Philonix fulvicollis Fitch, 1859(1858), sexual generation

Synonyms: Philonix fulvicollis Fitch (1859: 783), asexual females; Cynips (Teras) fulvicollis combination by Osten Sacken (1865: 379); Biorhiza fulvicollis combination by Ashmead (1885: 296); Biorrhiza fulvicollis combination and incorrect spelling by Dalla Torre (1893: 61); Philonyx fulvicollis combination and incorrect spelling by Smith (1910: 598); Cynips (Philonix) fulvicollis var. fulvicollis form fulvicollis combination by Kinsey (1930: 262); Cynips (Philonix) fulvicollis combination by Kinsey (1936: 207).
Philonix nigricollis Fitch, 1859. Synonymized by Kinsey (1930: 262).
Acraspis niger Gillette, 1889. Acraspis nigra corrected spelling by Dalla Torre (1893: 64); Philonix nigra combination by Beutenmueller (1909: 251); Philonyx nigra combination and incorrect spelling by Smith (1910: 598); Philonix niger correction by Felt (1918: 96). Synonymized by Kinsey (1930: 262).
Acraspis gillettei Bassett, 1900. Acraspis gillettii incorrect spelling by Beutenmueller (1904: 25); Philonix gillettei combination by Felt (1906: 711); Philonyx gillettei combination and incorrect spelling by Smith (1910: 598). Synonymized by Beutenmueller (1909: 252).
Kinsey (1930, 1936) described 6 additional species from the asexual generation and included them within his “fulvicollis complex”: Cynips (Philonix) canadensis Kinsey, 1930 on Q. alba, Q. macrocarpa, Q. michauxii, and Q. muehlenbergii; Cynips (Philonix) major Kinsey, 1930 on Q. alba, Q. macrocarpa, Q. michauxii, and Q. muehlenbergii; Cynips (Philonix) rubricosa Kinsey, 1930 on Q. alba and Q. stellata, Cynips (Philonix) vorisi Kinsey, 1930 on Q. bicolor, Q. michauxii, and Q. macrocarpa; Cynips (Philonix) insulensis Kinsey, 1936 on Q. gambelii and Q. macrocarpa; and Cynips (Philonix) latigenae Kinsey, 1936 on Q. gambelii. Kinsey also included in this complex the taxa Philonix gigas Weld, 1922 on Q. lyrata and Philonix lanaeglobuli (Ashmead, 1887) on Q. michauxii, both known only from an asexual generation, plus Philonix pallipes (Bassett, 1900), known only from a sexual generation on Q. alba. Since then, Melika & Abrahamson (2002) synonymized Philonix pallipes to Acraspis gemula (Bassett, 1881). Weld (1951) and Burks (1979) treated Philonix insulensis (Kinsey), Philonix latigenae (Kinsey), Philonix gigas Weld, and Philonix lanaeglobuli (Ashmead) as distinct valid species, while the rest were treated as varieties of P. fulvicollis. We follow this treatment, thus Philonix fulvicollis var. canadensis (Kinsey), syn. nov., Philonix fulvicollis var. major (Kinsey), syn.nov., Philonix fulvicollis var. rubricosa (Kinsey), syn. nov. and Philonix fulvicollis var. vorisi (Kinsey), syn. nov. are all considered synonyms of P. fulvicollis

Gall. The sexual generation galls (Fig. 204) occur singly in terminal buds of twigs of Q. macrocarpa. The gall is a relatively robust, seed-like, sub-ovoid cell approximately 2 mm long, dark brown when mature with longitudinal striations. Occupies the entire bud when mature, causing stunting or suppression of shoots and leaves produced from the bud.

Biology. See Fitch (1859), Weld (1922a, 1926), and Kinsey (1930, 1936) for the biology of the asexual generation. Asexual generation galls (Fig. 203) in Kinsey’s (1936) “fulvicollis” complex have been recorded on leaves of Q. alba, Q. bicolor, Q. chapmanii, Q. gambelii, Q. lyrata, Q. macrocarpa, Q. michauxii, Q. muehlenbergii, and Q. stellata, (Kinsey 1930, 1936; Burks 1979). Asexual generation females only begin to emerge from galls in the autumn of the year following the year of gall formation (Kinsey 1930), and we recorded females emerging up to four years after the year of gall formation. Adult females of the asexual generation (from galls collected in Manitoba but reared in Edmonton) emerged during the second half of October and early November. Sexual generation galls matured in May in Edmonton, and adults emerged 1–3 June 2008. Galls of the asexual generation appeared on leaves in Edmonton in July.

Distribution. USA: New York, Michigan, Tennessee, Illinois, Kansas (Burks 1979); Canada: Manitoba, Ontario, Quebec, New Brunswick.

