# species_source

description:
  Sphaeroteras zinzala (Kinsey) n. comb.

  Biorhiza zinzala Kinsey, 1937: 459.
  Trigonaspis zinzala (Kinsey) Pujade-Villar y Ferrer-Suay, 2015: 12.

  [Attending to the morphological characters and the host plants, we consider there to be three species of Sphaeroteras in Mexico: S solita, tanos, and zinzala. We cannot exclude the possibility that, in the future, the color forms S solita and S tanos will be considered the same species. For the moment, other than color, these species are separated by the host plants in which their galls are found; thus, S solita produces galls on multiple species of Quercus (arizonica, chihuahuensis, deserticola, jaralensis, magnoliifolia, microphylla, rugosa, and undulata [should be undata?]), while S tanos has only one host on which S solita is not found (Quercus repanda). Sphaeroteras zinzala has been collected in two hosts distinct from the other two species of Sphaeroteras mentioned (Quercus intricata and pringlei).

  The galls of the three species of Sphaeroteras from Mexico are indistinguishable. Kinsey (1937) mentioned that they are like those of B pulchripennis. They are characterized by being more or less circular, between 3-4.5 mm in diameter, being located in the lower side of the leaves sometimes in great numbers, being attached to the midrib or secondary veins, being yellow to light brown, having the surface smooth and glossy covered in a short pubescence easily rubbed off with a few short darker bumps, having the base more or less flat with a longitudinal depression corresponding to the point where it meets the leaf vein; the larval cell is eccentric, attached to the walls of the gall by fine white filaments.]


description:
  Trigonaspis ? quercusforticorne (Walsh, 1864) Incertae sedis

  Cynips q. forticornis Walsh, 1864: 490.
  Cynips (Teras) forticornis (Walsh) Osten Sacken, 1865: 340, 350, 353.
  Cynips forticornis (Walsh) Bassett, 1870: xvi.
  Xanthoteras forticornis (Walsh) Ashmead 1897b: 262. 
  Biorhiza forticornis (Walsh) Mayr, 1881: 32. 
  Biorrhiza forticornis Dalla Torre, 1893: 60 (error escritura).
  Xanthoteras quercusforticornis (Walsh, 1864) Weld, 1951: 625.
  Trigonaspis quercusforticorne (Walsh) Melika y Abrahamson, 2002: 181


description:
  Biorhiza ? eburnea (Bassett) Incertae sedis

  Dryophanta eburneus Bassett, 1890: 70. 
  Dryophanta similis Bassett, 1890: 71. (Sinonimia en Weld, 1951: 625).
  Dryophanta simillima Dalla Torre, 1893: 54. (Sinonimia en Weld, 1951: 625).
  Dryophanta glabra Gillette, 1894: 237. (Sinonimia en Beutenmüller, 1907: 466).
  Diplolepis eburnea (Bassett) Dalla Torre y Kieffer, 1910: 368.
  Xanthoteras eburneum (Bassett) Weld, 1951: 625.
  Biorhiza eburnea (Bassett) Melika y Abrahamson, 2002: 167.


description:
  Biorhiza ? emoryi (Ashmead) Incertae sedis

  Dryophanta emoryi Ashmead, 1896: 115. 
  Diplolepis emoryi (Ashmead) Dalla Torre y Kieffer, 1910: 367.
  Xanthoteras emoryi (Ashmead) Weld, 1951: 625.
  Biorhiza emoryi (Ashmead) Melika y Abrahamson, 2002: 167.


description:
  Erythres hastata, n. sp.
  Agamic form

  [Genus level description: Gall. — A separable, cone-like, terminal cluster of aborted, fasciated leaves and bracts enclosing a small, seed-like cell. The cluster composed of many, awl-shaped bracts and aborted leaves, the inner leaves elongate, narrow, awl-shaped, with sharply-pointed tips, each terminating in fine filaments, the outer bracts short, more broadly triangulate; the whole forming an elongate or swollen cone; fresh galls probably green in color, dried galls straw to medium brown, weathering more silvery gray; the galls up to 25,. mm. in length and 13. mm. in greatest diameter, internally the larval cell is small, up to 6.0 mm. in length and 2.0 mm. in greatest diameter, elongate spindle-shaped, sharply tipped, thin-walled, and closely embedded in the surrounding bracts. Highly separable galls, borne terminally on main or lateral stems.]

  Gall. — An elongate, slender, and rather loose cluster of fasciated leaves, up to 25. mm,, averaging near 20, mm. in length and 8.0 mm. in greatest diameter, the mature galls widely flaring and loose-leaved at top, with the greatest diameter near the tip. 

  Host. — Quercus serrulata [castanea] (or a close relative), one of the prickly-tipped, nearly entire-leaved black oaks which forms small to medium-sized trees. Possibly on other black oaks of the area. 

  Range. — Michoacan: Zacapu, 11 E, 8000’ (types). Probably restricted to a limited portion of Mexico which includes the eastern part of the state of Mochoacan

  Life History. — Adults: January 12, February 10, in the first year after the development of the galls.

  Hastata differs most from jaculi in having the conical gall very slender, loose and flaring open at the top, with the greatest diameter in mature galls there at the top.


description:
  Erythres jaculi, n. sp.
  Agamic form

  [Genus level description: Gall. — A. separable, cone-like, terminal cluster of aborted, fasciated leaves and bracts enclosing a small, seed-like cell. The cluster composed of many, awl-shaped bracts and aborted leaves, the inner leaves elongate, narrow, awl-shaped, with sharply-pointed tips, each terminating in fine filaments, the outer bracts short, more broadly triangulate; the whole forming an elongate or swollen cone; fresh galls probably green in color, dried galls straw to medium brown, weathering more silvery gray; the galls up to 25,. mm. in length and 13. mm. in greatest diameter, internally the larval cell is small, tip to 6.0 mm. in length and 2.0 mm. in greatest diameter, elongate spindle-shaped, sharply tipped, thin-walled, and closely embedded in the surrounding bracts. Highly separable galls, borne terminally on main or lateral stems.]

  Gall. — A swollen, very compact cone of fasciated leaves, with the greatest diameter near the base, with the apex of the gall drawn to a blunt point; up to 25. mm,, averaging near 20. mm. in length, up to 13.0 mm, averaging near 11. mm. in greatest diameter.

  Host. Q. Rossi;, a larger tree-form or black oak with large, coarsely reticulate leaves. Possibly on other black oaks of the area. 

  Rango. — Jalisco: Sayula, 13 SW, 7700’ (types). Possibly widely spread through a southwestern portion of Mexico which includes the state of Jalisco

  Life History. — Adults: March 20. 

  This insect, jaculi, is different enough from hastata. It is most distinct in its almost uniformly sooty black mesonotum, its conspicuously broader head and thorax, and the much more compact gall — a cone which is broadest -at base and printed at top.


description:
  Amphibolips bromus Pujade-Villar y Cuesta-Porta, 2020

  [A photo of the gall appears on page 14 of the pdf]

  [Gall: Monothalamous resulting from the deformation of a bud. Very thin and elongate, some 50 mm in length, reminiscent of the seeds of Bromus. The peduncle widens near the middle of the gall (4 mm in diameter), ending in a sharp point that tapers gradually from the midsection. The gall is thin-walled, with a smooth and hairless surface; a thin, spongy layer internally surrounds the central ovate larval chamber.]

  Hosts: Quercus affinis, crassipes

  [Biology: Only known from females. The mature gall was collected in May. The adults emerged at the end of the same month.]

  [Range: Reported from the states of Mexico City, Hidalgo


description:
  Amphibolips fusus Kinsey, 1937

  [A photo of the gall appears on page 14 of the pdf]

  [Gall: Monothalamous resulting from the deformation of a bud. Fusiform (9-13 mm in diameter), with a peduncle and a sharp point at the apex (resulting in a total length of 25-35 mm). Exterior wall thick and tough, light brown in color at maturity, with a smooth and glabrous surface (see comments). The internal tissue is relatively soft and spongy, composed of very dense radiating fibers, only visible at the base and the apex, dense and compact (but spongy) in the wide part of the gall where the larval chamber is found surrounded by a thin layer of spongy tissue. Central ovate larval chamber with relatively hard walls, and with its greatest length between 5-6 mm.]

  Hosts: Quercus eduardi, crassipes, laurina

  [Biology: Species known only from females. The mature galls were collected at the beginning of June. The two adults used for the description of the species were extracted dead from the gall in November; the only adult from Santa Fe emerged in July of 2014.]

  [Range: Mexico City, Zacatecas]

  [Comments: The maximum dimensions of the diameter of the gall are greater in the original description. After studying galls of A fusus collected by Kinsey and the type material, we deduce that this fact is erroneous. On the other hand, Kinsey mentions that on the surface of the gall there are small black protuberances that probably correspond to fungal pycnidia, abundant on the surfaces of old galls.]


description:
  Amphibolips cibriani Pujade-Villar, 2018

  [A photo of the gall appears on page 14 of the pdf]

  [Gall (Figs. 2b-c). Unilocular resulting from the deformation of a bud. Spherical (2-4 cm in diameter), often ending in a point and weakly stalked. The surface is rough highlighting a large number of protuberances. At first light green, light brown at maturity. Hard consistency. In a Cross section shows the lignified central larval chamber (3 mm in diameter + 1 mm thick the wall of the inner gill) surrounded by dense spongy tissue. 

  Hosts. Quercus crassipes (section Lobatae). 

  Biology. Species known only from the agamic generation . The adults of this generation emerge in the middle of May. 

  Distribution. Only known from Santa Fe (City of Mexico).


description:
  Femuros geniale, n. sp.
  Agamic form

  Gall. — Small, narrowly cylindric, with the walls of the upper half of the gall quite thin and often puckered, nearly or entirely shut at top, although the cavity thus enclosed occupies more than a third of the diameter of the whole gall; outside of gall silvery brown, with little or no bloom, diameter up to 8.0 mm, averaging nearer 6.0 mm.; length up to 11.0 mm., averaging nearer 8.5 mm. 

  Host. — Quercus undata, Possibly on other white oaks in the area. 

  Range. — Durango: Durango, 2 N, 6500' (types). Canatlan, 7 N, 7400' Probably restricted to a part of the Western Sierra of Mexico centering in western Durango.

  Life History. — Adults: in spring. 

  The present species is the most northern of the four herein described. We have it from two localities not far apart in the state of Durango. The galls of geniale are practically identical with those of integrum — these being the smallest and most regularly cylindric in the complex, 


description:
  Femuros integrum, n. sp.
  Agamic form

  Gall. — Small, narrowly cylindric, with the walls of the upper half of the gall quite thin and often puckered, nearly or entirely shut at top, although the cavity thus enclosed occupies more than a third of the whole gall; outside of gall silvery brown, with little or no bloom; diameter up to 7.0 mm., averaging nearer 6.0 mm.; length up to 10.0 mm., averaging nearer 8.0 mm. 

  Host. — Quercus potosina (types); Q. chihuahuensis; Q. undata; Q, reticulata [rugosa], Apparently one species on all these oaks; possibly found on still other white oaks in the area. 

  Range. — San Luis Potosi: San Luis Potosi, 15 W, 8000’ (types on Q. potosina. Also on Q. undata). Guanajuato: San Felipe, 20 SW, 8000’ (Q, chihuahuensis Leon, 20 NE, 9000’ (Q. reticulata). Probably restricted to a portion of the Western Sierra of Mexico including the western portion of San Luis Potosi and adjacent Guanajuato,

  Life History. — No data available. 

  Integrum is the wide-ranging species of more central Mexico, from the state of San Luis Potosi south for some undetermined distance, it is most closely related to its more northern neighbor, geniale. The galls of the two are identical, small, and quite strictly cylindric. 


description:
  Femuros lusum, n. sp.
  Agamic form

  Gall. — Very large, cylindric, with the thin-walled top half of gall wide open or flaring, making the gall widely-open cup-shaped with a thick and solid base, the open cavity almost as wide as whole gall; outside of gall silvery to yellow-brown, young galls quite scurfy, older galls naked. Diameter of body of gall up to 15.0 mm., averaging nearer 11,0 mm., the flared top sometimes as much as 19.0 mm. in diameter; length up to 19.0 mm., averaging near 15.0 mm. 

  Host. — Quercus conglomerata [rugosa]. 

  Range. — Michoacan; Morelia, 14 E, 7000’ (types). Possibly extending through the mountains of Michoacan and part of Jalisco, 

   The galls of lusum and perfectum are strikingly distinct, representing as extreme departure from the basic type of gall as one might readily conceive. In lusum the galls are scurfy, brown, very large, distinctly cylindric, with the cavity at top very large and broadly open, or even with its thin-walled boundaries widely flared open. The galls of perfectum are covered with a purplish-white scurf, of fair size, nearly spherical, with only a very small cavity at top which is more or less completely closed at the apex. 


description:
  Femuros perfectum, n. sp.
  Agamic form

  Gall. — Fairly large, nearly globular or slightly elongate or flattened, with the exceedingly small cavity in the upper half of the gall almost completely closed by very thick and solid walls, although a central pucker at the apex of the gall still indicates the opening to the cavity; cavity never wider than larval cell, often narrower; outside of gall silvery gray to purplish brown, conspicuously covered with a purplish-white bloom; up to 16.0 mm., averaging nearer 13.0 mm. in diameter, never more than 13.0 mm. in length. 

  Host. — Quercus macrophylla [magnoliifolia], the largest-leaved white oak of southwestern Mexico. 

  Range. — Guerrero; Chilpancingo, 6 W3 40007 (types). Tierra Colorado, 14 N, 3200? (galls). Probably restricted to a portion of the state of Guerrero, in southwestern Mexico. 

  Life History. — Adults: May 10, July, 8, in the first year after the development of the gall. 

  This insect, from the state of Guerrero in the southwestern corner o’ Mexico, is closest to F lusum. The galls of the two species are strikingly distinct — an interesting instance of divergent modifications of a single basic plan. See lusum for a comparison of the galls of the two. 


description:
  Femuros ocri, n. sp.
  Agamic form

  Gall. — Similar to those of F. repandae and F. ruidum. Much smaller, up to 12. mm., but averaging only 8,5 mm. in diameter. 

  Hosts. — Quercus sacame [arizonica] (types), Q, haematophlebia [magnoliifolia],. Q. rhodophlebia [rugosa]. Possibly on other white oaks of the area. 

  Range. — Aguascalientes: Pabellon, 20 W, 7000’ (types). Jalisco: Atequiza, 5 E, 6500’ (Q, haemalophlebia), Guadalajara, 25 S, 6000’ (Q. haematophlebia). Sayula, 13 SW, 7700’ (Q, rhodophlebia). Apparently one species through the states of Aguascalientes and Jalisco, in more southwestern Mexico.

  Life History. — Adults: May 10. 

  This species belongs to the same complex as F. repandae and F. ruidum. (described in Kinsey, 1937, Rev de Ent. 7:66-67). Ocri is found further north and west than the previously described species, this new form ranging through the states of Aguascalientes and Jalisco. The gall averages much smaller than those of the other species; but otherwise the gall is quite like those previously described. 


description:
  Found on Celtis reticulata in January in New Mexico [here](https://www.inaturalist.org/observations/104976905).


description:
  Erythres hastata

  [Genus level description:] The galls are monolocular and very peculiar in shape, represented by cone-like clusters, each gall of which enclosing a small larval cell (Fig. 3). They are some similar to the cone-like galls which are induced by some Cecidomyiidae (Diptera), Andricus quercusfoliatus (Ashmead, 1881), A. stropus Ashmead, 1887 and A. cinnamomeus Ashmead, 1887 from the Nearctic; A. foecundatrix (Hartig, 1840) from the Western Palaearctic and A. mukaigawae (Mukaigawa, 1913) from the Eastern Palaearctic, nevertheless, the adults of Erythres are quite distinct from all of mentioned species. The other very important difference between Erythres and mentioned Andricus species are the host oaks. Erythres species are known only from red oaks (section Lobatae of Quercus L.), while all the listed Andricus species are associated with white oaks only (section Quercus of Quercus). 

  Erythres includes two species: E. hastata and E. jaculi. Both species are very similar. The galls are different in shape but associated with the same oak host. Erythres hastata has an elongated gall and was collected by Kinsey on Q. castanea Née (= Q. serrulata Trel.) known from Michoacán and Zacatecas (Figs 3a, c) and E. jaculi has a thicker gall also on Q. castanea (= Q. rossii Trel.) known from Jalisco (Fig. 3b). 

  Erythres jaculi and E. hastata are morphologically very similar, as we have commented, and have the same host, then, it is possible that they could be the same species; nevertheless, we accept both species since they have different shape of galls (Fig. 3). More E. jaculi specimens must be studied in order to solve this problem. Since Kinsey descriptions, only E. hastata was collected from Q. eduardii Trel. (Fig. 3c) and Q. viminea Trel. (Pujade-Villar et al., 2013: Fig 3a). These two new host records with the earlier known one, Q. castanea, all belong to the section Lobatae of Quercus L., and thus Erythres related only to the red oaks. Quercus eduardii and Q. viminea are known only from Mexico, while Q. castanea is known also from Guatemala and El Salvador (Govaerts & Frodin, 1998) and thus it is possible that Erythres species can be found south of Mexico also. 


description:
  Erythres jaculi

  [Genus level description:] The galls are monolocular and very peculiar in shape, represented by cone-like clusters, each gall of which enclosing a small larval cell (Fig. 3). They are some similar to the cone-like galls which are induced by some Cecidomyiidae (Diptera), Andricus quercusfoliatus (Ashmead, 1881), A. stropus Ashmead, 1887 and A. cinnamomeus Ashmead, 1887 from the Nearctic; A. foecundatrix (Hartig, 1840) from the Western Palaearctic and A. mukaigawae (Mukaigawa, 1913) from the Eastern Palaearctic, nevertheless, the adults of Erythres are quite distinct from all of mentioned species. The other very important difference between Erythres and mentioned Andricus species are the host oaks. Erythres species are known only from red oaks (section Lobatae of Quercus L.), while all the listed Andricus species are associated with white oaks only (section Quercus of Quercus). 

  Erythres includes two species: E. hastata and E. jaculi. Both species are very similar. The galls are different in shape but associated with the same oak host. Erythres hastata has an elongated gall and was collected by Kinsey on Q. castanea Née (= Q. serrulata Trel.) known from Michoacán and Zacatecas (Figs 3a, c) and E. jaculi has a thicker gall also on Q. castanea (= Q. rossii Trel.) known from Jalisco (Fig. 3b). 

  Erythres jaculi and E. hastata are morphologically very similar, as we have commented, and have the same host, then, it is possible that they could be the same species; nevertheless, we accept both species since they have different shape of galls (Fig. 3). More E. jaculi specimens must be studied in order to solve this problem. Since Kinsey descriptions, only E. hastata was collected from Q. eduardii Trel. (Fig. 3c) and Q. viminea Trel. (Pujade-Villar et al., 2013: Fig 3a). These two new host records with the earlier known one, Q. castanea, all belong to the section Lobatae of Quercus L., and thus Erythres related only to the red oaks. Quercus eduardii and Q. viminea are known only from Mexico, while Q. castanea is known also from Guatemala and El Salvador (Govaerts & Frodin, 1998) and thus it is possible that Erythres species can be found south of Mexico also. 


description:
  Amphibolips cibriani n. sp.
  Agamic female

  [Photos of this gall appear on page 9 of the pdf]

  [Galls: Monolocular grown from a deformation of a bud. Spherical (2-4 cm in diameter), often tapered and weakly stalked. The surface is rough, with large number of protruding bumps. At first from green to be brown clear at maturity. Hard consistency. Transverse cross sections show the lignified central larval chamber (3 mm diameter + 1 mm thick internal gall wall) surrounded by dense spongy tissue.]

