larval chamber that shrivels with maturity; a bun- dle of longitudinal fibers present between larval chamber and gall apex. The gall of this species is one of only two Caryomyia galls found exclusively on the upper leaf surface. It is distinctive for its hairy spheroidal base and bumpy conical apex. Adult: Head: Antenna with 12 binodal flagel- lomeres, each of male flagellomeres (as in Fig. 239) with definite internode and neck, circumfila loops short, their bases widely separated; female flagellomeres with 2 horizontal circumfila con- nected by 2 vertical strands. Labella separated from one another anteriorly, hemispherical, short- setose, short-setulose. Palpus 4-segmented. Tho- rax: Wing length 4.2-4.3 mm in male (n=2), 2.5-3.3 mm in female (n=3). Male abdomen: Third to eighth tergites each with posterior setae continu- ous across width in mostly double row laterally, tapering to single row mesad, eighth tergite may have three setal rows laterally, the setae as strong as those of previous tergite. Genitalia (as in Figs. 256-257): hypoproct widened laterally to curve around sides of aedeagus, posterior margin se- tose, dorsal surface setulose only on posterior fourth; apex of aedeagus convex; gonostylus broad apically. Female abdomen: Third through eighth tergites as for male, setae of eighth tergite as long as on preceding tergite. Ovipositor: ninth segment pliable except for slender dorsal sclerite, setae sparse laterally, mostly as long as cerci, setae more numerous ventrally; cerci ovoid in lateral view, fused dorsally for about 1/2 length. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.8—-2.2 mm (n=6). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- ula (Fig. 310) with long, nearly parallel-sided, api- 84 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) cally notched tooth and long shaft slightly widened anteriorly and parallel-sided beyond. Papillae without setae, their bases not raised. Lat- eral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Ab- dominal segments with rounded verrucae cover- ing dorsum at midlength and smaller, horizontal rows of spicules on venter anterior to sternal papillae and posterior to ventral papillae. Termi- nal segment shorter than eighth. Second instar: Length 0.8-1.3 mm (n=9). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae not raised, asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. tomentosa, Entomology Road, BARC, Beltsville, MD, IX-26-1993, R.J. Gagné, deposited in USNM. Etymology—The name turbanella is a Latin noun meaning “little turban,” referring to the gen- eral shape of the gall. Affinities.—See under C. persicoides. Collections of galls —ALaBama: Greenville (to- mentosa). ARKANSAS: Piney (texana). CONNECTICUT: Plainfield (ovata). FLoriwa: Gainesville (glabra, to- mentosa). GEORGIA: 8 km E Fort Valley (pallida). MaryLANpD: Beltsville, USDA Agricultural Re- search Center (glabra, tomentosa); Brighton (tomen- tosa); Lusby (tomentosa); Ridgely (tomentosa). WasuinctTon, DC: National Arboretum (glabra). Specimens of C. turbanella examined.—ALa- BAMA: Greenville, [X-21—2000 (larva). ARKANSAS: Piney, IX-10-2004 (larvae). FLoripa: Gainesville (IX-21-2000). Georcia: 8 km E Fort Valley, X- 11-1993, J. & R. Payne (larvae). MARYLAND: Belts- ville, USDA Agricultural Research Center, IX- 26-1993, VIII-5-1998, and IX-28-1999 (larvae); Brighton, X-23-1999 (larvae); Lusby, IX-15-2005 & IX-15-2005 (larvae). WASHINGTON, DC: National Arboretum, VIII-9-1994 (larvae; 2 d,3 2, pupae, em. IV-1995). Caryomyia turbinata Gagné, new species Description.—Gall (Figs. 123-125): Occasional, found on Eucarya hickories; usually in groups, on lower leaf surface between veins; height 2.8-4.2 mm, obconic to spheroidal, widest at midlength or beyond, apical third eventually collapsing and becoming flat; surface brown to red, covered with both short and long brown hair not quite obscur- ing surface, the hair darker and denser apically; base with narrow, circular, shallow, central de- pression, sometimes barely noticeable, the leaf not exfoliated surrounding connection, usually with small convex, discolored area on reverse leaf sur- face; larval chamber basal, depressed-ovoid, lined with yellowish membrane, tissue beyond larval chamber red, soft, with large and spongy cells. This gall is unique among Caryomyia galls for its obconic shape, but in other characters appears most similar to the conical galls because of the ex- traneous, soft tissue outside the pellicle-lined lar- val chamber. Adult: Head: Antenna with 12 flagellomeres; male flagellomeres (as in Fig. 237) scarcely bin- odal, only slightly narrowed beyond basal cir- cumfilum, circumfila loops short, subequal in length; female flagellomeres with 2 horizontal cir- cumfila connected by 2 vertical strands. Labella (as in Fig. 242) separated from one another anteri- orly, hemispherical, short-setose, short-setulose. Palpus 4-segmented. Thorax: Wing length 1.92.2 mm in male (n=3); 2.2-2.6 mm in female (n=4). Male abdomen: Third through seventh tergites each with posterior setae continuous across width, in single row; eighth tergite weakly sclero- tized, without posterior setae. Genitalia (as in Figs. 247-248): hypoproct broadest near apex, posterior edge convex, margin setose except in middle third, dorsal setulae only on distal third; aedeagus furrowed apically and ventrally. Female abdomen: Third through eighth tergites each with posterior setae continuous across width, in mostly single row, usually double laterally; eighth tergite not less sclerotized than seventh and setae nearly as long. Ninth segment pliable except for slender dorsal sclerite that widens apically to invest part of cerci, setae sparse laterally, mostly as long as cerci, setae more numerous ventrally. Cerci elon- gate-ovoid in lateral view, pointed apically, fused dorsally for about 1/2 length. Pupa (as in Figs. 213-214): Antennal bases py- ramidal, separate, acutely pointed apicolaterally. Face with anterolateral pair of horizontal inci- sions, their posterior lip protruding to form a craterlike concavity. Frons and clypeus strongly protruded ventrally. Abdominal terga with sev- eral anterior rows of enlarged spinules. Larva: Third instar: Length 1.6-2.5 mm (n=10). Body cylindrical, anterior and posterior ends broadly rounded. Head directed anteriorly. Spat- RAYMOND J. GAGNE 85 ula (Fig. 305) with long, nearly parallel-sided, api- cally-notched tooth and long shaft. Papillae with- out setae, their bases not raised. Lateral papillae 2 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Abdominal segments with rounded and pointed verrucae covering dor- sum across midlength and horizontal rows of pointed spicules across venter anterior to sternal papillae and posterior to ventral papillae. Termi- nal segment shorter than eighth. Second instar: Not seen. Type material.—Holotype: Third instar (right- hand specimen of two on slide), from gall on C. to- mentosa, Entomology Rd, BARC, Beltsville, Mary- land, IX-26-1993, R.J. Gagné, deposited in USNM. Etymology—The name turbinata is a Latin ad- jective that means top-shaped or obconic and refers to the gall shape of this species. Affinities—The gall is unique in Caryomyia be- cause of its obconic shape, but the soft tissue of the distal half that eventually collapses brings to mind galls of, e.g., C. persicoides and C. turbanella. The male antennae and genitalia of the present species are quite different from the other two and more similar to the C. sanguinolenta group. The ovipositor is unlike that of either group, being similar to that of C. inclinata (Fig. 262). Collections of galls—ALAaBaAMa: Auburn (gla- bra, tomentosa); Greenville (tomentosa); Oak Mt. St. Pk, Shelby Co. (pallida). Arkansas: Beulah (tomen- tosa); London (tomentosa); Osage (texana). CON- NEcTIcuT: Amston (ovata); Phoenixville (ovata). Fioripa: Gainesville (glabra, tomentosa). GEORGIA: Carnesville (ovata); Lizella, J. & R. Payne (tomen- tosa). ILtLtiNois: Marshall (tomentosa). INDIANA: Spencer Co. (ovata). KeNTucky: Brownsville (to- mentosa). LOUISIANA: Lake Bistineau St. Pk, Web- ster Parish (texana). MARYLAND: Beltsville, USDA Agricultural Research Center, Beltsville (fomen- tosa); Clarksburg (tomentosa); Gaithersburg (tomen- tosa); Kemp Mill (glabra); Lusby (tomentosa); Ridgely (glabra, tomentosa); Lusby (glabra, tomen- tosa); Ridgely, (tomentosa); St. Leonard (tomentosa); Silver Spring (tomentosa); Wheaton (tomentosa). Massacuusetts: Great Barrington (ovata); Peter- sham (ovata). Mississippi: Natchez Trace Pkwy, Mile 123, Rankin Co. (tomentosa); Noxubee Natl Wildlife Ref, Oktibbeha Co. (tomentosa). Missouri: Conway (ovata, texana). NEw York: East Hampton (tomentosa). NORTH CAROLINA: Alamance (fomen- tosa); Kannapolis (tomentosa); Rowland (tomen- tosa). OKLAHOMA: Sallisaw (tomentosa). PENNSYLVA- NIA: Kirby (ovata). TENNESSEE: Dickson (tomentosa); Jackson (tomentosa). Texas: Daingerfield St. Pk, Morris Co. (tomentosa); New Waverly (texana). Vir- GINIA: Dinwiddie (tomentosa). WASHINGTON, DC: Walter Reed Army Medical Center (tomentosa). Specimens of CC. furbinata examined.— ARKANSAS: Osage, IX-9-2004 (larvae). FLORIDA: Gainesville, IX-21—2000 (larvae). GEorGiIA: Carnes- ville, IX-19-2000 (larvae). KENTUCKY: Brownsville, IX-21-1995 (larva). MARYLAND: Beltsville, USDA Agricultural Research Center, Beltsville, IX- 26-1993 (larvae); Gaithersburg, X-4-1994 (larvae; ?, pupae, em. IV-1995); Kemp Mill, X-1-2005 (larva); Lusby, IX-2003 (2 °, em. IV-2004); Ridgely, X-2-2000 (larvae), X-6-2005 (larvae; pupae, 12 °, em. IV-5 to 19-2006); Silver Spring, X-3-1993 (lar- vae; 6, 2 2, pupae, em. IV-25 to 27, 1994) & X- 23-1999 (larvae); Wheaton, IX-19-1993 (larvae). MassacuHusetts: Petersham, IX-6-—2000 (larvae). Missouri: Conway, IX-8—2004 (larvae). OKLaA- HOMA: Sallisaw, IX-9-—2004 (larvae). PENNSYLVANIA: Kirby, IX-6- 2004 (larvae). TENNESSEE: Dickson, IX- 11-2004 (larva). Texas: New Waverly, X-6-2001 (larva). WASHINGTON, DC: Rock Creek Park, IX- 22-1993 (larvae). Caryomyia urnula Gagné, new species Description—Gall (Figs. 28, 41-42): Late spring, early summer, common on the Eucarya section, rare on Apocarya section; found singly be- tween veins on either upper or lower leaf surface; 2.0-3.0 mm in height, cylindrical, slightly bulging at midlength, narrowed abruptly apically to con- vex areola surrounding prominent nipple, surface smooth to weakly lined longitudinally; hairless, not sticky, white to green, turning dark green or brown; base shallowly concave; wall firm, brittle, uniformly thin, larval chamber glabrous with lon- gitudinal ridges. This gall resembles that of C. cili- dolium and relatives in the caplike apex and the central indentation of the base, but it is smaller, and more cylindrical than spheroid. The apex does not break off easily as it does in galls of C. cilidolium and relatives. Adult: Unknown. Pupa (Figs. 195-196): Antennal bases oblique in ventral view parallel, apices closely adjacent, rounded anteriorly, dorsoventrally flattened. Face with longitudinal impression running along length, a slight anteromesal convexity posterior to each antennal base, and without anterolateral pair 86 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) of horizontal incisions. Frons and clypeus slightly convex. Abdominal tergal spinules all similarly small. Larva: Third instar: Length 1.6-2.1 mm (n=10). Body elongate-cylindrical, anterior and posterior ends rounded. Head directed anteriorly. Spatula (Fig. 276) with pointed anterior tooth and long lin- ear shaft barely wider than tooth. Lateral papillae 1-3 on each side of spatula. Only 4 dorsal papillae on first 7 abdominal segments. Papillae without setae, their bases not raised. Abdominal segments almost completely covered with rounded verru- cae dorsally and pointed verrucae ventrally. Ter- minal segment shorter than eighth. Second instar: Length 1.0-1.3 mm (n=10). Collar segment with patch of enlarged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, concave-sided shaft. Bases of dorsal and pleural papillae slightly raised, asetose. Only 4 dorsal papillae on first 7 abdominal segments. Type material.—Holotype: Third instar, from gall on C. tomentosa, Beltsville, Maryland, X- 8-2002, R.J. Gagné, deposited in USNM. Etymology.— The name urnula, a noun, is Latin for a small urn and refers to the shape of the gall. Affinities —The pupa is similar to that of the C. cilidolium group with the longitudinal central facial line and slight prominence posterior to the antennal bases, lack of a frontal projection, and lack of dorsal specialized spicules on the ab- domen. The larva of C. urnula, although it has a single-toothed spatula and is completely covered with verrucae, has the head directed anteriorly, a shorter terminal larval segment, and a much less verrucose integument. These differences may only reflect the fact that the gall of C. urnula is more cylindrical than spheroidal, but a more im- portant difference may be that the larva of C. ur- nula has only 4 dorsal papillae on the first through seventh abdominal segments while the C. cili- dolium group has 6. Biological notes.—This species is a generalist that occurs on both Eucarya and Apocarya sections and can be found on either leaf surface. Full- grown galls can be found any time from May through October and full-grown larvae anytime after early June, a time range unique for Cary- omyia. Development of galls is evidently stag- gered. In late May through October in central Maryland, galls can be found with first, second, or third instars inside, and second instars can be found in galls into late August. These galls appear to dehisce from the leaves as soon as the third in- stars mature, possibly because of the active larvae inside. Agitation by the full-grown larva in a de- hisced gall causes the gall to pop up and down and roll on a level surface. Early dehiscence may be the reason the galls are uncommon on leaves in late season, although occasional galls with live third instars can be found through September and into October, at least in Maryland. In one Septem- ber trip from Maryland through Arkansas to col- lect hickory galls, I found not a single example. Collections of galls—ALaBama: Auburn (ovata); Birmingham (pallida). Arkansas: S Foreman (illinoiensis). CONNECTICUT: Phoenixville (ovata); Thomaston (cordiformis). FLoripA: Alachua Co. (Hodges et al. 2006: bottom photo, upper half only). Georeta: Forsyth (glabra); 8 km E Fort Val- ley (pallida); Lizella (ovata). Kentucky: Grayson Lake St. Pk, Carter Co. (ovata); Pennyrile St. Pk, Christian Co. (ovata). LOUISIANA: Bistineau St. Pk, Webster Parish (tomentosa). MARYLAND: Beltsville, USDA Agricultural Research Center (tomentosa); Clarksburg (glabra, tomentosa); Ellicott City (cordi- formis, tomentosa); Gaithersburg (tomentosa); Lusby (tomentosa); Ridgely (cordiformis, tomentosa); Wheaton (cordiformis, glabra). Missouri: Cadet (cordiformis); Holly Ridge St. For., Stoddard Co. (tomentosa). Onto: Hocking St. For., Hocking Co. (glabra); Hocking Co. (tomentosa, Wells 1916: Fig. 28). TENNESSEE: Gatlinburg (ovata). VIRGINIA: Lees- burg (tomentosa). WASHINGTON, DC: National Ar- boretum (glabra). West VirGINIA: Beech Fork St. Pk, Wayne Co. (glabra, ovata, tomentosa); Falling Waters (cordiformis). Specimens of C. urnula examined.