Caryomyia viscidolium

Caryomyia aggregata

Caryomyia albipilosa

Caryomyia ansericollum

Caryomyia antennata

Caryomyia arcuata

Caryomyia asteris

Caryomyia biretta

Caryomyia caminata

Caryomyia caryae

Caryomyia caryaecola

Caryomyia cilidolium

Caryomyia conoidea

Caryomyia cucurbitata

Caryomyia cynipsea

Caryomyia deflexipili

Caryomyia echinata

Caryomyia eumaris

Caryomyia holotricha

Caryomyia flaticrustum

Caryomyia glauciglobus

Caryomyia glebosa

Caryomyia guttata

Caryomyia hirtidolium

Caryomyia hirtiglobus

Caryomyia inanis

Caryomyia inclinata

Caryomyia inflata

Caryomyia lenta

Caryomyia levicrustum

Caryomyia leviglobus

Caryomyia marginata

Caryomyia melicrustum

Caryomyia ovalis

Caryomyia persicoides

Caryomyia procumbens

Caryomyia purpurea

Caryomyia recurvata

Caryomyia sanguinolenta

Caryomyia shmoo

Caryomyia spherica

Caryomyia spiniglobus

Caryomyia spinulosa

Caryomyia stellata

Caryomyia striolacrustum

Caryomyia striolata

Caryomyia subulata

Caryomyia supina

Caryomyia thompsoni

Caryomyia tuberculata

Caryomyia tuberidolium

Caryomyia tubicola

Caryomyia tumida

Caryomyia turbanella

Caryomyia turbinata

Caryomyia urnula

Caryadiplosis biconvexa

Caryadiplosis venicola

Contarinia bulliformis

Contarinia cucumata

Gliaspilota glutinosa

Harmandiola nucicola

Caryomyia viscidolium Gagne, new species 

Hosts: Carya ovata, tomentosa, glabra, texana

Gall (Figs. 25, 43-45): Occasional, found on Eucarya hickories; on upper leaf surface, between veins; 3.0-4.1 mm in height, 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 surface of hickories. For notes on galls of related species, see under C. cilidolium. 

Affinities. — See under C. cilidolium. [C cilidolium: 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 circular 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, usually 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. tuberidolium 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. viscidolium, and the female for C. tuberidolium.]

Biological notes. — Pupae break out of the weak cap at the gall's apex. Emergence occurs in April. 

Range: AR, CT, IL, IN, KY, MD, MA, MS, NH, NJ, NY, NC, OH, PA, VA, DC,WV

Caryomyia aggregata Gagne, new species

Hosts: Carya tomentosa, texana, ovata, glabra, laciniosa

Gall (Figs. 21a-b, 156-157): Common, on most Eucarya hickories; on lower leaf surface on vein, usually clustered in a row along major 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 surface; base with small, irregular, shallow, central to off-center excavation; wall woody, thick, especially 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 apical intrusion of hairs in the larval chamber and occur between instead of on the veins.

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, coalescing 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. 

Range: AL, AR, CT, FL, GA, KY, IN, IL, MD, MA, MO, NH, NY, NC, OH, PA, TN, VA, DC, WV

Caryomyia albipilosa Gagne, new species 

Hosts: Carya tomentosa, glabra, cordiformis

Gall (Figs. 39-40): Common, a spring gall, on both Apocarya and Eucarya hickories; found singly or in groups on lower leaf surface between veins; 3.5-5.0 mm in height, elongate-conical, 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 concave; wall uniformly thin, becoming brittle; larval chamber rough with weak longitudinal ridges. This gall differs from that of C. striolata only in having a thick covering of long, 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. That the galls of these two species appear in spring and dehisce soon after further separates them from remaining Caryomyia spp. 

Biological notes. — In central Maryland this species is one of two spring galls with larvae developing between late April and early June. A series of adults emerged from caged previous year's galls on 16-18 April when hickory leaves were unfurling. 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 larvae. See under C. striolata for more details on timing of galls. 

Range: CT, MD, NY, VA, DC, WV

Caryomyia ansericollum Gagne, new name 

Hosts: Carya cordiformis

Caryomyia caryaecola Felt 1921: 114. Junior secondary homonym of Cecidomyia caryaecola Osten Sacken; Gagne 2004: 100, as new junior synonym of Caryomyia caryaecola (Osten Sacken) to avoid coining new name for C. caryaecola (Felt). 

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 recurved, 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, turning 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 between larval chamber and gall apex. 

This is the only gall on bitternut with thick, woody tissue; the others are thin-walled and brittle. Felt (1921) redescribed C. caryaecola Osten Sacken as a new species because he did not consider Osten Sacken's name to be valid, Osten Sacken having described it on the basis of the gall only. Unfortunately, Felt misidentified Osten Sacken's species, so Felt's homonym must be renamed 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. 

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 another. 

Range: IN, MD, NJ, NY, PA, VT, VA, WV

Caryomyia antennata Felt 

Caryomyia antennata Felt 1909: 292. 

Hosts: Carya ovata, laciniosa, glabra, tomentosa

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, covered 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 surrounding 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 excavation 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.

Affinities. — The gall of this species is unique for the larval chamber extending into the conical basal connection with the leaf. 

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. 

Range: AR, CT, IL, GA, IN, KY, MD, MA, NH, NY, TN, WV

Caryomyia arcuata Gagne, new species 

Hosts: Carya tomentosa, texana

Gall (Figs. 141-142): Rare, on Eucarya hickories; found singly on lower leaf surface 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 excavation; larval chamber located at base of gall, ovoid, lined with yellowish membrane, contrasting with brown to purple, large-celled, spongy, viscous tissue surrounding larval chamber, a bundle of longitudinal fibers present between larval chamber and gall apex. This gall shows some resemblance to that of C. sanguinolenta from which it differs in the sides of the gall being conspicuously flared upwards around the apical cone. 