  Hosts: Quercus crassipes

  Distribution: Mexico City

  Biology. Species known only from the agamic generation. Adults emerge in mid-May. These galls are frequently attacked by inquilines that are found in the larval chamber, greatly increasing its size and dividing it into compartments (Fig. 4c). The inquiline species is Synergus striatifrons Pujade-Villar and Lobato-Vila; the host of the Synergus obtained from Amphibolips sp (MEX-299) mentioned in Lobato-Vila & Pujade-Villar (2017), corresponds to A. cibriani n. sp


description:
  Aciurina bigeloviae
  Cotton-gall Tephritid

  Hosts: Ericameria nauseosa

  Galls occur singly or in pairs at axillary buds and appear as puffs of cotton attached to the stems.


externallink: https://www.biodiversitylibrary.org/item/54855#page/433/mode/1up
description:
  Estimating tephritid distribution from host-plant distribution is problematic because some gall-forming tephritid flies are known to be less widely distributed than their hosts, e.g., A. bigeloviae (Cockerell) and A. trixa.

  Both A. trixa and A. bigeloviae are known only from C. nauseosus; however, the galls they induce are readily distinguishable and is the best character for species recognition. Aciurina bigeloviae is known to form only large, cottony galls [see also Dodson and George (1986), Figure 2B] that sometimes bear small leaves studding the gall body, rather than a rosette of basal leaves as in A. semilucida (Goeden and Teerink 1996c). The tomentum of A. bigeloviae galls is thick and continuous over the entire gall, appearing like a wad of cotton. Aciurina bigeloviae galls have been collected from several sites in Idaho, Colorado, Arizona, New Mexico and Utah; however, most specimens in collections do not have the galls from which they were reared. The galls of A. maculata (Cole) also occur on C. nauseosus and are cottony; however, they are much smaller than A. bigeloviae galls and the adults are readily distinguishable.


description:
  Rhopalomyia bigeloviae

  Galls of this midge protrude 1-2 mm, look like knobs or fingerlike projections, and are glabrous and sticky. 


description:
  A gall midge which produces endogalls on Aciurina trixa and Aciurina bigeloviae. The endogalls take on the characteristics of the host gall, becoming sticky or hairy. The midge larvae apparently do not affect the fruit fly larvae in the host gall. 


description:
  <b>BENT-GALL MOTH   Gnorimoschema octomaculell[a]  Pl. 489</b>

  This moth induces pointed, monothalamous galls in terminal leaves of new shoots in spring on rubber rabbitbrush. Galls are typically bent at a 45-degree angle ... They are composed of swollen, succulent, distorted terminal leaves that are tightly bound together like the staves of a barrel ... 


description:
  Many moderately sized green, hairless galls, scattered across leaves of Salix reticulata. Some but not all galls have a prominent tuft of pink-white hairs sticking up from the top. Observed in Montana in July. 


description:
  Amphibolips kinseyi n. sp.

  [Photos of this gall appear on page 44 of the pdf]

  Gall (Fig. 16J). A relatively small, subglobose bud gall. The body of the gall reaches its greatest diameter near the middle, up to 20 mm in diameter. The gall is thick-walled, with lignified parenchyma, light brown when mature, with a smooth and naked surface; with spongious hard parenchyma internally; the hard-walled, ovate central larval chamber, with a mature length of 5.0–6.5 mm. 

  Host. Quercus crassipes.

  Biology. Only females are known. Mature galls were collected in December. 

  Distribution. Mexico: Ciudad de Mexico. 

  Remarks. This species was collected by Kinsey in 1932 and it was never described. Deposited in the Kinsey collection at the American Museum of Natural History (AMNH) as an ‘unidentified Amphibolips’. Characters are clearly differing from any other Amphibolips species (see diagnosis)


description:
  Amphibolips comini Cuesta-Porta & Pujade-Villar, nom. nov.

  Three days before the paper “Revision of the Amphibolips species of the ‘nassa’ species-complex from Mexico and Central America (Hymenoptera: Cynipidae)” was published in Zootaxa, another paper describing three new Amphibolips species from Mexico was published in ZooKeys. As a result, two different species have been described under the same name, ‘Amphibolips kinseyi’. To solve this homonymy, a new name Amphibolips comini Cuesta-Porta & Pujade-Villar, nom. nov. is proposed for replacing A. kinseyi Cuesta-Porta & Pujade-Villar, 2020, a junior homonym of A. kinseyi Nieves-Aldrey & Castillejos-Lemus, 2020. 


description:
  Aciurina michaeli
  Beaked-gall Tephritid

  This fly induces monothalamous, pear shaped, smooth bud galls on the current year's growth of rubber rabbitbrush and yellow rabbitbrush. 


description:
  Striatoandrius [sic] barriosi n. comb.

  The gall is a fused mass of larval chambers as also occurs in S. georgei, S. nievesaldreyi and S. sanchezi n. sp. Galls of S. barriosi occurs in Q. bumelioides Liebm. and Q. insignis M. Martens and Galeotti (section Quercus). 

  Also, the authors of the same paper mention that the gall of S. barriosi is similar to that of Andricus guatemalensis (Cameron 1883); this species has been considered incertae sedis by Pujade-Villar et al. (2011) because is not possible to differentiate the species according to the morphology of galls. In the original description, Cameron (1883: 71) mentions that the gall of Cynips guatemalensis is a hard mass covered of a long pilosity, but the adults obtained corresponds to an inquiline Synergus dorsalis Cameron (see the dissertation in Ritchie and Shorthouse 1987).


description:
  Stiatoandricus [sic] georgei n. comb.

  Andricus georgei

  [No gall description given]

  Hosts: In Quercus, section Quercus: Q. deserticola Trel., Q. glabrescens × obtusata, Q. glaucoides M. Martens and Galeotti, Q. laeta Liebm., Q. magnoliifolia Née, Q. resinosa Née and Q. rugosa Née (Quercus section); after a reexamen of host by S. Valencia-A, the host mentioned in Pujade-Villar et al. (2011, Q. mexicana Humb. and Bonpl.) is wrong, the correct host is Q. deserticola Trel. All host are new records from this species.

  Distribution: México, described from material collected in the Parque Nacional del Bosque del Pedregal (Bosque de Tlalpan, Ciudad de México). Its distribution is extended to have collected it in the states of Hidalgo, Jalisco, Michioacan, Morelos, Puebla and Zacatecas.


description:
  Striatoandricus maesi n. comb.

  Andricus maesi

  [No image or description of the gall is given]

  Hosts: Q. segoviensis Liebm., Q. glaucoides Mart. and Gal. and Q. obtusata Humb. and Bonpl. (section Quercus); also probably in Q. potosina Trel., Q. microphylla Née and Quercus nr rugosa (section Quercus, see remarks). 

  Distribution: Nicaragua, described from material collected in the cerro de Tisey (Estelí, Reserva Natural Tisey Estanzuela); México (Oaxaca) in this paper and probably also in the following states (see below): Jalisco, Michoacán, Puebla and Zacatecas. First record from México. 

  Remarks: Similar galls without any emergence were collected in Zacatecas (México): Mesa de San Gabriel (Q. potosina Trel.) and La Cumbre (Q. microphylla Née); also in Puebla (México): Camotepec (Q. obtusata); in Michoacán (México): Urapan (Quercus sp); and in Jalisco (México): Las Guayabas (Quercus nr rugosa).


description:
  Stiatoandricus [sic] nievesaldreyi n. comb.

  Andricus mexicanus Kinsey 1920: 309 [non Andricus mexicana Bassett, 1890]
  Andricus nievesaldreyi Pujade-Villar 2011; in Pujade-Villar et al. 2011: 32 [new name]

  [No description or photos of the gall are included]

  Hosts: The first hosts for this species are mentioned here: Q. laeta Liebm., Q. magnoliifolia Née and Q. rugosa Née (Quercus section).

  Distribution: México, described from material collected in the Sierra de Nayarit (Jalisco, México). Its distribution also includes in the states of Ciudad México, Hidalgo, Michoacán, Morelos and Tlaxcala. 

  Remarks: Kinsey (1920) described Andricus mexicanus from four adults that emerged in 1919 from galls on the upper side of leaves. Those galls were collected in 1900 by Diquet in Sierra de Nayarit (Jalisco, México) presumably from Q. magnifolia Née (= Q. macrophylla Née). Kinsey (1920) assigned to this species the individuals determined as Andricus mexicana Bassett, 1890 and Cynips guatemalensis Cameron, 1883 from the collections of Bassett (1890) at Guadalajara mountains (México) and Cameron (1883) at San Jerónimo (Guadalajara, México), respectively. A solution for the homonymy between Kinsey’s and Bassett’s species was proposed in Pujade-Villar et al. (2011) by changing the name of Andricus mexicanus Kinsey 1920 to Andricus nievesaldreyi Pujade-Villar 2011. The species A. mexicana Bassett and C. guatemalensis Cameron were only described through velvety galls on leaves. Both adults with smooth metasoma (Andricus tecturnarum group) and with striate metasoma (Striatoandricus) emerge from this type of galls from USA and México. Thus, Pujade-Villar et al. (2011) considered those two species incertae sedis until similar galls are collected at the type localities and the adults emerge. 


description:
  Striatoandricus sanchezi n. sp.
  Agamic generation

  [A photo of this gall appears on page 17 of the document]

  Gall (Fig. 10b): Adheres to the central nerve on the lower face of the rarely leaves on the lower face. It is presented as a mass of brown wool with circular-looking shape, with a non-brittle pubescence, 2.0–2.5 cm diameter. The pubescence is located above the central core forming a layer of 5–7 mm thick. The central polythalamous, light brown, dense and hard, without separating the larval chambers, which are slightly ovoid (1 × 1.5 mm). The number of larval cameras ranges from 7–15.

  Biology: Only asexual females are known. The galls appear in August and the adults emerge in March.

  Host: It occurs on Quercus magnoliifolia Née (section Quercus), endemic species from México.

  Distribution: México (Jalisco)


description:
  "... was reared ... from larvae collected ... the previous year from a burrlike gall (cf. Gagne´ 1989) on the basal part of the leaf petiole of American chestnut, Castanea dentata (Fagaceae)."


externallink: https://www.biodiversitylibrary.org/page/33538414#page/19/mode/1up
description:
  [As Cecidomyia? castaneae]:

  "An ellipsoidal, monothalamous gall found singly or in small groups near the base of leaves of chestnut, Castanea dentata, projecting from under side of the principal veins, occasionally from the midrib, seeming to be an enlargement of the vein.... The surface is smooth like the surface of the leaf. The walls are succulent .... light yellow-green, a few tinged with red ... collected in June..."


description:
  "It infests leaves of Allegheny chinkapin, Castanea pumila (L.) Mill. (Fagaceae), in eastern North America."


description:
  Summer galls occur only on *Solidago patula* (Asteraceae) at the apex of the main stem. Galls typically occur in tight clusters but also as singlets. Adults emerge from these galls in August and females deposit their eggs in living meristematic tissue at the base of the host plant where they overwinter. The spring crown gall forms at this location.


description:
  ... females deposit their eggs in living meristematic tissue at the base of the host plant where they overwinter. The spring crown gall forms at this location.


description:
  This midge was reared in May, 1913, by Mr, P. H. Timberlake from a pubescent, variable, conical, stem gall on rabbit brush, Chrysothamnus graveolens [Ericameria nauseosa], collected in the vicinity of Murray, Utah. The species approaches closely R. utahensis, from which it may be most easily separated by its decidedly smaller size, the fewer antennal segments and particularly by the peculiar type of gall.
  Gall. — A variable, conical outgrowth of the stem with a height of about 4 mm. and a basal diameter of 2 to 3 mm. The gall contains basally, an oval, thin-walled cell with a length of 2.75 mm. and a diameter of 1.75 mm., the hollow, distal portion being filled with a slender, thread-like growth of delicate cells placed end to end and forming a rounded or, in some cases, nearly truncate plug. 


description:
  Rhopalomyia chrysothamni
  Stem-gall Midge

  This midge induces small, conical-tubular monothalamous galls that project out at right angles to the stems of rubber rabbitbrush.


description:
  Several specimens of this species found in New Mexico from October-January were lacking any hair, so it seems that as the galls age they lose their hairiness.  


description:
  This species of leafy mistletoe induces abrupt stem swellings on Forestiera pubescens, serviceberry. Galls are the same color and texture as the branch it grows on and mistletoe grows from the galls. This species also has been observed inducing galls on cultivated Eastern cottonwoods, but the galls are much smaller and more subtle than on Forestiera. 


description:
  Woolly white mass that occurs at the tip of Artemisia ludoviciana stem. Engulfs some leaves but some stick out from the sides of the gall.


description:
  Aciurina idahoensis
  Leafy-cone-gall Tephritid

  Galls have numerous relatively thin, leafy bracts (bud scales) that stick out and sometimes recurve from the main gall body.


description:
  Aciurina idahoensis Steyskal is a univoltine, monophagous fruit fly that forms distinctive, leafy, axillary bud galls on Chrysothamnus viscidiflorus (Hooker) Nuttall, its only known host plant, throughout the western United States.
  Wangberg (1981) remarked that "... this species was common throughout southern Idaho and often abundant at various sites, ..." and that its distribution "... appears continuous and co-extensive with its host plant." Aciurina idahoensis has only been found in Deep Spring Valley in California.
  The gall of A. idahoensis is a stunted axillary branch, or rarely a branch apex, reminescent of a pine cone (Steyskal 1984). As such, it is subspheroidal when young (Fig. 7A), elongating into an ovoid as it grows and matures (Fig. 7D, E). The gall also bears basally expanded, sessile, linearly tapered, and apically pointed, smooth-margined leaves, which are shortened and more densely packed apically (Fig. 7E). The oldest, basal, overwintered leaves were dead and tan (Fig. 7 A, B); the new leaf growth light green (Fig. 7B). The gall body is largely composed of whitish, expanded leaf bases, nodes, and expanded pith parenchyma (Fig. 7D, F). The gall is uniloculate (monothalmus) and characteristically contains a small, initially subspheroidal (Fig. 7C), later narrow, elongate-ellipsoidal, central open chamber within which the larva feeds (Fig. 7D). This gall is quite unlike the axillary bud galls of A. ferruginea and A. michaeli, both externally and internally (Goeden and Teerink 1996), and indeed, more closely resembles the galls of an apparently unnamed Procecidochares, misidentified as P. stonei Blanc and Foote (Tauber and Tauber 1968, Green et al. 1993), and certain Cecidomyiidae, all of which form fleshy, leafy, bud galls with narrow, elongate, cylindrical central cavities on C. viscidiflorus in southern California (Goeden and Teerink, unpublished data). 


description:
  Aciurina ferruginea
  Medusa-gall Tephritid

  The galls are characterized by numerous long, thin leafy bracts, which appear to develop in fascicles.


externallink: https://www.biodiversitylibrary.org/page/28254596#page/429/mode/1up
description:
  ...the gall of A. ferruginea is subspheroidal and covered by many thin, threadlike leaves, its shape apparently unlike other tephritid galls described to date.

  ...galls of both species were pictured by Wangberg (1981) from Idaho, so these two tephritids [A. michaeli and ferruginea] probably occur in sympatry throughout the range of C. viscidiflorus. 

  By this time, the galls already appeared very leafy and consisted of stunted axillary branches, which continued to produce leaves in fascicles from many tiny axillary buds as the galls grew (Fig. 8D). 

   Its inner surface is shallowly pitted and thinly coated with sap upon which the larvae principally feed, as described for A. thoracica by Headrick and Goeden (1993).


description:
  Valentibulla californica
  Elliptical-stem-gall Tephritid

  These stem-colored galls may taper gently or expand abruptly and may be symmetrical or lopsided.


externallink: https://www.biodiversitylibrary.org/page/16152770#page/569/mode/1up
description:
  The known distribution of V. californica includes the western third of the U.S. as mapped by Foote et al. (1993), who also noted that it had been collected in Baja California Norte. Its known hosts are Chrysothamnus nauseosus (Novak et al. 1967, Wasbauer 1972, Wangberg 1978, Foote et al. 1993) and C. parryi (Gray) Greene (Wangberg 1978), a high-altitude species in southern California (Munz 1974) which we have not examined for the presence of V. californica.
  The larvae develop in short, subsurface mines mostly formed basally in current season's branches, and are not easily detected on infested plants. They cause limited swellings, not readily detectable galls.
  However, unlike A. thoracica and contrary to Wangberg (1978), we could not bring ourselves to call the feeding sites of V. californica "galls." Instead, like Foote et al. (1993), we prefer the description, slight "swelling," or better yet, as Dodson (1987b) described the "gall" of V. dodsoni Foote "a rather inconspicuous swelling" of a branch, because this is the only external sign of the larva or puparium within (Fig. 5A, B, C, D, E).
  Masked by normal variations in thickness of uninfested branches and stems of C nauseosus at our study site, and by the conspicuous galls caused by other insects, e.g. Aciurina trixa Curran (Wangberg 1981, Dodson and George 1986, Dodson 1987b) and several species of Cecidomyiidae (Gagne 1989), the slight swellings symptomatic of V. californica are, indeed, "inconspicuous."


externallink: https://www.biodiversitylibrary.org/page/37074622#page/144/mode/1up
description:
  Disholcaspis mellifica, new species

  Host: Quercus garryana Dougl.

  Gall: Bursting out of cracks in the bark, usually in rows, like galls of Callirhytis excavata (Ashm.) but extending only one or two millimeters up above the flaps of bark. On current year's growth in fall, often on sprouts close to the ground. The top is flat, rugose, and secretes honeydew. Larval cell basal and occupying most of the interior, the only thick dense tissue being directly above. Single galls from above measure up to 3 by 5 mm., but those in a row are much distorted by mutual pressure. 

  Habitat: The types are from galls collected Sept. 8, 1922, above Cedar Creek checking station in Sequoia National Park, Calif. The galls then contained larvae and secreted so much honeydew that it dripped to the ground and in the sun was even evaporated to a white solid. Pupae were found in galls on October 1. Dead adults were cut out Mar. 17, 1923. This is the undescribed gall whose secretion has been described more fully by the author (Bull. Brooklyn Ent. Soc, vol. 20, p. 177, 1925).


description:
  Disholcaspis mamillana Weld, 1957, sexual generation

  [A photo of this gall appears on page 13 of the pdf]

  Gall (Fig. 20): Length 2.4–2.9mm, buff-colored, oblong grain-shaped, bluntly pointed, thin walled, and glabrous. Arising from the buds at the base of new leaves during spring flush. Emergence holes are rough edged and near the apex. 

  Host plant remarks: The recorded host for the asexual generation wasps is Q. douglasii (Weld 1957). All sexual generation specimens here identified as D. mamillana were taken from a field identified Q. douglasii. No host voucher is available for these specimens


description:
  Disholcaspis prehensa Weld, 1957, sexual generation

  [A photo of this gall appears on page 17 of the pdf]

  Gall (Fig. 35): Length 2.2–3.5mm, buff to light brown colored, oblong grain-shaped, bluntly or sharply pointed, thin walled, and glabrous. Arising from the buds at the base of new leaves and flowers when they flush in the spring. Emergence holes are rough edged and near the apex. 

  Host plant remarks: Quercus dumosa is the host plant listed in the original description of D. prehensa (Weld 1957). However, the name Q. dumosa has been applied to almost all species of scrub oak in central and southern California. Asexual generation D. prehensa have also been photo-documented (Gross J. 2007 and 2012) and reared from Q. berberidifolia (listed as host for specimen D_prehensa_Dispre3). The host plant voucher for the sexual generation specimens USNMENT00961761, USNMENT00961924, and USNMENT00961763 is deposited at MARY and identified as Q. berberidifolia. This shows that the sexual generation of D. prehensa is found on the same host species as the asexual generation. 


description:
  Disholcaspis erugomamma Cooke-McEwen, sp. nov.

  [A photo of this gall is included on page 19 of the pdf]

  Gall (Fig. 47): Reddish brown, globular with a short, obtusely pointed nipple apically or few may lack the nipple entirely. Dispersed or more commonly tightly clustered and deforming around each other. Internally with dense, rather hard, corky texture; larval cell not free in galls examined. Gall is most similar to that of D. quercusmamma, but the surface is wrinkled in that species and relatively smooth in D. erugomamma. It is uncertain if the galls are nectar secreting, but the black sooty mold that is typical for mature nectar secreting galls was lacking. The galls were covered in dust when found but it is uncertain if that dust was a trait of the gall itself as opposed to its dusty arid environment near a road.