—GEoRGIA: 8 km E Fort Valley, X-11-1993, J. & R. Payne (larva). MaryYLANpD: Beltsville, USDA Agricultural Re- search Center, V-30-1977, V-22-1994, V-26 & VI- 4-1997, V-29-1998, VI-18-1999, V-16 & 28-2000, X- 8-2002, & V-26-2004 (larvae), and VI-9—-2004 (pupae taken from galls [V-1—2005); Clarksburg, V-23-2000 & V-26-2004 (larvae); Ellicott City, VI- 8-2005 (larvae); Lusby, IX-16-2003 (larvae); Wheaton, V-22-2000 (larvae). WASHINGTON, DC: National Arboretum, V-24—1994 (larvae). Caryomyia viscidolium Gagné, new species Description —Gall (Figs. 25, 43-45): Occa- sional, found on Eucarya hickories; on upper leaf surface, between veins; 3.0-4.1 mm in height, RAYMOND J. GAGNE 87 spheroidal, longer than wide, base broadly rounded to truncate in profile, apex with small central nipple surrounded by extensive areola that eventually separates from gall; surface resinous, especially on young galls, sticky, smooth except for scattered minute resin dots, areola paler than surrounding surface; base with wide, deep, circular indentation; wall firm, brittle, al- most uniformly thin, larval chamber glabrous, with longitudinal ridges. This gall and that of the unrelated C. turbanella are the only Caryomyia galls that occur exclusively on the upper leaf sur- face of hickories. For notes on galls of related species, see under C. cilidolium. Adult: Head: Antenna with 12 flagellomeres, in male (as in Fig. 240) with short necks and evenly cylindrical, girdled by 8-9 more or less parallel, occasionally interconnected, appressed circumfila interspersed with setae and covered with setulae; in female (Fig. 241), circumfila anastomozing on venter. Labella (Fig. 243) fused, the pair united, semicircular, setulose but asetose. Thorax: Wing length 2.3 mm in male (n=1) 3.1-3.3 mm in female (n=2). Male abdomen: Third through seventh ter- gites each with posterior setae continuous across width in mostly double, partly triple row; eighth tergite weakly sclerotized, with small posterior setae. Genitalia (as in Fig. 251): hypoproct broad- est posteriorly and expanded ventrally to partly cover sides of aedeagus, each of the 2 lobes with 2-3 apical setae, dorsal surface completely cov- ered with setulae on anterior half; aedeagus taper- ing gradually from wide base to narrow apex, the tooth large, dorsoventrally compressed. Female abdomen: Third through seventh tergites each with posterior setae in 3 horizontal rows laterally, thinning to 2, then one, with slight medial inter- ruption in setae; eighth tergite more weakly scle- rotized than preceding tergite, with weak poste- rior setae numerous laterally, continuing mesally in single, sparse row. Ninth segment (Fig. 261) pli- able except for slender dorsal sclerite, with nu- merous apicolateral and ventral setae, none longer than the cerci, the two groups separated laterally. Cerci elongate-ovoid in lateral view, pointed apically, fused dorsally for about 1/2 length. Pupa (as in Figs. 193-194): Antennal bases elon- gate, in ventral view parallel, closely adjacent, rounded anteriorly, the apices dorsoventrally flat- tened. Face with longitudinal impression running along length, a slight anteromesal convexity pos- terior to each antennal base, and without antero- lateral pair of horizontal incisions. Frons and clypeus slightly convex. Abdominal tergal spin- ules all similarly small. Larva: Third instar: Length 2.5-3.2 mm (n=10). Body slightly dorsoventrally flattened, elongate- ovoid, broadly rounded anteriorly, spindleform posteriorly. Head directed ventrally, surmounted dorsally by collar segment. Spatula (Fig. 279) with pointed anterior tooth and long, broad shaft. Papillae on dorsum and pleura prominently raised above surrounding surface, without setae. Lateral papillae 2 on each side of spatula. Six dorsal papil- lae on first 7 abdominal segments. Body com- pletely covered with verrucae. Terminal segment as long as eighth, rounded. Second instar: Length 1.0-1.8 mm (n=4). Collar segment with patch of en- larged spicules on each side. Spatula with acutely triangular anterior tooth and longer, wider, con- cave-sided shaft. Six dorsal papillae on first 7 ab- dominal segments. Bases of dorsal, pleural, termi- nal, and ventral papillae elongate, asetose. Type material—Holotype: Third instar (right- hand specimen of two on slide), from gall on C. to- mentosa, Brunett Ave., Silver Spring, Maryland, IX-12-1999, R.J. Gagné, deposited in USNM. Etymology—The name viscidolium, a noun, combines the Latin viscidus (sticky) and dolium (jar) and refers to the resinous, barrel-like gall. Affinities —See under C. cilidolium. Biological notes.—Pupae break out of the weak cap at the gall’s apex. Emergence occurs in April. Collections of galls—Arkansas: Osage (tex- ana); Palestine (ovata). CONNeEcTICUT: Danbury (ovata); Hartland (ovata); Harwinton (ovata); Litch- field (ovata); Meriden (glabra, ovata); Phoenixville (ovata). ILLINOIS: Marshall (tomentosa); Rock Island (ovata, tomentosa). INDIANA: New Castle (glabra). Kentucky: Grayson Lake St. Pk, Carter Co. (ovata, tomentosa); My Old Kentucky Home St. Pk, Nel- son Co. (ovata). MARYLAND: Beltsville, USDA Agri- cultural Research Center (glabra, ovata, tomentosa); Brighton (ovata); Ridgely (glabra, ovata, tomentosa); Silver Spring (tomentosa); Spencerville (tomentosa); Wheaton (glabra). MassacuusetTtTs: Bolton (ovata); Great Barrington (ovata); Magnolia (Carya sp.); Merrimac (ovata); vic. Worcester (Carya sp., Thompson 1915: 89, Fig. 218). Mississippi: nr. Ses- sums, Oktibbeha Co. (ovata). NEw HAMPSHIRE: Barrington (ovata); Seabrook (ovata). NEW JERSEY: Columbia (ovata). New York: Herkimer (ovata). 88 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) NortH Caro ina: Alberta (glabra). Onto: Hocking St. For., Hocking Co. (ovata); Hocking Co. (glabra, Wells 1916: Fig. 25); St. Clairsville (tomentosa). PENNSYLVANIA: Julian (glabra). VirGINIA: Huntley Meadows Pk, Fairfax Co. (tomentosa). WASHING- TON, DC: National Arboretum (glabra). West Vir- GINIA: Beech Fork St. Pk, Wayne Co. (ovata, tomen- tosa); 6 mi NW Hedgesville (ovata, tomentosa). Specimens of C. viscidolium examined.—Con- NectTicuT: Hartland, IX-7-2005 (1 3, pupal exu- viae, em. IV-19-—2006). ILLinois: Marshall, IX- 7-2004 (larva). MARYLAND: Beltsville, USDA Agricultural Research Center, IX-26-1993 & VII- 1-2000 (larvae); Brighton, IX-6-2002 (¢, pupa, em. IV-2003); Silver Spring, IX-12-1999 (larvae) & X-2003 (larvae; 2 2, pupae, em. IV-10—-2004); Wheaton, VII-3-1994 and IX-4-1994 (larvae). MassacuuseTts: Great Barrington, IX-6—2000 (larva). Onto: St. Clairsville, IX-6—2004 (larva). WASHINGTON, DC: National Arboretum, X- 13-1997 (larvae). West VIRGINIA: 6 mi NW Hedgesville, IX-19-1999 & VII-23—2000 (larvae). GENUS CARYADIPLOSIS GAGNE, NEW GENUS Type-species, Caryadiplosis venicola Gagné, by present designation. Diagnosis.—This genus differs from other gen- era of Cecidomyiidi by the following combination of characters: Adult without dorsal protuberance on occiput; costal wing vein unbroken at juncture with Rs; gonostylus short, broad, fully setulose, with wide tooth; protrusible ovipositor short; fe- male ninth tergite dorsally with narrow, pig- mented, stiff, dermal structure; cerci discrete, bi- laterally flattened, discoid, without distal pair of trichoid sensilla. Larva Contarinia-like, with full complement of papillae for supertribe (Gagné 1994), terminal segment with 1 pair of corniform, recurved papillae and three pairs with short setae. Description.—Adult: Head: Eyes connate. Oc- ciput without dorsal protuberance. Palpus with 3 to 4 segments. Labrum with short setae on venter. Labella discrete, nearly hemispherical but blunt at apex, setulose, setose. Antenna with 12 flagellom- eres: male flagellomeres (Fig. 328) binodal, inter- nodes and necks short, with 3 circumfila, 1 on basal node and 2 on distal node, loops of circum- fila subequal in length; female flagellomeres (Fig. 329) with cylindrical node and, except for last fla- gellomere, short necks, each node with 2 horizon- tal, appressed circumfila connected by 2 vertical strands. Thorax: Scutum with 4 longitudinal rows of setae. Scutellum with lateral group of setae on each side. Pleura hairless except for anepimeron. Wing: length 2.0-2.3 times width; C unbroken at juncture with Rs (as in Fig. 245); Rs curved apically, joining C near wing apex; Rs detectable only as stump on R; and situated closer to arculus than to R,. Legs 0.9-1.1 length of wings; tarsal claws un- toothed, curved beyond midlength; empodia as long as claws; pulvilli about 1/4 length of claws. Male abdomen (Fig. 330): First through seventh tergites entire, rectangular, about 5 times as wide as long; with single row of setae along posterior margin, interrupted mesally on seventh tergite; an extensive lateral group of setae on each side near midlength with sparse, setiform scales intermixed, the two lateral groups almost joining mesally, and anterior pair of trichoid sensilla; eighth tergite weakly sclerotized, short, 2 lateral papillae on each side and anterior pair of trichoid sensilla the only vestiture; sternites rectangular, about as wide as long, second through seventh with single horizon- tal row of setae posteriorly except seventh with double row, a large group of setae near midlength, and 2 closely adjacent trichoid sensilla anteriorly; eighth sternite with several rows of setae posteri- orly, bare anteriorly except for 2 widely separated trichoid sensilla. Genitalia (Figs. 332-333): cerci convex at apex, setose along posterior margin; hy- poproct slightly longer than cerci, divided into 2 rounded, posteriorly setose lobes, with ventral se- tulae throughout; horizontal ridge at midlength; aedeagus broad, slightly longer than hypoproct; gonocoxite bulbous; gonostylus short-ovoid, evenly setose, completely setulose. Female ab- domen (Fig. 331): First through seventh tergites as for male, eighth tergite with complete, mostly sin- gle row of posterior setae; sternites also as for male, except eighth not well defined, the pair of anterior trichoid sensilla of that sclerite widely separated, the posterior setae scattered; ovipositor short, protrusible, abruptly tapered beyond exten- sion of eighth segment; ninth tergite with narrow, pigmented sclerite, otherwise moderately covered with short, scattered setae; cerci ovoid, covered with dense setae but without peglike sensilla; hy- poproct short, broad, with 2 posterior setae. Pupa: Unknown. Larva: Third instar (Figs. 334-337).—Cylindri- RAYMOND J. GAGNE 89 cal, anterior end tapered, posterior end broadly rounded. Head directed anteriorly, apodemes longer than head capsule. Antenna about twice as long as wide. Spatula clove-shaped. Lateral papil- lae on each side of midline in two triplets, two of each group with short setae, the third slightly larger in diameter, without seta. Dorsal and pleu- ral papillae with minuscule setae. Terminal seg- ment with four pairs of papillae, one pair corni- form, recurved, one pair situated between corniform pair with short setae, two pairs placed laterad, with short setae. Second instar (not seen). Etymology.—The name Caryadiplosis combines Carya, the name of the host genus, with “diplo- sis,” meaning “double,” with reference to the bin- odal male flagellomeres. The suffix is commonly used for genera of the supertribe Cecidomyiidi. Affinities.—Caryadiplosis belongs to the super- tribe Cecidomyiidi, chiefly because of the two dis- tinct nodes and separate many-looped circumfila of the male flagellomeres and the lack of setae on the ventral papillae of the larval abdomen. It be- longs to the tribe Cecidomyiini because the adult has simple tarsal claws that are curved beyond midlength and the larval terminal segment has one pair of corniform, recurved papillae and three setiform pairs. The new genus shares with Caryomyia the fol- lowing character states that appear to be synapo- morphic: Wing unbroken at junction of R5 and C; anepisternum without vestiture; gonostylus ovoid and entirely covered with setulae; oviposi- tor short-cylindrical and abruptly narrowed be- tween eighth and ninth segments, the ninth segment with a dorsal, elongate, stiff, dark-pig- mented sclerite along its entire length, and cerci lacking a distal pair of peglike sensilla. Caryadiplosis differs from Caryomyia in the fol- lowing ways: adult abdominal tergites have lat- eral setae (a group separate from the posterior se- tae) that are absent in Caryomyia; adult sternites have the anterior pair of trichoid sensilla that are lost in Caryomyjia; larvae have six lateral papillae on each side of the midline, in two groups of three each, as do most Cecidomyiini, but the discrete grouping and at least three of the papillae are lost in Caryomyia; larval terminal papillae comprise a pair of corniform papillae and three pairs of se- tose papillae, as do most Cecidomyiini, but that are all merely flat or convex and asetose in Cary- omyia; larvae drop to the soil to pupate unlike those of Caryomyia whose larvae pupate in the galls. It is fortunate that this genus was found dur- ing this study because it serves as a bridge be- tween Caryomyia and Contarinia and relatives. De- spite the many striking differences between Caryadiplosis and Caryomyia, the two genera share the presumably synapomorphic features outlined above. If one assumes a close affiliation between the two genera, it appears that Caryomyia sepa- rated from Caryadiplosis while already on hicko- ries and that the greatly modified larvae of Cary- omyia are a response to their passage from simple to complex galls and from pupating in the soil to pupating in the galls. In the key to Nearctic genera of Cecidomyiidae in Gagné (1981), Caryadiplosis will run to Paradiplo- sis in the first half of couplet 146, except for the host indication, as Paradiplosis spp. are pests of fir and spruce. The two genera differ in the setation of the second through seventh adult abdominal tergites: those of Caryadiplosis have 10 or more lateral setae on each side in one or two rows that almost meet medially and are intermixed with setiform scales; the tergites of Paradiplosis have only 3-5 lateral se- tae close to the lateral margin and no scales. Differ- ences between females are more striking: in Caryadiplosis the ovipositor is barely longer than the seventh tergite, while in Paradiplosis it is as long as the rest of the abdomen. In addition, female cerci of Paradiplosis have an apical pair of setae dif- ferented from the remainder by their thickness and larger socket, but cerci of Caryadiplosis have no such differentiated cerci. Interestingly, in both gen- era the costal wing vein is unbroken at its juncture with R; and the gonostylus is completely setulose, similarities that beg the question of affinity be- tween the two genera. SPECIES TREATMENTS Caryadiplosis biconvexa Gagné, new species Description.—Gall (Figs. 33, 167-169): Com- mon, on both Eucarya and Apocarya sections; circu- lar swelling, ca. 3.5 mm in diameter, on lamina, showing on both sides of leaflet; similar in color and texture to leaf, but some mature galls may turn black before larva escapes; wall of uniform thick- ness, larval chamber circular, flattened, reflecting 90 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) gall shape; full-grown larva exits gall from small hole in center of gall on abaxial leaflet surface. Adult (only female known): Head: Eyes about 6 facets long at vertex; facets mostly circular, closely adjacent except slightly farther apart at midheight of head. Frons with 5 setae per side. Thorax: Pleura hairless except for anepimeron with 8-10 setae. Wing length 1.9-2.0 (n=2). Fe- male abdomen: As for C. venicola. Pupa: Unknown. Larva: Third instar: White at first, turning to yellow during third instar, body cylindrical, ante- rior end tapered, posterior end broadly rounded. Head directed anteriorly, with apodemes longer than head capsule. Antenna about twice as long as wide. Spatula (Fig. 337) with anterior tooth com- posed of two elongate, rounded lobes, and with long posterior shaft. Integument on dorsum with- out sculpturing, on pleura with pointed verrucae anterior to spiracles, and on venter with horizon- tal rows of sharp spicules anterior to sternal papil- lae. Lateral papillae on each side of midline in two triplets, two of each group with short seta, the third larger and without seta. Dorsal and pleural papillae with very short setae. Terminal segment (Fig. 336) covered dorsally with verrucae, with four pairs of papillae, one pair corniform, re- curved, one pair situated between corniform pair with tiny setae, two pairs placed laterad with short setae and situated on short lobes. Type material.—Holotype: Larva, ex biconvex gall on C. tomentosa, National Arboretum, Wash- ington, DC, X-8-1991, R.J. Gagné, deposited in USNM. Etymology.—The name biconvexa is an adjec- tive from the Latin with reference to the lens- shaped gall visible on both sides of a leaflet. Affinities —The larval terminal segment has dorsal verrucae in contrast with the smooth pre- ceding segments. This is an effective character for separating this species from C. venicola. The two species are further separated by the terminal papillae all being situated on lobes in C. biconvexa and the lack of such lobes on C. venicola. Biological notes.—Small circular galls with first instars were found as early as May 26 in central Maryland. On both June 5 and 9 yellow galls were found with full grown larvae or from which larvae had already escaped, indicating that development is swift. Possibly this species has more than one generation per year because galls with first instars were found as late as Sept. 11 and second and third instars on Sept 26, but the galls seemed not as com- mon in late summer as in spring. Larvae exit from the lower surface of the leaflet. Collections of galls.