Affinities. — See under C. sanguinolenta, from which this species differs in the shape of the gall. [C sanguinolenta: 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 extensive 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 deteriorates 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.]

Range: AL, FL, IL, LA, MO, SC

Caryomyia asteris Gagne, new species 

Hosts: Carya tomentosa, ovata

Gall (Figs. 32, 100-101): Rare, on Eucarya hickories, not found east of the Appalachian Mountains or Ohio, except for one specimen 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 laterally 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 present) equal to more than 1/3 expanded gall diameter; surface covered with fine, short, brown hairs not obscuring gall surface; base tapering to small, conical pedicel, leaf with some irregular exfoliation surrounding connection with gall and showing 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. 

Affinities. — See under C. inanis. [C inanis: This species and C. asteris are similar for their fusiform larvae with raised, but asetose 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 instar. 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 continuous in C. inanis but horizontal and starlike in C. asteris.]

Range: IL, IN, MS, OH

Caryomyia biretta Gagne, new species 

Hosts: Carya ovata, laciniosa, tomentosa

Gall (Figs. 16, 144): Rare, on Eucarya hickories, but not E of Appalachian Mrs.; on lower leaf surface between veins; 2.5-3.0 mm high, depressed-spheroid, broad apical surface flat to slightly concave with central elongate nipple; 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, especially 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. 

Affinities. — See under C. sanguinolenta, from which this species differs in the shape of the gall. [C sanguinolenta: 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 extensive 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 deteriorates 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.]

Range: CT, GA, IL, MA, NH, OH

Caryomyia caminata Gagne, new species 

Hosts: Carya ovata, glabra

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 corresponding convexity present on opposite leaf surface; two-chambered, the basal larval chamber about 1/4 length of gall, ovoid, smooth, longitudinally ridged, the apical false chamber also ridged and open or closed apically, the partition between chambers with minute, central perforation. This gall is known from only two specimens, one a figure in Wells (1916). 

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, foliaceous, 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. 

Range: IN, OH

Caryomyia caryae (Osten Sacken) 

Diplosis caryae Osten Sacken 1862: 191; Felt 1909:293 (Caryomyia). 
Hormomyia arcuaria Felt 1908: 388; Felt 1918: 46 (Caryomyia). New synonym. 

Hosts: Carya ovata, glabra, tomentosa, laciniosa, texana, floridana, pallida

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 often gall subcylindrical and constricted near basal third; green or yellow, becoming tan to red or brown, hairless, more or less sticky, variably covered with small resin glands; base with weak to prominent conical pedicel in center of circular indentation, leaf in most specimens slightly exfoliate 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 spheroidal. 

Affinities. — As mentioned in the gall description, 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. 

Biological notes. — In central Maryland, galls were first noticed in late May when they contained first and second instars, but the galls were immature, soft and very sticky. Small, barely noticeable adjacent galls with first instars were recognizable but smaller and stickier. Through the first week of August most galls are brittle and contain second instars, but during this period occasional galls may contain a third instar. After early August galls usually contain third instars. Adults were reared in early April the following year. 

Range: AL, AR, CT, FL, GA, IL, KY, LA, MD, MA, MS, MO, NH, NY, NC, OH, OK, Ontario, PA, SC, TN, Tamaulipas, TX, VA, DC, WV

Caryomyia caryaecola (Osten Sacken) 

Cecidomyia caryaecola Osten Sacken 1862: 192; 
Felt 1915: 222 (Caryomyia). 

Hosts: Carya glabra, laciniosa, texana, tomentosa, ovata

Gall (Figs. 12-13, 70-71).— Common, on Eucarya hickories; single or in groups, attached 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.

Affinities. — This species may be related to C. caryae (q.v.) and its two close relatives. The gall of the present species shares with the other three a similar recessed, 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 partially 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. 

Range: AL, AR, CT, GA, IL, IN, KY, ME, MD, MA, MS, MO, NH, NY, NC, OH, OK, TN, VA, DC, WV

Caryomyia cilidolium Gagne, new species 

Hosts: Carya cordiformis

Gall (Figs. 51-52). — Occasional, known only from C. cordiformis of the Apocarya section; found singly, usually on lower leaf surface, rarely on upper, usually between but occasionally on veins; 3.1-3.9 mm in height, spheroidal, 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, circular indentation; wall firm, brittle, almost uniformly thin, larval chamber glabrous, with longitudinal ridges. For notes on galls of related species, see below under affinities. 

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 circular 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, usually 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. tuberidolium 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. viscidolium, and the female for C. tuberidolium.

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.

Range: AR, CT, MD, MO, NJ, NY, PA, TN, VT, WV

Caryomyia conoidea Gagne, new species

Hosts: Carya glabra, ovata, tomentosa, laciniosa

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 adjacent to leaf uniformly hard, conical and often with cylindrical extension at apex, sides developing longitudinal ridges and furrows upon maturity; green to red or brown surface smooth, with sparse, short, erect, white to tan hairs not obscuring surface; base with central, circular, shallow depression, leaf with short, irregular exfoliation surrounding basal connection to gall; larval chamber 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 traversed by bundle of fibers between larval chamber and gall apex. This gall is most similar to that of C. sanguinolenta, q.v. for remarks.

Affinities. — See under C. sanguinolenta, from which this species differs in the shape of the gall. [C sanguinolenta: 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 extensive 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 deteriorates 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 note. — In spring the pupa emerges from the base of the gall (as in Fig. 136). 