  Biology: Only the asexual generation is known at this time. All wasps reared for this study emerged toward the end of December or early January. 

  Etymology: Named for the gall morphology. From Latin, erugo meaning to free of wrinkles (noun in apposition) and mamma as a reference to the mammiform gall shape (Brown 1956). 

  Distribution: Known only from the single collecting event in western Texas. Collecting site was revisited in spring 2020 and empty fall galls were present. 

  Host plant remarks: The host plant voucher is deposited at MARY and identified as Q. havardii. It has a small range from northwestern Texas to New Mexico and Oklahoma, with disjunct populations in northern Arizona and southern Utah (Nixon, 1997).


description:
  [The author mistakenly described this gall as being induced by Aciurina ferruginea, and his description perfectly matches what is now known as Aciurina michaeli] It is monothalamous, smooth, pear-shaped and occurs at the axillary buds on the current year's growth. A few modified leaves usually arise from the gall surface. The color of galls is green, becoming pale green to yellow with age. Some may have pinkish hues. Twenty-five mature galls collected near the Craters of the Moon National Monument, Butte County, in 1975 ranged from 8.5 to 16.0 mm (x = 12.9 mm) in length and 4.5 to 8.0 mm (x = 6.4 mm) in width. The young gall resembles the mature gall in form. The stem and most of the leaves that would normally arise from the axillary bud are incorporated as gall tissue. A few unmodified leaves may remain and grow from the surface. In some instances stem growth occurs from the gall and many leaves will develop from such a stem above the gall. Larval feeding appears to be the stimulus for A. ferruginea [A. michaeli] gall growth. Growth continues up to the pupal stage. If a larva dies early in development, the gall ceases growth. A mature gall shows evidence of desiccation. Following emergence of the gall-inhabitant the gall desiccates rapidly and will abscise from the host plant. No portions of the host plant's foliage appear suitable for gall growth as long as growing stems are present. Galls are usually concentrated on the lower half of branches rather than toward the tips.


externallink: https://www.biodiversitylibrary.org/item/43955#page/124/mode/1up
description:
  Holcaspis weldi, sp. nov.

  Gall. In clusters on the terminal twigs and leaves of Quercus reticulata. 

  Monothalamous. Globular, thin-shelled, yellowish and tinged various shades of pink and red, covered with a dense rusty brown, pubescence, which may be rubbed off with the fingers. Inside there is a rounded kernel held in position by radiating fibers which are covered with a moss-like substance. The inner side of the shell is densely covered with a rusty brown moss-like substance. Diameter 20-40 mm. 

  Habitat: Mexico. 

  The galls were purchased by Mr. Lewis H. Weld from a woman at a fruit stand on the street in the Mexico City and the specimens evidently came from the nearby mountains. The galls were bought September 12th, 1910 and the adults emerged between October 2nd and 9th. The oak was kindly identified for me by Dr. N. L. Britton.


description:
  Cynips (echinus) douglasii
  Agamic form douglasii

  Holcaspis douglasii Ashmead, 1896, Proc. U.S. Nat. Mus. 19:127. 
  Cynips echinus douglasii agamic form douglasii Kinsey, 1930, Ind. Univ. Studies 84-86:186, figs. 23, 154-159, 175. 
  Cynips quercus-echinus Douglasi Houard, 1935, Marcellia 28:108, fig. 40-43. 

  To the data given in Kinsey 1930, add: 

  RANGE.-California: Island Mountain (E. R. Leach in Kinsey coll.). Visalia (galls, C. D. Michener in Kinsey coll.). Santa Rosa (Weld acc. Houard 1935). 

  LIFE HISTORY..—Adults: December 8, 9-14, 16-30. Most of the emergence late in December. 

  The new locality in Trinity County is somewhat northwest of the previously known range for douglasii. The material may average darker than that from Central California. Twenty-one of the 684 insects are very small (1.5 mm. or less) and very dark, some of the specimens being almost entirely black. It remains to determine whether these are abnormally developed individuals, a local, mutant race, or stray individuals from an undescribed species of the complex.


description:
  Cynips (echinus) dumosae Kinsey 
  agamic form 

  Cynips echinus dumosae Kinsey, 1930, Ind. Univ. Studies 84-86:199, figs. 26, 147-149, 179. 
  Cynips echinus douglasi err. det. Hoffmeyer, 1933, Ent. Meddell. Kbhvn. 18: 248. 
  Cynips dumosae Kinsey, 1930, Workbook in Biol.: 35. 

  To the data given in Kinsey 1930, add: 

  RANGE.-California: Santa Anita Canyon (north of Monrovia) (C. D. Michener in Kinsey coll.). 

  LIFE HISTORY..—Adults: March 25, and April (without day). These dates are still later than the previously recorded dates for January and February. 

  PARASITE.-Callimome warreni Cockerell (acc. Hoffmeyer 1933)


description:
  Cynips (echinus) echinus 
  agamic form echinus Osten Sacken 

  Cynips quercus echinus Osten Sacken, 1870, Trans. Amer. Ent. Soc. 3:56. 
  Cynips echinus echinus agamic form echinus Kinsey, 1930, Ind. Univ. Studies 84-86:191, figs. 24, 156-158, 163, 176, 191.
  Diplolepis echina Balduf, 1932, Proc. U.S. Nat. Mus. 79 (28):60. 
  Cynips echinus Kinsey, 1934, Workbook in Biol.: 35. 

  To the data given in Kinsey 1930, add: 

  PARASITE.-Decatoma occidentalis Balduf (acc. Balduf 1932). 

  Cynips (echinus) vicina 
  agamic form vicina Kinsey 

  Cynips echinus vicina agamic form vicina Kinsey, 1930, Ind. Univ. Studies 84-86:196, figs. 25, 177. 
  Cynips vicina Kinsey, 1934, Workbook in Biol.: 36. 

  To the data given in Kinsey 1930, add: 

  RANGE.-California: Sequoia National Park (galls, C. D. Michener in Kinsey coll.). 

  LIFE HISTORY..—Galls: mature September 4. 


description:
  Cynips (dugèsi) brevipennata (Gillette) 
  agamic form 

  [No photos of this gall are included]

  [no name] Osten Sacken, 1873, Hayden Rpt. U.S. Geol. Surv.: 567, No. 1. Kinsey, 1926, Introd. Biol., fig. 277a. 
  Holcaspis brevipennata Gillette, 1893, Ent. News 4:31. Gillette, 1896, Trans. Amer. Ent. Soc. 23:93, 96. Dalla Torre and Kieffer, 1902, Gen. Ins. Hymen. Cynip.: 53. Beutenmüller, 1909, Journ. N.Y. Ent. Soc. 17:46. Beutenmüller, 1909, Bull. Amer. Mus. Nat. Hist. 26:42, pl. 9 figs. 4-5. Thompson, 1915, Amer. Ins. Galls: 17, 39. 
  Disholcaspis brevipennata Dalla Torre and Kieffer, 1910, Das Tierreich 24:373, 632, 634, 636, 811. Felt, 1918, N.Y. Mus. Bull. 200:100, fig. 63 (4-5). Kinsey, 1920, Bull. Amer. Mus. Nat. Hist. 42:398. Houard, 1928, Marcellia, 24:106. 
  Andricus pellucidus Kinsey, 1920, Bull. Amer. Mus. Nat. Hist. 42:309, 384, pl. 23 figs. 19-21. 
  Diplolepis brevipennata Weld, 1922 (not all records), Proc. U.S. Nat. Mus. 61 (18):7. Weld, 1926, Proc. U.S. Nat. Mus. 68 (10):19. 
  Cynips dugès i var. A Kinsey, 1927, Field and Lab. Man. in Biol.: 110. 
  Cynips dugès i brevipennata Kinsey, 1930, Ind. Univ. Studies 84-86:282, figs. 44, 262, 273, 278. 

  GALL.-Similar to all galls of the C. dugèsi and C. bella complexes; mature galls light rosy tan, unspotted; often shining; of moderate size, up to 19. mm., averaging nearer 14. mm. in diameter. 

  HOSTS.—White oaks of the Q. Gambelii complex (= Q. undulata Gillette and other older authors, not undulata of recent authors). Also on Q. fendleri (acc. Weld), and Q. grisea (acc. Weld). 

  RANGE.-Colorado: Manitou (Gillette; brevipennata types. Also galls, Kinsey coll.). Colorado Springs (Carpenter, pellucidus types). Colorado Springs, 25 SE (Holloway in Kinsey coll.). Happy Canyon (20 SE Denver). Palmer Lake (I. C. Campbell in Kinsey coll.). Trinidad (galls, Holloway in Kinsey coll. Also Weld). Morley, Wetmore, West Cliff, La Veta, Spanish Peaks (acc. Weld). 

  New Mexico: Raton, Wagon Mound, Shoemaker, Glorieta, Tijeras, Las Vegas Hot Springs (acc. Weld). Raton, 8 N (galls, Holloway in Kinsey coll.). Raton Pass (W. J. Hughes in Kinsey coll.). Raton, 28 E (C. Schwachheim in Kinsey coll.). Santa Fe, 12 E (galls, Holloway in Kinsey coll.). 

  Confined to an area in Colorado and New Mexico, east of the Continental Divide, extending from Denver, Colorado, south to a point (Tijeras) east of Albuquerque, N.M. Replaced in Colorado west of the Divide by C. cubitalis; replaced elsewhere in the Southwest by still other species of the dugès i complex. Figure 18. 

  INQUILINES.–Synergus similis Gillette (acc. Gillette 1896). S. atripes Gillette (acc. Gillette 1896). S. oneratus (Harris) (acc. Dalla Torre and Kieffer 1910). Doubtful determination. 

  LIFE HISTORY..—Larvae: August 22-25 (in most cases too small to breed). Pupae: October 4. Adults: in galls October 6, 31; November 9. Emerging October 31. November 15, 20, 26. December 15. January 10. Most of the emergence in November. See Kinsey 1930: 284 for further details. 

  This short-winged insect is the best-known species of the complex, occurring on the eastern front of the Rockies in Colorado and New Mexico in a region where no species of the Cynips bella complex is found. Brevipennata is, consequently, the only producer of the oak-apple galls in that area. 

  From the other distinctly short-winged species representing the dugèsi complex in the United States, brevipennata is distinct in having a generally bright rufous body color, a small but distinct areolet, and faint markings in the cubital cell. Of the two other short-winged species, cubitalis, in Colorado west of the Divide, has a nearly closed areolet and a nearly solid line of marks paralleling the cubital vein; and pupoides, in West Texas, has a generally dark rufous color with heavier marks in the cubital cell. These three short-winged insects must have originated one from the other, the first coming by direct mutation from such a long-winged type as C. simulatrix or C. subnigra in Southern Arizona and New Mexico. 


description:
  Cynips (dugèsi) capronae (Weld) 
  agamic form
  Figures 9, 18, 41, 56 

  Diplolepis capronae Weld, 1930, Proc. Ent. Soc. Wash. 32:29. 
  Diplolepis Capronae Houard, 1934, Marcellia 28:35, figs. 14, 15

  GALL.-Similar to other galls of the dugèsi and bella complexes, sometimes somewhat flattened basally, but sometimes with the vertical longer than the horizontal axis; up to 18. mm., averaging nearer 10. mm. in diameter; surface smooth and shining; older galls light brownish yellow, conspicuously striped with vertical, rarely anastomosing bands of purplish brown. Figures 9, 56. 

  HOST.-Quercus subturbinella, a desert scrub of Californian affinities. 

  RANGE.-Arizona: Camp Creek (Weld coll., types). Globe. Safford (galls only). Utah : Zion Park (questionable determ.). Possibly restricted to Quercus sub turbinella in an area in Central Ari zona, north of a line drawn from Safford to Phoenix. Possibly it is the same species ranging on the same oak into Southwestern Utah. Figure 18.

  LIFE HISTORY..—Immature larvae: September 8. 

  Adults mostly emerged November and December (acc. Weld), and January 20 (Kinsey). 

  In 1920 I found the conspicuously striped galls of this species abundant in the Apache Trail area of Central Arizona. The material from which Weld described the species was collected just north of that same area.

  From a gall which I collected in 1929 in the southwestern corner of Utah (Zion Park), I cut a well-developed adult which seems to differ from a paratype of capronae which Mr. Weld has lent me only in having darker wing veins and heavier spotting in the cubital cell. But in the original description it is noted that this spotting varies in the type series, and I should not question the identity of the Central Arizona and Southwestern Utah material if it were not that the gall from which my lone Utah insect was cut is without stripes, although somewhat marbled. My Utah specimens, however, were collected early in September, when the galls were very young and succulent; and it is possible (though by no means established) that the markings of the typical capronae gall do not develop until the structure is more mature. Weld's material and my 1920 material was collected when most of the insects had emerged, and my 1929 material when most of the larvae were too immature for breeding. We know some other cynipids which range from Central Arizona into Southwestern Utah, on Q. subturbinella, which is the host of capronae; but we cannot be entirely certain of the range of capronae until we can secure larger series of adults. 

  This oak on which capronae occurs is a peculiar, prickly leaved scrub that pushes further out onto the desert in Southwestern Utah and adjacent Arizona (and Nevada?) than any other oak which I know in the United States. All of the affinities of the oak are in California. Its isolated occurrence east of the Great Basin must be connected with the moist period which that region enjoyed during the late Pleistocene. At that time several members of the California flora and fauna advanced east ward across a region that is now too dry to support any oak. Every cynipid found on Quercus subturbinella is, therefore, of especial interest in the interpretation of our Southwestern faunas and floras. In this particular case the host is Californian, but the insect entirely Rocky Mountain in its affinities. It would appear that the morphologic mutations by which this insect was separated from the other complexes of Atrusca must have been accompanied by as unique physiologic mutation which led to the insect's selection of its host. Moreover, it isapparent that these mutations must be of recent date. They must have occurred since the above-mentioned period in the late Pleistocene. 

  Cynips capronae shows most of the characters found in the adjacent species of the dugèsi complex. It differs from most of the others in having a much shorter, more nearly triangulate hypopygial spine, a broad anterior depression of the scutellum which is divided by a fine but distinct carina, and a gall which is prominently striped. Note that the species occurs at one edge of the range of that complex, in an area and on an oak from which no other species of the complex is known. It is clearly a geographic segregate derived from C. simulatria by fairly direct though amaterial mutation. We have debated the desirability of recognizing these larger mutations by creating a distinct complex for capronae. If we did, it would certainly emphasize the fact that a complex may arise in the same fashion as species arise from each other within a group. To keep the species in the dugèsi complex is to emphasize the fact that degree of difference is no indicator of the number of times a species is removed from another in evolution. The phylogenetic interpretation seems clear; the manner in which we translate that story into the taxo nomic nomenclature is not so important. 

  Capronae is ruled out of the bella complex by its much lighter wing markings (including the virtual absence of spots in the radial cell), and by the important fact that there is a species, vanescens of the bella complex, in that same part of Arizona and Utah, on the same oak host.


description:
  Cynips (dugèsi) catena, new species
  agamic form

  GALL.-Similar to all galls of the C. dugèsi and C. bella complexes. Mature galls spherical, yellow-tan to rosy-tan, flushed rose when younger, touched with darker purple rose in age, rather well peppered with fine, purplish marks; more or less dull; up to 17. mm., averaging near 14. mm. in diameter. 

  HOST.-Quercus grisea, the evergreen white oak of the area. 

  RANGE.-Texas: Alpine, 10 E. Possibly restricted to the single, low mountain range east of Alpine. Figure 18.

  LIFE HISTORY..—Galls: nearly mature October 13. Adults: December 6, 14. January 9, 15. Most of the emergence in the first half of December. 

  It is a great surprise to find this distinct species within 14 miles of the type locality for C. pupoides. The types of pupoides were collected seventeen years ago in the brush-covered hills west of the town of Alpine. They are matched by material collected that year from Fort Davis and, within the last year, from the Chisos Mountain Park. The extreme localities for pupoides are fully 100 miles apart. The stands of oak in that area are anything but continuous, and in places they may be as much as 20 or 30 miles apart. The oak-free stretch east of the town of Alpine is hardly 8 miles wide, but it isolates the so-called Del Norte Mountains from which we have this perfectly distinct insect, catena. 

  While catena shows itself some relative of pupoides, it, as a matter of fact, is closer to the Central Texan C. cava than it is to the brevipennata-pupoides stock. The phylogenetic map will show how the brevipennata and cava stocks have come in from the west and east, respectively, to put their end products within 8miles of each other on either side of Alpine. Cateba differs from cava most obviously in its strictly spherical gall—the galls of cava being bulboid. The insect of cateba is generally darker rufous in color and has heavier wing veins.


description:
  Cynips (dugèsi) cava (Weld)
  agamic form

  Diplolepis cava Weld, 1926, Proc. U.S. Nat. Mus. 68 (10):22, fig. 5.
  Cynips cava Kinsey, 1930, Ind. Univ. Studies 84-86:293, figs. 46, 260-261,270, 283. Houard, 1934, Marcellia 28:38. Houard, 1935, Marcellia 28: 110

  GALL.-Similar to other galls of the C. bella and C. dugèsi complexes, but gourd-shaped instead of spherical, having a short, rather stout, drawn out, more or less cylindric base which is up to 3.0 mm. in length and 3.0 mm. in diameter; body of gall up to 22.0 mm. in diameter. Young galls light yellow with a rosy tinge, older galls becoming light russet brown, unspotted or only obscurely spotted. Figure 9. 

  HOST.-Quercus breviloba, Q. breviloba f. laceyi. 

  RANGE.-Texas: Austin (type, Weld coll.; also Patterson in Kinsey coll.). Round Rock. Leander. Boerne and Kerrville (acc. Weld 1926). Probably restricted to a limited area in Central Texas, extending mostly west from Austin. Figure 18

  LIFE HISTORY..—Galls: Young on June 7. Pupae: November 10, 20. Adults: November 25, 26. December 4, and before and after December 8. 

  The gall differs from those of all the northern species of dugèsi in being stalked to bulboid in shape. Taken together, these differences would seem enough to warrant recognition of the distinct complex in which we cataloged this species in our 1930 monograph. But the potentialities for all of the cava characters are in the dugèsi complex. This is evidenced by their appearance, singly, or in limited combinations, in one place or another within that complex; and the geographic position of cava also indicates that it is a geographic segregate of the dugèsi complex. Whatever we do with the species for the convenience of our cataloging, it seems certain that cava is, phylogenetically, no further removed from simulatria than simulatriæ is from the next species within the list. Such extreme developments of the dugèsi complex as cava, brevipennata, capronae, and others give evidence of the essential identity of the processes by which both species and higher categories come into existence. 

  Cynips catena is the closest relative of cava. Catena is clearly an offshoot of the same stock which has moved further west into a single mountain range east of the town of Alpine, in West Texas. Cava is distinct from catena in being brighter rufous in color, in having a shorter hypopygial spine, rather distinctly shorter wings, a slightly smaller areolet, and a stalked-bulboid gall. 


description:
  Cembrotia sp. 
  Needle-lip-gall Midge

  The midge induces short or stunted, bifurcated needle galls on Colorado pinyon pine from Colorado westward, with limited distribution in California in the eastern Mojave Desert.


description:
  Cynips (dugèsi) conexa, new species
  agamic form

  GALL.-Similar to all galls of the C. bella and C. dugèsi complexes. Mature galls in part light yellow, in part rosy or rosy brown, conspicuously marked with numerous, small, purplish spots, sometimes with a trace of a bluish puberulence; galls small, up to 18. mm., but averaging nearer 13. mm. in diameter. Figure 67. 

  HOST.-Quercus nudinervis. NOT on Q. macrophylla in the same locality. [these are both Q magnoliifolia in modern terms]

  RANGE.-Guerrero: Taxco, 8 NE, 8000' (types). Probably restricted to Q. mudimervis in the mountains of Northeastern Guerrero. Figure 18. 

  LIFE HISTORY..—Adults: April 2, 15. 