— (The small number of col- lections does not reflect how common this species is because I did not always make a note of old galls). ALABAMA: Auburn (aquatica); Greenville (to- mentosa). ARKANSAS: London (cordiformis). GEOR- GIA: Forsyth (ovata). INDIANA: Richmond. (cordi- formis). MaryLanp: Beltsville, USDA Agricultural Research Center (glabra, tomentosa); Calverton (to- mentosa); Clarksburg (tomentosa); College Park (glabra); Ellicott City (cordiformis, tomentosa); Long Beach (cordiformis); Lusby (tomentosa); Piney Grove (ovata); Ridgely (glabra, tomentosa); Spencerville (to- mentosa); Wheaton (glabra). MASSACHUSETTS: Peter- sham (ovata). New York: Hyde Park (ovata); Nas- sau (Carya sp.). NoRTH CAROLINA: Alberta (glabra). Kannapolis (tomentosa); Providence (tomentosa). Onto: Vinton Co. (cordiformis, Wells 1916: Fig. 33). OKLAHOMA: Sallisaw (tomentosa). PENNSYLVANIA: Millsboro (glabra). TENNESSEE: Dickson (tomentosa); Jackson (tomentosa). WASHINGTON, DC: (tomentosa). WEST VirGINIA: Falling Waters (cordiformis). Specimens of C. biconvexa examined.—Mary- LAND: Beltsville, USDA Agricultural Research Cen- ter, VI-14 & 29-2000 & VII-12-2000 (larvae), VI- 15-2004 (2 2, em. IV-20-2005); Calverton, X-10- 2005 (larvae); Clarksburg, X-4—2005 (larvae); Elli- cott City, IX-2-2000 (larvae). New York: Nassau, X- 3-1907, E.P. Felt (larva). WASHINGTON, DC: Na- tional Arboretum, X-8-1991 (larva); Walter Reed Army Medical Center, IX-26-2005 (larvae). WEST VirGInia: Falling Waters, VI-21—2004 (larvae). Caryadiplosis venicola Gagné, new species Description.—Gall (Figs. 165-166): Occasional, on both Eucarya and Apocarya sections; an elon- gate, hard swelling of midvein or petiole, usually showing on both sides of leaflet, 5 mm long, 2mm high when single chambers, longer and broader when several chambers are in tandem; exterior tan to brown, rough; wall of uniform thickness; larval chamber elongate; an exit hole eventually develops on lower leaf surface through which the full-grown larva falls to ground. Adult: Head: Eyes about 6 facets long at vertex; facets mostly circular, closely adjacent except slightly farther apart at midheight of head. Frons RAYMOND J. GAGNE 91 with 3-5 setae per side. Thorax: Pleura hairless except for anepimeron with 7-9 setae. Wing length 2.0 mm in male (n=1); 2.3-2.5 mm in female (n=3). Male abdomen: Segments 6-8 as in Fig. 330; genitalia, Figs. 332-333. Female abdomen: Seg- ments 8 to end, Fig. 331. Pupa: Unknown. Larva: Third instar: Orange. Spatula (Fig. 335) with anterior end deeply divided into two elon- gate, subtriangular lobes and with long posterior shaft. Integument on dorsum without sculptur- ing, on pleura with pointed verrucae anterior to spiracles, and on venter with horizontal rows of sharp spicules anterior to sternal papillae. Termi- nal segment (Fig. 334) smooth on dorsum, with- out verrucae, with four pairs of papillae, one pair corniform, recurved, one pair situated between corniform pair with short setae, two pairs placed laterad, with short setae, not situated on short lobes. Type material——Holotype: Larva, middle of 3 specimens on slide, ex midrib swelling on C. cordi- formis, Ridgely, Maryland, IX-23-1999, R.J. Gagné, deposited in USNM. Etymology.—The name venicola, noun from the Latin meaning vein inhabitant, refers to the gall site. | Affinities.—See under C. biconvexa for charac- ters to separate larvae of that species from C. venicola. Biological notes.—My earliest-season sighting of galls in central Maryland was on July 22. I no- ticed them more commonly in Maryland and else- where in late August and September. Larvae left the galls on collected leaves in autumn, burrowed into the soil, and spun cocoons. In early April the following year, full-grown pupae found their way to the surface of the soil and adults emerged. Collections of galls—Arkansas: Beulah (to- mentosa); Chester (texana); Osage (tomentosa); Piney (texana). FLoriwa: Gainesville (glabra, tomen- tosa). GEORGIA: Forsyth (glabra). MARYLAND: Beltsville, USDA Agricultural Research Center (to- mentosa); Cabin John (cordiformis); Ellicott City (to- mentosa); Long Beach (cordiformis); Lusby (tomen- tosa); Ridgely (cordiformis, tomentosa). Missouri: (tomentosa). NORTH CAROLINA: Wise (tomentosa). Ouro: Yellow Springs (glabra). OKLAHOMA: Salli- saw (tomentosa). TENNESSEE: Crossville (pallida); Jackson (tomentosa); Mohawk (tomentosa). WEST VirGINIA: Falling Waters (cordiformis). Specimens of C. venicola examined.— ARKANSAS: Osage, IX-9—-2004 (larvae); Piney, IX- 10-2004 (larva; ¢,3 2, em. IV-6/10-2005). MaAry- LAND: Cabin John, VIII-26—2004 (larvae); Ellicott City, IX-2-2000 (larva); Long Beach, IX-15-2005 (larvae); Lusby, IX-15-2005 (larvae); Ridgely, Ix- 23-1999 & IX-11-2003 (larvae). Onto: Yellow Springs, IX-7-2004 (larvae). West VIRGINIA: Falling Waters, IX-10-2005 (larvae). GENUS CONTARINIA RONDANI Contarinia Rondani 1860: 289. Type species, Tipula loti De Geer, by original designation; Gagné 2004: 108 (catalog). Contarinia is a large genus of more than 300 species, most of which are found in flower heads or marginal leaf rolls. Very few of them are found in complex galls. Two species are placed in Con- tarinia here only in the broad sense because adults, not yet known for both, are needed for strict generic assignment. Various other genera of Cecidomyiini, including Caryadiplosis and Har- mandiola, have larvae generally similar to Con- tarinia (Gagné 1989). SPECIES TREATMENTS Contarinia bulliformis Gagné, new species Description —Gall (Figs. 170-171): On pecan, Apocarya section; a bubble-like swelling of mid- vein or petiole, usually showing on both sides of leaflet, about 1 cm long, 3 mm high; green, smooth; wall of uniform thickness, larval cham- ber of same shape as gall, containing one to sev- eral larvae. Adult and pupa: Unknown. Larva: Third instar: Yellow, body cylindrical, anterior end tapered, posterior end tapering but truncate at end. Head directed anteriorly, apodemes as long as head capsule. Antenna about twice as long as wide. Spatula (Fig. 342) with ante- rior teeth pointed, their bases bulging slightly at lateral angles and with long posterior shaft. In- tegument on dorsum without sculpturing but with horizontal rows of verrucae anterior to dor- sal papillae, on pleura with pointed verrucae an- terior to spiracles, and on venter with horizontal 92 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) rows of sharp spicules anterior to sternal papillae. Lateral papillae on each side of midline in two triplets, two of each group with short seta, the third of slightly larger diameter and without seta. Dorsal and pleural papillae with setae no longer than wide. Terminal segment (Fig. 341) mostly covered dorsally with pointed verrucae, with four pairs of papillae all situated at the end of cylindri- cal lobes, one pair large, corniform, recurved dor- sally, the pair situated between corniform pair with well-developed setae 3 times as long as wide, one lateral pair with setae 4 times as long as wide, the remaining lateral pair with minute setae no longer than wide. Second instar: (not seen). Type material——Holotype: Larva, ex bubble swelling on leaf vein of C. illinoiensis, Green- sleeves Orchard, Caddo Parish, Louisiana, V- 5-1982, Hall & Stoetzel, deposited in USNM. Etymology.—The name bulliformis, an adjec- tive from the Latin meaning bubble-shaped, refers to the gall form. Affinities.—This species does not appear for now to fit in Caryadiplosis because papillae of the terminal segment are all situated on cylindrical lobes and four of the papillae have fairly long se- tae. In Caryadiplosis the terminal setae are minute and the subtending lobes are small or nonexistent. In addition, the spatula lobes of C. bulliformis are pointed rather than rounded. This is one of only two species of Contarinia (sensu lato) taken from hickories. It differs from more typical Contarinia spp. by the distinct lobes subtending all of the ter- minal papillae. Biological notes.—This species was found by Manya B. Stoetzel in the course of her field work on Phylloxera on pecan. Nothing further is defi- nitely known of the biology of this species, but the full-grown larvae probably escape the galls to drop to the soil. Collections of galls and specimens of C. bulli- formis examined.—LouIsIANA: Greensleeves Or- chard, Caddo Parish, V-5-1982, Hall & Stoetzel, 4 larvae (including holotype). Contarinia cucumata Gagné, new species Description.—Gall (Figs. 29, 172-174): Infre- quent, on Eucarya and Apocarya sections; on lower leaf surface, adjacent to vein; 3.6-5.0 mm in length, constricted at point of attachment to leaf, cylindrical and decumbent beyond, convex api- cally, not dehiscent; surface rough, green becom- ing yellow, then brown, with many scattered, long, fine, white hairs not obscuring surface; base initially crimped, opening to upper surface of leaf when mature; wall thin, at first soft, later brittle, larval chamber rough. Adult and pupa: Unknown. Larva: Third instar: White, body cylindrical, anterior end tapered, posterior end broadly rounded. Head directed anteriorly, with apo- demes longer than head capsule. Antenna about twice as long as wide. Spatula (Fig. 339-340) with anterior end divided into two triangular lobes (Fig. 339) that become rounded (Fig. 340) with use, and long posterior shaft. Integument on most of dorsum without sculpturing, on pleura with pointed verrucae anterior to spiracles, and on ven- ter with horizontal rows of sharp spicules anterior to sternal papillae. Lateral papillae on each side of midline in two triplets, two in each triplet with short seta, the third slightly larger in diameter and without seta. Dorsal and pleural papillae with very short setae. Terminal segment (Fig. 338) cov- ered dorsally with pointed verrucae, with four pairs of papillae, one pair corniform, recurved, one pair situated between corniform pair with tiny se- tae, two pairs placed laterad, situated on short lobes, with tiny setae. Second instar: (not seen). Type material.—Holotype: Third instar (middle of 3 specimens on slide), ex leaf gall, C. tomentosa, Clarksburg, Maryland, V-26—2004, R.J. Gagné, de- posited in USNM. Etymology.—The name cucumata, an adjective, is a shortened version of the Latin cucumerata, meaning cucumber-like, with reference to the gall’s general resemblance to a cucumber. Affinities—Larvae of Caryadiplosis venicola, also on hickory, are generally similar to this species, but young third instars of C. venicola have rounded spatula teeth while those of C. cucumata have pointed teeth. Contarinia.cucumata is further distinguished by the complex gall it causes; most Contarinia spp. make no gall or simple ones, such as leaf rolls. Biological notes.—This species shows wear of its spatula while still in the gall. Figs. 339-340 show the spatula of a newly molted third instar and an older one. The spatula of the newly molted larva is not yet entirely hardened or pigmented. In central Md, galls were first noticed on May 23 when still only bubble-like and containing first in- RAYMOND J. GAGNE 93 stars. Galls were greenish white to light green, somewhat sticky, and covered with close, long white hairs. In subsequent days these hairs be- came farther apart as the gall quickly elongated along the leaf axis and became darker green. Lar- vae developed quickly and began to leave the galls by June 9 through the newly opened exit aperture on the upper side of the leaf. Mature lar- vae are white and active. Collections of galls —ALaBaAma: Auburn (ovata, tomentosa); Oak Mt. St. Pk, Shelby Co. (tomentosa). FLoripa: Gainesville (tomentosa). GEORGIA: 8 km E Fort Valley (pallida). Lizella (tomentosa). KENTUCKY: Brownsville (tomentosa). MaryLAND: Beltsville, USDA Agricultural Research Center (tomentosa); Brighton (tomentosa); Clarksburg (tomentosa); Gaithersburg (tomentosa); Lusby (tomentosa); Ridgely (cordiformis); Wheaton (tomentosa). Mis- sissipPi: 7 mi E Starkville (tomentosa). Missouri: Cadet (tomentosa, C.V. Riley). New York: Albany (Carya sp.). Onto: Hocking Co. (glabra, Wells 1915: Fig. 29). TENNESSEE: Dickson (tomentosa). VIRGINIA: Rt. 81, mi. 13.9, Washington Co. (ovata). WEST ViR- GINIA: Beech Fork St. Pk, Wayne Co. (tomentosa) Specimens of C. cucumata examined.—Manry- LAND: Clarksburg, V-23-2000, V-26-2004 & VI- 9-2004 (larvae). Missouri: Cadet, VI-8-1890, B.D. Walsh (larva). WEST VIRGINIA: Beech Fork St. Pk, Wayne Co., VII-18-1988 (larva). GENUS GLIASPILOTA GAGNE, NEW GENUS Type-species, Cecidomyia glutinosa Osten Sacken, by present designation. Diagnosis.—This genus belongs to the ce- cidomyline supertribe Lasiopteridi because of the presence of setae on the ventral papillae of the lar- val eighth tergum. This genus is unique for the dissimilar tarsal claws, which may or may not have teeth, regardless of the leg they are on. The third instar is strongly modified for life in a liquid medium and is unique among Lasiopteridi for its dorsal abdominal lobes and the long setae of the dorsal and pleural papillae. Description.—Adult (only female known): Head: Eyes connate. Occiput without dorsal pro- tuberance. Frons with setae. Labrum without se- tae. Labellum separate, hemispherical, setose, and covered with setulae. Palpus 4-segmented. An- tenna: scape short-spheroid, with single horizon- tal row of setae on venter; pedicel short-spheroid, with single horizontal row of setae near base on venter, and scattered distal setae laterally and dor- sally; with 10 flagellomeres (Figs. 343-344): mostly short-pyriform, setose mainly ventrally, ringed with 2 appressed circumfila connected by two lon- gitudinal bands. Thorax: Scutum with 4 longitudi- nal rows of setae mixed with scales. Scutellum with lateral group of setae mixed with scales on each side. Anepimeron setose on dorsal angle, anepisternum setose, pleura otherwise bare. Wing with R; thick, slightly curved anteriorly to join C before wing apex, C broken at juncture with Rs, r- m not apparent. Legs thickly covered with scales; tarsal claws (Fig. 345) rounded beyond midlength, toothed or untoothed; empodia as long as claws; pulvilli nearly half length of claws. Abdomen (Figs. 346-347): First through seventh tergites rec- tangular, each with pair of anterior trichoid sen- silla, single, sparse row of posterior setae, and cov- ered with scattered scales, these sparse on seventh tergite; eighth tergite triangular or trapezoid, with anterior pair of trichoid sensilla and scattered short setae posteriorly and laterally beyond sclero- tized portion; eighth segment ringed posteriorly by single row of short setae. Second through sev- enth sternites rectangular, with anterior pair of tri- choid sensilla, single row posterior setae except seventh sternite with single row of setae at mi- dlength and scattered scales elsewhere. Ovipositor (Fig. 346) short, the protrusible half no longer than twice length of seventh tergite and with setae pres- ent laterally and ventrally; cerci (Fig. 337) fused into single lobe, with 8 peglike setae apically and scattered setae elsewhere; hypoproct long, narrow, with 2 posterior setae. Pupa: Unknown. Larva: Third instar (Figs. 232-234, 349-350).— Body depressed-cylindrical, ends tapering. Head directed anteriorly, with short apodemes. An- tenna less than twice as long as wide. Spatula evi- dent only as short, triangular sclerite. Dorsal papillae 6 on each segment, with long setae, those of first thoracic and eighth abdominal segments set on slightly raised papillae, those of remaining segments set at ends of long lobes: the inner pair on separate triangular lobes, the outer pairs each on a single, broad, apically bifid lobe. Pleural papillae with long setae, as long as dorsals, set on slightly raised lobes. Sternal papillae without se- 94 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) tae. Lateral papillae very small, in two sets on each side of midline, an outer group of 3 papillae, each with seta, two of them smaller than the third, and an inner singlet with short seta. Ventral papil- lae each with short seta. Terminal segment with eight papillae, each with seta somewhat shorter than dorsals, one pair shorter than remaining three. Second instar (Fig. 348): Body cylindrical. Spiracles raised on prominences. Papillae with se- tae as for third instar but setae not appreciably longer than papillar bases and none with lobes. Etymology.