Range: CT, KY, ME, MD, MA, NH, NY, NC, OH, Ontario, PA, DC, WV

Caryomyia cucurbitata Gagne, new species 

Hosts: Carya tomentosa, ovata, texana

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 circular, 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. ansericolla 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. 

Range: AL, AR, KY, MD, MO, NC, OH, TN, WV

Caryomyia cynipsea (Osten Sacken) 

Cecidomyia cynipsea Osten Sacken 1862: 193; 
Felt 1918: 47 (Caryomyia); 
Gagne 2004: 276, as unplaced sp. of Cecidomyiinae. 

Hosts: Carya ovata, tomentosa, glabra

Gall (Figs. 162-164): Rare, on Eucarya hickories; on lower leaf surface, along midrib; 2.6-5.0 mm in height, irregularly spheroid, smooth, hairless, yellow turning brown; base with irregular, shallow, central excavation; woody, polythalamous, with numerous, discrete, irregularly ovoid, smooth walled, larval chambers. This is the only polythalamous gall formed by a Caryomyia. Some other species, e.g., C. aggregata, may appear to be so because their galls will occasionally be found so crowded as to be partially 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. 

Affinities. — This species forms the only polythalamous gall in the genus. 

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 polythalamous (Fig. 162). Third instars were found in galls from the same site the same year in mid-September. The following spring 3 females were reared, all emerging from separate exit holes from one of the galls. 

Range: GA, MD

Caryomyia deflexipili Gagne, new species 

Hosts: Carya texana, tomentosa, laciniosa, pallida, ovata

Gall (Figs. 66-67): Occasional, on Eucarya hickories, but not east of the Appalachians, except for New England and on the Georgia Piedmont; single or dispersed, on lower leaf surface between veins; 2.8-3.7 mm in height, spherical with small umbo at apex; green, becoming tan to brown, surface matte, non-sticky, with short, sparse, white, decumbent hairs not obscuring surface; base with small conical pedicel, leaf irregularly exfoliate surrounding base of gall; wall firm, brittle, uniformly thin, enclosing glabrous, longitudinally ridged larval chamber of same shape as gall. This gall is most similar to that of C. glauciglobus (Fig. 68), which, instead of having a sparse covering of deflexed hairs, is naked and covered with a bluish bloom. 

Affinities. — 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. 

Biological notes. — Pupae exit from the side of the gall in April and May after winter diapause.

Range: AR, CT, GA, IL, IN, KY, ME, MA, MS, MO, NH, NJ, NY, OH, PA, TN, WV

Caryomyia echinata Gagne, new species 

Hosts: Carya tomentosa, ovata, pallida, texana

Gall (Figs. 14, 152-153): Rare, on Eucarya hickories west of the Appalachian Mts; on lower leaf surface, between veins; length 2.4-5.0 mm, spheroidal 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, central 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 between instead of on veins.

Range: AR, IL, MS, MO, OH, TX, Ontario

Caryomyia eumaris Gagne, new species 

Hosts: Carya tomentosa, ovata, glabra, pallida

Gall (Figs. 9a, 88-89): Common, found on Eucarya hickories; usually single or scattered, attached to small veins on lower leaf surface; 2.2-2.7 mm in length, recumbent, subovoid except for apical, curved, tapered extension; rough textured, white, green, to tan, with short, sparse, white hairs not obscuring surface; base of gall with short, central conical projection surrounded 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. 

Affinities. — Galls of several other species, C. procumbens, C. recurvata, C. spinulosa, and C. supina, 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. 

Range: AL, AR, CT, FL, GA, IL, KY, LA, MD, MA, MS, MO, NY, NC, OH, OK, SC, TN, TX, VA, DC

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); Gagne 2004: 100, as junior synonym of C. caryae (Osten Sacken). 

Hosts: Carya ovata, texana, pallida, glabra, tomentosa

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 obscuring gall surface; base with circular, deep, central excavation, leaf without exfoliation surrounding connection; larval chamber basal, ovoid, floccose, green to brown, occasionally with slight 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. aggregata 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 lectotype 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 discrete and grow between veins, those of C. aggregata 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. 

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. 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 second instars, as well as young hymenopterous parasitoids. By June 22 and through July 16, all galls were at full size and hard. On August 3, galls contained 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. 

Range: AL, AR, CT, FL, GA, IL, IN, KY, LA, MD, MA, MS, MO, NH, NY, NC, OH, OK, PA, SC, TN, TX, VA, DC, WV

Caryomyia flaticrustum Gagne, new species 

Hosts: Carya ovata, tomentosa, glabra, laciniosa, texana, pallida

Gall (Figs. 16a, 78-81): Common, on Eucarya hickories; on lower leaf surface between veins, usually clustered; 2-4 mm in height, usually longer than wide, conical to cylindrical, deeply concave apically, walls encircling concavity flared outwards, cylindrical, or ingrown 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 extensive, thick leaf exfoliation. 

Affinities. — Caryomyia flaticrustum, C. levicrustum, C. melicrustum, and C. striolacrustum are four common, widely distributed species on the Eucarya section with diminutive, more or less discoid 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. melicrustum, except that galls of the latter are subtended 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. 

Range: AL, AR, CT, FL, GA, IL, IN, KY, ME, MD, MA, MO, MS, MO, NH, NJ, NY, NC, OH, OK, PA, SC, TN, TX, VA, DC, WV

Caryomyia glauciglobus Gagne, new species 

Hosts: Carya tomentosa, ovata, laciniosa, glabra

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 becoming tan to brown, smooth, not sticky, hairless, covered with blue-gray powdery bloom; base with small conical pedicel, leaf with irregular exfoliation around base of gall; wall firm, brittle, uniformly thin, larval chamber glabrous with longitudinal ridges. This gall appears superficially similar to those of C. deflexipili and C. caryae because of their spherical shape without conspicuous 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. 