  In the mountains near Taxco, in Northeastern Guerrero, the prevailing oaks are Q. macrophylla and Q. nudinervis. Cynips spadia represents the dugèsi complex on Q. macrophylla; conexa is segregated on Q. nudinervis. Out of 64 insects from Q. nudinervis, all are typically conexa except one. 

  From all other galls of the dugèsi complex in Southern Mexico, conexa galls are distinguished by their conspicuous covering of small, purplish spots. 

  That conexa is a host-isolated relative of C. (dugèsi) spadia: near Taxco, in Guerrero, cannot be questioned. Its hypopygial spine, however, is some departure from the normal for the sub genus Atrusca, although it is not so elongated that one would consider placing it in another complex. Conexa is, however, also close to spiculi (near Mexico City), although this later insect has a still longer spine which might well lead to its placement in another complex or subgenus. 


description:
  Cynips (dugèsi) cubitalis, new species
  agamic form

  GALL.-Similar to all galls of the C. dugèsi and C. bella complexes. Mature galls light rosy tan, unspotted; often shining; of moderate size, up to 22. mm., averaging nearer 16. mm. in diameter. Figure 52. 

  HOSTS.—Several oaks of the Q. Gambelii complex. 

  RANGE.-Colorado: Glenwood Springs. Aspen, 6 E. Gunnison, 20 W (galls only). Colona. Ouray, 3 S. Durango, 12 N (types). Mesa Verde. Lewis. Utah : Monticello, 10 E (galls only). La Sal (galls only). Probably restricted to an area in the Rocky Mountains west of the Continental Divide, known from Western Colorado and easternmost Utah. Figure 18.

  LIFE HISTORY.—Larvae: August 23 to September 2, in most cases too young to breed. Adults: October 29, November 4, 6, 9, 20, 21, 24. 

  This short-winged species, cubitalis, is the producer of all the oak apples found in Colorado west of the Continental Divide. There is no species of the Cynips bella complex in that area. On the eastern slopes of the Colorado Rockies oak climbs hardly above 6,000 feet, falling some 4,000 feet short of most of the passes. On the western slopes, another oak of the Gambelii complex climbs much higher, reaching to about 9,500 feet, although it also falls short of the passes. The Western Colorado cynipids are, in consequence, well isolated from the Eastern Colorado species by a considerable oak-free area which extends over the Divide and well down on the eastern slopes of the range. Cubitalis is best distinguished by its nearly closed areolet and by the fusion of the marks in the cubital cell which form a line parallel to the cubital vein. Although the insect is in other respects hardly separable from brevipenmata, the characters noted are constant throughout the several localities, well distributed over the western half of Colorado, from which we have bred insects. 


description:
  Cynips (dugèsi) deceptrix, new species
  agamic form

  GALL.-Similar to all galls of the C. bella and C. dugès i complexes. Mature galls rosy or brownish tan, unspotted; always dull, minutely rough ened; moderately large, up to 22. mm., averaging under 17. mm. in diameter. Figure 70. 

  HOSTS.—Quercus undata, Q. chihuahuensis. Insects from other oaks, especially the dwarf oaks, Q. depressipes and Q. striatula in the area, not clearly belonging to this species. 

  RANGE.-Chihuahua: Parral, 12 W, 6800' (Q. undata; also galls on Q. saca me). Parral, 20 S (Q. undata). 

  Durango: Villa Campos, 21 S, 6700' (Q. undata). Indé, 20 E, 6300' (Q. chihuahuensis). Patos, 15 W, 8000' (Q. sacame; also galls, Q. chihuahuen sis, Q. durangensis). Otinapa, 8500' (galls, Q. reticulata, Q. durangensis). Canatlan, 7 N, 7400' (Q. undata, types; also on Q. chihuahuensis). Durango, 2 N, 6500' (Q. undata). 

  Zacatecas: Cantuna, 6 N, 6700' (Q. undata)

  Probably restricted to the Western Sierra of Mexico, centering in the state of Durango, and extending northward only into the edge of Chihuahua and southward into the edge of Zacatecas. Figure 18

  LIFE HISTORY.—Adults: November 9, 10, 17. December 5. February 1, 5, 8, 10, 18, 22, 26. March 2. Most of the emergence in the first half of February. 

  The range of deceptrix centers in the state of Durango. The break between its range and that of emergens to the north occurs in the desert area between Chihuahua City and Parral, in the southern end of the state of Chihuahua. The southern limits of the range are marked by the high grass plateaus and astonishingly barren mountains of more northern Zacatecas. Through its range, deceptrix occurs on the common oaks of the region, both Q. undata and Q. chihuahuensis, but insects which we have from the dwarf oaks, Q. depressipes and Q. striatula, from Patos and Otinapa in Durango, do not appear to be deceptrix. Our limited material from the dwarfs is, however, not in condition to warrant description. 

  Deceptrix is distinguished from its very close relative, emergens, by its more rufous color, especially its antennae which are largely bright rufous. From all of the more southern species of the complex, it is more strikingly distinct. 

  In the northern half of the range of deceptrix, C. (bella) aspera produces an oak-apple gall which, however, is usually to be distinguished from the large, dull-surfaced, unspotted gall of deceptrix; for the gall of C. (bella) aspera is smaller and more heavily mottled.


description:
  Cynips (dugèsi) dugèsi (Mayr)
  agamic form

  Dryophanta Dugès i Mayr, 1886, Verh. zoo.-bot. Ges. Wien 36:370, pl. 12 fig. 1a, 1b, 2. Mayr, 1902, Verh. zoo.-bot. Ges. Wien 52:290. Dalla Torre and Kieffer, 1902, Gen. Ins. Hymen. Cynip.: 52.
  Dryophanta dugesii Dalla Torre, 1893, Cat. Hymen. 2:50.
  Dryophanta Dugesii Kieffer, 1901, André Hymen. d’Europe 7 (1):621.
  Diplolepis dugesi Dalla Torre and Kieffer, 1910, Das Tierreich 24:344, 355, 811; figs. 76-77.
  Dryophanta Dugesi Küster, 1911, Die Gallen der Pflanzen: 170.
  Dryophanta dugesi Beutenmüller, 1911 (in small part only), Bull. Amer. Mus. Nat. Hist. 30:345, pl. 12 figs. 6-7. Felt, 1918, N.Y. Mus. Bull. 200:98, fig. 97 (6-7).
  Cynips dugèsi dugèsi Kinsey, 1930 (in small part only), Ind. Univ. Studies 84–86: 281-282.

  GALL.-Similar to all galls of the C. bella and C. dugèsi complexes. Mature galls rosy tan, unspotted; weakly shining; of moderate size, up to 21. mm., averaging nearer 17. mm. in diameter. Figure 71.

  Original description Translation: The gall, which is produced by an as yet undescribed species of cynipid, is found on Quercus mexicana Humbolt and Bonplandt, and belongs to that group of American leaf galls which are connected with the under surfaces of the leaves only by a single point, have thin walls, and show in cross-section a central, egg-shaped larval cell from which radiating threads go to the inner surfaces of the outer walls of the gall. It is entirely spherical and measures 6. to 15. mm. in diameter; has a very thin wall which is for the most part not shining, largely a beautiful red in color, in part light yellow, or even with a light yellow ground finely marbled red; in the early stages it is covered with soft and short downy hairs which later must fall off because of the interference of the enlarging galls with each other or through rubbing. By cutting open the gall, one sees numerous, very fine, radially placed hairs extending from the centrally-placed, monothalamous larval cell to the outer wall, while, for instance in the galls of Dryophanta bella Bassett and D. polita Bassett these hairs are not very numerous and more or less fine. If one directs the cut through the new species so that it does not go through the larval cell, one may never or rarely see the cell because of the many hairs arising from it (see the illustration showing the cross-section of the gall).

  HOST.S.–G) wercus microphylla, an alpine, dwarf white oak. Less often on Q. jaralensis, a scrub white oak. Not on the black oaks, Q. mexicana or Q. hypoleuca to which the types were mistakenly ascribed. 

  RANGE.-Guanajuato: León, 20 NE, 9000' (Q. microphylla, Q. jara lensis). San Felipe, 20 SW, 8000' (Q. fara lensis). Type locality not known, but probably from near León. Chiefly restricted to the alpine dwarf oak, Q. microphylla, in that por tion of the Western Sierra of Mexico which centers in Guanajuato. Figure 18.

  LIFE HISTORY..—Adults: February 15, 16, 20. March 2. 

  All previous determinations of insect and gall material as Cynips dugèsi (in Beutenmüller 1911, Kinsey 1930, etc.) prove to have been incorrect. Through the courtesy of Dr. F. Maidl of the Vienna Museum I have now been able to examine a type insect and galls, and discover dugèsi to be an insect which we, in our own collections, found nearly confined to the little alpine dwarf, Q. microphylla, in the high mountains which lie northeast of León, in the Central Mexican Sierra. The twigs and leaves with the type galls have been variously mis-determined as belonging to the black oak, Quercus mexicana, or to the black oak, Q. hypoleuca ; but to one who has studied Mexican oaks in the field they are unmistakably those of Quercus microphylla, the same host on which we found the insect. Although the type collection was without locality record, other than “Mexico,” Dr. Dugès lived in León, and it is not improbable that his collections were made in the same high mountains from which we collected the species just northeast of León. We also have the insect from nearer San Felipe, also in the state of Guanajuato, but found it nowhere outside of that state. 

  Practically all of the insects bred from all of the other oaks which occur in the Sierra near León represent Cynips (dugèsi) vulgata, but a few of the insects from the scrub oak, Q. jaralensis, prove to be C. dugèsi, indicating that the host isolation which normally segregates dugèsi on Q. microphylla may, on occasion, break down. No species of the C. bella complex occurs anywhere near the area in which dugèsi is found.


description:
  Cynips (dugèsi) emergens, new species
  agamic form

  GALLS.–Similar to all galls of the C. bella and C. dugèsi complexes; apparently indistinguishable from galls of C. (bella) bella which occur in the northern half of the range of emergens. Mature galls rosy or brownish tan, unspotted; more often dull (?); often large, up to 22. mm., averaging near 18. mm. in diameter. Figure 60. 

  HOST.S. Quercus undata, Q. chihuahuensis, Q. sacame [arizonica], Q. Gambelii. Apparently on all the oaks of the area. 

  RANGE.-Chihuahua: Pacheco, 13 NE, 7400' (Q. Gambelii, Q. saca me). Pacheco, 20 E, 5400' (Q. saca me). San Buenaventura, 3 E, 5600' (Q. chihuahuensis, Q. saca me). Las Cruces, 15 N, 6500' (Q. chihuahuensis). Namiquipa, 30 W, 5200 (Q. chihuahuensis). Madera, 10 SE (Q. chihua huensis, galls only). Matachic, 3 N, 6700' (Q. chihuahuensis). Pedernales, 2 E, 7500' (Q. chihuahuensis). Santa ſsabel, 6 E, 6000' (Q. undata; types). Santa ſsabel, 16 NE, 6000' (Q. undata; galls, Q. chihuahuensis). Probably restricted to the Western Sierra of Mexico in the state of Chihuahua, from Santa Ísabel north. Figure 18.

  LIFE HISTORY..—Adults: December 10. January 20, 25, 29, 30. February 1, 5, 6, 9, 10, 17, 18, 28. Most of the emergence late in January and early in February. 

  Emergens, distributed throughout the Sierra of Chihuahua, from Santa Ísabel and north, is one of the three closely related species which as a group range through our Southwest and south ward through the Mexican state of Durango. From the American species simulatrix, emergens is distinguished by its heavier veins and larger areolet; from deceptrix in Durango it is distinct in having antennae which are dark even on the basal segments. The break between the ranges of the two Mexican species comes in the hundred miles (largely desert and entirely oak-free, as far as we saw them) between Chihuahua City and Parral, in the southern end of the state of Chihuahua. Some of the Parral specimens were not clearly assignable to one or the other species, indicating some hybridization of emergens and deceptrix in that area, but only a few of the several hundred individuals in our collections were difficult to place. 

  It must be noted that the galls of this species are apparently indistinguishable from the galls of C. (bella) bella, which occur in the northern part of the range of emergens (from Las Cruces, Chihuahua, and north). The galls of C. (bella) pomifera, which occur in the southern portion of the range of emergens, are prominently mottled and hence almost always separable from those of emergens. It is interesting to find that the ranges of the species in the bella complex in this area do not coincide with the ranges of the species of the dugèsi complex. 


description:
  Cynips (dugèsi) longa, new species
  agamic form

  GALL.-Similar to all galls in the C. bella and C. dugèsi complexes. Mature galls very light olive tan, less often rosy, unspotted; slightly tuberculate, not pubescent, of moderate size (never large), up to 18. mm., averaging nearer 15. mm. in diameter. Figure 58.

  HOST.-Quercus rhodophlebia. Not on the dwarf oak, Q. repanda, in the same area.

  RANGE.-Hidalgo: Pachuca, 7 N, 8700' (types). Probably throughout the mountains rimming the Valley of Mexico,
  possibly occurring further east and north, but not southwest of those mountains. Figure 18.

  LIFE HISTORY..—Adults: March 6, 18.

  In the mountains to the north of the Valley of Mexico, this insect produces the common oak apples on the taller tree oaks, but not on the little alpine dwarf, Q. repanda. See C. pumilio, the species on the dwarf oak, for a further discussion. 

  Longa is remarkable in having a nearly normal radial cell. In this respect, the insect is matched only by its Guatemalan relative, C. luminata. This normal cell is, of course, consequent on the fully normal wing of the insect (awing-body ratio of 1.35). The insect is probably close to the place of origin of the whole complex. For a further discussion of this point, see the introduction to the whole dugèsi complex. 

  The galls of longa are slightly more distinctive than is usual in this complex, being more olive tan in color, with a minutely tuberculate surface. 

  No species of the C. bella complex is found near the range of C. longa. 


description:
  Rhopalomyia medusa
  Medusa-gall Midge 

  A subtle characteristic of these galls that separates them from galls of R. medusirassa (Plate 534) is the reddish-brown, curled leafy bracts that stand above underlying hairs covering the gall body. 


description:
  Rhopalomyia sp. A.
  Spiny-cone Gall Midge

  The flat top and green, pointed, recurved bracts, which are sparsely arranged along the sides, distinguish this gall.


description:
  Rhopalomyia ampullaria 
  Urn Gall Midge

  These tubular galls may occur singly or in coalesced clusters of two or three, usually protruding at right angles from the host leaf. 


description:
  Rhopalomyia anthoides 
  Leaf-bud Gall Midge

  This midge induces monothalamous bud galls on Great Basin sagebrush that are composed of numerous compact, narrow bracts. 


description:
  Rhopalomyia calvipomum
  Plum Gall Midge

  This midge induces ovoid monothalamous leaf galls that protrude mostly from the lower surface of leaves of Great Basin sagebrush.


description:
  Atrusca dumosae Melika & Pujade-Villar, new species
  Asexual female

  [Photos of the gall appear on page 7 of the pdf]

  Gall (Fig. 14). Galls are nearly spherical, slightly tapering at the base, monolocular, located usually on the underside of leaves, attached by a single fine point to the main vein. Young galls are greenish to yellowish; mature galls brownish tan or dark reddish brown, unspotted or heavily mottled with brownish purple. The surface of the gall smooth, only microscopically roughened, shiny, thin-walled, with radiating fine filaments, supporting the central larval chamber. Galls are 18-25 cm in diameter.

  Biology. The asexual generation is only known, inducing galls on Quercus dumosa Nutt [this ID is likely mistaken; Quercus dumosa does not occur at the type locality, and similar galls have been observed on Quercus turbinella, which does]. (section Quercus of Quercus, white oaks), which is distributed only in California (USA) and Baja California Norte, Mexico (Govaerts & Frodin 1998). Galls were collected in late October and beginning of November. Adults emerged right after galls were collected. 

  Etymology. The species is named after the host oak, Q. dumosa Nutt., it was found on. 

  Distribution. Currently known from the type locality only: Mexico, Agua Amarilla, San Pedro Mártir, Baja California Norte [N 32°08´28´´ and E 115°56´08´´], 1729 m a.s.l. This is the first species from the genus Atrusca known from Baja California, Mexico.


description:
  Cynips (dugèsi) lucaris, new species
  agamic form

  GALL.-Similar to all galls of the C. bella and C. dugèsi complexes. Mature galls light yellow to rosy tan, flushed rose when younger, darkening with age; up to 28. mm., averaging nearer 14. mm. in diameter. 

  HOST.-Quercus pilicaulis, a widespread tree with smoother, obovate, and crenate leaves—a “chestnut oak.” 

  RANGE.-Guatemala: Huehuetenango, 14 S, 7500'. Huehuetenango, 3 S, 7300'. Quiché, 2 N, 7500'. Sololá, 11 N, 8500': Sacapulas, 9 S, 6000' (galls). Guatemala City, 5 W, 6500'. Restricted to the one oak, but probably ranging throughout the high mountain mass of Southwestern Guatemala; possibly extending into the connecting mountains of Chiapas. Figure 18.

  LIFE HISTORY..—Adults: January 7, 8, 14, 15. February 1. 

  This insect and C. luminata are responsible for the oak apples of the main mountain mass of Southwestern Guatemala. Lucaris is confined to the smoother, crenate-leaved oak of the region, Q. pilicaulis. Luminata is in the same localities on the coarser leaved oak of the decipiens complex. The host of luminata usually occurs some hundreds or thousands of feet higher than the host of lucaris in the same mountain ranges. Both of the insects belong to the main line of the dugèsi complex, its geographically closest relative among our described species being C. longa from the southern end of the state of Hidalgo in Mexico. These Guatemalan insects are not related to the long-spined species of the complex. 

  Lucaris is readily distinguished from luminata by its generally dark color, especially by the large amount of black on the thorax and abdomen, by its slightly shortened wing with its com plex spotting in the cubital cell, the heavy blotch at the base of the cubital cell, the heavy infuscation and frequent spot in the radial cell, the shorter length of the whole insect (which averages near 2.9 mm.), and the smaller gall (which averages only about 1.4 mm.). 


description:
  Cynips (dugèsi) luminata, new species
  agamic form

  GALL.-Similar to all galls of the C. dugès i complex. Mature galls light rosy tan to bright rose, becoming light to dirty brown with age; some times unspotted, sometimes obscurely spotted or mottled with purplish; fairly large galls up to 32. mm., averaging nearer 22. mm. in diameter.

  HOST.-Quercus decipiens [rugosa], the white oak with the largest and coarsest leaves in the region. 

  RANGE.-Guatemala: Huehuetenango, 3 S, 7300'. Huehuetenango, 14 S, 7500': Quezaltenango, 5 N, 8800'. Sololá, 11 N, 8500'. San Marcos, 5 E, 8000'. San Marcos, 3 W, 8200'. Restricted to the one oak, but probably ranging throughout the high mountain mass of Southwestern Guatemala; possibly extending into the connecting mountains of Chiapas. Figure 18.

  LIFE HISTORY.-Galls still soft, not wholly mature in last week of December. ADULTS: January 7, 8, 14, 21, 29. 

  This insect and C. lucaris account for the oak apples of the main mountain mass of Southwestern Guatemala. Luminata is confined to the coarse-leaved oak which belongs to the Q, decipiens complex. Lucaris is on the “chestnut oak,” Q. pilicaulis, in the same area, although pilicaulis usually occurs a few hundred or thousand feet below the other oak. Both of the insects show relations to C. longa, from the southern end of the state of Hidalgo, north of Mexico City, nearly 300 miles from the Guate malan border. They are not closely related to the long-spined insects which break off of the dugèsi stocks near Mexico City. 


description:
  Cynips (dugèsi) occidua, new species
  agamic form

  GALL.-Similar to all galls of the C. bella and C. dugèsi complexes. Mature galls yellowish or bright rosy tan, younger galls richer rose; unspotted; smooth and shining to dull matte; very thin walled; moderate to very large, up to 24. mm., averaging near 21. mm. in diameter. Figure 57. 

  HOSTS.—Quercus depressipes, Q. potosina. Probably restricted to scrub oaks. 