—The name Gliaspilota is coined from the Greek glia (= glue) and spilos (= spot), with reference to the sticky leaf spot formed by the only included species. The name is feminine. Affinities —The new genus belongs to the ce- cidomyiine supertribe Lasiopteridi because of the presence of setae on the ventral papillae of the lar- val eighth tergum. The adult female (the male is unknown) is superficially similar to Dasineura ex- cept for its very short ovipositor and undivided eighth tergite. Most Dasineura spp. have a long ovipositor and a longitudinally divided eighth ter- gite. The female of one species, Dasineura rhois Co- quillett, only tentatively assigned to Dasineura, also has a short ovipositor and the eighth tergum shaped as in Gliaspilota, but differs from G. gluti- nosa in having separate cerci. Fused cerci and sepa- rate cerci do not necessarily preclude these two species from belonging in the same genus, inas- much as Oligotrophus (sensu stricto) includes a pair of closely related species, one with fused cerci, the other separate (Sylvén & Tastas-Duque 1993), but the larvae of G. g/utinosa and C. rhois are too differ- ent to consider as close relatives. Third instars of G. glutinosa are unique among Lasiopteridi for their dorsal abdominal lobes and the long setae of the dorsal and pleural papillae. These are charac- ters evidently derived for life in resinous liquid and are analogous to the dorsal lobes seen in some species of Cecidomyia that live immersed in pine resin but belong to the supertribe Cecidomyiidi. The tarsal claws of G. glutinosa are unique among Cecidomyiinae in that the presence of teeth on the tarsal claws follows no rules: the fore, mid, and hind claws may be toothed or not, and both states may be present on separate claws of the same acropod. Larvae of another genus of Lasiopteridi, Ac- ericecis, also live in plant exudate on open galls of leaves of Acer spp. (Gagné 1983, Harris 2004), but that genus shows no close connection to Gliaspi- lota. The wings of Acericecis are very different: its R5 vein is narrow, curved posteriorly on its distal half to join the costa at the wing apex, and no break is present at the junction of R5 and the costa. The female cerci of Acericecis are discrete, those of Gliaspilota fused. Gliaspilota glutinosa (Osten Sacken), new combination Cecidomyia glutinosa Osten Sacken 1862: 193; Felt 1909: 115 (Caryomyia); Gagné 1989: 211 (Cecidomyia); Gag- né 2004: 277, as unplaced species of Cecidomyiinae. Description.—Gall (Figs. 175-177): Occasional on both Eucarya and Apocarya sections, single or several on the underside of a leaflet; a discolored, mustard-yellow, irrregularly ovate spot on the leaf lamina, 5-8 mm in diameter; presence appar- ent from top of leaf only by discoloration, but on bottom of leaf covered with a colorless mucilagi- nous substance. Adult (female only): Head: Eye bridge about 5 facets long; facets circular, closely adjacent except slightly farther apart at midheight of head. An- tenna with 10 flagellomeres, on one specimen the tenth nearly divided into two separate flagellom- eres; third flagellomere as in Figs. 343-344. Tho- rax: Wing length 1.4-1.5 mm (n=3). Tarsal claws (Figs. 345) toothed or untoothed, sometimes one toothed and the other untoothed on same acropod. Abdomen: Sixth segment to cerci, Figs. 346-347. Pupa: Unknown. Larva: Third instar (Figs. 232-234, 349-350).— Yellow to orange. Spatula barely evident, only apical tooth showing, triangular. Dorsal papillae situated on long, flat lobes. Terminal segments as in Figs. Second instar (Fig. 348): Dorsal papillae not situated on lobes. Eighth and terminal abdom- inal segments as in Fig. 350. Etymology.—The name glutinosa is an adjec- tive from the Latin meaning sticky, with reference to the resinous substrate in which the larva lives. Type material.—Syntypes: Three leaf spots char- acteristic of G. glutinosa damage, on leaflet frag- ments of Carya sp., Washington, DC, Osten Sacken, deposited in USNM; larva(e), same data, lost. Biological notes.—Usually one, up to three free-living larvae can be seen in the liquid on each spot on the lower surface of leaflets. After full- RAYMOND J. GAGNE 95 grown larvae drop to the ground, the tissue under the mucilage dries, and the leaf is left with an ir- regular, brown spot. In northeastern United States fresh galls can be found as early as mid-April. Larvae have all dropped to the soil by mid-June. In Florida I found fresh galls in September, so it is possible this species has more than one generation per year there. Collections of galls.—(This species is much more common than apparent from the short list of collections; I did not always note every time I saw the characteristic damage, which, by mid-June ex- cept in Florida, the larvae have already vacated). ARKANSAS: Palestine (ovata). FLORIDA: Gainesville (tomentosa). MARYLAND: Beltsville, USDA Agricul- tural Research Center (glabra, tomentosa); Brighton (tomentosa); Calverton (tomentosa); Gaithersburg (tomentosa); Clarksburg (tomentosa). MASSACHU- sETTS: Great Barrington (cordiformis, ovata). Mis- sourt: Cadet (Carya sp.). New York: Albany (Carya sp.), New York (Carya sp.). Texas: Browntown (illinoiensis). WASHINGTON, DC: unspecified (Carya sp.). West VirGINIA: Falling Waters (cordiformis); 6 mi NW Hedgesville (ovata). Specimens of C. glutinosa examined.—FLoRIDA: Gainesville, IX-21—2000 (larva). MARYLAND: Belts- ville, USDA Agricultural Research Center, V- 30-1993 (larvae), V-22-1994 (larvae), V-28-1994 (larvae; 3 2, em. IV-1995). Missouri: Cadet, VI- 18-1883, B.D. Walsh (larva). New York: Albany, VI-1912, E.P. Felt (larva); New York, VI-20-1907, E.P. Felt (larva). Texas: Browntown, V-13-1982, M.B. Stoetzel (larva). West VirGINIA: Falling Wa- ters, VI-21—-2004 (larvae); 6 mi NW Hedgesville, VII-26—2000 (larvae). GENUS HARMANDIOLA KIEFFER Harmandia Kieffer 1896: 5. Type species, Cecidomyia tremulae Winnertz, by original designation. Preocc. Rochebrune 1882. Harmandiola Skuhrava 1997: 166, new name for Harman- dia Kieffer; Gagné 2004: 163 (catalog). Inclusion of Cecidomyia nucicola Osten Sacken enlarges the scope of this genus that heretofore comprised some gall-formers of poplars and chestnuts (Gagné & Payne 1992). In the key to Nearctic genera of Cecidomyiidae in Gagné (1981), Harmandiola runs to the first half of couplet 103 because the species known up to 1981 had toothed tarsal claws. Now that some species in the genus are known to have untoothed claws, the genus will also run to the first half of couplet 146 with Paradiplosis, except for the host indication. Wings of Harmandiola and Paradiplosis are differ- ent: the costal wing vein is broken at its junction with Rs; in Harmandiola but not in Paradiplosis. Harmandiola nucicola (Osten Sacken), new combination Cecidomyia caryae nucicola Osten Sacken 1870: 53, in- valid polynomial; caryaenucicola Jarvis 1909: 84 (Ce- cidomyia). Cecidomyia nucicola Osten Sacken 1878: 6; Felt 1918: 49 (Caryomyia); Gagné 1973: 868 (Contarinia). Description.—Gall (Figs. 178-180): Rare, so far known only from Eucarya section; a single-cham- bered blister of the nut husk, up to 5mm wide and 4 mm high, with same color and texture as husk; larval chamber ovoid, each with a single larva. Adult (male only): Head: Eyes connate, about 6 facets long at vertex; facets mostly circular. closely adjacent except slightly farther apart at midheight of head. Occiput without dorsal protuberance. An- tenna: scape cylindrical, shorter than wide, with 4 long, apicodorsal setae and 2 mesobasal trichoid setae; pedicel short-spherical, partially ringed with long setae ventrally and laterally; with 12 fla- gellomeres, the twelfth with short elongation to distal node; flagellomeres (Fig. 357) foreshortened, binodal but internodes and necks short, with 3 cir- cumfila, 1 on basal node and 2 on distal node, loops of circumfila subequal in length, short, not attaining bases of next distal circumfilum, setulae on nodes sparse. Frons with 3-4 setae per side. Labrum with short setae on venter. Labellum sepa- rate, nearly hemispherical but blunt at apex, setu- lose, setose. Palpus 4-segmented. Thorax: Scutum with 4 longitudinal rows of setae. Scutellum with lateral group of setae on each side. Pleura hairless except for anepimeron with 9-10 setae. Wing 2.1 mm long; C broken at juncture with Rs; Rs curved apically, joining C posterior to wing apex; Rs de- tectable only as bump on Rs, about equidistant be- tween arculus and R;. Legs about 1.5 times as long as wings; tarsal claws (Fig. 36) untoothed, curved beyond midlength; empodia as long as claws; pul- villi about 1/3 length of claws. Male abdomen: 96 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) First through seventh tergites entire, rectangular, about 5 times as wide as long; with 1 row of setae along posterior margin, another horizontal row on each side near midlength, with anterior pair of tri- choid sensilla, and elsewhere covered with scales, the sixth and seventh tergites each with fewer scales and the seventh tergite with only 2 lateral se- tae; eighth tergite weakly sclerotized, short, ante- rior pair of trichoid sensilla the only vestiture. Ster- nites rectangular, about as wide as long, second through seventh with 1 or 2 horizontal rows of se- tae posteriorly, 3 groups of setae near midlength, these merging on seventh sternite, and 2 closely adjacent trichoid sensilla anteriorly; eighth stern- ite with several rows of setae posteriorly, bare an- teriorly except for 2 widely separated trichoid sen- silla. Genitalia (Figs. 352-355): cerci large, broad for greatest length, each abruptly narrowed near apex to evenly rounded lobe, the lobe setose pe- ripherally and ventrolaterally; hypoproct slightly longer than cerci, linear, completely setulose, the apex bilobed with several apical and ventroapical setae, sides at midlength slightly ventrally curved, with bulbous ventral prominence before mi- dlength; aedeagus elongate-linear, slightly longer than hypoproct; gonocoxite elongate-cylindrical; gonostylus long, evenly setose, the setae longer at base, setulose on basal third, carinate beyond. Pupa: Unknown. Larva: Third instar: Body cylindrical, anterior end tapered, posterior end broad. Head directed anteriorly, with apodemes longer than head cap- sule. Antenna about twice as long as wide. Spat- ula (Fig. 357) with anterior end deeply divided into two roughly equilateral triangular lobes, the lateral sides expanded and weakly erose, and with long posterior shaft. Integument on dorsum without sculpturing, on pleura with pointed ver- rucae anterior to spiracles, and on venter with horizontal rows of sharp spicules anterior to ster- nal papillae. Lateral papillae on each side of mid- line in two triplets, 2 of each group with short seta, the third of slightly larger diameter, without seta. Dorsal and pleural papillae with very short setae. Terminal segment (Fig. 356) mostly covered dorsally with pointed verrucae, with four pairs of papillae, one pair corniform, recurved, one pair situated between recurved pair short-corniform and situated on lobes, two pairs placed laterad, on lobes, with setae short, no longer than wide. Sec- ond instar: (not seen). Type material—Syntypes: Galls on nut husk, from C. ovata, Rhinebeck, NY, VIL-9-1869, Osten Sacken, in USNM; larva(e), same data as galls, lost. Etymology—The name nucicola is a Latin noun meaning nut inhabitant. Affinities —This species is tentatively placed in Harmandiola because its elongate hypoproct has a definite ventral projection near midlength and its aedeagus is long, parallel-sided, and blunt-tipped. Originally in Cecidomyia (Osten Sacken 1870), this species was later placed in Contarinia based on lar- val characters (Gagné 1973). The male, reared sub- sequently, does not fit in Contarinia because of its genitalia and the presence of three circumfilar loops on each flagellomere. The male differs from other species in Harmandiola by the regular- in- stead of irregular-length circumfila loops of the male flagellomeres. Other known congeners have the middle circumfila shorter than the remaining two. Until now, Harmandiola contained 13 species (Gagné 2004); the 11 species with known hosts form galls on poplars and chestnuts (Gagné & Payne 1992). Larvae of this genus could pass for many other genera of Cecidomyiini, including Contarinia and Macrodiplosis, because of the makeup of the papillae of the terminal segment, particularly the large, corniform, recurved pair. Biological notes.—The full-grown larva cuts a straight, cylindrical exit passage to the exterior of the nut husk and falls to the ground where it over- winters. Collections of galls —Georaia: Macon (glabra). MarYLAND: Silver Spring (tomentosa). NEw YORK: New York (ovata); Rhinebeck (ovata). ONTARIO: vic. Guelph (Jarvis 1909, ovata). Specimens of C. nucicola examined.—GEORGIA: Macon, VI-2-1986, D. Ring (larvae). MARYLAND: Silver Spring, VII-2-1985, (larvae), VII-3-1992 (6, em. V-1993). New York: New York, I-23-1884, E.B. Southwick (larva); Rhinebeck, VIII-9-1869, C. Os- ten Sacken (larva, excised from syntype gall). GENUS PARALLELODIPLOSIS RUBSAAMEN Parallelodiplosis Rtiibsaamen 1910: 297. Type species, Diplosis galliperda Low, monotypy; Gagné 2004: 217 (catalog). This genus of 22 species is widely distributed but mainly Holarctic and includes inquilines in galls of insects from various orders (Gagné 2004). RAYMOND J. GAGNE 97 Parallelodiplosis caryae (Felt) Cecidomyia caryae Felt 1907: 45; Felt 1908: 411 (Clinodiplo- sis); Felt 1921: 168 (Parallelodiplosis). Description.—Adult: Head: Eyes connate, 11-12 facets long at vertex; facets hexagonoid, all closely adjacent. Occiput with dorsal protuber- ance with 2 apical setae. Frons with 6-10 setae. La- bellum ellipsoid, pointed apically, with several lat- eral setae. Palpus 4-segmented, with prominent palpiger. Male antennal flagellomeres binodal, distal node of each constricted at basal third; dis- talmost flagellomere with elongate, constricted terminus; 1 circumfilum present on basal node of each flagellomere, 2 on distal; loops of circumfila subequal in length, long, but not quite reaching next distal circumfilum. Female flagellomeres suc- cessively shorter from base to apex of antenna, the 6 first flagellomeres constricted and elongated be- tween basal setae and basal band of circumfilum, remaining 6 flagellomeres evenly cylindrical, the last with elongate, constricted terminus; each fla- gellomere surrounded by two appressed circum- fila connected by two longitudinal bands. Thorax: Wing unmarked, 2.3 to 2.9 mm long, Rs; strongly curved toward apex, joining C posterior to wing apex. Tarsal claws (Fig. 360) untoothed, bent be- yond midlength; empodia reaching only about half distance to bend in claws. Male abdomen: Third through sixth tergites horizontally divided laterad between posterior and lateral setae, with anterior pair of trichoid sensilla, posterior setae continuous across sclerite. Seventh tergite similar in shape, but some specimens lack lateral setae and posterior row of setae discontinuous mesally; eighth tergite weakly pigmented, the only vesti- ture the anterior pair of trichoid sensilla. First through eighth sternites pigmented only on poste- rior two-thirds, anterior pair of papillae situated on basal, membranous third, setae present along posterior margin and covering midlength of scle- rite. Genitalia (Figs. 361): cerci with posterior mar- gin straight but slightly longer on lateral than mesal angle, with setae along posterior margin; hypoproct much longer than cerci, parallel-sided, nearly as wide as aedeagus, with 4-6 setae on pos- terior margin, and covered with long setulae on both dorsum and venter; aedeagus elongate, longer and slightly wider than hypoproct, distal margin convex; gonocoxite elongate-cylindrical, splayed; gonostylus elongate-cylindrical, with se- tulae near base and covered beyond with minute carinae and widely scattered short setae. Female abdomen: First through sixth tergites as for male; seventh tergite undivided, longer than sixth; eighth tergite unpigmented with anterior pair of trichoid sensilla and posterior row of weak setae. First through sixth sternites as in male; seventh sternite fully pigmented, narrower and longer than sixth; eighth sternite unpigmented, with an- terior pair of trichoid sensilla and scattered, short setae posteriorly. Ovipositor (Fig. 362) slightly protrusible, venter but not dorsum of ninth seg- ment with setae, cerci elongate-ovoid, with pair of apical peglike setae and scattered setae elsewhere; hypoproct short, narrow, with 2 short, posterior setae. Pupa: Head: Antennal base rounded apically, not conspicuously projecting anteriorly; cervical sclerite with two elongate setae; face without ven- tral projections, with 2 pairs of long-setose papil- lae, one on frons, one on labrum. Prothoracic spir- acle elongate, pointed apically. Abdomen evenly covered with uniformly small, pointed spicules, none enlarged and spiniform. Larva: Third instar (Fig. 48): Length 2.5-2.9 mm. Orange. Integument covered with mostly pointed verrucae. Antenna less than twice as long as wide. Spatula (Fig. 359) elongate, parallel- sided, with 2 rounded anterior teeth. Lateral tho- racic papillae in 2 groups of 3 on each side of cen- tral line, 2 papillae in each group with tiny seta, the third without. Dorsal and pleural papillae with elongate setae. Terminal segment (Fig. 358) with 8 papillae as follows: 1 pair large- corniform, anteriorly recurved, pigmented; 1 pair with long setae, as long as dorsal setae of previous segment; 2 pairs with very short setae not surpassing width of socket but setae of outer pair slightly the larger. Second instar (not seen). Type material.