Affinities. — See under C. deflexipili. [C deflexipili: 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.]

Range: CT, GA, IN, KY, ME, MD, MA, MS, NY, OH, TN, WV

Caryomyia glebosa Gagne, new species 

Hosts: Carya ovata, tomentosa, glabra

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, traversed 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. 

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. 

Range: AR, CT, IL, MD, MO, NC, TN, WV

Caryomyia guttata Gagne, new species 

Hosts: Carya floridana

Gall (Figs. 154-155): Found commonly but only on the central Florida endemic, 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 resemble 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.

Range: FL

Caryomyia hirtidolium Gagne, new species 

Hosts: Carya tomentosa, glabra, ovata

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 truncate in profile, apex with small central nipple surrounded 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. 

Affinities. — See under C. cilidolium. [C cilidolium: 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 circular 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, usually 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. tuberidolium 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. viscidolium, and the female for C. tuberidolium.]

Range: AL, AR, CT, GA, MD, OK, SC, VA, DC

Caryomyia hirtiglobus Gagne, new species 

Hosts: Carya tomentosa, pallida, glabra, texana, ovata

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 indentation, the corresponding leaf connection surrounded 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 longitudinal 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 longitudinally, the gall's uniformly thin wall and its conical connection to the leaf will serve to distinguish it from otherwise superficially similar galls. 

Affinities. — This species forms a natural group with C. caryae and C. shmoo because of the general shape of the gall. See under C. caryae for further discussion. [C caryae: As mentioned in the gall description, 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.]

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 second 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 August 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. 

Range: AL, AR, CT, FL, GA, KY, MD, MA, MS, MO, NY, NC, OK, TN, VA, DC, WV

Caryomyia inanis Felt 

Caryomyia inanis Felt 1909: 292. 

Hosts: Carya tomentosa, ovata, laciniosa

Gall (Figs. 8, 98-99): Occasional, on Eucarya hickories, not found east of the Appalachian 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 diameter; green to brown with sparse, long, white or brown hairs not obscuring gall surface; base with prominent conical projection in center of circular indentation, leaf with thin exfoliation surrounding gall base; larval chamber ovoid, wall firm, brittle, uniformly thin, smooth with longitudinal ridges. This gall has a similar structure to that of C. asteris, although that is not at first apparent. The foliate extension of the gall of C. inanis is vertical and tubular, while that of C. asteris is horizontally disposed.

Affinities. — This species and C. asteris are similar for their fusiform larvae with raised, but asetose 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 instar. 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 continuous in C. inanis but horizontal and starlike in C. asteris. 

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 ins tars. In early July, galls were still green but with third instars that were very active when the galls were opened. In August the galls had turned brown. 

Range: CT, KY, ME, MA, NH, NJ, NY, OH, PA, TN, WV

Caryomyia inclinata Gagne, new species 

Hosts: Carya laciniosa, tomentosa, ovata

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 incline, 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 pedicel; 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.

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. 

Biological notes. — Between June and beginning 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 instars. Pupae emerge through the side of the gall near the base. 

Range: AR, CT, IL, IN, KY, ME, MA, NH, NJ, NY, NC, OH, Ontario, Tamaulipas, WV

Caryomyia inflata Gagne, new species 

Hosts: Carya pallida, tomentosa, laciniosa, glabra, ovata

Gall (Figs. 5a, 102-103): Occasional, on Eucarya hickories, but not found on Atlantic 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 conical 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 similarly shaped except that the gall of C. tumida is completely covered with short, decumbent hair. 

Affinities. — This species and C. tumida come from similarly shaped, two-chambered galls, but that of C. inflata is smooth, the other thickly covered with short, decumbent hairs. Beyond that, the galls have certain similarities to that of C. asteris. Although the latter gall is not two-chambered, second instars of all three species have setose dorsal and pleural papillae. The rays of the C. asteris gall may indicate some midpoint development of a strictly two-chambered gall. 

Biological note. — Pupae emerge from a hole at the base of the gall next to the leaf connection. 

Range: AL, CT, GA, IL, IN, KY, ME, MA, MS, NJ, MY, OH, TN, WV

Caryomyia lenta Gagne, new species 

Hosts: Carya tomentosa, laciniosa, ovata

Gall (Figs. 5, 106-107): Occasional, 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 conical, with small nipple or tiny opening at center 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 surrounded laterally and apically by much larger, false chamber, the chambers separated by hemispherical, thin, sticky wall, both chambers longitudinally 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 except 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. 

Affinities. — The two-chambered gall has certain 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. 

Range: CT, IL, IN, KY, ME, MA, PA, NY, OH, WV

Caryomyia levicrustum Gagne, new species 

Hosts: Carya ovata, pallida, texana, glabra, tomentosa

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; surface glabrous, not sticky, green to tan; base conical, 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. 

Affinities. — Galls of this species appear most like those of C. flaticrustum but instead of being soft and resinous, are hard and glabrous. For further remarks, see under C. flaticrustum. [C flaticrustum: Caryomyia flaticrustum, C. levicrustum, C. melicrustum, and C. striolacrustum are four common, widely distributed species on the Eucarya section with diminutive, more or less discoid 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. melicrustum, 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.]

Biological notes. — In central Maryland galls were noticed as early as mid-June. Galls with second 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. 