  RANGE.-Zacatecas: Zacatecas, 22 SW (galls only, Q. potosima). La Mesa, 7000' (types, Q. depressipes). Probably restricted to an area in the Western Sierra of Mexico, in Zacatecas (and the adjacent corner of Ja lisco?). Figure 18. 

  LIFE HISTORY.—Adults: November 21, 24, 29, 30. January 10. February 5, 20, 24. Most of the galls gathered on November 21 still too young to breed. Most of the emergence in February. 

  This insect is known from only two collections on the high plateau that lies southwest of the city of Zacatecas. Whether the species is a geographic segregate occurring on all of the oaks of the region, or whether it is restricted to the alpine scrub oaks, remains to be discovered. Its closest relatives are oriens, found in the eastern mountains of San Luis Potosí, and vasta and oriunda in the eastern Sierra of Tamaulipas. From all of these others occidua is distinguishable by its nearly uniform, bright brownish rufous. From C. vulgata, the next more southern species in the Western Sierra, occidua differs in its shorter wing body ratio (1.15), in its lack of black on the thoracic lines, and in several other characters. 

  No species of the C. bella complex is known from the area in which occidua occurs, so galls alone are determinable in this area.


description:
  Cynips (dugèsi) oriens, new species
  agamic form

  GALL. Similar to all galls in the C. bella and C. dugèsi complexes. Mature galls rosy tan or brighter rose, unspotted; usually not large, up to 21. mm., but averaging nearer 14. mm. in diameter. Figure 53

  HOSTS.—Quercus potosina, Q. jaralensis [potosina], Q. intricata, Q. Pringlei, and Q. cordifolia, all of which are scrub oaks, though not all closely related oaks. Also on the larger trees, Q. polymorpha (?) and Q. sacame [arizonica]. 

  RANGE.-San Luis Potosí : Cerritos, 21 W, 6000' (types). Tamau lipas : Miquihuana, 7 SE, 6000'. Probably restricted to scrub oaks in an area in the Eastern Sierra of Mexico, centering in the eastern part of San Luis Potosí and an adjacent southern portion of Tamaulipas. Figure 18. 

  LIFE HISTORY.—Adults: November 23, 27, 30. December 6, 7, 12, 14, 15, 21. January 2, 9, 10. February 1, 5, 15, 24, 25, 26, 27, 28. March 1, 15. The bulk of the emergence in January and throughout February. 

  Cynips oriens, known from the Eastern Sierra of Mexico near Cerritos, S.L.P., and from the adjacent mountains of Southern Tamaulipas, has its closest relatives in C. vasta, C. occidua, and C. oriunda. Oriens, vasta, and occidua occur on scrub oaks, while oriunda occurs on the large tree, Q. polymorpha. Vasta is found in a more eastern ridge of the Eastern Sierra (near Tula, in Tamaulipas) about 40 miles northeast of Cerritos. The closest relative of oriens is oriunda, from the tall tree, Q. polymorpha, in the southwestern portion of Tamaulipas. 

  Thirty-five miles south of Cerritos, in the Eastern Sierra near Rio Verde, we found C. (dugèsi) pulea, an insect with a much reduced wing averaging only 0.90 times the body in length. In that group with reduced wings are the species puler, pumilio, pusa, and pictor, ranging through a great length of the Eastern Sierra both north, south, and east of oriens. From all of these oriens is distinct in its wing-body ratio of 1.10; but since the galls of both groups of species are essentially the same, insects must be had before determinations can be made. No species of the bella complex is known to occur near the range of oriens. 

  Cynips (dugèsi) oriunda, new species
  agamic form

  GALL.-Similar to all galls of the C. bella and C. dugès i complexes. Mature galls yellow or more rosy tan, unspotted, up to 12. mm. in diameter. 

  HOST.-Quercus polymorpha. 

  RANGE.--Tamaulipas: Ciudad Victoria, 21 W, 4500'. San Luis Potosí: Cerritos, 25 W, 6000' (galls only). Probably restricted to the one oak, Q. polymorpha, in the Eastern Sierra of Southern Tamauli pas and adjacent San Luis Potosí. Figure 18. 

  LIFE HISTORY..—Adult: December 7

  This insect is very close to oriens which occurs on the scrub oaks of an adjacent portion of the Eastern Sierra. Oriunda is probably confined to the larger and quite distinct oak, Q. polymorpha.


description:
  Cynips (dugèsi) pictor, new species
  agamic form

  GALL.-Similar to all galls of the C. bella and C. dugès i complexes. Mature galls rather rich reddish or brownish tan, lightly but in most cases distinctly spotted; not shining; moderately small, up to 17. mm., averaging nearer 12. mm. in diameter. 

  HOSTS.—Quercus cordifolia [striatula], Q. Pringlei. At the type locality (Miqui huana, 25 NW, Tams.) the first is the very dwarf scrub oak at the highest elevations (8600' or more); and the second is the head-high or taller shrub covering the slopes (7500' to 8500'). 

  RANGE.--Tamaulipas: Miquihuana, 25 NW, 8500'. Probably extending northward, from west of Miquihuana, on the west ern slopes of the range of the Eastern Sierra which marks the southern portion of the boundary between Tamaulipas and Nuevo Leon. Replaced by C. oriens on the eastern slopes of the same range (nearer and east of Miquihuana). Figure 18. 

  LIFE HISTORY..—Adults: November 26. December 6, 7, 14. January 1, 2, 8, 9, 16, 17, 23. February 16. Most of the emergence between Decem ber 6 and January 2. 

  In the remote and inaccessible area which forms the very high mountain boundary between the southern extensions of the states of Nuevo Leon and Tamaulipas, the present species, Cynips pictor, accounts for the oak apples on the western slopes of the Divide west of the town of Miquihuana. Directly east of the Divide, with its elevation of around 9,000 feet, it is C. oriens which represents this complex; and in an isolated canyon a little further east (22 miles north of Tula), C. vasta is the species producing similar oak apples. Our material of pictor comes from a single locality which is between the lumber station and mill of the hacienda known as “Aceradera”; but the range continues northward for some distance, past Soledad and west of Galeana, and pictor may extend for a hundred miles over the western slopes of this very high and inaccessible range. There seems no constant distinction between the material from the very low, alpine scrub, Q. cordifolia, and the taller shrub of the slopes, Q. Pringlei. 

  This insect is one of the Eastern Sierran, short-winged group of species which includes C. pumilio, C. pulex, and C. pusa. Pictor is the most northern member of the group. From pumilio, which occurs in Southern Hidalgo, pictor is further to be distinguished by having a gall which is distinctly even if lightly spotted. Pictor, puler, and pusa are nearer the same color and are similarly hairy on the thorax, but pictor is dis tinguishable by its peculiar venation and spotted galls. 

  Cynips (dugèsi) pulex, new species
  agamic form

  GALL.-Similar to all galls of the C. bella and C. dugès i complexes. Mature galls light yellow tan, unspotted; rather small, up to 14. mm., aver aging near 12. mm. in diameter. Figure 65.

  HOSTS.—Quercus macrophylla, Q. potosina.

  RANGE.-San Luis Potosí: Rio Verde, 14 W, 6500' (Q. macrophylla, types. Also on Q. polymorpha and Q. potosina). Not occurring to the north or west of the known locality; possibly occurring further east or south; possibly limited to the mountains west of Rio Verde. Figure 18.

  LIFE HISTORY.—Adults: Early spring, before March 2

  The type localities of pulex (from Rio Verde, in the state of San Luis Potosí) and of pusa (from Jacala, in Northern Hidalgo) are nearly a hundred miles apart, and it is nearer 125 miles to the type locality of pictor to the north (in Southwestern Tamaulipas). It is not impossible that additional species of the same group may occur in the largely impassable mountain areas that lie between Rio Verde and the nearest localities for these other species.

  The type locality of pulex is only 35 miles from the type locality of the longer-winged species oriens (from Cerritos, also in San Luis Potosí); but the affinities of pulex are entirely with pictor, pusa, and pumilio, and not with oriens. Pulea is represented in our collections from the low trees, Q. macrophylla, the large tree, Q. polymorpha, and from the quite unrelated scrub oak, Q. potosina.

  No species of the C. bella complex occurs near the area in which C. pulex is found.

  Cynips (dugèsi) pusa, new species
  agamic form

  GALL-Similar to all galls of the C. bella and C. dugèsi complexes. Mature galls light olive tan, sometimes browner, but not rosy; unspotted; medium in size, up to 16. mm., averaging 13. mm. in diameter.

  HOST.-Quercus repanda var., the lowest dwarf oak encountered along the Laredo-Mexico City highway east and west of Jacala.

  RANGE.-Hidalgo: Jacala, 7 NE, 5000'. Jacala, 24 SW, 6800". Probably restricted to a portion of the Eastern Sierra in the northern end of the state of Hidalgo. Figure 18

  LIFE HISTORY..—Adults: December 10, 14. January 1

  This short-winged insect forms the common oak apples on the low-tufted dwarf oaks that are to be found both northeast and southwest of the town of Jacala, in the high mountain mass negotiated by the new Laredo-Mexico City highway. This is the main range of the Eastern Sierra, and so continuous that the species may spread for a good many miles. Its closest relative is C. pulex, found in an isolated range west of Rio Verde—nearly a hundred miles by airline from the type locality of pusa. Pulex: and pusa are clearly in the same evolutionary line as pumilio, which is found on Q. repanda, a close relative of the host of pusa, in the mountains at the southern end of the state of Hidalgo. The known localities for pusa and pumilio are still 50 miles apart. The intervening mountain mass is largely inaccessible, for it is avoided by the built highways, and the cynipid fauna between the recorded localities for the two species remains unknown and unpredictable.

  The galls of pusa are distinctly olive, instead of rosy tan as in pumilio or yellow tan as in pulex.

  Cynips (dugèsi) pumilio, new species
  agamic form

  GALL.—Similar to all galls of the C. bella and C. dugès i complexes. Mature galls light rosy tan, unspotted; always smooth and shining, large, up to 25. mm., averaging nearer 18. mm. in diameter. Figure 59.

  HOST.-Quercus repanda, a low, alpine dwarf oak. Not on the taller oaks of the same area.

  RANGE.-Hidalgo: Pachuca, 7 N, 8700'. Mexico: Mexico City, 32 E, 10,600'. Mexico City, 12 W, 8500'. Mexico City, 20 S, 9400' (galls only). Puebla: El Seco, 5 SW, 8200' (gall). Michoacán: Morelia, 14 E, 7000'. Probably wherever the alpine dwarf, Q. repanda, occurs. The recorded range, from Morelia to Pachuca and the mountains of Puebla represents the known range of the oak, but it may occur more widely, especially east and north. Figure 18

  LIFE HISTORY..—Adults: January 12, 15, 25. February 2. March 7, 17, 25. Most of the emergence in March.

  This is the short-winged producer of the oak apples on the dwarf oaks of the Mexican Cordillera, from Morelia to Pachuca and the mountains of Puebla. It may occur still further north and east. On the taller oaks in the mountains rimming the Valley of Mexico, in galls which may hang square over the car pet of dwarf oaks which bear Cynips pumilio, there is a long winged member of the dugèsi complex, namely Cynips longa. The galls of the two are nearly identical and one, gathering them without separating the material from the two hosts, might be surprised to find the longest-winged species in the complex, and the smallest-known dwarf in the complex coming from the one collection. It must convince one that the long-winged and short-winged species are relatives, and demonstrate the significance of mutation and isolation in the origin of species.

  Pumilio is one of a group of short-winged species which also includes pictor, pusa, and pulex—all of them found in the East ern Sierra of Mexico. Geographically the species comes nearest C. pusa, pumilio being recorded from the southern end of the state of Hidalgo, while pusa is found in the northern end of the same state. Both insects occur on the same, very dwarf oak, Q. repanda, or species of that complex of oaks. The main highway connecting the two ends of the state of Hidalgo largely avoids the oak-inhabited mountain area, so we do not know whether any other species of this short-winged group occur between the ranges of pumilio and pusa.

  No species of the C. bella complex occurs near the range of C. pumilio.


description:
  Elongate swellings along the lamina of leaves of Salix amygdaloides, parallel with the midrib or lateral veins, with deep incisions in the surface of the gall where they are crossed by lateral veins, similar to those observed on [Euura proxima](https://www.gallformers.org/gall/3629). Observed in Minnesota in July.


description:
  Euura lasiolepis Smith

  The stem galling sawfly, Euura lasiolepis Smith, hereafter called the stem galler, oviposits in the petioles of very young leaves and initiates a gall in the stem below the leaf. The mature gall is a greatly swollen portion of the new shoot (Price and Craig 1984). Larvae feed in tissue derived from the pith in the developing stem (Smith 1970). The stem galler overwinters and then pupates in its gall. 

  Range: along streams and near springs from 1200-2400 m in northern Arizona


description:
  Rhopalomyia medusirrasa
  Woolly-bud Gall Midge

  These large galls are composed of numerous leaflike structures covered with long forked hairs, which give the galls an overall white-green color. 


description:
  Pontania sp. 

  an undescribed species near P pacifica that occurs on the same host plant

  [Photos of the galls appear on page 5 of the pdf. Matching Smith's [type-iv leaf galls](https://www.gallformers.org/gall/3660), with thin walls and projecting entirely from the lower side of the leaf]

  The Pontania sp. leaf-galler on arroyo willow in northern Arizona appeared to be P. pacifica Marlatt based on morphological criteria, but the galls, life history, and gall growth patterns were different, suggesting that the Pontania sp. leaf-galler in Arizona is an undescribed species that occurs on the same host plant (H. Goulet, personal communication). 

  The galls formed by Pontania sp. in Arizona were spherical, thin walled, and hung from the underside of the leaf midrib (Fig. 5 and 6). The galls formed by P. pacifica in California were thick walled and protruded through both sides of the leaf (Caltagirone 1964, Smith 1970). 

  Other aspects of Pontania sp. biology distinguished it from P. pacifica. In California, P. pacifica was multivoltine and bred year-round on S. lasiolepis, which is evergreen in sheltered places, and produced at least a few new leaves every month (Smith 1970). Pontania sp. in Arizona is primarily univoltine. The eggs and larvae of Pontania sp. were distinctly smaller than P. pacifica eggs and larvae (Caltagirone 1964). Larval development for Pontania sp. was > 2.5-fold longer than for P. pacifica (Caltagirone 1964). Pontania sp. leaf galls in Arizona continued to grow long after the leaf had stopped growing, and the presence of a sawfly larva in the gall was required in most cases for the gall to attain maximum growth. P. pacifica galls grew only as long as the leaf grew, and galls without a live sawfly larva attained normal size (Caltagirone 1964)

  Range: northern AZ


description:
  Pontania new species (near pacifica)

  The leaf galler, Pontania new species (near pacifica) (see Clancy et al. 1986), oviposits in the leaf midrib and a gall forms adjacent to the midrib and below the leaf

  Range: along streams and near springs from 1200-2400 m in northern Arizona


description:
  Euura sp [this may not be the same species found on this host in CA]

  The petiole galler, Euura sp., oviposits in the petiole of young leaves and produces an elongated and swollen petiole in which the larva feeds. 

  Range: along streams and near springs from 1200-2400 m in northern Arizona


description:
  Euura sp. (an unnamed leaf-midrib galling sawfly)

  Host: Salix lasiolepis

  In northern Arizona, Euura sp. females usually initiate one gall per leaf in the midrib of coyote willow leaves during May. Larvae hatch from eggs and feed inside the gall through June. Larvae then emerge from the galls and fall to the ground, pupate and over-winter. The following April, adults emerge from the ground, mate, and initiate a new generation. In Euura gallers, final gall volume generally results from an interaction of the ovipositing female and the host plant so that galls reach full size before larvae hatch or commence feeding (Smith 1970, and demonstrated by Price and Clancy 1986 for E. lasiolepis).

  Range: Colorado River at Lee's Ferry, Arizona, 25 km downstream from Glen Canyon Dam


description:
  Rhopalomyia clinata
  Leaf Gall Midge

  The midge induces small, tubular, thin-walled, monothalamous galls on leaflets of California sagebrush.


description:
  Euura sp.

  Host: Salix fragilis

  The specific identity of the species of Euura in this study is uncertain. 

  This particular species of Euura produces stem galls on crack willow, S. fragilis L. Apparently, this species of willow was originally planted from nursery stock and has become established along drainages in the Front Range of Colorado (Harrington, 1954).

  Gall development occurred rapidly. By the end of May gall diameter reached 7.1 mm (Fig. 1). However, average gall size varied greatly throughout the season, ranging from a diameter of 5.8 mm (wet weight 0.22 g) to 10.5 mm (wet weight 3.3 g) (Fig. 2). 

  Range: East Plum Creek, near Castle Rock CO


description:
  Rhopalomyia hirtibulla
  Bead Gall Midge

  This gall midge induces small, hairy, monothalamous, round to slightly conical bead-like galls on leaves of Great Basin sagebrush.


description:
  Euura sp

  Host: Salix scouleriana

  Euura sp. attack the axillary buds of S scouleriana. Oviposition begins during the first week of July and can last for up to 2 wk. Galls are initiated by injection of colleterial fluid during egg deposition by the adult female and consist of undifferentiated parenchymatous tissue that provides resources to the developing larvae. Approximately 7-14 d after oviposition, eggs hatch and, we assume, pass through five instars for males and six instars for females, as in another Euura species. From the second week in October to the end of November, late-instar larvae chew an emergence hole in the gall wall and drop or crawl to the ground, where they spin cocoons and overwinter. They remain in the soil over the winter months and emerge as adults the following summer.

  Range: northern AZ


description:
  Rhopalomyia hirtipomum
  Apple Gall Midge

  This midge induces round, densely hairy, solitary, monothalamous galls on either surface of leaves of Great Basin sagebrush.


description:
  Rhopalomyia nucula 
  Hairy-bud Gall Midge

  While some are round, others are pear shaped; both possess liplike openings at the apex. [All iNaturalist observations for this species lack the openings at the apex though they do narrow at the top. Russo states in his book that some of this species are round, not all narrow at the apex.]


description:
  Rhopalomyia pomum
  Sponge Gall Midge

  This midge induces large, irregularly shaped, hairy, spongy galls on Great Basin sagebrush. 


description:
  Rhopalomyia tumidibulla 
  Blister Gall Midge

  This midge induces small, integral, lenticular, monothalamous galls in leaves of Great Basin sagebrush.


description:
  These galls are superficially similar to Rhopalomyia hirtibulla, but R. tumidibulla occurs singly or in pairs, in contrast to the large groups that hirtibulla seem to occur in. R. tumidibulla are also 3-4 mm in length in contrast to 1mm in hirtibulla [Russo 2021].


description:
  The integral, polythalamous stem galls of this midge develop on new growth of Great Basin sagebrush in summer, reaching maturity in early fall. 


description:
  Rhabdophaga strobiloides

  [Photos of this gall appear on page 2 of the pdf]

  The pine cone willow gall midge is univoltine in Michigan. Adults emerge from the galls on Salix eriocephala Michx. and other willows on warm days in late April or early May. Eggs are laid singly on the leaves and stems of the host. Head capsule measurements indicate three larval instars. The first-instar larva emerges in early May and penetrates the soft tissues at the base of the developing shoot tip. Shortly thereafter, the gall begins to develop and is about half grown by the time the second-instar larva appears in late May. The last-instar larva appears in July. Before overwintering, the larva constructs a cocoon which may or may not be closed at the top. Pupation occurs in early April. Numerous inquilines and parasites inhabit the developing and mature gall. 

  The gall (Fig. 1) is monothalamous and forms by extensive hypertrophy of a developing shoot tip of willow. It is usually apical the first season and sub-apical the next year if it remains on the stem. 

  When young, the gall resembles a miniature cabbage because it is green, nearly spherical, and enshrouded with leaves. The inner leaves become scale-like and the outer leaves abort before the gall matures. At maturity in August, the stem at the gall base dies, then the developing scales become heavily setaceous, dry, and turn gray. 