Comments. Digweed (2010) treated Philonix gigas (Weld), Philonix insulensis (Kinsey), and Philonix nigra (Gillette) as synonyms of P. fulvicollis, although without formal synonymization. Molecular analysis, however, shows that P. fulvicollis and P. nigra are distinct species.

Philonix nigra

Quercus alba, prinoides
Leaf galls, detachable
Globular, 5-8 mm, covered with grey felt, on under side Sept-Oct. Deciduous. One-celled.

Quercus bicolor
Leaf galls, detachable
Globular, 5-8 mm, covered with greyish felt, scattered on the under side of leaf in Sept-Oct. Central cell is supported by coarse radiating fibers.

Quercus macrocarpa, muehlenbergii
Leaf galls, detachable
Globular, 5-8 mm in dia, with gray pubescence, dropping in the fall.

[Photo caption]
On Q alba. Also on Q bicolor, prinoides, macrocarpa, muehlenbergii. Galls drop before the end of July altho some as late as Oct. Emergence in Nov and Dec distributed over three seasons beginning the first fall. In Baltimore MD 30 adults emerged Nov 27-Dec 9 the first fall from 100 galls. Emergence mostly in daylight. Some placed on ground a few inches from base of a white oak tree simply wandered away.

[Fitch speculatively associates these two species as well as Biarhiza nigra with root galls because he found the adults on the ground or snow, and he thought the adults resembled wasps known to induce root galls on oaks in Europe. They are described based on differences in the adult wasp and none of the descriptions mention a gall.]

Philonix fulvicollis, new species

Philonix nigricollis, new species

Acraspis niger, n. sp.

Hosts: Quercus alba

Galls.--Small, brown, globular bodies, densely covered with a grayish pubescence which gives them the appearance of felt on their outer surface, attached to the under surface of the leaves of the white-oak in September and October. Galls exactly similar are very common on the leaves of the bur-oak Quercus macrocarpa, but from these I have not succeeded in rearing the flies. Internally these galls have a fragile central cell surrounded and held in place by a dense growth of dark brown radiating fibres. The galls resemble very closely those in my collection of Acraspis lanaeglobuli Ash.

Range: MI, IO

Acraspis gillettei n. sp.

Galls three-tenths of an inch in diameter. Perfectly round, slightly uneven or pimply, each pimple crowned with a tuft of hair- like pubescence. They are of a pale ash gray color, but brown or black if long exposed to the weather. The larval cell is central and is kept in place by a spongy mass that is loosely fibrous on the inner surface of the gall. The cell is oval, .15 by .18. The shell very thin and hard. The galls affect the leaves of Q. alba, and, as a rule, the tops of tall old trees. They are rarely abundant, but occasionally occur in great numbers on an isolated tree or a small grove of oaks. I took this species at first for A. niger Gill., but it differs from that species in several particulars.

Cynips (fulvicollis) fulvicollis (Fitch)
agamic form

Philonix fulvicollis Fitch, 1859, 5th Rpt. Nox. Ins. N.Y.:783.
Cynips fulvicollis fulvicollis agamic form fulvicollis Kinsey, 1930, Ind. Univ. Studies 84-86:262, figs. 41, 228, 234, 242-243, 252, 255.
Cynips fulvicollis Houard, 1934, Marcellia 28:44. Kinsey, 1934, Workbook in Biol.:42. Houard, 1935, Marcellia 28:117.
Philonix niger Britton, 1934, Conn. State Ent. Rpt. 33:395.

To the data given in Kinsey 1930, add:

RANGE.-Maine: Kezar Falls (galls, Mrs. D. Tenney in Kinsey coll.). Connecticut: Westport (gall, acc. Britton 1934). Maryland: Oakland, 9 S (galls). Virginia: East Falls Church (Weld acc. Houard 1935). West Virginia: Wolf Summit. Parkersburg (galls). Ohio: Chillicothe. Eaton, 6 E. Coolville (galls). Zanesville, 7 S. Kent (galls, Erlanson in Kinsey coll.). West Union. Malta, 2 N (with much major). Xenia, 5 W. South Bloomingville, 4 E. Michigan: Grayling. Big Rapids (galls). Maple Rapids (galls, E. S. Anderson in Kinsey coll.). Indiana: Kentland (R. Kessler in Kinsey coll.). Greensburg (galls, G. F. Hyatt in Kinsey coll.). Ladoga (galls, Hugh Lee in Kinsey coll.). Mongo. Hartsville, 4 W. Aurora (galls). Wyandot (galls). English (galls). Romney (galls). St. John (galls). Crossplains (galls). Chandler, 1 W. Belmont, 5 S (galls). Westfield, 9 N (galls). Winchester, 8 SW (galls). Bloomington, 8 W. Bloomington, 7 SW (Bugbee in Kinsey coll.). Illinois: Mahomet (galls, S. L. Neave in Kinsey coll.). Mt. Carmel, 12 W (gall). Seneca (galls). Green Valley in Du Page Co. (gall). Stock ton, 5 W. Michigan: West Branch (galls). Pentwater (gall, F. Payne in Kinsey coll.). Interlocken (galls, R. Voris in Kinsey coll.). Kentucky: Lebanon (gall). Whitley (galls). Carrollton, 2 S. Missouri: Marshall, 11 S (galls). Kansas City (gall). Mineola (galls). Kimmswick. Wisconsin: Dodgeville (galls). Stevens Point, 7 W (gall). Stevens Point, 5 N. Sparta, 7 W (galls). See Figure 120.