Range: AL, AR, CT, FL, GA, IN, KY, ME, MD, MA, MS, MO, NH, NJ, NY, NC, OH, OK, SC, TN, TX, VA, DC, WV

Caryomyia leviglobus Gagne, new species 

Hosts: Carya tomentosa, ovata, texana, glabra, pallida

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 areola; hairless, not sticky, glabrous, light green, becoming 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. 

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. As with other species (viz. the C. caryae and C. cilidoliurn groups) that live unconstrained in a rather large, spherical larval chamber, the integument is completely covered with spiny verrucae that evidently allow 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 Maryland 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. 

Range: AL, AR, CT, FL, GA, KY, LA, MD, MA, MS, MO, NH, NY, NC, OH, OK, TN, TX, VA, DC, WV

Caryomyia marginata Gagne, new species 

Hosts: Carya texana, ovata, tomentosa

Gall (Figs. 139-140): Infrequent, on Eucarya hickories; on lower leaf surface, between or on veins; length 3.2-5.0 mm, conical, the base flat against leaf surface, circular, with flattened 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 exfoliation 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 collapses. 

Affinities. — See under C. sanguinolenta, from which this species differs in the shape of the gall. [C sanguinolenta: 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 extensive 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 deteriorates 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.]

Range: AL, AR, CT, KY, ME, MA, MS, MO, NH, NY, OH, Ontario, WV

Caryomyia melicrustum Gagne, new species 

Hosts: Carya ovata, pallida, texana, glabra, laciniosa, tomentosa

Gall (Figs. 30, 82-83): Common, on Eucarya hickories; on lower leaf surface between 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 mucillaginous; subtending leaf tissue with conspicuous, 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. flaticrustum galls, but galls of the latter lack the conspicuous leaf exfoliation surrounding the base of galls of C. melicrustum. 

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. [C flaticrustum: Caryomyia flaticrustum, C. levicrustum, C. melicrustum, and C. striolacrustum are four common, widely distributed species on the Eucarya section with diminutive, more or less discoid 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. melicrustum, except that galls of the latter are subtended 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.]

Biological notes. — In late May in Central Maryland, one can find young galls not yet completely broken out of the leaf covering and simulating young galls of C. hibicola (Fig. 74) and containing 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. 

Range: AL, AR, CT, FL, GA, IN, KY, ME, MD, MA, MS, MO, NH, NJ, NY, NC, OH, Ontario, SC, TN, TX, VA, DC, WV

Caryomyia ovalis Gagne, new species

Hosts: Carya glabra, tomentosa

Gall (Figs. 110-111): Rare, on Eucarya hickories; on lower leaf surface, between veins; 2.7-3.5 mm in height, ovoid to elongate ovoid, usually longer than wide, slightly bilaterally 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. caminata that are brittle, foliaceous, and matte. 

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. 

Range: MD

Caryomyia persicoides (Osten Sacken) 

Cecidomyia persicoides Osten Sacken 1862: 193; Felt 1918: 46 (Caryomyia). 

Hosts: Carya pallida, ovata, texana, tomentosa, glabra

Gall (Figs. 17, 126-129): Common, found on Eucarya hickories; single or clustered, 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 hair obscuring surface; connection to leaf circular and shallowly concave or flush with leaf; larval chamber located basally, ovoid, lined with yellowish pellicle different in texture from surrounding 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. 

Affinities. — Galls somewhat resemble those of the C. sanguinolenta species group because of their extensive soft tissue and the presence 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 addition 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 central 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 August and September. Pupae break through the side of the gall. I once noticed a sciarid pupal exuviae sticking out of a hole near the gall's apex; possibly some other organism made the hole. 

Range: AL, AR, CT, FL, GA, IL, IN, KY, MD, MA, MO, NH, NY, NC, OH, OK, Ontario, PA, SC, TN, TX, VA, DC, WV

Caryomyia procumbens Gagne, new species 

Hosts: Carya ovata, glabra, laciniosa, tomentosa

Gall (Figs. 94-95): Occasional, found on Eucarya hickories, but not east of the Appalachians, except in New England; usually found singly, attached to vein on lower leaf surface; 4-6 mm in length, horizontal, elongate, straight, narrowing gradually from rounded base to pointed apex; white, green, to tan, with stiff, sparse white hairs not obscuring surface; base of gall with central conical pedicel in shallow, circular indentation; wall firm, brittle, uniformly thin, larval chamber 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. 

Affinities. — See under C. eumaris. [C eumaris: Galls of several other species, C. procumbens, C. recurvata, C. spinulosa, and C. supina, 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.]

Range: CT, IL, IN, KY, MS, MO, OH, PA, TN, WV

Caryomyia purpurea Gagne, new species 

Hosts: Carya ovata, texana, glabra, tomentosa

Gall (Figs. 15, 120-122): Common, 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, conical prominence at apex, surface covered with long, thick, mostly dark brown hair obscuring gall surface; base with deep, circular, central excavation, the leaf without exfoliation surrounding connection; wall woody, of even thickness but thicker apically if peak present, green to brown; larval chamber at base of gall ovoid, violet to purple. Exteriorly, 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. 

Range: AL, AR, CT, GA, IL, MD, MA, MO, NH, NJ, NY, NC, OH, Ontario, SC, TN, VA, DC

Caryomyia recurvata Gagne, new species 

Hosts: Carya texana, ovata, tomentosa, glabra, pallida

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 narrowed 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 following gall axis from leaf vein connection to recurved 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. 

Affinities. — See under C. eumaris. [C eumaris: Galls of several other species, C. procumbens, C. recurvata, C. spinulosa, and C. supina, 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.]

Range: AL, AR, CT, IL, KY, LA, MD, MS, OH, MO, TN, DC

Caryomyia sanguinolenta (Osten Sacken) 

Cecidomyia sanguinolenta Osten Sacken 1862: 192; Felt 1909: 293 (Caryomyia). 