  The mature gall is top-shaped, being prolate above and flattened below. It has numerous (60 or more) regularly arranged scales-hence the resemblance to an immature pine cone (Fig. 1A). These scales are broadened and thickened leaves tightly arranged on a barely elongated stem. Brodie (1909) postulated that the base of the gall represents about 1 m of normal branch reduced to 6 mm. The larva or pupa (Fig. 1B) is in a central cavity surrounded by a ring of stiff elongated scales. Galls found on S. eriocephala and containing healthy larvae averaged 22X 19 mm (Table I); parasitized galls are generally smaller and more spherical. Brodie (1909) gives mean measurements of 12 X 15 mm for galls on S. humilis. 


description:
  Rhabdophaga sp.

  Host: Salix discolor

  [A photo of this gall appears on page 2 of the pdf]

  The insect could not be identified with certainty from its gall either. The gall, however, does resemble one made by the nodule willow gall midge, [Rhabdophaga nodula](https://www.gallformers.org/gall/3558) (Felt 1940).

  Rhabdophaga sp. makes an exit hole and damages the stem in the same manner as several "shot hole" species of Rhabdophaga that attack willows. (Shot hole refers to the insect's exit hole in the stem.) But it cannot be included in the shot hole group of midges because they do not cause swellings on the stem (Barnes 1935), whereas this midge does--at least on the stems of Salix discolor. 

  The gall on S. discolor develops from hypertrophy of the stem tissues during early summer. The larva enters the stem through the leaf bases during early shoot development, so that the gall formed later occurs on the proximal half of the completed shoot (Fig. 1). Some galls may be monothalamous, but most of them are polythalamous because generally more than one larva attacks the same shoot at the same time. 

  The young gall is soft and green like the developing shoot; the old gall is hard, has a woody interior, and is colored externally like the bark of the mature shoot. A mature gall is prolate or spindle-shaped and may be partly covered with buds (Fig. 1). Fine elongate striations rib the sides of the gall. The size of the mature gall varies considerably (Table I) depending upon the number and distribution of the larvae within. 

  Occasionally an adult midge emerges from the host in an atypical location. Two emerged from large buds that had not developed, and four adults emerged from the sides of a large gall of the willow beaked gall midge, Mayetiola rigidae (O.S.). Without the protruding pupal cases after emergence, the abnormal gall harboring these insects would never have neen noticed. 

  Vacated galls remain on the tree indefinitely and increase in diameter each year as long as the shoot puts on diameter increment. Galls dissected on living shoots 1 and 2 years after being vacated showed the larval cells nearly or completely filled with callus tissues. 

  Heavily populated shoots often break off at the mid-gall where the larvae are congested and the interior mined by larval cells. Early yellowing of foliage and premature leaf drop also occur on shoots that are heavily galled. Such shoots die before winter, and sometimes the gall on a dead shoot shrinks and kills the larvae within. 

  Adult eclosion begins in mid-April shortly after the pupa breaks through and protrudes two-thirds of its body from its cell in the gall. About 12 minutes elapse between the time the pupal skin splits and the time the adult is dry and able to fold its wings over its abdomen. The meconium is voided after that. The adult then walks up and down the stem. 

  Nearly all emergence occurs between 0700 and 1130 hours. The peak emergence day in 1964 was 23 April when 768 adults, or more than 70% of the total that year emerged from four small shrubs. Newly emerged adults began flying at 1300 hours and left the shrubs by mid-afternoon. No adults were seen on the shrubs at night. 

  The gall develops rapidly during July and August, and is nearly full sized and complete, except for final color changes, by late August when the third-instar larva appears. 

  Range: East Lansing, MI


description:
  Mayetiola rigidae

  Hosts: Salix discolor

  Mayetiola rigidae attacks many species of willows. It has been reported from Salix discolor Miihl., S. humilis Marsh., S. rostrata Richards, S. cmdata Miihl., S. lucida Miihl., S. petiolaris Sm., and S. vimirzalis L.

  The gall (Fig. 1) is monothalamous and develops by extensive hypertrophy of a willow bud. Galls are either apical or subapical depending on whether the affected bud is a terminal or lateral. First evidence of gall formation occurs when the developing bud becomes swollen; at this time several of the leaves have flushed and ring the upper part of the gall. When the gall matures, however, the leaves attending it die. Most of the leaves fall off, but occasionally one may persist for several months or over the winter. 

  The mature gall is subglobular to fusiform and varies considerably in size and shape depending upon the host species and whether larvae are parasitized. Parasitized galls are small. Galls on S. discolor are larger than those on most other species. The size of galls observed in this study is indicated in Table I. 

  Each mature gall terminates in a slender hollow beak with an open tip which leads to an interior chamber or cell where the insect develops. The gall is hard and woody. It is green during development but turns partly to completely red in the fall. A vacated subapical gall will continue to enlarge yearly as long as its supporting branch makes diameter growth. Occasionally, a gall on a short slender twig breaks off during the winter. Sometimes a small branch above a subapical gall dies and breaks off. 

  Adults emerge from the galls soon after the first few warm days in April. Nearly all adults emerge in the morning between 0700 and 1200 hours on sunny days; the peak occurs between 0900 and 1000 hours. Males appeared first each year; females appeared 1-3 days later. The sex ratio in 1963 as determined from 85 reared adults was nearly 1 : 1. 

  Eclosion starts within a few minutes after the pupa wriggles up the chamber and protrudes from the tip of the beak. About 3 minutes elapse between the time the pupal skin splits and the adult is dry. About 10 minutes after eclosion the meconium is ejected from the tip of the abdomen; it is usually released in several short bursts of fine droplets. Considerable walking up and down the stem follows. 

  In mid-May, at the onset of the second instar, the gall is just discernible as a swollen leafy bud. By the time the insect reaches the third instar in early July the gall has attained over 85% of its growth but is still shrouded with green leaves. The gall rounds up, hardens, turns red, and drops most of the attending leaves during the first few weeks of the third larval stadium. 

  Range: East Lansing, MI


description:
  Cynips (dugèsi) pupoides Kinsey
  agamic form

  Diplolepis brevipenmata err. det. Weld, 1922 (in part), Proc. U.S. Nat. Mus. 61 (18): 7 (Teacas record only).
  Cynips dugès i var. B Kinsey, 1927, Field and Lab. Man. in Biol.: 110.
  Cynips dugès i pupoides Kinsey, 1930, Ind. Univ. Studies 84-86: 284, Figs. 44, 265, 267, 279.

  GALL.-Very similar to all galls of the C. dugèsi and C. bella com plexes, apparently not to be distinguished from the galls of C. (bella) congesta which occur in the same region. Mature galls rosy tan, obscurely freckled or unspotted; more or less shining; up to 23. mm., averaging near 17. mm. in diameter. 

  HOSTS.—Quercus grisea, Q. intricata (?), the evergreen white oaks of the area

  RANGE.-Texas: Alpine, 4 W. Fort Davis. Chisos Mt. Park (Q. grisea and Q. intricata ?). Probably confined to the mountain ranges of West Texas. Figure 18.

  LIFE HISTORY.—Adults: December 6, 14, 16. Most of the emergence still earlier. 

  The gall of pupoides is indistinguishable from the gall of Cymips congesta of the bella complex; and since the two occur in the same region, the galls cannot be assigned even to their complexes unless insects are available for study. 


description:
  Cynips (dugèsi) simulatrix Kinsey
  agamic form

  Cynips dugès i var. C. Kinsey, 1927, Field and Lab. Man. in Biol.: 110.
  Cynips dugesi simulatric Kinsey, 1930, Ind. Univ. Studies 84-86: 280, figs. 44, 263, 264, 266, 276, 287.
  Cynips dugès i or simulatrix Hoffmeyer, 1933, Ent. Meddel. Kbhvn. 18: 247, 248.

  GALL.-Very similar to all galls of the C. bella and C. dugèsi complexes, apparently indistinguishable from the galls of C. (bella) bella which occur in the same area. Mature galls rosy or brownish tan, unspotted; dull to shining; up to 27. mm., averaging near 20. mm. in diameter. 

  HOSTS.—Q. undulata, Q. grisea, Q. oblongifolia, Q. arizonica, Q. Gambelii. Apparently on all of the white oaks of the area. Records for Q. grisea and Q. undulata often interchangeable because of poor determinations of New Mexican oaks. 

  RANGE.-New Mexico: Magdalena. Near Alamogordo, 7000'. High rolls. Hillsboro (types). Kingston. (Above in more detail in Kinsey 1930: 280). Mountain Park (galls. Q. undulata, L. H. Bridewell in Kinsey coll.). Embudo Canyon in Sandia Mts. (E. B. Williamson in Kinsey coll.). Burro Mts. (Weld coll.). San Marcial, 24 S (galls, Q. grisea, Holloway in Kinsey coll.). Mayhill (galls, Q. Gambelii, Holloway in Kinsey coll.). Arizona: Prescott (Kinsey coll., and E. R. Leach, Jr., in Kinsey coll.). Safford. Oracle. Bisbee. Sabino Trail in Santa Catalina Mts., 4000'. Santa Rita Mts. Courtland. Fort Huachuca. (Above in more detail in Kinsey 1930: 280.) Oracle (galls, Q. arizonica, Holloway in Kinsey coll.). Ash fork, Chiricahua Mts., Patagonia (Weld coll.). Probably confined to the western half of New Mexico and the state of Arizona. Figure 18.

  LIFE HISTORY..—Adults: November 15, 20, 27. December 14. January 6, 23. Most of the emergence late in November or early in December.

  This wide-ranging species, found in Western New Mexico and in Southern Arizona, is distinct enough from the four other species found in our Southwest, but it has very close relatives in the two species, emergens and deceptria, which are found in the Sierra of Chihuahua and Durango. 

  The gall of simulatrix is indistinguishable from the gall of Cynips bella of the bella complex; and since the two occur in much the same region, the galls cannot be assigned even to their complexes unless insects are available for study. 


description:
  Cynips (dugèsi) spadix, new species
  agamic form

  GALL.-Similar to all galls of the C. bella and C. dugèsi complexes. Mature galls yellow to light rosy tan, unspotted, usually shining, sometimes puberulent or even powdered; large to very large, up to 32. mm., averaging near 22. mm. in diameter. Figure 54. 

  HOST.-Quercus macrophylla. Not on Q. nudinervis in the same area. 

  RANGE.-Guerrero: Taxco, 8 NE, 8000' (types). Chilpancingo, 6 S, 4000'. Tierra Colorada, 14 N, 3200'. Probably restricted to Quercus macrophylla in a southwestern portion of Mexico, involving most of the state of Guerrero. Figure 18

  LIFE HISTORY..—Adult: January 20. February 1, 10. March 10.

  Spadix is known to extend from the northern boundary of Guerrero to the southwestern limit of oak, not 25 miles from the Pacific Coast, on the road from Mexico City to Acapulco. The galls of spadix are more or less distinctive in a complex of mostly uniform galls. They are light yellow tan and in some cases very large in size and pruinose in surface. The material from our most southern locality, Tierra Colorada in Guerrero, includes more large and distinctly pruinose specimens than the material from the type locality (Taxco, in northeastern Guerrero); but the insects from these two collections appear identical, and we credit the differences in the two lots of galls to the fact that one is a larger and more inclusive series. 

  Near Taxco, Quercus macrophylla and Q. nudinervis may be found growing together. Spadix is confined to macrophylla. On Q. nudinervis the conspicuously spotted galls of C. conexa represent the same complex. See conexa for the distinctions between the two. 

  In Morelos there is another insect, C. spinalis, which matches spadix in color, but it matches spinifera in its long spine, smooth mesopleuron, and light olive tan galls. 


description:
  Cynips (dugèsi) spiculi, new species
  agamic form

  Diplolepis n. sp. Trotter, 1910, Boll. Lab. Zoël. Portici 5:129 [syn. based on Trotter material in Kinsey collection].

  GALL.-Similar to all galls of the C. bella and C. dugèsi complexes. Mature galls peculiarly rosy brown or darker, prominently bluish with a puberulence, sometimes faintly, less often conspicuously spotted, mottled, or irregularly striped with purplish marks; smooth, shining; large galls, up to 25. mm., averaging near 20. mm. in diameter. Figure 69. 

  HOSTS.—Quercus texcocana [deserticola], Q. conglomerata [rugosa], Q. decipiens [rugosa]. Probably on all of the white oaks of the area except the alpine dwarf, Q. repanda.

  RANGE.-Mexico: Mexico City, 25 E, 7000' (Q. texcocama; types). Mexico City, 17 S, 8200' (Q. texcocana, Q. conglomerata). Mexico City, 12 W, 8500' (Q. texcocana, Q. conglomerata). Puebla: El Seco, 5 SW, 8000' (Q. texcocana). Tehuacán, 17 NE, 7900' (Q. decipiens). Probably ranging from the Valley of Mexico, through the mountains to the southeast, to the borders of the state of Vera Cruz. Replaced at the northern end of the Valley (near Pachuca) by C. longa. Figure 18.

  LIFE HISTORY.—Adults: December 8, 16, 23. January 2, 25. Feb ruary 1, 25. March 10, 30. Some pupae alive April 13. The bulk of the emergence early in March. 

  This is the dugèsi representative on all of the white oaks except the alpine dwarf, Q. repanda, in the mountains rimming the Valley of Mexico. The gall is larger than those of most of the Southern Mexican species, and often obscurely spotted or striped with dull purple. On the dwarf oak, repanda, the short-winged C. pumilio is found in the same area. At the northern end of the Valley, near Pachuca, C. (dugèsi) longa is found on Q. rhodo phlebia. Along the Mexico City–Acapulco road across the Divide at the southwestern end of the Valley, C. (dugèsi) spinalis is found on the westerly facing slopes of the same mountain range. Spiculi, then, does not range west of the Valley of Mexico, but it extends southeast well toward the borders of the state of Vera Cruz. 


description:
  Cynips (dugèsi) spinalis, new species
  agamic form

  GALL.-Similar to all galls of the C. bella and C. dugèsi complexes. Mature galls very light olive tan, less often rosy, unspotted, smooth, shining, often bluish puberulent, rather small, up to 16. mm., averaging near 13. mm. in diameter. 

  HOST.-Quercus texcocama [deserticola]

  RANGE.-Morelos: Cuernavaca, 8 N, 8700' (types). Probably restricted to the western slopes of the mountains which separate the Valley of Mexico from the valleys of Morelos. Figure 18. 

  LIFE HISTORY.—Galls: Still fresh and succulent, January 7. Adults: January 25.

  This long-spined insect is very close to C. spinifera of the adjacent state of Michoacán, matching it in its long spine, smooth mesopleuron, and olive tan galls. In all these characters it differs from the otherwise very similar species, spadix, in Guerrero. The three species must be segregates of the same stock, spinalis and spinifera being closer than either is to spadix. 

  Our collections of spinalis were made on the Mexico City Acapulco road on the mountain slopes above the city of Cuerna vaca. This was only a few miles—perhaps not 8 or 10—from a locality from which we collected the very different insect, C. spiculi, on the eastern slopes of the same mountain overlooking the Valley of Mexico. The two localities, however, are separated by a high divide, at about 10,500 feet, to which white oaks do not ascend, and spiculi and spinalis are thus effectually isolated. In our study of Cynipidae we have found no other area in which so many narrow-ranging species are as effectively isolated as in the high mountain and sub-tropic valley country of the Mexican Cordillera.


description:
  Cynips (dugèsi) spinescens, new species
  agamic form

  GALL.—Very similar to all galls of the C. bella and C. dugèsi complexes. Mature galls in part light yellowish tan and in part rich rosy or purplish brown, usually well marked with purple spots or streaks, with traces of a bluish bloom; of moderate size or smaller, up to 19. mm., aver aging nearer 14. mm. in diameter. Figure 68. 

  HOST.-Quercus Purpusi, the rusty, pubescent white oak of the region. 

  RANGE.-Mexico: Jilótepec, 7 NW, 9500' (types). Probably restricted to a small area in the high mountains about 45 miles northwest of Mexico City. Figure 18

  LIFE HISTORY..—Adults: March 6, 17

  In the mountains east, south, and west of Mexico City, the dugèsi complex is represented by C. pumilio (on Quercus repanda) and C. spiculi (on all other white oaks), but in the high mountains 45 miles northwest of Mexico City we found this long spined species, spinescens, as the segregate of the complex. The locality is similarly about 45 miles east of Pachuca, where pumilio and longa are the dugèsi species, and perhaps 80 miles from a locality north of Querétaro where we found C. (dugèsi) vulgata. Spinescens is, therefore, probably restricted to a limited mountain area in and adjacent to the northwestern corner of the state of Mexico. It is a long-spined species very close to the long spined spiculi and the short-spined longa. It is to be distinguished from spiculi in averaging smaller in size, in having a radial cell which is relatively longer because the second abscissa of the radius is not so strongly curved in its terminal third, and in having galls which are at least in part yellow tan with a rather obvious spotting of purple. The gall characters alone would mark it as a distinct species. 


description:
  Cynips (dugèsi) spinifera, new species
  agamic form

  GALL.-Similar to all galls in the C. bella and C. dugès i complexes. Mature galls light straw tan to yellowish brown, less often tending to brownish or purplish olive, rarely rosy, practically unmarked, with a con spicuous bluish puberulence; rather small galls, up to 17. mm., averaging near 13. mm. in diameter. Figure 55. 

  HOST.-Quercus conglomerata [rugosa], the larger, common white oak of the region. 

  RANGE.-Michoacán: Morelia, 14 E, 7000' (types). Purépero, 5 W, 8500'. Probably restricted to a limited mountain area in the Mexican Cordil lera of Michoacán. Figure 18.

  LIFE HISTORY..—Adults: January 2, 25. February 15. March 22. April 10, 15. Most of the emergence late in March.

  This long-spined insect is a very close duplicate of the short spined C. (dugèsi) spadia and of the long-spined C. spiculi which occur in the adjacent states of Guerrero and of Mexico, respectively.  The galls of spinifera are distinctly small and never as rosy brown as the galls of spadix. 

  Spinifera is still more closely matched by C. spinalis which occurs in another adjacent area, namely in Morelos. In spinifera and spinalis the spine, mesopleuron, and gall characters are the same, but the areolet of spinifera is always smaller, and the thoracic lines darker than in spinalis.

  In the same area with spinifera, the alpine dwarf oak, Q. repanda, bears a short-winged species of the dugèsi complex, C. pumilio. 


description:
  Cynips (dugèsi) subnigra, new species
  agamic form

  Cymips dugèsi simulatrix, err. det. Kinsey, 1930, Ind. Univ. Studies 84-86: 280 (Organ Mt. record only)

  GALL.-Very similar to all galls of the C. bella and C. dugèsi complexes, apparently indistinguishable from galls of C. (bella) bella which occur in the same region. Mature galls rosy or brownish tan, unspotted or obscurely mottled, usually shining, often large, up to 25. mm., averaging near 20. mm. in diameter. Figure 62. 

  HOST.-Quercus grisea. 

  RANGE.-New Mexico: Las Cruces, 16 E (= Organ Mts.) (types). Soldedad Canyon in Organ Mts. (galls, L. H. Bridewell in Kinsey coll.). Possibly restricted to the Organ Mts. in Southeastern New Mexico; or possibly occurring in all the eastern mountains of New Mexico, south of Albuquerque. Figure 18.

  LIFE HISTORY.—Adults: January 10, 26; February 5, 28

  Subnigra has been taken only in the Organ Mountains of Southeastern New Mexico (just north of El Paso). It is one of the most surprising discoveries in the complex. While C. simulatrix, which is the wide-spread representative of the dugèsi complex in Southern Arizona and Southwestern New Mexico, is matched by 2 species (emergens and deceptrix) in Mexico which are distinguished from simulatrix only with difficulty, the present species, subnigra, is abundantly distinct from everything else in the complex. Subnigra is the only species of the 23 which has a spot in the radial cell. It is a large-bodied, strikingly dark in sect with other distinctive characters. It is a long-winged insect interpolated between the short-winged brevipennata to the north and the short-winged pupoides to the south and east. It now remains to discover how many of our previously published records for simulatrix (Kinsey 1930: 280–281), based on galls alone, really apply to subnigra. Is subnigraactually restricted to the Organ Mountains; or is it the wide-spread species throughout New Mexico east of the Rio Grande and south of Albuquerque? 