—Holotype: Male, taken on hick- ory, Albany, New York, VI-19-1906, Felt #331, de- posited in New York State Museum, Albany. Remarks.—The first published mention of this species was by Osten Sacken (1862). He wrote in his discussion under C. holotricha that he had found in some galls a pale orange larva with the anterior of the spatula ending in two triangular points. Larvae of Caryomyia species are white, so the orange color would describe a Parallelodiplosis larva (Fig. 48), and the forked spatula would pre- 98 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) clude it from belonging to Lestodiplosis. Felt (1907) based his initial description on a male caught on hickory, and his subsequent, fuller description (Felt 1921) on that male and also of a female caught at the same time as the male. He later ob- tained adults reared by a colleague from Cary- omyia inanis galls (Felt 1908, 1921) and eventually succeeded in rearing some from at least Caryomyia aggregata galls (Felt 1921). The species has to date been associated with galls of ten species of Cary- omyia. The larva and pupa are described here for the first time and the adults redescribed in full. Affinities.—Of the described North American species in this genus, only one other, Parallelodiplo- sis acernea (Felt), has tarsal claws with a similar slight curve beyond midlength. Both species have generally similar male genitalia, but the hy- poproct of P. acernea is definitely narrower with relation to the aedeagus than is that of P. caryae. The basal flagellomeres of the female antenna are regularly cylindrical in P. acernea rather than greatly constricted as in P. caryae. Larvae of the two species appear identical. Interestingly, P. acernea is also an inquiline but in galls of Pachypsylla spp. (Hemiptera: Psyllidae) and possibly also Celticecis sp. nr. spiniformis (Ce- cidomyiidae) on hackberry, Celtis occidentalis L. (Moser 1965). First instars of P. acernea evidently bore into galls and enter cavities occupied by psyllid nymphs; nymphs of the psyllid always die after larvae of P. acernea reach them, possibly be- cause of competition for food (Moser 1965). Biological notes.—Parallelodiplosis caryae has now been found inside or reared from galls of 10 species of Caryomyia (see list of collections below). The very active, orange larvae can occasionally be seen in September and October inside Caryomyia galls (Fig. 48). On two occasions, once in Mary- land, once in Massachusetts, I found third instars crawling on a leaf in the vicinity of C. holotricha galls. It is possible they had recently exited galls and were on their way to drop to the soil, al- though leaving the galls does not appear neces- sary because on several occasions I reared adults in spring from Caryomyia galls kept through win- ter. When P. caryae larvae are found inside galls, the Caryomyia larvae are missing or dead and the surface of the larval chamber appears to be scraped, the chamber littered with the resulting particles. Access into the galls is evidently through a hole at the base or side of a gall, but I do not know precisely what makes the hole; it might have been caused by rot beneath a gall in which the gallmaker was already dead or possibly by some boring insect. A gall of C. viscidolium found in Piney, AR, had a small circular hole in its side. In another case, a predaceous Lestodiplosis larva also had presumably entered through a hole at the base of a gall of C. tuberidolium and was feeding on the P. caryae larva (Fig. 48). Collections of galls and specimens of P. caryae examined.—ARKANSAS: Beulah (tomentosa), in gall of C. tuberidolium, IX-10-2004, larva; Chester (tex- ana), in gall of C. holotricha, IX-9-2004, larva; Piney (texana), in gall of C. hirtiglobus, X-10—2004, larva. GeorGIA: Forsyth (ovata), in galls of C. holotricha, IX—22-2000, larvae. MARYLAND: Beltsville, USDA Agricultural Research Center (tomentosa), VILI- 22-2000, larvae on leaf among galls of C. holotricha, and found in and reared from galls of C. sanguino- lenta and C. caryaecola, X-2002, 2, em. IV—2003; Clarksburg (tomentosa), in gall of C. tuberidolium (Fig. 48); Ellicott City (tomentosa), in gall of C. pur- purea, IX-2-2000, larvae; Ridgely (cordiformis), in galls of C. tubicola, X-2-2000, 3, 2, pupae, em. IV- 2001; and (tomentosa) in galls of C. purpurea, X- 2-2000, larva. MASSACHUSETTS: Petersham (ovata), on leaf among galls of C. holotricha, IX—6-2000, lar- vae; Worcester (Carya sp.) ex galls of C. inanis, M.T. Thompson, 2 6, pupae. New York: Albany (Carya sp.), VI-19-1906, E.P. Felt, 5 (holotype), 2; Nassau (Carya sp.) ex galls of C. holotricha, VII-29-1906, E.P. Felt, larva; New York City (Carya sp.), ex galls of C. aggregata (reported as C. persicoides in Felt 1921: 169), E.P. Felt, 2 2. Texas: New Waverly (tomen- tosa), ex galls of C. leviglobus, X-6-2001, 2 3, em. IxX-2002. RAYMOND J. GAGNE 99 KEY TO HICKORY GALLS CAUSED BY CECIDOMYIIDAE Caryomyia galls are best keyed still attached to their leaflets because their position on the lamina or leaf and any surrounding leaf exfoliation are useful characters. Some galls may need to be cut open in order to see their internal structure. For this key as well as in the illustrations, Caryomyia galls are ori- ented apex upwards without regard to which leaf surface they occur on. — Phylloxerids (Hemiptera: Phylloxeridae: Phylloxera) and eriophyids (Acarina: Eriophyidae) also form galls on hickories but their galls are distinct from those made by gall midges. Phylloxera galls (Figs. 181-182) contain aphidlike insects, often in large numbers. Mite galls (Figs. 1, la, 2) have no broadly open chamber but are instead made up of a series of tight wrinkles that enfold the many mi- nuscule mites. Closed cecidomyiid galls on hickory will have one or more ovoid chambers, each con- taining a legless larva with a very small but definite head capsule. 9(8). 10(8). 11(10). 12(6). 13(12). 14(13). 15(13). 16(15). 17(16). 18(17). 19(18). Slickysleatispot withiexposed larval(Figs: 179-177) s).0. 3.15 ects oysjasljoce eos ns et oe Gliaspilota glutinosa pimple orcomplex leafor nut husk gall with enclosed larva <2. . 2.5.2.2 -.0.2- sees sss: 2 Gall a simple swelling of petiole, leaflet, or nut husk (Figs. 165-171, 178-180) ....................... 3 Callkcomplexonileaflet(Figs;o—32; SOS MOA) wv. tery ct coed) ole ap ie oes ASN elo cise cic nas ciaiasbedas eae lctsredsnswotiogs 6 Iemisphericaliswelling of nut husk (Figs. 178=180) oo. ere. oe eee et ee Harmandiola nucicola Swellineronperoleonleatiet{ (Eigsil GS-17) ie vy waver scu Mere terctsrene) sounkeusie) se ysKs ain in leyeye ceed sislsia'cLoyo¥sl.o © sucyoke 4 Circular, biconvex swelling of lamina (Figs. 33, 167-169) ...................... Caryadiplosis biconvexa SwvellingronperOleiormleak Velm) (ig Sel OS 1G, lA O17.) aes age seh) hol eas o05) 5 slp\ osha =) once 8 sashes feisaod BT) 3) 34: Fok 5 Hard, elongate swelling of petiole or leaf vein (Figs. 165-166) .................... Caryadiplosis venicola Soft, bubblelike swelling of lamina, known only from pecan (Figs. 170-171) ....... Contarinia bulliformis Recumbent, main axis parallel or nearly parallel to leaflet (Figs. 88-97, 172-173) ................2205. 7 iRerpendicular to leaflet or, only slightly leaning (Figs.36, 145, 147) 20 oo... nee ee ety 12 Apex convex; exit apparent on opposite leaf surface; indehiscent (Figs. 29, 172-173) . . . Contarinia cucumata Apex pointed; without apparent exit hole on opposite leaf surface; dehiscent (Figs. 88-97) ........... 8 NOT LVvEnAnrOWeGIDelOLela PeXG (hi gS yOGe Il) ee wa i ERM hecho p ACs Sis of Sfeuedsieush sa cyeieishsi sie ees) 9 Cradtially;marrowed from base tojapex (EIS 929. ysey are vehi «eho te kf) ope obeys aK hse eee osheel sles 10 Sunfaceroughy with sparse Nain (Figs) 9a,, 68-39)! wei persis rrels ici cisiehisiese cashiers 5 6 <1 Caryomyia eumaris Surtacesmoothnainless (Hiss 19,90 —9il ie ee cee srye somere ie tus ein le ete sae ean os Caryomyia recurvata Hairless; base not bulbous, flattened against leaf surface (Figs. 96-97) ................ Caryomyia supina Rainy Abaseib ul bOUS) (ETO S 502-95) ape acer seo eee E NR ool oia ees uP agock Roto hal cgeuckcuo iGo atope\ouo. « SioleBsia os nts 11 amlong, crinkly, (Figs: 92-93) om bittermUtor rasp) jc iin iors locas loss: Caryomyia spinulosa Hlainshort, straight (Figs: 94-95) (on Eucaryd SeCHOM) 9 oj. yo sei oe oi eke ss os Caryomyia procumbens Surtacenainy srhemalrstniclOr sparse\(HIESiO9 OM) essen to Faye estal sic) coe alelae, oksio) s evarn« ¥aloysi gece cealefayspoasl eens 13 Surface hairless except for possible tuft of sparse short hair at extreme apex (Figs. 41,43) ........... cy With partial or complete false chamber above larval chamber (Figs. 99,105) .............---.20005- 14 Withoutsralse: chamber (igs 740) 50) Maer yece pecans Leg skstchcroysts cic dais Sf5\0) foo bie 5 fuse) ngs saotey ip efehena egos Ke) oasal 15 Depressed spheroid, densely covered with flat, decumbent hair; false chamber closed (Figs. 104— HOS) Weenene eet mae eR A eee Yo oses hs est ce I aa yee cacce de dn yaye oa hoa, ilo ol sunjn'dageloiue colonels Caryomyia tumida Upstanding, sparsely covered with erect hair; false chamber open apically (Figs. 8, 98-99) ............. Soe bo DOGS OO TA CSTE STOWE S oO DIOG ONE Bale Hs on cee ar eanints ke Dee rT SSC oc Caryomyjia inanis lattened discoid stellatel(Rigss 32, lOO VOM) Me oie rcpets cies eicte spers eels o)on8' 95) oye ausiensl shots Caryomyia asteris Flongateorspheroidalynotstellatel(Higs339) 64) act cicyiels) iar <1. al ol stoke ekepensioes ous teetogsis reverence ckansteVe 16 Wall almost uniformly thin, brittle, gall of same shape as larval chamber (Figs. 40,65) .............. 17 Wall thick, soft or woody, extraneous to shape of ovoid larval chamber (Figs. 132,159) ............. 23 Elongate-conical, gradually narrowed from base to apex; hair as long as gall diameter; dehiscing in spring GHEE Se) ipa Sisal ado Oa oe Om OOO a NER ee OO OnE E In Om a nEOG aoe aor Caryomyjia albipilosa Spheroidal; hair not half as long as gall diameter; dehiscing in summer/fall (Figs. 49,53) ........... 18 Apex with dimple; larval chamber with intrusion of hair from apex (Figs. 23, 112-115) ................ seca tsi s eteusbere ern ene teh aevet tobe ore eee can eee vase in teme enets sie) nioeetr awe Grae dy ela ogs cu Supe testes Caryomyia thompsoni Apex without dimple; larval chamber without intrusion of hair from apex (Figs.50,54 ............. 19 Rlarmthickyobseunnpisuntace (bios 49— 50) Meme vate Cer barratry seal tier ere cusp eee Iker nate Noite a 20 100 20(19). 21(19). 22(21). 23(16). 24(23). 25(24). 26(25). 27(26). 28(26). 29(28). 30(29). 31(30). 32(28). 33(32). 34(32). 35(34). 36(35). 37(12). 38(37). 39(38). 40(42). THE GALL MIDGES (DIPTERA: CECIDOMYIHDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Hair sparse, motiobscuring surtace\(Figs.ol—52)) iia. 4-.eaierie ine eras ieee ee 21 Tuft of hair at apex thicker and longer than remaining hair, tuft covering circular, apical, thin areola (Figs. Fao] )) ESOC CREE POOIE en PE ROC Oe OCA Oso Oe oo On ooo OraE haa o goes Caryomyjia hirtidolium Hair uniformly distributed; surface texture uniform (Figs. 53-54) ................ Caryomyjia hirtiglobus Base with deep, circular central impression; apex with circular, thin areola surrounding nipple; on bitter- MULE PISSTOUKOZ) ta erccs eae e ary elope tee sce ay otclaterets oar ateas entanee! ce Paces ete ete eee eee Caryomyia cilidolium Base conical at center; apical texture uniform except for nipple; on Eucarya section (Figs. 64-67) ...... 22 With prominent apical nipple; hair brown, stiff, erect (Figs. 64-65) .............. Caryomyia spiniglobus With slight apical nipple; hair white, soft, decumbent (Figs. 66-67) ................ Caryomyia deflexipili Larval chamber with conspicuous tuft of hair intruding apically (Figs. 116-117) ....Caryomyia holotricha Larval chamber without intrusion of hair igs! 124-125)132=133)) chs .ae aes ae eee 24 Obconic, apex flattened; hair appreciably thicker and darker at apex than elsewhere (Figs. 123- LOPS) pera eRe ae RSE SEE rE BOS EO EER ERA at RU MPMREIRT SRSA ences cA RIEALR DCMU Rr AAR R era RAR ocd C8 Bs Caryomyia turbinata Spheroidal or widest near base or middle; hair uniformly distributed throughout or thicker at base (Figs. WS 2—133, VO7SRAO) ico, sah cobs ay acoks Sceyare 6 Race, 545k psa) ayecncd nade oa se reas Gusts aR 25 On upper leaf surface; bulbous base smooth and hairy, conical apex bumpy and nearly hairless (Figs. POZA UDS) ssc n chess ayeca) coe eles Sony sats cle oh ore ge acvon yaclcsia aes wea uel Sa ia ofl cies a ey ee ea Caryomyia turbanella On lower leaf surface; not conspicuously bumpy apically, uniformly hairy throughout (Figs. 137-140, A a A 0) Dieser iE esha el Ue ES ie en Lunes intel Civiy a omdro-0/6 0 0 0 0.0.0 00 26 Surface smooth, easily visible through sparse hair (Figs. 137,139) ................-0.0-2se ese o-ee 27 Surface rough and/or covered with resinous dots, the hair usually thick (Figs. 145, 147, 152)......... 28 Spheroidaliatbase/becomingiconicall (Figss137—138) avin... cisa terete ee Caryomyia conoidea Discoid at base, narrowed abruptly beyond (Figs. 139-140) ...................... Caryomyia marginata Narrowed abruptly from large base; apex digitate (Figs. 145,147) 22... 0)... aie see cee 29 Spheroidal or conical and narrowed gradually; apex not digitate (Figs. 154-158) ................... 32 Base:variously, lobed (Bigs. T45—146) ie ss ssi: peeps neyotest geusdoush Cael seksi ache ee COE Caryomyia inclinata Base evenly spherical (Figs.147,,149) 152) io cc.6 can. sreysnsitie nyetein si gcue 0, o-nieye ie io oii eee 30 On lamina; some hair stiff and as long as gall diameter (Figs. 14, 152-153) ........... Caryomya echinata On or adjacent to veins; hair bowed, less than half gall diameter (Figs. 147, 149) .................... 31 Surface with dark, resinous dots; on Eucarya section (Figs. 149-150) .............. Caryomyia cucurbitata Surface without resinous dots; on bitternut (Figs. 147-148) .................... Caryomyia ansericollum Surface’ visible:through hair (Figs 154/158) 2. re he ere wise to nce os sy ot acne eee eae 33 Surface not visible through hair (Figs. 121,156) 2.2200. es se oe | eee eee 34 Base with central hemispherical connection to leaf; surface smooth, hair extensive (Figs. 158- 1 Leh) PASI nea a Ann Ree Laie Died ieee Nay A aR itne re cianinl aN mote hBler a tec5.0 b'5 tho Caryomyia antennata Base with circular central impression; surface rough, covered with resinous dots, hair sparse (Figs. PSFK NSS) ove ouneteainairpayake Myce tyats aera taec spel abet: sce ah opr au egaegotsas, ako otal eye ates ee eee Caryomyia guttata On lamina; base with deep, circular, central impression; larval chamber usually purple (Figs. 15, 121-122) sci ulislagatel(ahin'tua pgm sy o' Spe) Slane usnbas Susgsy seme vee ash. Aree ean te fede ac Es PRES Lees shee gh gee Caryomyia purpurea On or closely adjacent to leaf vein; base without central depression; larval chamber green or brown (Figs. 126-129) 156-157, W60-16L) ose aces as olcle idle «dr elerive casas cueuere, o1a/oyes. «ove 2 «lo 3c ee eee 35 Unevenly spheroidal, usually found closely compressed into groups; larval chamber irregularly ovoid (Figs 21a2iby SOT 5 7) as tere rset sites a. ayss ie earstons onaeiaee Ore cr eaet stay sparse na Caryomyia aggregata Spherical’ to:conical, always discrete; larval chamber ovoid 2. 2 i)020 ee wee oss eckson 36 Spherical, woody; larval chamber at center of gall (Figs. 21, 160-161) ............6.. Caryomyjia spherica Spheroidal or conical, soft, the top portion eventually shrinking somewhat; larval chamber at base of gall leaf easts IZA ALAC IPAS) ss iad BOHASS Sic. ciisl Os hin RAMS G05 GiGi hak Soke lel hoo ION Bipot bs Caryomyjia persicoides Surface sticky to touchy (ysis sn ses gae aie ushers c eioungore. 61a eve oily eid ciara sverela ove asst eee eee 38 Surface NOE SHC Ye ese ese esse aps os cco -aberaneth s the iasar oh agecdiand taps lube el ela yahagelseeet isn ska ae eee eile 44 sheathed at base by conspicuous leaf exfoliation (Figs.83, 107): 2... Jochen. 3 yo ce ene eke 39 Not sheathed at base by conspicuous leaf exfoliation (Figs. 41,55) ........ 0. cece e cece eens 40 Spherical, rubbery; larval chamber surrounded laterally and above by large, false chamber (Figs. 5, 11 Ser L070 SMa ses NN ta URN ag iA rr es SA ME ve, oS Cah er sonic OS Caryomyia lenta Short-cylindrical, hard; without false chamber (Figs. 30, 82-83) ................ Caryomyia melicrustum With'apical nipple surrounded by large areola’ (Figs. 43,47) oo... 12 21 2 minpes eleye 0 oon «oo eileen 41 Surface uniform or apex concave or with simple umbo (Figs. 41, 55, 78) «2.0.6... cece ee eee eee eee 42 41(40). 