Hosts: Carya tomentosa, texana, pallida, glabra, ovata

Gall (Figs. 10, 134-136): Common, found on Eucarya hickories; single or clustered, 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 convex at first, becoming concave, then furrowed upon further drying, the apex obtuse to acute; hairless, green, turning brown, crimson, or purple, 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 purple 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. 

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 extensive 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 deteriorates 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).

Range: AL, AR, CT, FL, GA, IL, KY, LA, MD, MA, MO, NH, NJ, NY, NC, OH, Ontario, PA, TN, VA, DC, WV

Caryomyia shmoo Gagne, new species

Hosts: Carya tomentosa, texana, pallida, glabra, ovata

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-spheroidal at base, apical third abruptly narrowed, set to one side, and tipped with an umbo; green to yellow, turning tan to brown, smooth, hairless, not sticky; base with prominent conical pedicel set into circular indentation, leaf without conspicuous exfoliation surrounding base of gall; wall firm, brittle, thin, larval chamber shaped as for gall, glabrous, with longitudinal ridges. This gall resembles 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. 

Affinities. — This species forms a natural group with C. caryae and C. hirtiglobus because of the general shape of the gall.

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. 

Range: AL, AR, CT, FL, GA, LA, MD, MS, MO, NH, NY, NC, OH, PA, SC, TX, VA, DC, WV

 Caryomyia spherica Gagne, new species 

Hosts: Carya glabra, laciniosa, ovata, pallida, tomentosa

Gall (Figs. 21, 160-161): Infrequent, 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 narrow, shallow, circular depression; larval chamber at center of sphere, ovoid, tan to brown, the surrounding thick walls large-celled, woody. Galls of this species are superficially similar to the rounder galls of C. persicoides, but the larval chamber 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. 

Affinities. — Galls of C. spherica and the more spherical galls of C. persicoides can be superfically similar in gross aspect. 

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. 

Range: AL, CT, FL, GA, IL, MD, MS, NC, TN, TX, VA

Caryomyia spiniglobus Gagne, new species 

Hosts: Carya tomentosa, glabra, laciniosa, ovata

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 areola; green to yellow, becoming tan to brown with widely spaced, short, stiff light brown hairs not obscuring surface; base with small conical pedicel, leaf without exfoliation; wall firm, brittle, uniformly thin, larval chamber glabrous with longitudinal ridges. This gall differs from that of C. leviglobns in that it has hair. Both galls are distinctive for the raised apical knob.

Affinities. — See under C. leviglobus, from which the present species differs only in the hairy galls. [C leviglobus: 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. As with other species (viz. the C. caryae and C. cilidoliurn groups) that live unconstrained in a rather large, spherical larval chamber, the integument is completely covered with spiny verrucae that evidently allow these species purchase in crawling across the curved surface of the larval chamber.]

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 beginning of August galls were regularly found with third instars. Feeding larvae lie prone against the concave surface of the larval chamber. 

Range: AL, AR, CT, FL, GA, KY, LA, MD, MA, MS, NJ, PA, SC, TN, VA, DC, WV

Caryomyia spinulosa Gagne, new species 

Hosts: Carya cordiformis

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, recumbent, base bulbous, tapered beyond into elongate, 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 longitudinal ridges following gall axis from leaf vein connection to apex. This gall is superficially similar 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. 

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. [C eumaris: Galls of several other species, C. procumbens, C. recurvata, C. spinulosa, and C. supina, 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.]

Range: AR, IN, MD, MA, MO, PA, NY, VA, WV

Caryomyia stellata Gagne, new species 

Hosts: Carya ovata, laciniosa, glabra, tomentosa, floridana, pallida

Gall (Figs. 11a, 143): Occasional, on Eucarya hickories; single or clustered, on lower leaf surface between veins; 5.0-9.0 mm in height, elongate-conical, base flared laterally, the flattened edge often stellate, sometimes merely irregularly 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 chamber 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 tentatively 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. sanguinolenta except for a much higher crown and flared sides. 

Affinities. — See under C. sanguinolenta, from which this species differs in the shape of the gall. [C sanguinolenta: 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 extensive 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 deteriorates 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.]

Range: AL, AR, FL, GA, IN, KY, ME, MD, MA, MS, MO, NH, NY, OH, OK, TN, WV

Caryomyia striolacrustum Gagne, new species 

Hosts: Carya tomentosa, laciniosa, floridana, glabra, texana, ovata

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 and deeply set into leaf swelling detectable on top surface of leaf as tan to black, conspicuous convexity, apical half of gall short-cylindrical with funnel-like, deep, narrow to wide concavity surrounded by correspondingly narrow to thick walls, center of funnel with short umbo; surface not sticky, yellow to brown, minutely longitudinally striated, the striations partly fibrous on distal half; gall often surrounded and occasionally partially covered by conspicuous, brown, irregularly 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 resembles that of C. melicrustum but the surface of the latter is mucilaginous and smooth. 

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 remarks, see under C. flaticrustum. [C flaticrustum: Caryomyia flaticrustum, C. levicrustum, C. melicrustum, and C. striolacrustum are four common, widely distributed species on the Eucarya section with diminutive, more or less discoid 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. melicrustum, except that galls of the latter are subtended 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.]

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 contained second instars or third instars. 

Range: AL, AR, CT, FL, GA, IN, KY, ME, MD, MA, MS, MO, NH, NJ, NY, NC, OH, OK, SC, TN, TX, VA, DC, WV

Caryomyia striolata Gagne, new species 

Hosts: Carya aquatica, cordiformis, glabra, ovata, tomentosa

Gall (Figs. 36-38): Common, a spring gall, on both Eucarya and Apocarya hickories; 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 especially on young galls, white to green, occasionally 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. 