  The galls of C. (dugèsi) subnigra are indistinguishable from the galls of C. (bella) bella. Both species occur on the same oaks in the Organ Mountains; but while subnigra does not occur south of that range, bella extends for 200 miles into the Sierra of Chihuahua. 


description:
  Cynips (dugèsi) vasta, new species
  agamic form

  GALL.—Similar to all galls of the C. bella and C. dugès i complexes. Mature galls peculiarly, deep rosy brown tan, practically unspotted; rather small, up to 21. mm., but averaging nearer 14. mm. in diameter. Figure 64. 

  HOSTS.–Scrub oaks, Quercus intricata and Q. Pringlei (the two species not well separated in our collections). Also on Q. sillae [vaseyana] ?

  RANGE.--Tamaulipas: Tula, 22 N, 6100' (types, Q. intricata, Q. Pringlei). Tula, 26 N, 5300' (Q. sillae, galls only). Possibly restricted to a limited portion of the Eastern Sierra of Mexico, in the southern end of the state of Tamaulipas. Figure 18. 

  LIFE HISTORY..—Adults: February 2, 16. Most of the emergence on the later date.

  This insect is closest to C. oriens, which we found 40 miles to the south in the Eastern Sierra near Cerritos, and to C. oriumda, which we found west of Ciudad Victoria in Tamaulipas, hardly 35 miles from the type locality of vasta. Vasta differs from oriens and oriunda in having more bright rufous over the whole body, the antennae and legs being similarly bright rufous. Vasta also differs from oriens in having lighter wing veins and less abundant cubital cell markings. C. occidua, in the Western Sierra of Zacatecas, is also a relative from which vasta is best distin guished by its black thoracic lines and small areolet. 

  Our limited collections in the Eastern Sierra of Mexico give us no basis for judging the extension of the ranges of these several species, but the taking of C. oriens and C. oriunda so near the type localities of vasta shows that vasta cannot extend very far either northeast or southwest. It is possible that the largely barren mountains and valleys in this portion of Mexico isolate some species with ranges not larger than a single slope of a single mountain ridge. 

  No species of the C. bella complex is yet known to occur in the range of vasta, but since the bella complex is known not far away, gall determinations for this species should be verified by insect material. 

  Cynips (dugèsi) vulgata, new species
  agamic form

  Dryophanta dugès i err. det. Beutenmüller, 1911, Bull. Amer. Mus. Nat. Hist. 30:345 (in large part). Err. det. Kinsey, 1920, Bull. Amer. Mus. Nat. Hist. 42: pl. 32 fig. 25.
  Cynips dugès i dugès i err. det. Kinsey, 1930, Ind. Univ. Studies 84-86:281 282, figs. 44, 268, 277 (in large part).

  GALL.-Similar to all galls of the C. bella and C. dugès i complexes. Mature galls rosy tan, unspotted; dull to weakly shining; of moderate size, up to 20. mm., averaging near 14. mm. in diameter. Figure 66. 

  HOSTS.—Quercus potosina, Q. jaralensis [potosina], Q. chihuahuensis, Q. undata, Q. sacame [arizonica], Q. reticulata [rugosa]. Possibly on all oaks of the area except the alpine dwarf oaks, Q. microphylla, etc. 

  RANGE.-Aguascalientes: Pabellón, 20 W, 7000' (Q. sacame, Q. un data, Q. chihuahuensis). Pabellón, 23 W, 8500' (Q. potosima). Jalisco: Aguascalientes, 35 E, 7000' (Q. potosima). San Luis Potosí: San Luis Potosí, 15 W, 8000' (Q. potosina, types. Also Q. potosina, Ed. Palmer, 1878, in Kinsey coll.; and Q. unda'a, Kinsey coll.). Guanajuato: Mountains of Guanajuato (Dugès in U.S. Nat. Mus.). San Felipe, 8 NE, 7000' (Q. undata, galls only). San Felipe, 20 SW, 8000' (Q. chihuahuensis. Also Q. macrophylla 2 galls only). León, 20 NE, 9000' (Q. faralensis, Q. reticulata). Querétaro: Querétaro, 28 N, 7300' (Q. chihuahuensis; galls, Q. jara lensis). Wide-spread, from Aguascalientes to Guanajuato and Querétaro, known from all the oaks of the area except dwarfs of the Q. microphylla complex. Figure 18. 

  LIFE HISTORY..—Adults: December 4, 5, 6, 9, 10, 11, 13, 15, 16, 19, 26. February 10, 15, 20, 24, 25, 28. March 1, 2, 3. Most of the emergence in February.

  Vulgata produces the common oak apple in the Western Sierra of Central Mexico, from Aguascalientes and the western mountains of San Luis Potosí to Guanajuato and Querétaro. Although we have bred material from 7 of the oaks of that area, we find vulgata on all of them except the little alpine dwarf, Q. microphylla. It is C. dugèsi on that oak, an insect which is close to vulgata. 

  No species of the C. bella complex is known to occur near the range of C. vulgata, so galls from this area need not be confused with material of the other complex. 


description:
  Cynips (bulboides) bulboides, new species
  agamic form

  GALL.-Bulboid, as described for the complex. Mature galls peculiarly dark rosy brown, not noticeably puberulent, usually unspotted; younger galls more often spotted, with some of the mature galls obscurely spotted, but only a very few of them conspicuously spotted with purple; up to 23. mm., averaging nearer 15. mm. in diameter. Figures 9, 77. 

  HOSTS.—Quercus undata, Q. chihuahuensis, Q. sacame, Q. oblongifolia, Q. reticulata, Q. durangensis, Q. depressipes, Q. striatula. Apparently on all of the white oaks of the area. 

  RANGE.-Chihuahua: Parral, 20 S (Q. undata; also galls on Q. sacame). Durango: Villa Campos, 21 S, 6700' (Q. undata). Indé, 20 E, 6300' (Q. chihuahuensis). Patos, 15 W, 8000' (Q. chihuahuensis, Q. sacame, Q. oblongifolia. Also galls on Q. reticulata, Q. striatula). Canatlan, 7 N, 7400' (Q. undata). Durango, 2 N, 6500' (Q. undata). Otinapa, 8500' (Q. sacame, Q. durangensis, Q. depressipes, Q. reticulata). Zacatecas: Cantuna, 6 N, 7600' (Q. undata). Apparently restricted to the Sierra of Durango and immediately ad jacent portions of Chihuahua and Zacatecas. Figure 75

  LIFE HISTORY..—Adults: November 20, December 5, January 20, 29. February 4, 5, 10, 12, 15, 22, 26, 29. March 2, 3, 6, 16. April 10. The bulk of the emergence in the first half of February. 

  In the bulboides complex this is the only species represented by large series in our collections. The species occupies the center of the range of the complex. It probably does not represent the oldest species in the group, although it is the only one which (in insect characters) is precisely identical with one of the species (spiculi) of the dugèsi complex. Between spiculi and bulboides lie 3 other species (1 of the dugès i and 2 of the bulboides complex), and the identity of bulboides and spiculi must be taken to be coincidence, dependent upon the close relations of the two species. The bulboid gall of bulboides is quite different from the spherical galls of the dugèsi and bella complexes.

  In northern Durango (at Villa Campos, Indé, and possibly elsewhere) heavily mottled, spherical galls, indistinguishable from those of C. (bella) aspera, give a long-spined insect which is in every respect identical with C. bulboides. This insect, here listed as C. (bulboides) aequalis, may be a Mendelian form of the present species, bulboides, or it may represent a distinct species which is the connecting link between the bella and bulboides complexes.


description:
  Cynips (bulboides) aequalis, new species
  agamic form

  GALL.-Not bulboid but spherical, apparently indistinguishable from the galls of C. (bella) pomifera and C. (bella) aspera. Mature galls light straw yellow, sometimes touched with rose or brown, always conspicuously mottled with purplish brown; up to 15. mm. but averaging near 10. mm. in diameter. Figures 9, 76.

  HOSTS.—Quercus chihuahuensis, Q. undata.

  RANGE.-Chihuahua: Pedernales, 2 E, 7500' (det. uncertain; Q. chi huahuensis). Parral, 20 S (Q. undata). Durango: Villa Campos, 21 S, 6700' (Q. undata). Indé, 20 E, 6300' (Q. chihuahuensis; types). Apparently restricted to an area in the Western Sierra of southern most Chihuahua and Northern Durango. Figure 75.

  LIFE HISTORY.—Adults: January 20, 30. February 8, 9, 10, 19. Most of the emergence late in January and early in February.

  The insect of aequalis is to all appearances identical with the insect of C. (bulboides) bulboides which occurs on the same trees, in the same localities in which aequalis is found. But the galls of the two are very different. The bulboid gall typical of the present complex is replaced in aequalis by a small, spherical, heavily mottled gall indistinguishable from that of C. (bella) pomifera and C. (bella) aspera. On gall characters alone, aequalis belongs to the bella complex, but the insect is in many respects unlike bella and—again—identical with bulboides. I am inclined to consider aequalis a Mendelian form within the species bulboides, and it is not impossible that it is the connecting link between the bulboides and bella complexes. Data that may throw more light on the origin of higher categories may be unraveled if we can obtain further material of this insect. 

  The only localities in which we have definitely connected the insects and galls of aequalis are Villa Campos and Indé, in the northern end of the State of Durango. In our earlier collections from more northern localities we bred, in a single mixture, spherical galls which were mottled, spherical galls which were uniformly tan, and bulboid galls. We did this because the abund ance of intermediates and poorly defined galls in those localities misled us into believing that we had only one highly variable series. But in breeding material from Villa Campos and Indé we separated the 3 types of galls, securing insects of aequalis from the mottled galls, of dugºsi with a few stray adults of C. (bella) aspera from the uniformly tan galls, and of bulboides from the bulboid galls. 

  Determinations in this group must be based on both insect and gall characters. When based on insects alone they may apply to either of the identical insects, bulboides or aequalis; when based on galls alone they may belong to the identical galls of aequalis and aspera, or to those of several other species of the bella and dugès i complexes. Until further collections can be made in Southern Chihuahua and Northern Durango, when each and every type of gall will be bred separately, it is uncertain how far north aequalis actually ranges. The southern limit of the range is probably not far from Indé, because all spotted galls, both of aequalis and of the bella complex, disappear from our collections south of Indé. 


description:
  Cynips (bulboides) bulbacea, new species
  agamic form

  GALL.-A bulboid gall, as described for the complex. Mature galls rosy or (often) yellow brown, not as dark as in some other species; notice ably blue with a puberulence, occasionally unspotted but almost always well marked, in some cases conspicuously marked with slightly raised spots and ridges of purple; up to 25. mm., averaging nearer 18. mm. in diameter. Figure 80. 

  HOSTS.—Quercus laeta, Q. polymorpha, Q. macrophylla, Q. sacame. Possibly on all of the white oaks of the area. 

  RANGE.-Tamaulipas: Miquihuana, 7 SE, 6000' (Q. sacame). San Luis Potosí: Rio Verde, 14 W, 6000' and 6500' (Q. laeta, types; also Q. polymorpha and Q. macrophylla). Probably widely spread through one of the more western ranges of the Eastern Sierra. Figure 75.

  LIFE HISTORY.—Adults: January 8, 27. February 4, 10, 12, 15, 24. Most of the emergence early in February.

  The two locality records for this species indicate that it ranges a considerable distance north and south; but the several ridges of the Eastern Sierra often isolate distinct cynipids, so bulbacea may not be expected east or west of the one range which includes the mountains near Miquihuana and those west of Rio Verde. 

  The species is to be distinguished from bulboides by its smaller areolet, its wholly punctate anterior parallel lines, its smaller but more completely sculptured foveae, and (best of all) by its more bluish, puberulent galls with their conspicuous, often elevated mottling of purple spots and lines. 


description:
  Cynips (bulboides) bulbulus, new species
  agamic form

  GALL-A bulboid gall, as described for the complex. Mature galls rosy or yellow brown, not as dark as in some other species, not noticeably puberulent except on younger galls, usually unspotted, but sometimes well marked with purple; up to 23. mm., averaging nearer 17. mm. in diameter. Figure 78. 

  HOST.-Quercus chihuahuensis. Not recovered from any of the sev eral other white oaks of the area. 

  RANGE.-Chihuahua: Madera, 10 SE (gall only). Namiquipa, 30 W, 5200'. Matachic, 3 N, 6700'. Pedernales, 2 E, 7500' (types). Probably restricted to the Western Sierra in a more southern portion of Chihuahua. Figure 75.

  LIFE HISTORY.--Adults: February 1, 5, 12, 18.

  This insect produces the most northern of the bulboid oak apples, the complex reaching its northern limit in the middle of the state of Chihuahua, still 165 miles south of the International Boundary. Bulbulus was not commonly represented in our col lections in Chihuahua, certainly being much rarer (at least in 1931) than the next more southern species, bulboides. The in sect is best distinguished from the other species of the complex by the better developed line of spots which lies in the cubital cell parallel to the second abscissa of the radius, by the more fully sculptured foveae, and by the distinctly darker coloration of the whole body


description:
  Cynips (bulboides) bulla, new species
  agamic form

  GALL.-A bulboid gall, as described for the complex. Mature galls peculiarly dark rosy or dark yellow brown, not noticeably puberulent ex cept on younger galls; unspotted or only obscurely spotted; up to 25. mm., averaging nearer 18. mm. in diameter. Figure 79. 

  HOSTS.—Quercus chihuahuensis, Q. jaralensis, Q. undata, Q. reticulata, Q. macrophylla. Possibly on all the white oaks except the alpine dwarf, Q. microphylla, in the area. 

  RANGE.-San Luis Potosí: San Luis Potosí, 15 W, 8000' (Q. undata). Guanajuato: San Felipe, 8 NE, 7000' (Q. macrophylla). San Felipe, 20 SW, 8000' (Q. chihuahuensis, Q. macrophylla, Q. jaralensis). León, 20 NE, 9000' (Q. reticulata, Q. jaralensis). Querétaro: Querétaro, 28 N, 7300' (types); Q. chihuahuensis; also galls, (Q. faralensis). Probably restricted to an area in Central Mexico extending from San Luis Potosí through Guanajuato and Querétaro. Figure 75.

  LIFE HISTORY..—Adults: December 9, 10, 11, 16, 18. February 10, 15. Most of the emergence early in February

  Our material is scant, but it represents five localities and five hosts, ranging over a Central Mexican area 100 miles in diameter. From the adjacent state of Aguascalientes we have material obviously representing a different species of the complex, but our collections are not adequate for description. 


description:
  Cynips (aggregata) aggregata (Weld)
  agamic form

  Diplolepis aggregata Weld, 1926, Proc. U.S. Nat. Mus. 68 (10): 15, fig. 3. Kinsey, 1930, Ind. Univ. Studies 84–86: 446. Houard, 1935, Marcellia 28: 99.
  “Cynips howardiana Kinsey in litt., possibly C. plumbea Weld” Hoffmeyer, 1930, Ent. Meddel. Kbhvn. 17: 214.
  Andricus aggregatus Hoffmeyer, 1933, Ent. Meddel. Kbhvn. 18: 247.

  GALL.-Large, up to 33. mm., averaging nearer 28. mm. in diameter, with a slender, drawn-out, sharply-pointed pedicel at base; light pinkish to rose and light russet brown, often finely spotted (never striped), sometimes with a whitish or bluish puberulence; occurring singly or (more often) in large clusters; the galls inserted in cracks in the bark of young twigs. Figures 87, 89. 

  HOSTS.—Quercus oblongifolia, Q. arizonica, Q. sacame, Q. diversicolor [rugosa] (?), Q. grisea, Q. toumeyi (the last record acc. Weld 1926). 

  RANGE.-New Mexico: Hillsboro (galls, Q. grisea). Arizona: Globe. Safford (galls, Q. arizonica). Fort Grant (galls, Morrison acc. Weld 1926). Oracle (Q. arizonica, Kinsey coll. Also acc. Weld 1926, and also Holloway in Kinsey coll.). Sabino Trail in Santa Catalina Mts., 4000' (Q. oblongifolia, Q. arizonica, Q. diversicolor. Also Weld coll., and types, Chrisman coll.). Santa Rita Mts. (Q. oblongifolia, Q. ari zonica). Whetstone (Q. arizonica). Tumacacori Mts. (galls., acc. Weld 1926). Fort Huachuca (galls, Q. arizonica. Also Jacot acc. Weld 1926). Chiricahua Mts. (Q. arizonica, Weld and Hubbard, acc. Weld 1926). Pata gonia (galls, Q. tourneyi, acc. Weld 1926). 

  Apparently confined to the oak-inhabited mountain ranges of South eastern Arizona and Southwestern New Mexico. Figure 84 

  LIFE HISTORY..—Adults: Alive in gall November 26. Emerging November 5, 22, 27. Also before December 3, after January 1, on January 11, and on later dates. An insect cut from a gall on November 26 was still alive in a pill box on December 31 (acc. Weld 1926). The bulk of the emergence probably early in January.

  This is the only species of the complex known from the United States. It is found only in the southeastern quarter of Arizona and in the immediately adjacent portions of New Mexico. On all of the evergreen white oaks in that area, the stem galls of aggregata are conspicuous objects forming clusters two thirds as large as a man's fist. My own collections of the species were made in 1919, but it is difficult to understand why the species was not collected earlier and described before 1926. 

  Aggregata finds its closest relative in C. finitima, which occurs in the adjacent state of Chihuahua, in Mexico. From finitima, aggregata is to be distinguished by having a gall which is unspotted or only finely spotted with purple. The differences between the two insects are not great but enough, especially when fortified by the gall characters, to indicate that two populations are isolated in the two regions. 


description:
  Cynips (aggregata) cucurbita, new species
  agamic form

  Cynipide No. 24 Houard, 1928, Marcellia 24:47, Figs. 52-55.

  GALL.-Rather large, up to 30. mm., but averaging nearer 22. mm. in diameter, the globular body tapering gradually (more so than in other species) into a long, drawn-out, sharply-pointed pedicel at base; light rose or yellow, older galls becoming a peculiar, light rosy tan, less often browner in color, with very little puberulence; occurring usually in dense clusters, the galls inserted in cracks in the bark of young twigs. Figure 90. 

  HOST.S.—Quercus haematophlebia [magnoliifolia], Q. rhodophlebia [rugosa]

  RANGE.-Jalisco: Atequiza, 5 E, 6500' (Q. haematophlebia). Gua dalajara, 25 S, 6000' (Q. haematophlebia). Sayula, 13 SW, 7700' (Q. rhodophlebia). Probably restricted to a portion of the state of Jalisco (and adjacent areas to the south?). Figure 84

  LIFE HISTORY.—Adults: April 1.

  This species, from the state of Jalisco in Southwestern Mexico, has a gall which is distinct in being peculiarly light rosy tan, not unlike some of the ripe cucurbs in color. The mesonotum of the insect has more bright rufous than is to be found on any of the other insects of the complex. We have bred cucurbita from three localities east and south of Guadalajara, in Jalisco, but insufficient material which we have from other localities indicates that the species may not spread north or west of that city


description:
  Cynips (aggregata) finitima, new species
  agamic form

  GALL.-Moderate in size, up to 25. mm., but averaging nearer 19. mm. in diameter, with a slender, drawn-out, sharply-pointed pedicel at base; light yellow tan to dark russet brown, usually spotted or irregularly striped with dull purple, these markings broader and more conspicuous than in aggregata; often with a whitish or bluish puberulence; occurring singly or (more often) in large clusters; the galls inserted in cracks in the bark of young twigs. Figure 85. 

  HOST.-Quercus sacame, a close relative of the more northern Q. arizonica. 

  RANGE.-Chihuahua: Pacheco, 13 NE, 7400', and 20 E, 5400' (types). Possibly wide-spread throughout the Western Sierra in the state of Chihuahua, at least in all but the southernmost hundred miles of that state. Figure 84.