These additional locality records are all within the limits of the range predicted in our previous publication on Cynips. The new dates for gall origin and development and for insect emergence also fall within the range previously published. We have over 800 insects of this species which we have bred since the previous publication. Of these, 250 were bred in the third year after the collection of the galls.

Our new material has been the inspiration for complete re determinations of the several thousand insects and galls which we have previously studied and reported. It should be said, frankly, that the re-determinations were undertaken with numerous misgivings of our previous interpretations of the group, due to the small number of clear-cut taxonomic characters available on these highly simplified, wingless insects. After several weeks of study on this one species, we find our previously published conclusions well substantiated. We are dealing here with a population of hybrid origin, the product of the meeting of northern and southern species during the late Pleistocene glaciation. Unfortunately our collections, large as they are, are still too scant to warrant a statistically genetic analysis (which I have attempted) of the nature of the material. For such a study we need many more large series from many more localities.

Cynips (fulvicollis) major Kinsey
agamic form

Cynips fulvicollis major Kinsey, 1930, Ind. Univ. Studies 84-86:257, figs. 39, 227, 239, 249.

To the data given in the above, add:

RANGE.-Ohio: West Union, 4 S. Gallipolis, 6 N. Indiana: Charleston (H. Spieth in Kinsey coll.). Kentland, 4 N (R. Kessler in Kinsey coll.). Crawfordsville (Hugh Lee in Kinsey coll.). Fort Wayne. Chandler. Illinois: Marshall (galls, H. Spieth in Kinsey coll.). Olney. Danville (S. L. Neave in Kinsey coll.). Missouri: Allenton (E. S. Anderson in Kinsey coll.). Springfield (gall, R. Voris in Kinsey coll.). Sullivan, 1 NE (in part fulvicollis). Arkansas: Forrest City (gall). Camden, 7 N (gall). Busch, 4 W. (gall). Sheridan, 4 N (gall). Newport, 3 E (gall). Louisiana: Bernice. See Figure 120.

LIFE HISTORY..—Adults: October 20. November 22, 27. December 1, 4, 6, 8, 9, 10, 12, 14, 15, 16, 17, 18, 19, 20, 22, 23, 24, 26, 28. January 2, 3, 4, 5, 7, 8, 10, 15, 20, 21. February 2, 6, 20. April 25. Some emergence during the first year, most of the emergence in late December of the second year, some emergence in the third winter after the formation of the galls.

Most of the new locality records lie within the previously recorded range. Three of the new localities lie further north on the Indiana-Illinois boundary. Although this latter area is nearly 200 miles north of the southern front of the Wisconsin glaciation, it is an area from which we have this and some other southern species which are not ordinarily found within the glaciated territory. Three of the records extend the range through Arkansas into Northern Louisiana.

Our newly bred material gives us some more first- and second-year emergence records, and the first third-year records which we have for this species.

Cynips (fulvicollis) vorisi Kinsey
agamic form

Cynips fulvicollis vorisi Kinsey 1930, Ind. Univ. Studies 84-86:255, figs. 38, 230, 238, 248.

To the data given in the above, add:

RANGE.-Illinois: Sparland (Q. macrocarpa, galls). Missouri: Kansas City (Q. macrocarpa). See Figure 120.

These new records for vorisi make its range more nearly accord with the range of typically Ozark species of Cynipidae. The galls collected at Kansas City on August 14 were for the most part small and immature, but some of them had attained the size and fibrous internal structure characteristic of the species. Since our previous publication we have bred hundreds of insects of vorisi in the second winter after the collection of the galls, and no less than 98 adults in the third winter after collection. Careful examination of the unemerged galls fails to show any larvae that remain to emerge in a fourth winter, and the third-year emergence will probably stand as the most delayed emergence known in the genus Cynips

Cynips fulvicollis Fitch [not certain this is a synonym]
Agamic form, fulvicollis Fitch

Gall, globular, smooth to pubescent, on leaves.

Adults Nov, Dec

On Q alba, michauxii