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 further discussion. [C albipilosa: 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. That the galls of these two species appear in spring and dehisce soon after further separates them from remaining Caryomyia spp.]

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 develop 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 because 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 parasitized larvae. Such galls could be found through September. 

Range: AL, CT, GA, KY, MD, NJ, DC, WV

Caryomyia subulata Gagne, new species 

Hosts: Carya ovata, laciniosa, tomentosa, texana

Gall (Figs. 27, 72-73): Occasional, on Eucarya hickories but not east of Appalachian 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 superficially 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. 

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. tubicola (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. 

Range: AR, CT, IL, IN, KY, ME, MA, MS, NH, NJ, NY, OH, Ontario, PA, TN, WV

Caryomyia supina Gagne, new species 

Hosts: Carya ovata, glabra, tomentosa, cordiformis, texana

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, gradually narrowing along length to pointed apex; green, yellow, to tan, hairless; base of gall with central conical pedicel in shallow, circular indentation; wall firm, brittle, uniformly thin, larval chamber glabrous with longitudinal ridges following gall axis from leaf vein connection to apex. See remarks about similar galls under C. spinulosa. 

Affinities. — See under C. eumaris. [C eumaris: Galls of several other species, C. procumbens, C. recurvata, C. spinulosa, and C. supina, 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.]

Range: AL, AR, CT, GA, KY, MD, MA, MO

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. 

Hosts: Carya ovata, laciniosa, floridana, texana, pallida, glabra, tomentosa

Gall (Figs. 23, 112-115): One of the commonest Caryomyia galls, on Eucarya hickories; usually in clusters, on lower leaf surface between 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, shallow depression, the leaf without exfoliation surrounding connection; wall of uniform thickness, firm, woody, the larval chamber invaded from both apex and base by vertical cylindrical intrusions, 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 examples 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 apical 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. 

Affinities. — In adult, pupal, and larval characters this species is similar to C. holotricha. The two species are the only congeners with an apical intrusion 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 greenish 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. 

Range: AL, AR, CT, FL, GA, IL, KY, LA, ME, MD, MA, MS, MO, NH, NJ, NY, NC, OH, OK, Ontario, PA, SC, TN, VA, DC, WV

Caryomyia tuberculata Gagne, new species 

Hosts: Carya texana, ovata, glabra, tomentosa

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 excavation, leaf without exfoliation surrounding connection; wall uniformly thick, woody, larval chamber ovoid, floccose, white, green, to purplish. The gall is most like that of C. purpurea because of its shape, the thick, woody wall, and the leaf connection but, unlike that of C. purpurea, is bumpy and hairless. 

Affinities. — The closest relative among species with long male flagellomeres and an apically convex 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.

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 regularly found. The full-grown larva fills the larval chamber. Adults emerge through a hole in the side of the gall. 

Range: AL, AR, CT, GA, IN, KY, LA, MD, MA, MS, MO, NH, NC, OK, PA, SC, TN, VA, DC, WV

Caryomyia tuberidolium Gagne, new species 

Hosts: Carya glabra, texana, tomentosa, pallida, ovata

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 surrounded 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, almost uniformly thin, larval chamber glabrous, with longitudinal ridges. For notes on galls of related species, see under C. cilidolium. 

Affinities. — See under C. cilidolium. [C cilidolium: 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 circular 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, usually 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. tuberidolium 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. viscidolium, and the female for C. tuberidolium.]

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. Second and third instars are active when disturbed. 

Range: AL, AR, CT, GA, KY, MD, MA, MS, NH, NJ, NY, NC, OK, PA, SC, TN, VA, DC, WV

 Caryomyia tubicola (Osten Sacken) 

Cecidomyia tubicola Osten Sacken 1862: 192; Felt 1909:293 (Caryomyia). 

Hosts: Carya glabra, laciniosa, texana, pallida, tomentosa, ovata, cordiformis

Gall (Figs. 26, 74-77).— Common and widespread (Map 1) on Eucarya hickories and occasional on C. cordiformis of Apocarya section; found singly or in groups; 3-7 mm in height, on lower leaf, usually between veins, occasionally 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 resembles that of C. subulata 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. striolacrustum. 

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.

Biological notes. — This may be the most common 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. 

Range: AL, AR, CT, FL, GA, IL, IN, KY, LA, ME, MD, MA, MS, MO, NH, NJ, NY, NC, OK, Ontario, PA, SC, TN, TX, VA, DC, WV

Caryomyia tumida Gagne, new species

Hosts: Carya ovata, tomentosa

Gall (Figs. 104-105): Rare, known from only 2 collections, on Eucarya hickories; on lower leaf surface, between veins; 3 mm in 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 conical 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 striate. 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. 

Affinities. — See under C. inflata. [C inflata: This species and C. tumida come from similarly shaped, two-chambered galls, but that of C. inflata is smooth, the other thickly covered with short, decumbent hairs. Beyond that, the galls have certain similarities to that of C. asteris. Although the latter gall is not two-chambered, second instars of all three species have setose dorsal and pleural papillae. The rays of the C. asteris gall may indicate some midpoint development of a strictly two-chambered gall.]