42(40). 43(42). 44(37). 45(44). 46(45). 47(44). 48(47). 49(48). 50(48). 51(47). 52(51). 53(52). 54(52). 55(52). 56(55). 57(55). 58(57). 59(58). 60(59). 61(57). RAYMOND J. GAGNE 101 Surface bumpy, conspicuously covered with resin glands; usually on under surface of leaflet (Figs. 46— AS) rer sea ven te peeve te eiese sy Wl, SNE Ty teh his Contra Seo staal. oeeare We a way enya aul Caryomyjia tuberidolium Surface smooth, with sparse resin glands; on upper surface of leaflet (Figs. 25,43-45) ...............0.. 910 00057900404 DEN tao OT ONG FO Ob EO GO n Utp Oa eee IOn Dis oCOnr nero ORen Erma Caryomyia viscidolium Surface bumpy; base with deep, circular, central impression (Figs. 118-119) ....... Caryomyjia tuberculata Surface smooth except for minute resin glands; base conical (Figs. 55-56, 78-80) ................--- 43 Spheroidal, apex convex; surface smooth, hard, brittle (Figs. 18, 20-20a, 55-58) ....... Caryomyia caryae Short-cylindrical, apex concave; corona surrounding concavity soft at first, later contracting and forming rough, browMErUst(EigswlGat7C—Sll) irk jy easy wer seyoc afl Resell tel viele etelele eee Caryomyia flaticrustum With\false|chamberiabovellarvalchamber|(Figss102>103/ 108-191) -20)2.. sti eae eisse.. cot. wae 45 WWithoutifalsechamiben demmcrishy tain mysinrretiely eiritentontanieletcitrs asi inilsn mere aay ik heiph ava onal 47 Nearly three times as long as wide, sheathed at base by leaf exfoliation; false chamber open at apex (Figs. Hepa S— N09) arcs yaks ayhecsy shay. clay avage (ote casPahee.h achcriy weclws ystlegs’ Aereheh te weal atisoptianel tbh» Sa Caryomyia caminata No more than twice as long as wide, without leaf extension surrounding base; false chamber closed (Figs. LOZ SUNOS re OU) paca scorers tslueh sci Mey yap areas ys’ ACh eestooy st Aasto ss Ess ls ty weenie alee’ sildgascer elas est eeioIeNeNe Aly 46 Texture rubbery; about as wide as long; with umbo at apex (Figs. 5a, 102-103) ........ Caryomyia inflata Woody; about twice as high as wide; without umbo at apex (Figs. 110-111) ............ Caryomyia ovalis Elongate-cylindrical, usually at least twice as long as wide (Figs. 37, 41, 72,75) ...............0000. 48 spheroidal orconical,aboutiasjlongiasiwide (Figs: 59/62) ..2 2) taissystyey palsies ispsiel os loieielt yt - 51 Base with shallowly concave, circular, central impression, not set into leaf socket (Figs. 37-38, 41- AD) ANE VEY pie tet tT A kate ny oh ystems st aac hs Gps VOM, Sleiilaode dies ie aids S aisbdiste’ eae Heaney 49 Base conical or deeply recessed, set into surrounding leaf socket (Figs. 72-77) .............-..0005. 50 SUnracemunutelysiiateapexpointed (Figs. 36-38)! . 2... 26.5505. eessce sss. ee ee Caryomyia striolata Surface smooth; apex blunt, with umbo at center (Figs. 28, 41-42) ................... Caryomyia urnula Evenlycylindnicalbaseiconicall(Eigse26),/4—77)) sas mete far =) 222) fishes avsiee oo ie oo Caryomyia tubicola Tapered from base to pointed apex; base deeply recessed (Figs. 27, 72-73) .......... Caryomyia subulata Wall uniformly thin, of same shape as larval chamber (Figs. 60,63) ............... 000 e eee e eee eee o2 Wall thick, extraneous to shape of larval chamber (Figs. 135, 164) ....................2....00 000s 55 Spherical, with umbo or nipple at apex (Figs. 62,68) ...................0.. ts fepetg gs ae sesh 53 Spueroidal marrowedibeyondymidlengthy)(Figs759)770) ja. % sec eects cis aie ishoyee ofepehe eye) fetes “ciel 54 Surface minutely striate, green; with prominent nipple at apex (Figs. 19, 62-63) ....Caryomyia leviglobus Surface smooth, glaucous; with small umbo at apex (Figs. 68-69) .............. Caryomyia glauciglobus Abruptly narrowed beyond midlength to blunt apex; on lamina (Figs. 59-61) ......... Caryomyia shmoo Gradually narrowed beyond midlength to pointed apex; on vein (Figs. 12-13, 70- TW" ya Eos AHO. OBO NOOO POO ATCO Oe OOO OA OAT RCD ee OOS aM nes Se oe Caryomyia caryaecola Short-cylindrical, apex flat to concave; base sheathed by leaf exfoliation (Figs. 84-87) .............. 56 Conical to spheroidal; base not sheathed by leaf exfoliation (Figs. 135, 163) ..................20005. BY/ Surface minutely striate laterally, rugose apically (Figs. 31, 84-85) ............ Caryomyia striolacrustum Slitiia ceIsmMOOLUK(FISShSO—87) arc romeerisly. bic breuirae mulitSveb dl stejalanraoeyecaiils- Bane lh dligieas Caryomyia levicrustum Conicall(RigswlS0 134 Mle WAS) ieey cits cicneloriaye.oleleretiils sone (oy 2 aie aiuiasisishete oo.s peed die gee dest ne keeles 58 Splheroidali(Bis si AA G3) dix. oascud copes tesla rccsy atta itis cry skeyelos, opiwustel ch ofatorabe eresolens loa. o.shs aces sae 61 Surface covered with conspicuous bumps (Figs. 130-131) .................0..ee0e. Caryomyia glebosa Surfaceismoothiexcept forlongitudinal furrows (Figs. 134, 141)) occ co. cee yee ee 59 Base curved upwards, apex set in resulting concavity (Figs. 141-142) ................ Caryomyia arcuata Baseyparallelstopeata(kigss (S41 Ag) iy ka cata users alec spe csp soe efsuexesy cheney vice voueyaust oy acus va, Rope eweoe este 60 Base expanded horizontally, usually stellate; cone elongate-attenuate (Figs. 11a, 143) ... Caryomyia stellata Broadlysconical\ (Ries slO 34136) are cteyar ticle eek. eleisrsve cin cis + ais ois ous aisles isle a Caryomyia sanguinolenta Spheroidal, attached to vein; multi-chambered (Figs. 162-164) .................... Caryomyjia cynipsea Depressed-spheroidal, on lamina; single chambered (Figs. 16, 144). ................. Caryomyia biretta 102 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) KEY TO MATURE LARVAE OF CECIDOMYTIDAE ON HICKORIES Mature (third instar) cecidomyiid larvae can usually be characterized by the presence of a spatula, a sclerotized dermal structure on the venter of the prothorax (Fig. 269). This structure usually serves to separate a third instar from a second instar, which usually has none. There are, however, exceptions, in- cluding Caryomyia, in which the second instar has secondarily developed a spatula. A couplet is added here that will separate those two instars of Caryomyia. The key to galls is still the best way to key Cary- omyia spp. because galls are distinctive, unlike larvae of some species that can only be keyed to group. This key to larvae nonetheless points out those differences that do occur among species of Caryomyia and serves as validation when used in combination with the gall key. 4(3). 5(4). 6(5). 7(6). 8(7). 10(1). 11(10). 12(11). Terminal abdominal segment with 6-8 papillae each bearing a seta or short, thick corniform structure (Figs, 334; 348) corinne scacs aielules Siang tious duie alata) cueue sora age avele apel clots: ete et ae 2 Terminal abdominal segment with 8 flat or domed papillae (Fig. 271), none with seta or corniform struc- CURE; SEMNUS|CAPYOMIYIA noc een close as oiaya maces sn ulahe lls ie odegeusts uepetels Gets e;aae' eS eee eRe eee 10 Dorsum with conspicuous pseudopods (Figs: 232,350)" ov.) unt ssc elo. 3 ree Gliaspilota glutinosa Dorsum without pseudopods coco) oie oo oles te ses w sue seniare oi cys Segakeituere 0/0 ws 97s. eee Cee 3 Antennae more than 3 times as long as wide; venter with pseudopods; anus dorsal .................. ce NRA AMEND Jar ell SU ular atone lec Ru RUN sina Lestodiplosis spp., not keyed further, see p. 9 Antennae no more than twice as long as wide (Fig. 228); venter without pseudopods; anus ventral (Fig. DIY) oiesuie ests sisia alsiocel ey okslunals sie aneuistanregats « eersiolt's ele ebeegae lire Si) 4s tacit RCE eee 4 Terminal abdominal segment with papillae all similarly setose (as in Fig. 348) ....................05- wis ole’ Jani oo! AUS6e Gy aia soen sepa RSOD SV ononaal staaiec Aled Mad kay ene taua patos Dasineura sp., not keyed further, see p. 3 Terminal abdominal segment with dissimilar papillae, at least one pair corniform (Figs. 334,358) ..... 5 Terminal abdominal segment with 1 pair of papillae with long setae, 2 pairs with short setae, and one pair of large, recurved, corniform papillae\(Fig. 358) ooo ..c cece sos ese ate seers Parallelodiplosis caryae Terminal abdominal segment with 3 pairs of short, setiform papillae and 1 pair of corniform papillae (Figs.'334, 356) ios. jreia otis ageusve sys 8 oie una) aacue Menemensnast sa atare S eloyslavelts esse ae nee pee eer 6 Terminal segment completely smooth, without verrucae, the 3 pairs of setiform papillae not situated on lobes (Eig 334) eee Nore ee ne cl yer custiera tata ued Sera waMeee ct ageL sy eMe Ete Ne ICSE tee fla Seats Renee gee Caryadiplosis venicola Terminal segment with some verrucae, the 3 pairs of setiform papillae situated on lobes (Figs 336, BOG) aoe eae a eaueleamacsse ee mestaiele Maeno sttietene iltyed cure miintevergeeierd eseksre tern fcc) sls fps noe oe ae 7 Spatula with bilaterally symmetrical teeth (Figs. 337, 339) ooo asso. S class otic oto poe eRe RSE 8 Spatula with serrated, laterally lobed teeth’ (Figs. 342) 357 ic. oo ciiuce tute ote ee cee ei Ren eR eee 9 Spatula teeth triangular (Fig. 339); terminal segment with verrucae between corniform papillae (Fig. S16 fo} URR ema RED ute CALE Mc AAT ae Aras 8 ae ies hic a a ee AN Contarinia cucumata Spatula teeth rounded (Fig. 337); terminal segment without verrucae between corniform papillae (Fig. BBO) sorseny te todo seqaiet ls eusss eeu camveriatee Sketieeeehe Me TG Vets sence a eae SEAS Gator ecne eee eae Caryadiplosis biconvexa Setae of setiform terminal papillae mostly conspicuous, two pairs appreciably longer than third pair (Fig. BAT) 2, orayan : ecauaue acetone si ceeehes uae ceneteti orci Reece cater cea En Ce te Ren erect ee eee Contarinia bulliformis Setae of setiform terminal papillae barely longer than sockets, all of approximately same length (Fig. BOG) cei cicio skit ay cde oss RENT eon ne RED te AES Sc EY MC OP REPRE enc d iage nears Dane Ate ct ce Harmandiola nucicola Collar segment with group of conspicuously enlarged spicules on each side (Fig. 273); tooth of spatula acutely triangular, usually half as long as rectangular shaft (Fig. 273)... 2.2... cece eee eee eee ee eee ES AR oa ea eae areca rane eee Wn re Abs Bales al hia (8 Caryomyia, second instars; not keyed further Collar segment either without spicules (Fig. 269) or spicules uniform in size (Fig. 228); anterior tooth of spatula, if acutely triangular, usually much less than one third as long as spatula (Figs. 316-327); Cary- OMY IA, EMI! IAS EATS Is mtieveics.ohe.cosarsyeushon ceri dswontieteuckeva toy ome venenoRa ahi was CaUske BRAD CRO ER Rare ACR RAE AC ROCK eee 11 Spatula with 2 separate teeth (Figs. 274-275, 282-285) or tooth parallel-sided and notched at apex, (Figs. 28O= 289 29 Tro isvasage voll ocauil aOshdei seo ove tenty citcls Ma Niamey ala caOhie aisle ois t/a bars eae aera a ceenRGT aero a ane eee 12 spatula with single triangular tooth (Figs. 276-281, 290)) or oe ss lecleit ca eterdiessiclels tle e oreiy sienna 25 Spatula teeth separated to shaft, occasionally with denticles between (Figs. 274-275, 282-285) ....... 13 Spatula tooth more or less parallel-sided with notch at apex (Figs. 286-289, 291) ..............e 00s AW, 13(12). 14(13). 15(14). 16(15). 17(12). 18(17). 19(17). 20(17). 21(20). 22(21). 23(22). 24(20). 25(11). 26(25). 27(26). 28(27). 29(28). 30(25). 31(30). 32(31). RAYMOND J. GAGNE 103 Spatula teeth equilateral, the space between with one or more denticles (Figs. 274-275); abdomen with ex- tensivielsmoothtareas\ (Rigsw2lo— 2,7) ew isrvers sie ert taster cts ey. tet Aer ie ran eens C. albipilosa, C. striolata Spatula teeth appreciably longer than wide, the space between evenly concave (Figs. 282-285); abdomen completelyrcovered withiverrucaeionspicules\(Bigs..218-219) iy ianrs 4. eels tacmiencielc are tele 14 Head directed anteriorly (as in Fig. 224); dorsal and pleural papillae of abdomen without setae C. subu- lata Head directed ventrally (Fig. 219); dorsal and pleural papillae of abdomen with setae .............. 15 Dorsal papillae of prothorax without setae or, if setae appear to be present, no longer than width of socket PFT eee OR Men Cont SEN seis SVE ESIOL es sdenehais' GRopsusatan Geel relat Altera i allamslaiatels Sumeiats Secnehue mess C. caryae Dorsallipapillac of prothoraxiwatintSetae yen .ty-c.jscsiore ee Hrd Netn ever se see aye ec wus ens ote ier aouaye a stele daretons 16 Spatula teeth separated from one another by distance greater than their length (Fig. 283); pleural papillae OLE POIROT NOLEDK WWM SAIS 8's nin'olks into ba Socio RO MOb CA os so ah cower Dano nag Ae mbonS.onnt C. hirtiglobus Spatula teeth separated from one another by distance shorter than their length (Fig. 284); pleural papillae Ohprothoraxawithoubisetacmererscee eet «oleate sree a ree errr ci-tohcel re epee es eis a sesh C. shmoo Terminal segment tapering, pointed, as long as or longer than eighth (Fig. 229); dorsal and lateral papillae situatedton\prominentibasesthigc228) prt. inte reeeW ate tote cie Pay -vaveuat- teeter Vere seal secreted otsfe tus shepenerev eh 18 Terminal segment convex, not pointed, shorter than eighth (Fig. 220); dorsal papillae not situated on PLoOMUnentybases; (aspnyktoy2iG) irs A Muar. s aesieestelatte ty. cP het lalate ty snarth sare sPsaestircyseysrd splays sieioces 20 Terminal segment approximately as long as eighth (as in Fig. 230); terminal setae situated posterolaterally (LES OTA) leet 5 0:8 615 Cho ida SOND, ita Geet 8. Sta, fs CaP MPaR ROCCO OLA han BUR a Ree era C. asteris, C. inanis Terminal segment at least 1 1/2 times as long as eighth segment (Fig. 226-227); terminal setae situated wemtralll yg (1 P229) rronst sitar se cetcodnieet st PNsks (ce ho PN gers cn oe llap polis ofa s Say ay ajlayalce Seva, a aveteya Seara apse Chott 19 Spatulashattmomwaiderthamtooth (Fig2298) ite. ory i te ete sista cicisies seiiercicielccistays sie C. deflexipili Spatilarshatimuchtwiderthantoothi(Big#299)) ses sk arts tenia = sie kernel C. glauciglobus ADdominalsermentsa—/Awithmno dorsal papillae. rwlntar weber ereicderc testers rer eleven lssacionstener yates) sean 21 Abdonminalisesmentsu—/with4idorsalipapillac) {52 eels se eee Ses slots oll aide licle elles os 24 Spatulatoothuminutelysserrate’attapexc(Fig3297)) Seatrack lca ss i eclstel selvsks sickvortenersbons ote C. caminata Spatula tooth strongly strongly notched at apex ((akeas2ctoy ILD. Ric eno aan DRS Om ch OM RAS AG A nas 22 Bodyacompletelyicoveredywathiverrucaely+. 0... deere fonts seem orete Mic te Naa C. caryaecola Bodyawithvextensivelsmoothyareasir. pachsyy. 6 «1 - chist cieteislssie Gs si aels cielo inty~ «. Sielvisiay.teldet als 6hskoays ciate 23 Abdominal segment 8 with extensive verrucae between spiracles ....................5--- C. cynipsea Abdominal segment 8 entirely smooth between spiracles ..................00-005 C. inflata, C. tumida Spatula cruciate, the shaft very broad anteriorly, tooth conspicuously narrowed from base to notched APOGRISSHZO— 209M. vorravne niece whe else OR Rolain (enya ay batts C. flaticrustum, C. melicrustum, C. striolacrustum Spatula, if broad at anterior end of shaft, with more or less parallel-sided tooth (Figs. 291,306) > ....... C. arcuata, C. biretta, C. conoidea, C. cucurbitata, C. echinata, C. eumaris, C. glebosa, C. marginata, C. persi- coides, C. procumbens, C. purpurea, C. recurvata, C. sanguinolenta, C. spinulosa, C. stellata, C. supina, C. turbinata, C. turbinella Head directed ventrally, prothorax hooded above head capsule (Figs. 230-231); dorsal and pleural papil- laefonyprominentraisedibasesy ampere eit... eden ttt iter ere erect ne elena UNA Srareasnls 26 Head directed anteriorly; dorsal and pleural papillae not on prominent, raised bases ............... 30 Papillae, except on terminal segment, all with setae (as in Fig. 281) .......... C. leviglobus, C. spiniglobus Fapillaewwithoutsetael(asim Big 2 79) wie pues... heya wy b neuer ere tee hos cl cteun tite cldisyetcyoieveraqctCotpste aaioes-s 27 Terminal abdominal segment acute; spatula tooth about 1/2 length of shaft (Fig. 279) ................. oo} 6 cgolnd oEoIgo es GOURD DdabuLOUGe CR AOOD hOB L ooo nDadM Cao uN RUD oo od ood ao OBE Oot C. viscidolium Terminal abdominal segment rounded (Fig. 230); spatula tooth 1/3 or less length of shaft (Fig. 280) . . . 28 mamterionendiorfspatulaishaftnarrow,rounded (Fig.277)) 7.5 see. le oe ve eee oe C. cilidolium —AMterionendonrspatilawiderane ular (Gissy2S0) vomits satis itor. slobolore ehedeuseesier- lees sia) okeisieneleleysists 29 Spatula shaft twice as broad anteriorly and posteriorly than at midlength (Fig. 280) ..... C. tuberidolium Spatula shaft only slightly boader anteriorly and posteriorly than at midlength (Fig. 278) . C. hirtidolium Porsalfandipleuralipapillaeiwithishortsetae) ys eciyyass - -iericlejores siete srtereere ayeisisielo elses C. spherica Dorsaltandipletralipapillacwithoutsetaeyse 1) ceraey ello c lpr ee ciieieiotele eistantere chao) scl feler eet 31 Bodyzcompletelyzcoverediwith verrucae (asim Figs: 221-222) me irae seit aerelsy sists )a) )sbe chs telete oe 32 Body partly smooth, at least at midlength of ventral abdominal segments (as in Figs. 215-217) ...... 