Range: IL, PA

Caryomyia turbanella Gagne, new species 

Hosts: Carya tomentosa, ovata, pallida, glabra, texana

Gall (Figs. 132-133): Rare but widely distributed, on Eucarya hickories; on veins of upper leaf surface; length 4.0-5.0 mm, spheroidal, abruptly narrowed to conical apex; tan to purple, base with long, sparse, white hairs not obscuring surface, apical cone all or partly covered with small bumps and mostly hairless; base with circular, shallow, central excavation, leaf not exfoliate surrounding connection; larval chamber near base of gall, depressed-ovoid, lined with yellowish membrane, contrasting with brown to purple, large-celled, spongy, viscous tissue surrounding larval chamber that shrivels with maturity; a bundle 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. 

Affinities. — See under C. persicoides. [C persicoides: Galls somewhat resemble those of the C. sanguinolenta species group because of their extensive soft tissue and the presence 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 addition to these differences, adults of the two groups are different enough that it appears soft-tissued galls evolved at least twice in Caryomyia.]

Range: AL, AR, CT, FL, GA, MD, DC

Caryomyia turbinata Gagne, new species 

Hosts: Carya glabra, ovata, texana, pallida, tomentosa

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 obscuring surface, the hair darker and denser apically; base with narrow, circular, shallow, central depression, sometimes barely noticeable, the leaf not exfoliated surrounding connection, usually with small convex, discolored area on reverse leaf surface; 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 extraneous, soft tissue outside the pellicle-lined larval chamber. 

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). 

Range: AL, AR, CT, FL, GA, IL, IN, KY, LA, MD, MA, MS, MO, NY, NC, OK, PA, TN, TX, VA, DC

Caryomyia urnula Gagne, new species 

Hosts: Carya illinoinensis, tomentosa, glabra, cordiformis, ovata, pallida

Gall (Figs. 28, 41-42): Late spring, early summer, common on the Eucarya section, rare on Apocarya section; found singly between veins on either upper or lower leaf surface; 2.0-3.0 mm in height, cylindrical, slightly bulging at midlength, narrowed abruptly apically to convex 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 longitudinal ridges. This gall resembles that of C. cilidolium 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. 

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 Caryomyia. Development of galls is evidently staggered. 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 dehisced 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 September trip from Maryland through Arkansas to collect hickory galls, I found not a single example. 

Range: AL, CT, FL, GA, KY, LA, MD, MO, OH, TN, VA, DC, WV

Caryadiplosis biconvexa Gagne, new species 

Hosts: Carya ovata, glabra, cordiformis, tomentosa, aquatica

Gall (Figs. 33, 167-169): Common, on both Eucarya and Apocarya sections; circular 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 thickness, larval chamber circular, flattened, reflecting gall shape; full-grown larva exits gall from small hole in center of gall on abaxial leaflet surface. 

Affinities. — The larval terminal segment has dorsal verrucae in contrast with the smooth preceding 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 common in late summer as in spring. Larvae exit from the lower surface of the leaflet. 

Range: AL, AR, GA, IN, MD, MA, NY, NC, OH, OK, PA, TN, DC, WV

Caryadiplosis venicola Gagne, new species

Hosts: Carya cordiformis, pallida, glabra, texana, tomentosa

Gall (Figs. 165-166): Occasional, on both Eucarya and Apocarya sections; an elongate, hard swelling of midvein or petiole, usually showing on both sides of leaflet, 5 mm long, 2 mm 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. 

Biological notes. — My earliest-season sighting of galls in central Maryland was on July 22. 1 noticed them more commonly in Maryland and elsewhere 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. 

Range: AR, FL, GA, MD, MO, NC, OH, OK, TN, WV

Contarinia bulliformis Gagne, new species 

Hosts: Carya illinoinensis

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 chamber of same shape as gall, containing one to several larvae.

Biological notes. — This species was found by Manya B. Stoetzel in the course of her field work on Phylloxera on pecan. Nothing further is definitely known of the biology of this species, but the full-grown larvae probably escape the galls to drop to the soil. 

Range: LA

Contarinia cucumata Gagne, new species 

Hosts: Carya ovata, tomentosa, pallida, cordiformis, glabra

Gall (Figs. 29, 172-174): Infrequent, 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 apically, not dehiscent; surface rough, green becoming 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. 

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 instars. Galls were greenish white to light green, somewhat sticky, and covered with close, long white hairs. In subsequent days these hairs became farther apart as the gall quickly elongated along the leaf axis and became darker green. Larvae 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 larvae are white and active. 

Range: AL, GA, KY, MD, MS, MO, NY, OH, TN, VA, WV

Gliaspilota glutinosa (Osten Sacken), new combination 

Cecidomyia glutinosa Osten Sacken 1862: 193; Felt 1909: 115 {Caryomyia); Gagne 1989: 211 {Cecidomyia); Gag- ne 2004: 277, as unplaced species of Cecidomyiinae. 

Hosts: Carya cordiformis, glabra, tomentosa, ovata

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 apparent from top of leaf only by discoloration, but on bottom of leaf covered with a colorless mucilaginous substance. 

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-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. 

Range: AR, FL, MD, MA, MS, NY, TX, DC, WV

Harmandiola nucicola (Osten Sacken), new combination 

Cecidomyia caryae nucicola Osten Sacken 1870: 53, invalid polynomial; caryaenucicola Jarvis 1909: 84 (Ce- cidomyia). 
Cecidomyia nucicola Osten Sacken 1878: 6; Felt 1918: 49 (Caryomyia); Gagne 1973: 868 (Contarinia). 

Hosts: Carya ovata, tomentosa, glabra

Gall (Figs. 178-180): Rare, so far known only from Eucarya section; a single-chambered blister of the nut husk, up to 5 mm wide and 4 mm high, with same color and texture as husk; larval chamber ovoid, each with a single larva. 

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. 

Range: GA, MD, NY, Ontario

The Gall Midges (Diptera: Cecidomyiidae) of Hickories (Juglandaceae: Carya)