33 Spatulaitoothvabout d/silensthiofishaft(Eigsoill6) -eeaee = ceirce cis joc poley telsl sets tee re C. thompsoni Spatula tooth about 1/5 length of shaft; 2, occasionally 3, lateral papillae on each side of spatula (Fig. 276) SR Ne PP eaten ERR EW SRE NEA Ta 5 con Wan ollayal sv ghaoaed, & uisrdin okey alstale i nhe:WWo, zecaananehenele rane C. urnula THE GALL MIDGES (DIPTERA: CECIDOMYITDAE) OF HICKORIES (JUGLANDACEAE: CARYA) 104 33(31). Abdominal segments without posterior spicules om vemter 2.30.00. eo eles oor eee ene oasis 34 Abdominal segments with posterior/spiculesion;venter) @iyirc-)-iise eae elciehie. «toon ae een 36 34(33). Abdominal segments with verrucae at midlength on dorsum .................0020000000- C. tubicola Abdominal segments with verrucae only laterally on dorsum ...................--005- C. levicrustum 36(33). Abdominal first through seventh segments with 4 dorsal papillae .C. aggregata, C. ansericollum, C. guttata Abdominal first through seventh segments with 6 dorsal papillae “a ES SE eee C. antennata, C. holotricha, C. inclinata, C. lenta, C. ovalis, C. tuberculata ACKNOWLEDGMENTS I thank the many people who helped advance this long project, first of all Sally Gagné, whose fa- vorite gall midges these are, for her interest and constant support. Lucrecia Rodriguez of the Sys- tematic Entomology Laboratory (SEL) deserves special mention for her multivaried computer as- sistance and making plates out of the color and SEM photographs, all with effusive good humor. Diana Marques expertly made plates out of the line drawings. I acknowledge also Robert W. Carl- son, SEL, for his help with computer programs and equipment. Nit Malikul continued her much appreciated, painstaking task of rendering perma- nent slide mounts, so indispensable for gall midge work. Scott D. Whittaker, SEM Lab Manager, Smithsonian Institution, patiently assisted my sessions with the SEM. For their interest and hos- pitality, leading me to good collection sites, and in some cases, making additional collections on their own, I am grateful to: Wayne Brewer, University of Alabama, Auburn, AL; Richard L. Brown, Uni- versity of Mississippi, Starkville, MS; Denis and Barbara Gagné, Amston, CT and Murrells Inlet, SC; Kenneth L. Hibbard, Florida Department of Agriculture and Consumer Services, Ft. Pierce, FL; Tim L. McCabe, New York State Museum, Al- bany, NY; Jerry and Rose Payne, Lizella, GA; Gary J. Steck, Florida State Collection of Arthropods, Gainesville, FL; and Manya B. Stoetzel, Long Beach, MD, whom I also thank for Figs. 170-171. Larry L6f, Director of the Gorgas Science Founda- tion, Inc., Brownsville, Texas, graciously allowed me to accompany a group visiting the Founda- tion’s field station, Rancho del Cielo, in the Sierra Madre Oriental, SW of Victoria, Tamaulipas, Mex- ico, to search for Caryomyia there. L.J. Grauke, USDA Pecan Breeding and Genetics Laboratory, College Station, Texas, kindly gave advice and in- formation concerning hickories. Kevin Tunison, USDA National Arboretum, Washington, DC, granted me permission to collect in the National Arboretum, Washington, DC. Hymenopterist col- leagues in the SEL, Matthew Buffington, Michael W. Gates, Eric E. Grissell, and Terry P. Nuhn, en- thusiastically received and identified reared para- sitoid Hymenoptera. Ronald A. Ochoa of the same Laboratory kindly identified the mite galls in Figs. 1-2. Allen L. Norrbom, Lead Scientist, Diptera Unit, SEL, was unstintingly supportive of my work. I thank John Updike for kindly granting me permission to use as epigraph part of his poem Condo Moon that evokes so beautifully the history of life, and the Ohio Academy of Science for per- mission to republish the two fine plates from Wells 1916 that serve here as an additional practi- cal aid for identifying the hickory gall midges. Iam grateful also to both the in-house and journal reviewers for their careful reading, corrections, and suggestions: Michael W. Gates and Allen L. Norrbom, both of SEL; Netta Dorchin, Buck- nell University, Lewisburg, Pennsylvania; Keith M. Harris, Ripley, United Kingdom; and Peter Kolesik, University of Adelaide, South Australia, Australia. LITERATURE CITED Beutenmiiller, W. 1904. The insect-galls of the vicinity of New York City. Journal of the American Museum of Natural History 4: 89-124. [Reprinted as Guide Leaflet No. 16. American Museum of Natural His- tory. 38pp.] Brown, R.G. & Brown, M.L. 1972. Woody Plants of Mary- land. Port City Press, Baltimore, Maryland. i-lx, 1-347. Buhr, H. 1964. Bestimmungstabellen der Gallen (Zoo- und Phytocecidien) an Pflanzen Mittel- und Nordeuropas. Band I. VEB Gustav Fischer Verlag, Jena. i-xv, 1-761. Buhr, H. 1965. Bestimmungstabellen der Gallen (Zoo- und RAYMOND J. GAGNE 105 Phytocecidien) an Pflanzen Mittel- und Nordeuropas. Band II. VEB Gustav Fischer Verlag, Jena. 763-1572 & 25 pls. Csoka, G., Stone, G.N. & Melika, G. 2005. Biology, ecol- ogy, and evolution of gall-inducing Cynipidae. Pp. 573-642 in Raman, A.,Schaefer, C.W., & Withers, T.M. (eds.). Biology, Ecology, and Evolution of Gall- inducing Arthropods. Volume 2. Science Publishers, Inc. Enfield, New Hampshire, USA & Plymouth, England. Davis, M.B. 1983. Quaternary history of deciduous forests of eastern North America and Europe. An- nals of the Missouri Botanical Garden 70: 550-563. Delcourt, H.R. & Delcourt, P.A. 1984. Ice age haven for hardwoods. Natural History 1984 (9): 22-28. Dziurznski, A. 1961. The inhabitants of the galls of Mikiola fagi Htg. Part I. Materials for the morphol- ogy and development of Mikiola fagi Htg. (Itonidi- dae), as well as of its endophagous primary parasite Secodes coactus Ratzb. (Chalcididae). Acta Zoologica Cracoviensia 6: 9-41, pls. VI-X. Elias, T.S. 1980. The Complete Trees of North America: Field Guide and Natural History. Van Nostrand Reinhold: Outdoor Life Book Division, Times Mirror Maga- zines, Inc., New York. i-xii, 1-948 pp. Felt, E.P. 1906. Insects affecting park and woodland trees. New York State Museum Memoir 8 (vol. 1), 1-457, 48 pls. Felt, E.P. 1907. New species of Cecidomyiidae. New York’ State Education Department, Albany. 1-53. Felt, E.P. 1908. Appendix D. Pp. 286-422, 489-510, pls. 33-34. In his 23d report of the State Entomologist on injurious and other insects of the State of New York 1907. New York State Museum Bulletin 124: 5-541, 44 pls. Felt, E.P. 1909. Additional rearings in Cecidomyiidae. Journal of Economic Entomology 2: 286-293. Felt, E.P. 1911. A generic synopsis of the Itonidae. Jour- nal of the New York Entomological Society 19: 31-62. Felt, E.P. 1915. Appendix: A study of gall midges II. Pp. 79-242, pls. 1-14. In his 30th Report of the State En- tomologist on injurious and other insects of the State of New York 1913. New York State Museum Bul- letin 175: 5-257, 16 pls. Felt, E.P. 1916. Appendix: A study of gall midges IV. Pp. 101-172, 199-203, pls. 14-18. In his 31st Report of the State Entomologist on injurious and other insects of the State of New York 1915. New York State Museum Bulletin 186: 7-215, 18 pls. Felt, E.P. 1918. Key to American insect galls. New York State Museum Bulletin 200: 5-310, 16 pls. Felt, E.P. 1921. Appendix: A study of gall midges, VII. Pp. 81-240, 255-280, pls. 8-20. In his 34th Report of the State Entomologist on injurious and other in- sects of the State of New York 1918. New York State Museum Bulletin 231-232: 7-288, 20 pls. Felt, E.P. 1940. Plant Galls and Gall Makers. Comstock Publishing Co., Ithaca, New York. i-viii, 1-364. Felt, E.P. 1958. II. Subfamily Itonidinae (= Cecidomyi- inae) (with revisions by C.L. and J.E. Remington). Pp. 93-206. In Guide to the Insects of Connecticut. Part VI: The Diptera or True Flies of Connecticut. Sixth Fascicle: March Flies and Gall-Midges. State Geo- logical and Natural History Survey, Storrs, Con- necticut. Foote, R.H. 1965. Family Cecidomyiidae (Itonididae). Pp 241-295. In A. Stone, Sabrosky, C. W. Wirth, W.W., Foote, R.H. & Coulson, J.R. (eds.). A catalog of the Diptera of America north of Mexico. United States Department of Agriculture Handbook 276: i-iv, 1-1696. [Reprinted in 1983 by Smithsonian Institu- tion Press, Washington, D.C.] Fowells, H.A. (ed.). 1965. Silvics of Forest Trees of the United States. U.S. Department of Agriculture, For- est Service, Agriculture Handbook No. 271: i-iv, 1-762. Gagné, R.J. 1968. A taxonomic revision of the genus As- teromyia (Diptera: Cecidomyiidae). Miscellaneous Publications of the Entomological Society of America 6: 1-40. Gagné, R.J. 1973. A generic synopsis of the Nearctic Ce- cidomyiidi (Diptera: Cecidomyiidae: Cecidomyi- inae). Annals of the Entomological Society of America 66: 857-889. Gagné, R.J. 1978. A systematic analysis of the pine pitch midges, Cecidomyia spp. (Diptera: Cecidomyiidae). U.S. Department of Agriculture Technical Bulletin 1575: 1-18. Gagné, RJ. 1981. Cecidomyiidae. Pp. 257-292, in McAlpine, J. F., Peterson, B. V., Shewell, G. E., Teskey, H. J., Vockeroth, J. R., & Wood, D. M. (eds.). Manual of Nearctic Diptera. Volume 1. Monograph No. 27. Research Branch, Agriculture Canada, Ot- tawa. (4), i-vi, 1-674. Gagné, R.J. 1983. Acericecis Gagné, a new genus for Ce- cidomyia ocellaris Osten Sacken (Diptera: Cecidomyi- idae), the maple leaf ocellate gall maker in North America. Proceedings of the Entomological Society of Washington 85: 704-709. Gagné, R.J. 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ithaca, New York and London. i-xi, 1-356. Gagné, RJ. 1994. The Gall Midges of the Neotropical Re- gion. Cornell University Press, Ithaca, New York and London. i-xv, 1-352. Gagné, RJ. 2004. A catalog of the Cecidomyiidae (Diptera) of the world. Memoirs of the Entomological Society of Washington 23: 1408. Gagné, R.J., Balciunas, J.K., & Burrows, D.W. 1997. Six new species of gall midges (Diptera: Cecidomyi- idae) from Melaleuca (Myrtaceae) in Australia. Pro- ceedings of the Entomological Society of Washington 99: 312-334. Gagné, RJ. & Marohasy, J. 1993. The gall midges (Diptera: Cecidomyiidae) of Acacia spp. (Mi- mosaceae) in Kenya. Insecta Mundi 7: 77-124. 106 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Gagné, R.J. & Payne, J.A. 1992. A new species of Har- mandia (Diptera: Cecidomyiidae) damaging leaves of Allegheny chinkapin in eastern United States and a redescription of the genus. Journal of Entomological Science 27: 383-391. Gleason, H.A. 1963. The New Britton and Brown Illus- trated Flora of the Northeastern United States and Adja- cent Canada. Vol. 2: The Choropetalous Dicotyle- doneae. Hafner Publishing Co. Inc., New York and London. i-iv, 1-655. Graham, A. 1999. Late Cretaceous and Cenozoic History of North American Vegetation. Oxford University Press, Oxford. i-xviii, 1-350. Grauke, L.J. 1988. A cultivar list for hickory. Seventy- ninth Annual Report of the Northern Nut Growers Asso- ciation, Inc. Lebanon Valley College, Annville, PA, Aug. 7-10. 1988: 131-143. Grauke, L.J. 2003. Hickories. Pp. 117-166, in Fulbright, D. (ed.). Nut Tree Culture in North America. Northern Nut Growers Association, Inc., Hamden, Connecti- cut. Grauke, L.J. & Pratt, J.W. 1986. A Field Guide to the hickories of North America. Seventy-seventh Annual Report of the Northern Nut Growers Association, Inc. Augustana College, Rock Island, IL, Aug. 10-14. 1986: 123-153. Hanson, P.E. & Gomez-Laurito, J. 2005. Diversity of gall-inducing arthropods of Costa Rica. pp. 673-692 in Raman, A., Schaefer, C.W., & Withers, T.M. (eds.). Biology, Ecology, and Evolution of Gall-inducing Arthropods. Volume 2. Science Publishers, Inc. En- field, New Hampshire, USA & Plymouth, England. Harris, K.M. 2004. A new species of gall midge, Acerice- cis campestre, inducing leaf galls on field maple (Acer campestre). Cecidology 19: 41-44. Hodges, A., Buss, E., & Mizell, R.F.,, IIL. 2006. Insect Galls of Florida. University of Florida, Institute of Food and Agricultural Sciences, Gainesville, Florida. i-x, 1-130. Jarvis, T. 1909. A catalogue of the gall insects of Ontario. Thirty-Ninth Annual Report of the Entomological Soci- ety of Ontario 1908: 70-98, pls. A-P. Jones, R.G., Gagné, R.J., & Barr, W.F. 1983. Biology and taxonomy of the Rhopalomyia gall midges (Diptera: Cecidomyiidae) of Artemisia tridentata Nuttall (Compositae) in Idaho. Contributions of the American Entomological Institute 21: (2), 1-79, (11). Kieffer, J.-J. 1896b. Quatre nouveaux genres du groupe Diplosis. Miscellanea Entomologica 4: 4—5. Krombein, K.V., Hurd, P.D., Jr., Smith, D.R., & Burks, B.D. 1979. Catalog of Hymenoptera in America North of Mexico. Vol. 1. Symphyta and Apocrita (Parasitica). Smithsonian Institution Press, Washington, D.C. i- xvi, 1-1198. Manchester, S.R. 1987. The fossil history of the Juglan- daceae. Monographs in Systematic Botany from the Missouri Botanical Garden 21: 1-137. Mohn, E. 1961. Gallmticken (Diptera, Itonididae) aus El Salvador. 4. Zur Phylogenie der Asphondyliidi der neotropischen und holarktischen Region. Sencken- bergiana Biologica 42: 131-330. Moser, J.C. 1965. The interrelationships of three gall makers and their natural enemies, on hackberry (Celtis occidentalis L.). New York State Museum and Science Service Bulletin 402: 1-95. Osgood, E. A., & Gagné, RJ. 1978. Biology and taxon- omy of two gall midges (Diptera: Cecidomyiidae) found in galls on balsam fir needles with descrip- tion of a new species of Paradiplosis. Annals of the Entomological Society of America 71: 85-91. Osten Sacken, C.R. 1862. V. On the North American Ce- cidomyidae. Pp. 173-205, Pl. I in Loew, H. (ed.). Monographs of the Diptera of North America. Part I. Smithsonian Miscellaneous Collections 6 (1 [pub. 141]): 1-221, pls. I-II. Osten Sacken, C.R. 1870. Biological notes on Diptera. (Article 2nd.). Transactions of the American Entomo- logical Society 3: 51-54. Osten Sacken, C.R. 1878. Catalogue of the described Diptera of North America. Smithsonian Miscellaneous Collections 270: i-xlvi, 1-276. Osten Sacken, C.R. 1904. Record of My Life and Work in Entomology. The Author, Cambridge, Massachu- setts. i-vii, 1-240, (2). Roininen, H., Nyman, T., & Zinovjev, A. 2005. Biology, ecology, and evolution of gall-inducing sawflies (Hy- menoptera: Tenthredinidae and Xyelidae. pp. 467— 494 in Raman, A., Schaefer, C.W., & Withers, T.M. (eds.). Biology, Ecology, and Evolution of Gall-inducing Arthropods. Volume 2. Science Publishers, Inc., En- field, New Hampshire, USA & Plymouth, England. Rondani, C. 1860. Stirpis cecidomyarum. Genera revisa. Nota undecima, pro dipterologia italica. Atti della Societa Italiana di Scienze Naturali (1859-1860) 2: 286— 294, pl. D. Roskam, J.C. 1977. Biosystematics of insects living in fe- male birch catkins. I. Gall midges of the genus Semu- dobia Kieffer (Diptera, Cecidomyiidae). Tijdschrift voor Entomologie 120: 153-197. Rtibsaamen, E.H. 1910. Ueber deutsche Gallmticken und Gallen. Zeitschrift fiir wissenschaftliche Insekten- biologie 6: 283-289. Sato, S. & Yukawa, J. 2001. Absense record of Fagus gall midges (Diptera: Cecidomyiidae) on Ulleung Is- land, Korea and in North America. Esakia 41: 17-25. Sears, P.B. 1914. The insect galls from Cedar Point and vicinity. The Ohio Naturalist 15: 377-388, pls. XVIII- XXI. Skuhrava, M. 1997. Family Cecidomyiidae. Pp. 71-204 in Papp, L. & B. Darvas, B. (eds.). Contributions to a Manual of Palaearctic Diptera (with Special Reference to Flies of Economic Importance). Volume 2: Nematocera and Lower Brachycera. Science Herald, Budapest. Stebbins, F.A. 1910. Insect galls of Springfield, Massa- RAYMOND J. GAGNE 107 chusetts and vicinity. Springfield Museum of Natural History Bulletin 2: 1-138. Stone, D.E. 1990. Juglandaceae. Pp. 348-358 in Kubitzki, K., Rohwer, J.G., & Bittrich, V. (eds.). The Families and Genera of Flowering Plants. Vol. 2. Dicotyledons: Magnoliid, Hamameliid and Caryophyllid Fami- lies. Springer-Verlag, Berlin and New York. Stone, D.E. 1997. 28. Juglandaceae A. Richard ex Kunth. Walnut Family. Pp. 416-428 in Flora of North Amer- ica Editorial Committee. Flora of North America. Vol. 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford University Press, New York and Oxford. Sylvén, E. & Tastas-Duque, R. 1993. Adaptive, taxo- nomic, and phylogenetic aspects of female abdomi- nal features in Oligotrophini (Diptera, Cecidomyi- idae), and four new Dasineura species from the western Palearctic. Zoologica Scripta 22: 277-298. Thompson, M.T. 1915. An Illustrated Catalogue of Ameri- can Insect Galls. Nassau, Rensselaer County, New York. 1-116. Tsuda, K. 1982. Midge galls produced on the leaves of Fagus-species in Kyushu. Satsuma 31: 117-128. Wells, B.W. 1916. A survey of the zoocecidia on species of Hicoria caused by parasites belonging to the Erio- phyidae and the Itonididae (Cecidomyiidae). Ohio Journal of Science 16: 37-57, pls. I-II. Wolfe, J.A. & Leopold, E.B. 1967. Neogene and early Quarternary vegetation of northwestern North America and Northeastern Asia. Pp. 193-206. In Hopkins, D.M. (ed.). The Bering Land Bridge. Stan- ford University Press, Stanford, California. 1-495. Yukawa, J. & Masuda, H. 1996. Insect and Mite Galls of Japan in Colors. Zenkoku Noson Kyoiku Kyokai, Tokyo. 1-826. 108 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) OHLO JOURNAL OF SCIENCE. Vou. XVI, PLATE I. Bertram W. Wells, Figs. 1-16. Plate I of Wells, 1916, reproduced through the courtesy of The Ohio Academy of Science. Galls of hick- ory. Figs. 1-2, Eriophyidae mite galls. 1, 1a, Eriophyes sp. 2, Aceria caryae (Keifer). Figs. 3-4, Nonexistent: these numbers referred in Wells’s text to species for which he had no figures. Figs. 5-16, Cecidomyiidae, Caryomyia spp. 5, C. lenta. 5a, C. inflata. 6, possibly C. ovalis or young gall of C. inflata. 7, C. caminata. 8, C. inanis. 9, C. recur- vata. 9a, C. eumaris. 10, C. sanguinolenta. 11, Possibly C. stellata. 11a, C. stellata. 12 & 13, C. caryaecola. 14, C. echi- nata. 15, C. purpurea. 16, C. biretta. RAYMOND J. GAGNE 109 OHIO JOURNAL OF SCIENCE. VOL. XVI, PLATE II. ; MMM ayy SONG Bertram W. Wells. Figs. 16a—33. Plate II of Wells, 1916, reproduced through the courtesy of The Ohio Academy of Science. Galls of hickory, all by Cecidomyiidae. Figs. 16a—28, Caryomyia spp. 16a, C. flaticrustum. 17, C. persicoides. 18, C. caryae. 19, C. leviglobus. 20-20a, C. caryae. 21, C. spherica. 21a—21b, C. aggregata. 22, C. sp. possibly caryae. 23, C. thomp- sont. 24, Possibly young gall of C. tuberidolium. 25, C. viscidolium. 26, C. tubicola. 27, C. subulata. 28, C. urnula. Fig. 29, Contarinia cucumata. Figs. 30-33, Caryomyia spp. 30, C. melicrustum. 31, C. striolacrustum. 32, C. asteris. Fig. 33, Contarinia biconvexa. 110 THE GALL MIDGES (DIPTERA: CECIDOMYITIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Figs. 34-47. Immature stages and galls of Caryomyia spp., the galls entire and in longitudinal section. 34-38, Cary- omyjia striolata. 34, Egg on leaflet (globules are resinous exudates). 35, First instar initiating gall. 36, Group of galls. 37, Gall. 38, Gall with second instar. 39-40, C. albipilosa galls. 40, With second instar. 41-42, C. urnula galls. 42, With second instar. 43-45, C. viscidolium galls. 44, With third instar. 45, Overwintered galls, one with de- tached apical cap. 46-47, C. tuberidolium galls. 46, Young, rapidly growing galls, thickly covered with resin gland exudates. Lines indicate approximate length of mature galls. RAYMOND J. GAGNE 111 Figs. 48-59. Galls of Caryomyia spp., entire or in longitudinal section. 48, C. tuberidolium, with second instar of in- quiline Parallelodiplosis caryae preyed upon by early instar Lestodiplosis sp. 49-50, C. hirtidolium. 51-52, C. cili- dolium. 53-54, C. hirtiglobus. 53, With larva of Lestodiplosis sp. near base. 55-58, C. caryae. 55-56, Weak-walled form. 57-58, Robust form. 59, C. shmoo. Lines indicate approximate length of mature galls. 112 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Figs. 60-71. Galls of Caryomyia spp., entire or in longitudinal section. 60-61, C. shmoo. 61, Arrow shows pupal exit on overwintered gall. 62-63, C. leviglobus. 64-65, C. spiniglobus. 65, with third instar. 66-67, C. deflexipili. 67, with second instar. 68-69, C. glauciglobus. 70-71, C. caryaecola. 71, with second instar. Lines indicate approximate length of mature galls. RAYMOND J. GAGNE 113 Figs. 72-83. Galls of Caryomyia spp., entire or in longitudinal section. 72-73, C. subulata. 74-77, C. tubicola. 74, Young galls, some not yet erupted from basal collar. 77, Overwintered galls, one with pupal exit, the other with excavated area prepared by larva. 78-81, C. flaticrustum. 80, apical corona partially collapsed. 81, Overwintered galls, that on left with basal excavated areas prepared by larva; gall on right with completely collapsed corona. 82-83, C. melicrustum. Lines indicate approximate length of mature galls. 114 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Figs. 84-95. Galls of Caryomyia spp., entire or in longitudinal section. 84-85, C. striolacrustum. 86-87, C. levicrustum. 88-89, C. eumaris. 90-91, C. recurvata. 91, With parasitoid larvae. 92-93, C. spinulosa. 94-95, C. procumbens. Lines indicate approximate length of mature galls. RAYMOND J. GAGNE 115 Figs. 96-107. Galls of Caryomyia spp., entire or in longitudinal section. 96-97, C. supina. 98-99, C. inanis. 99, With third instar. 100-101, C. asteris. 102-103, C. inflata. 104-105, C. tumida. 106-107, C. lenta. Lines indicate approxi- mate length of mature galls. 116 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Figs. 108-120. Galls of Caryomyia spp., entire or in longitudinal section. 108-109, C. caminata. 110-111, C. ovalis. 112-115, C. thompsoni. 112, Pair varying in hairiness. 113, Sparsely haired form. 114-115, Thickly haired form. 116-117, C. holotricha. 118-119, C. tuberculata. 119, With second instar. 120, C. purpurea. Lines indicate approxi- mate length of mature galls. RAYMOND J. GAGNE 117 Figs. 121-132. Galls of Caryomyia spp., entire or in longitudinal section. 121-122, C. purpurea. 122, With second in- star. 123-125, C. turbinata. 126-129, C. persicoides. 126-127, Spheroid form. 128-129, Conical form. 130-131, C. glebosa. 132, C. turbanella. Lines indicate approximate length of mature galls. 118 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Figs. 133-144. Galls of Caryomyia spp., entire or in longitudinal section. 133, C. turbanella. 134-136, C. sanguinolenta. 136, Base of overwintered gall showing pupal exit. 137-138, C. conoidea. 139-140, C. marginata. 141-142, C. arcu- ata. 143, C. stellata. 144, C. biretta. Lines indicate approximate length of mature galls. RAYMOND J. GAGNE 119 Figs. 145-156. Galls of Caryomyia spp., entire or in longitudinal section. 145-146, C. inclinata. 147-148, C. ansericol- Tum. 149-151, C. cucurbitata. 152-153, C. echinata. 154-155, C. guttata. 156, C. aggregata. Lines indicate approxi- mate length of mature galls. 120 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Figs. 157-168. Galls of hickories, entire or in longitudinal section. Figs. 157-164, Caryomyia spp. 157, C. aggregata. 158-159, C. antennata. 159, With third instar. 160-161, C. spherica. 162-164, C. cynipsea. 162, Young gall with sev- eral larval chambers. 164, With third instars in separate chambers. 165-166, Caryadiplosis venicola. Figs. 167-168, Caryadiplosis biconvexa. 167, Upper leaf surface. 168, Lower leaf surface. Lines indicate approximate length of mature galls. RAYMOND J. GAGNE 121 Figs. 169-180. Galls of hickories, entire or in section. 169, Caryadiplosis biconvexa, larval exit on upper leaf surface. 170-171, Contarinia bulliformis. 172-174, Contarinia cucumata. 174, Larval exit on upper leaf surface. 175-177, Gliaspilota glutinosa 175, Upper leaf surface. 176, Lower leaf surface. 177, Detail of lower surface, third instars in- dicated by arrows. 178-180, Harmandiola nucicola. Lines indicate approximate width of mature galls. 122 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Figs. 181-186. Galls of Phylloxera caryaevenae Fitch, entire and in section; line indicates approximate length of gall. Figs. 183-185. Female of Caryomyia conoidea. 184, Postabdomen, lateral. 185, Same, dorsal, showing pigmented sclerite on ninth segment. Fig. 186. Male postabdomen of Caryomyia aggregata, dorsal on left, ventral on right. 123 RAYMOND J. GAGNE ventral. 190, lateral. 192, Second to , Same lateral. 189, J Second and third abdominal segments Lateral view. 188, Head and thorax ventral. 191 y , pupa. 187 192. Caryomyia tubicola Figs. 187— s , dorsal. J Third to fifth abdominal segments fourth abdominal segments 124 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) 148 pm 186 pm i 4 104 yr 104 ym Figs. 193-198. Caryomyia spp., pupal anterior segments, ventral and lateral views. 193-194, C. tuberidolium. 195-196, C. urnula. 197-198, C. caryae. 125 RAYMOND J. GAGNE 202, lobus. 201- & 1 irt ch G y 199-200 1eWws. pupal anterior segments, ventral and lateral v iglobus. lev ia spp., 199-204. Caryomy C. shmoo. 203-204, C Figs. 126 THE GALL MIDGES (DIPTERA: CECIDOMYIHDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Figs. 205-210. Caryomyia spp., pupal anterior segments, ventral and lateral views. 205-206, C. spherica. 207-208, C. holotricha. 209-210, C. inflata. - RAYMOND J. GAGNE 127 aan 216 aS Figs. 211-216. Caryomyia spp., pupal anterior segments, ventral and lateral views. 211-212, C. marginata. 213-214, C. sanguinolenta. Figs. 215-216. Caryomyia striolata, larva. 215, Lateral. 216, Anterior segments, lateral. 128 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) 188 ray wre Figs. 217-222. Caryomyia spp., larvae. 217, C. striolata, posterior segments, lateral. 218-219, C. caryae. 218, Dorsal. 219, Anterior segments, ventral. 220-222, C. sanguinolenta. 220, Dorsal. 221. Anterior segments, ventral. 222, Posterior segments, dorsal. RAYMOND J. GAGNE 129 Taam DOA 1 mm 1 mm 228 Too jm Figs. 223-228. Caryomyia spp., larvae. 223-225, C. glebosa. 223, Dorsal view. 224, Anterior segments, ventral. 225, Posterior segments, dorsal. 226-228, C. deflexipili. 226, Dorsal view, with parasitoid larva attached. 227, Ventral. 228, Anterior segments, ventral. 130 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) {QB jim mm 108 ym Figs. 229-234. Caryomyia spp., larvae. 229, C. deflexipili, posterior segments, ventral. 230-231, C. cilidolium. 230, Dor- sal. 231, Anterior segments, ventral. Figs. 232-234, Gliaspilota glutinosa, larvae. 232, Lateral. 233. Anterior seg- ments, dorsal. 234, Posterior segments, dorsal. RAYMOND J. GAGNE 131 Figs. 235-246. Caryomyia spp. Figs. 235-241, third antennal flagellomeres. 235, C. albipilosa, male. 236, C. albipilosa, female. 237, C. tubicola, male. 238, C. tubicola, female. 239, C. caryae, male. 240, C. tuberidolium, male. 241, C. vis- cidolium, female. Figs. 242-244. Labella and palpus. 242, C. inflata. 243, C. viscidolium, female, only first and sec- ond palpal segments shown. 244, C. spherica. Fig. 245. C. tubicola, wing. Fig. 246. C. tubicola, acropod. 132 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Figs. 247-251. Caryomyia spp., male genitalia: a = sclerite on tenth tergum. 247, C. tubicola, cerci not shown, dorsal. 248, C. tubicola, lateral. 249, C. albipilosa, dorsal. 250, Same, hypoproct and aedeagus only. 251, C. cilidolium, dorsal. RAYMOND J. GAGNE 133 1 ' ! | MMe, a! 1 SG ( ‘ \ CEEOL ed Figs. 252-257. Caryomyia spp., male genitalia: a = sclerite on tenth tergum; b = posterior extent of setulae on dorsal surface of hypoproct. 252, C. thompsoni, dorsal. 253, Same, cerci and aedeagus only. 254, C. purpurea, cerci not shown, dorsal. 255, Same, aedeagus and hypoproct, lateral. 256, C. persicoides, aedeagus and hypoproct, dorsal. 257, Same, lateral. 134 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) gee Ses eo Ci Cacats ea ro paliaas Oa C ek — wo FZ ~—. (Sa ——— eee Se eS Si 259 = i COCK Kig i eee ae — ae SAC ~~ = Ne eS ae LL SSS See é zi 260 NEE alg 7 a Se coe <2 See a | oe \\ \\ RSC ee we thik Ne ee ae oe as 6 War RS ee 261 tens ah ae X ye {SS Figs. 258-264. Caryomyia spp., female postabdomens, lateral. 258, C. tubicola, seventh segment to cerci. Figs. 259-264. Ninth segment and fused cerci. 259, C. tubicola. 260, C. albipilosa. 261, C. viscidolium. 262, C. inclinata. 263, C. ansericollum. 264, C. cynipsea. RAYMOND J. GAGNE 135 267 268 Figs. 265-268. Caryomyia spp., female ninth segment and fused cerci, lateral. 265, C. persicoides. 266, C. inanis. Figs. 267-268. C. albipennis, head of pupa. 267, Ventral. 268, Lateral. 136 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) spatula “ventral papilla lateral papillae ventral papillae spiracle =o dorsal papilla on pleural papilla y terminal papilla Figs. 269-271. Caryomyia striolata, third instar. 269, Anterior segments, ventral. 270, Posterior segments, ventral. 271, Same, dorsal. RAYMOND J. GAGNE 137 enlarged spinules antenna bucca spiracle collar segment sternal papilla ventral papilla a ine pail spatula Figs. 272-273. Caryomyia tubicola, second instar. 272, Dorsal. 273, Detail of anterior segments, ventral. 138 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) f 276 per - 7 @ 2 a 2, eras a ia Ce 28 fp. ss 286 2 579 580 281 @ ; y) /\ f \ a \ 6 2 \ ee a aye Saale, af Ne Ft zo / 6 ah ag 284 285 ‘ ; 7 , = 2 2 is 287 288 289 290 291 Figs. 274-291. Caryomyia spp., larval spatulas with sternal and lateral papillae. 274, C. striolata. 275, C. albipilosa. 276, C. urnula. 277, C. cilidolium. 278, C. hirtidolium. 279, C. viscidolium. 280, C. tuberidolium. 281, C. leviglobus. 282, C. caryae. 283, C. hirtiglobus. 284, C. shmoo. 285, C. subulata. 286, C. caryaecola. 287, C. melicrustum. 288, C. strio- lacrustum. 289, C. flaticrustum. 290, C. levicrustum. 291, C. eumaris. 292 293 @€ ®) @ \l ) P Le. 297 298 ® ° cae > RAYMOND J. GAGNE see. a Ws 139 & » 296 | 301 306 Figs. 292-311. Caryomyia spp., larval spatulas with sternal and lateral papillae. 292, C. supina. 293, C. procumbens. 294, C. recurvata. 295, C. spinulosa. 296, C. stellata. 297, C. caminata. 298, C. deflexipili. 299, C. glauciglobus. 300, C. asteris. 301, C. inanis. 302, C. biretta. 303, C. sanguinolenta. 304, C. arcuata. 305, C. turbinata. 306, C. purpurea. 307, C. glebosa. 308, C. cucurbitata. 309, C. marginata. 310, C. turbanella. 311, C. cynipsea. 140 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) vat hao sen ye \ 319 323 iI 327 Figs. 312-327. Caryomyia spp., larval spatulas with sternal and lateral papillae. 312, C. conoidea. 313, C. inflata. 314, C. persicoides. 315, C. echinata. 316, C. thompsoni. 317, C. spherica. 318, C. lenta. 319, C. ansericollum. 320, C. inclinata. 321, C. guttata. 322, C. tubicola. 323, C. aggregata. 324, C. antennata. 325, C. ovalis. 326, C. tuberculata. 327, C. holotricha. RAYMOND J. GAGNE 141 J Figs. 328-333. Caryadiplosis venicola. 328, Male third flagellomere. 329, Female third flagellomere. 330, Male ab- domen, segments 6-8. 331, Female abdomen, segments 8 to end (arrow denotes sclerite on ninth segment). 332, Male genitalia, dorsal. 333, Same, ventral, cerci removed (arrow denotes horizontal ridge at hypoproct mi- dlength). 142 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Figs. 334-342. Larval postabdomens and spatulas with sternal and lateral papillae. 334-335, Caryadiplosis venicola. 336-337, Caryadiplosis biconvexa. 338-340, Contarinia cucumata, 339-340 showing difference in wear between teeth on young larva and on fully mature larva, respectively. 341-342, Contarinia bulliformis. RAYMOND J. GAGNE 143 Figs. 343-350. Gliaspilota glutinosa. 343, Female third flagellomere, ventral. 344, Same, dorsal. 345, Acropod. 346, Fe- male postabdomen, 6th segment to end. 347, Detail of female fused cerci and hypoproct. 348, Second instar postabdomen, dorsal. 349, Third instar, anterior segments, ventral. 350, Same, posterior segments, dorsal. 144 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) 353 Figs. 351-357. Harmandiola nucicola. 351, Male third flagellomere. 352-355. Male genitalia. 352, Dorsal. 353, Detail of hypoproct, dorsal. 354, Ventral. 355, Detail of cerci, hypoproct, and tip of aedeagus, dorsal. Figs. 356-357, Larva. 356, Posterior segments, dorsal. 357, Spatula with sternal and lateral papillae. RAYMOND J. GAGNE 145 Figs. 358-362. Parallelodiplosis caryae. 358, Third instar postabdomen, dorsal. 359, Spatula with sternal and lateral papillae. 360, Acropod. 361, Male genitalia, dorsal. 362, Female postabdomen, seventh segment to end. 146 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF HICKORIES (JUGLANDACEAE: CARYA) Map 1. Distribution of Caryomyia tubicola (circles) superimposed on natural range of hickories (gray area, based on Fowells 1965 and Elias 1980) in North America. A circle may cover several adjacent collection sites. RAYMOND J. GAGNE 147 Map 2. Distribution of Caryomyia inanis (circles), C. inflata (squares), and C. lenta (triangles), superimposed on nat- ural range of hickories (gray area, based on Fowells 1965 and Elias 1980) in North America. ‘ = i j . vr y Ns 1 i beh bor NE Ga MEMOIRS OF THE AMERICAN ENTOMOLOGICAL SOCIETY Available Back Issue List The Blattidae of Panama. Morgan Hebard. 1919. 148 pp. 6 plates. $8.00. 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