  LIFE HISTORY.—Adults: February 4 and later (unrecorded) dates

  This insect, from Northern Chihuahua, is very close to aggregata of Southern Arizona and New Mexico. Finitima is generally darker, especially on the sides of the thorax and abdomen, it averages smaller, and—most certain distinction of all—it has a gall which is more conspicuously spotted and covered with purple markings which are broader than those of aggregata. 


description:
  Cynips (aggregata) sierrae, new species
  agamic form

  GALL.-Large, up to 32. mm., averaging nearer 28. mm. in diameter, with a slender, drawn-out, sharply-pointed pedicel at base; pinkish rose to brownish tan and darker brown, only obscurely mottled with purple, some times with a bluish puberulence, occurring singly or (more often) in large clusters; the galls inserted in cracks in the bark of young twigs. Figure 88. 

  HOSTS.—Quercus sacame [arizonica], Q. chihuahuensis. 

  RANGE.-Durango: Patos, 15 W, 8000' (Q. sacame, Q. chihuahuensis). Zacatecas: La Mesa, 8000' (galls, Q. saca me). Aguascalientes: Pabellón, 20 W, 7,000' (galls, Q. rhodophlebia.). Probably occurring throughout the more central portion of the West ern Sierra of Mexico, although the determinations are certain only from Southwestern Durango. Figure 84

  LIFE HISTORY. —Adults: Mature in galls in first half of November. Emerging November 13, 18, 19. December 30. January 15. February 5, 10, 15, 20, 22, 23, 25. March 2, 10. April 1. Most of the emergence in the first half of February, the emergence dates previous to the first of February and after the last of February applying to a few stray insects only. 

  We have bred the insect from the galls taken at only a single locality (west of Patos in more southern Durango), but we have galls of probably the same species from other and more southern locali ties in the Western Sierra. Our host records for Quercus sacame and Q. chihuahuensis may apply to one oak, for we were not able to make satisfactory differentiation of the two species in the field at this locality. Although both of the extreme types of oak were present, there also appeared to be every intermediate and recombination of the two types.


description:
  Cynips (aggregata) tigrina, new species
  agamic form

  GALL.-Moderate in size, up to 28. mm., but averaging nearer 20. mm. in diameter, the globular body drawn out into a very short, abruptly truncate stalk at base; light yellow to pinkish or rosy tan, very conspicuously spotted and irregularly striped with rich purple, usually dusted with a bluish puberulence; occurring singly, rarely in twos or threes, attached to the main veins of the leaves. Figures 9, 86. 

  HOST.-Quercus nudinervis. 

  RANGE.-Guerrero: Taxco, 8 NE, 8000' (types). Possibly extending over a much wider area in Southwestern Mexico, but so far known from only the single locality in Northeastern Guerrero. Figure 84. 

  LIFE HISTORY..—Adults: March 18, 25. April 1, 2. The bulk of the emergence late in March and early in April.

  The gall is spectacularly spotted with purple, and is unique in the complex in being located on leaves instead of on stems. This leaf gall provides the evidence that the stem-oviposition common in the complex was derived from leaf-ovipositing ancestors. See the introduction to the whole complex for a further discussion of the relation of this insect to the bulboides and dugèsi stocks. 


description:
  Cynips (bella) aspera, new species
  agamic form

  GALL.-Similar to all galls of the C. bella and C. dugèsi complexes. Mature galls rosy or (more often) light yellowish tan, always conspicuously mottled with purplish brown; sometimes large, up to 17. mm., but averaging much smaller, nearer 13. mm. in diameter. Figure 93. 

  HOSTS.—Quercus undata, Q. chihuahuensis, Q. sacame [arizonica]. Probably on all of the white oaks of the area. 

  RANGE.-Chihuahua: Parral, 12 W, 6800" (galls, Q. undata, Q. sacame). Parral, 20 S (Q. undata; types). Durango: Villa Campos, 21 S, 6700' (Q. undata). Indé, 20 E, 6300' (Q. chihuahuensis). Canatlán, 7 N, 7400' (Q. undata; one female only). Apparently confined to a portion of the Western Sierra of Mexico in southernmost Chihuahua and Northern Durango. Figure 96

  LIFE HISTORY.—Adults: November 8. January 29, 30. February 9, 10, 22. Most of the emergence early in February.

  This is the southernmost species of the bella complex. The northern limit of the range of aspera (Parral in southernmost Chihuahua) coincides with the northern limit of the range of C. (dugèsi) deceptrix, but deceptrix extends for 175 miles south of the southernmost point (east of Indé) from which we have aspera. Aspera is distinguished from the related pomifera to the north by having bright or rich rufous on the head, thorax, antennae, and legs, and an areolet of smaller size. These characters are, however, highly variable, and since the galls of aspera and pomifera agree in being light tan with a conspicuous purplish brown mottling, there is some reason for considering these two as one species. 

  The mottled galls of aspera are almost always readily distin guishable from the uniformly tan, dull-surfaced galls of C. (du gèsi) deceptria. They differ in no respect from the mottled galls of C. (bulboides) aequalis (q.v.) which is known from several localities in Northern Chihuahua. 


description:
  Cynips (bella) bella Bassett
  agamic form

  Cynips bella Bassett, 1881, Canad. Ent. 13:93.
  Holcaspis maculipennis Gillette, 1894, Canad. Ent. 26:236. Beutenmüller, 1909, Journ. N.Y. Ent. Soc. 17:46.
  Cynips bella bella Kinsey, 1930, Ind. Univ. Studies 84-86; figs. 45, 263, 272, 280. See the last reference for a more extended bibliography.
  [NOT Holcaspis maculipennis err. det. Beutenmüller et al., in references to Pacific Coast material which is correctly Cynips mirabilis Kinsey.]

  GALL.-Similar to all galls of the C. bella and C. dugèsi complexes, apparently indistinguishable from galls of C. (dugèsi) simulatrix, C. (dugèsi) subnigra, and C. (dugèsi) emergens which occur in various parts of the range of the present species. Mature galls rosy or brownish tan, uniformly colored or only obscurely mottled; often large; up to 27. mm., averaging nearer 20. mm. in diameter. 

  HOSTS.—Quercus grisea, Q. undulata, Q. chihuahuensis, Q. oblongifolia, Q. arizonica, Q. sacame [arizonica], Q. diversicolor [rugosa] (acc. Weld), Q. Gambelii, and Q. Toumeyi (acc. Weld). Apparently on all of the white oaks of the area. 

  RANGE.-Arizona: Safford. Oracle. Courtland. Fort Huachuca, Bis bee. Santa Catalina Mts. near Tucson (types of bella). Santa Rita Mts. Tumacacori Mts. Patagonia Mts. Chiricahua Mts. Mule Mts. Nogales. New Mexico: Tijeras. Blue Canyon near Socorro. Abo Pass. Nogal Canyon. Burro Mts. Fierro. Near Alamogordo at 7000'. Highrolls. Moun tain Park. Hillsboro. Kingston. Organ Mts. (types of maculipennis). Soledad Canyon in Organ Mts. (Above records in detail in Kinsey 1930: 290.) Embudo Canyon in Sandia Mts. (galls. E. B. Williamson in Kinsey coll.). San Marcial, 24 S (galls, Q. grisea, Holloway in Kinsey coll.). Mayhill (galls, Q. Gambelii, Holloway in Kinsey coll.). Magdalena (Q. undulata, Kinsey coll.). Las Cruces, 16 E (= Organ Mts.; Q. grisea, Kinsey coll.). Chihuahua: Pacheco, 20 E, 5400' (Q. saca me). Pacheco, 13 NE, 7400' (Q. sacame, galls on Q. Gambelii). San Buenaventura, 3 E, 5600' (Q. sa came, Q. chihuahuensis). Las Cruces, 15 N, 6500' (Q. chihuahuensis). Apparently one species throughout the southern halves of Arizona and New Mexico and the northern half of Chihuahua. Known from Oracle and Safford (possibly from Prescott), Arizona, and from the Sandia Mountains, New Mexico, southward to Las Cruces in Chihuahua. Figure 96. 

  LIFE HISTORY..—Galls: Nearly mature July 9; mature August 16, 22. Adults: October 25, 30. November 1. December 2, 10, 21, 26. Janu ary 1, 2, 10, 20, 25, 26, 29, 30. February 1, 5, 6, 10, 14, 15, 20, 28. Only stray adults before December. Most of the emergence in mid-December in Arizona and New Mexico, but in January in Chihuahua.

  Throughout the whole of the southern half of Arizona and New Mexico and the northern half of Chihuahua, there seems to be but a single species of the bella complex, namely the typical species bella. We do not know of any other cynipid which ranges over as large a portion of our Southwest and Mexico. Within nearly the same area, there are three species of the dugèsi complex, producing galls which are apparently indistinguishable from the galls of bella; but repeated study of our more than 800 insects fails to show a basis for recognition of more than one species of bella in this area. Statistical studies might show differences in averages in the size of the areolet, and in the sizes of the spots in the radial and the cubital cells; but the limits of individual variation and the averages discernible on ready in spection do not show any differentiation into species. At one time it appeared to us that the insects from the Organ Mountains in New Mexico (= maculipennis Gillette) were, together with the Northern Chihuahua material, distinct from the rest of the Southwestern material in having larger areolets and more fused spots in the radial and cubital cells; but subsequent restudy of the material does not confirm that first impression. It is to be noted, however, that our Arizona series are not as large as those from Chihuahua, and additional material from Southern Arizona would provide a better basis for determining the limits of true bella. 

  A physiologic item is in part responsible for some doubt as to the identity of the Arizona and Chihuahua material. The Arizona insects emerge for the most part in mid-December; our Organ Mountain and Chihuahua material agreed in emerging, for the most part, in January and even early February. This may or may not be associated with specific distinctions. 

  C. (dugèsi) emergens, in Chihuahua, ranges as far south as Santa Ísabel. C. bella does not go as far south. It is C. (bella) pomifera, which is found in the Sierra from Namiquipa to Santa Ísabel, in Chihuahua.

  From all other species of the complex, bella is distinguished by the more abundant and finer spots in the cubital cell, by the more massive, more complex spot in the radial cell, and by the larger areolet. While any one of these characters may fail to characterize individuals, the combination of characters clearly differentiates series of any size.

  Cynips (bella) vanescens Kinsey
  agamic form

  Diplolepis bella err. det. Weld, 1930 (northern Arizona records only), Proc. U.S. Nat. Mus. 68 (10): 18.
  Cynips bella bella err. det. Kinsey, 1930 (Prescott, Ariz. record only) Ind. Univ. Studies 84–86: 289.
  Cynips bella vanescens Kinsey, 1930, Ind. Univ. Studies 84-86; 292, figs. 45, 282.
  Diplolepis bella err. det. Weld, 1930, Proc. Ent. Soc. Wash. 32:29.

  GALLS.—Very similar to all galls of the C. bella and C. dugèsi complexes. Mature galls rosy or brownish tan, not mottled, up to 28. mm., averaging near 20. mm. in diameter.

  HOSTS.—Definitely on Quercus subturbinella. The record of Q. grisea for the types is less certain. No records available to show whether other white oaks in the area are also involved.

  RANGE.-Arizona: Globe (types). Apache Trail at Fish Creek (galls only, Kinsey coll.). Cave Creek (Weld coll.). Prescott (Kinsey coll. and E. R. Leach, Jr., in Kinsey coll.). Hackberry, Ashfork, Williams (Weld coll.). Utah : Leeds, 4 W (Q. subturbinella). Pine Valley (Q. subturbinella). Zion Park (Q. subturbinella). Apparently ranging from Southwestern Utah (and Southern Nevada?) through the northern half of Arizona as far south as the Apache Trail and Globe. Figure 96

  LIFE HISTORY.--Adults: November 20, 27. December 14. Mostly before January 20. Some after January 20

  Originally described from a single insect from Globe in Central Arizona, this species is now recovered from several localities in Northern Arizona and in Southwestern Utah. While the new material makes modifications in our description of the species, the identity of the Utah and Arizona material seems certain, for some of the Utah specimens fully match the type in having the spots faint and reduced in all cells, including the cubital cell. The available locality records for this insect now mark the limits of the range of the host, Q. subturbinella (and its relatives?). This is an interesting scrub oak with Californian affinities. On the same oak, in the same area, Cynips (dugèsi) capronae produces a gall which is (ordinarily?) distinguishable by its heavy purple striping. In series the insect of vanescens is recognizable from other species in the bella complex by its combination of a smaller size, a very black body, and a small areolet.


externallink: https://www.biodiversitylibrary.org/page/53516882#page/296/mode/1up
description:
  Cynips bella var congesta, new variety
  agamic form

  GALL. — As described for the species, apparently indistinguishable from the galls of the other varieties of bella or from the gall of Cynips dugesi pupoides which occurs in the same region. On leaves of Quercus grisea.

  RANGE. — Texas: Alpine and Fort Davis (Kinsey coll.). Probably confined to the mountain ranges of western Texas (and adjacent regions?). Figure 45.

  These insects were emerging on December 14 (in 1919) at Alpine, Texas, and two days later at Fort Davis, Texas. Cynips dugesi pupoides, whose galls also occur at Alpine and Fort Davis, is a small, short-winged insect that cannot be confused with congesta.


description:
  Cynips bella congesta Kinsey
  agamic form

  Cynips bella congesta Kinsey, 1930, Ind. Univ. Studies 84-86:292, figs. 45, 281.
  Cynips bella var. Hoffmeyer, 1930, Ent. Meddel. Kbhvn. 17:214.
  Cynips bella vanescens (error) Hoffmeyer, 1933, Ent. Meddel. Kbhvn. 18:247.

  GALL.-Very similar to all galls of the C. bella and C. dugèsi complexes, apparently indistinguishable from galls of C. (dugèsi) pupoides which occur in the same region. Mature galls rosy tan, not mottled, up to 23. mm., averaging near 17. mm. in diameter. 

  HOST.-Quercus grisea, the evergreen white oak of the area

  RANGE.-Texas: Alpine. Fort Davis. Marfa, 22 S. Probably confined to the mountain ranges of West Texas. Figure 96.

  LIFE HISTORY..—Adults: December 14, 16.

  Restudy of our material in connection with the Mexican material now available in the complex confirms our opinion that congesta, from West Texas, is a distinct species. 


description:
  Cynips (bella) pomifera, new species
  agamic forms

  GALL.-Similar to all galls of the bella and dugèsi complexes. Mature galls rosy or (more often) light yellow tan, always conspicuously mottled with purplish brown; sometimes large, up to 22. mm., but averaging much smaller, usually not over 14. mm. in diameter. Figure 92. 

  HOSTS.—Quercus chihuahuensis, Q. undata. Probably on the other white oaks of the region. 

  RANGE.-Chihuahua: Namiquipa, 30 W, 5200' (Q. chihuahuensis). Madera, 10 SE (Q. chihuahuensis). Matachic, 3 N, 6700' (Q. chihuahuensis). Pedernales, 2 E, 7500' (Q. chihuahuensis). Santa ſsabel, 6 E, 6000' (Q. undata, types). Santa Isabel, 16 E, 6000' (Q. chihuahuensis, Q. undata). Apparently restricted to a portion of the Mexican Sierra in Central Chihuahua, known from Namiquipa to Santa ſsabel. Figure 96.

  LIFE HISTORY..—Adults: January 20, 29, 30. February 1, 2, 5, 9, 10, 15, 28. March 2. Most of the emergence early in February.

  While Cynips (bella) bella ranges widely over Southern Arizona, New Mexico, and Northern Chihuahua, C. (bella) pomifera is restricted to the 90 miles of Sierra which lie between Las Cruces and Santa Ísabel in more southern Chihuahua. This area represents only the southern half of the range of C. (dugèsi) emergens, an insect which produces a gall indistinguishable from that of pomifera. Pomifera and aspera agree in having rather small, light yellowish tan galls which are conspicuously mottled with purplish brown, and in this character they are distinct from the more northern species of the complex. The mottled galls of these two more southern species of the bella complex are readily distinguishable from the uniformly colored, dull-surfaced galls of the dugèsi species which occur in the same area. 

  All of the galls of C. (bella) bella from north of Las Cruces, Chihuahua, are uniformly colored, unmottled; all of the galls of C. (bella) pomifera from south of Namiquipa are conspicuously mottled. However, the collections from Las Cruces and from Namiquipa include galls of both types, thus giving evidence of some interbreeding of bella and pomifera where the two ranges meet. The insects from those localities also show some hybridization, although their characters are too indefinite to give as clear evidence as the galls supply. Considering the proportional amounts of the characters appearing in each locality, we have listed Las Cruces, Chihuahua, in the range of C. bella, and the more southern locality, Namiquipa, in the range of C. pomifera; but it is to be understood that both of these are in a transition area between the two ranges. 


description:
  Cynips (centricola) centricola Osten Sacken
  agamic form

  Cynips quercus centricola Osten Sacken, 1861, Proc. Ent. Soc. Phila. 1:53, 58. 
  Holcaspis centricola Beutenmüller, 1909, Journ. N.Y. Ent. Soc. 17:46. Davis, 1918, Journ. N.Y. Ent. Soc. 26:48. 
  Cynips centricola centricola Kinsey, 1930, Ind. Univ. Studies 84-86:297, figs. 47, 271, 284 (omitting the Illinois, Virginia, and Western Missouri records). 
  Cynips centricola Davis, 1930, Bull. Brooklyn Ent. Soc. 25:233. 

  To the data given in Kinsey 1930, add: 

  RANGE.-Missouri: Mineola (galls only). North Carolina: Marshall (gall only). 

  LIFE HISTORY..—Galls: Mature at end of September. Adults: In galls early in October and early in November. Emerging November 17, 26. December 8.

  Cynips (centricola) rubrae (Karsch)
  agamic form

  Diplolepis quercus-rubrae Karsch, 1880, Zeitschr. gesammt. Naturw. 53:293, pl. 6 figs. 4a,b.
  Cynips centricola rubrae Kinsey, 1930, Ind. Univ. Studies 84-86:301, figs. 47, 274, 275.
  Diplolepis centricola err. syn. Weld, 1931, Proc. Ent. Soc. Wash. 33:224.

  To the data given in Kinsey, 1930, add:

  RANGE.-Texas: Greenville (galls only).

  Weld (1931) compared specimens of centricola from Washington, D. C., with the Karsch types of rubrae at the Berlin Museum, and concluded that the two were synonyms. They are clearly in the same complex, but Weld's synonymy cannot stand until the characters of the several species in the group are taken into consideration. It is more likely that my strians is a synonym of rubrae. See the note under strians in the present study. 


description:
  Cynips (plumbea) plumbea Weld
  agamic form

  Cynips plumbea Weld, 1926, Proc. U.S. Nat. Mus. 68 (10):64, fig. 12. Metcalf and Flint, 1932, Fundamentals of Ins. Life: fig. 223. Houard 1935, Marcellia 28:101.
  Cynips (Philonix) plumbea Kinsey, 1930, Ind. Univ. Studies 84-86:245–248, figs. 36, 200, 225-226, 235-236, 246.

  GALL.-Large, hemispherical, with a broadly flattened base; surface smooth and shining, shriveling; bluish gray scurf easily wiped off, exposing the dark tan to purple brown color of the older gall; up to 11.0 mm., averag ing 8.0 mm. in diameter. Figures 103, 109. 

  HOSTS.—Quercus oblongifolia, Q. arizonica. 

  RANGE.-Arizona: Globe. Whetstone. Oracle. Santa Catalina Mts. Santa Rita Mts. Fort Huachuca. Courtland. Chiricahua Mts. Nogales. Patagonia Mts. Tumacacori Mts. Data in more detail in Kimsey 1930:245. Apparently restricted to Southeastern Arizona (and to immediately adjacent portions of New Mexico?). Figure 102

  LIFE HISTORY.—Pupae: December 7. Adults: November 22. December 14, 30. January 6, 8, 12, 13, 24, 25. February 3, 4, 6. (See Kinsey 1930:247). Most of the emergence early in January.

  This is the only species of the complex that occurs within the United States. It is apparently confined to the island mountains of Southeastern Arizona (and immediately adjacent New Mexico?). We have no data further than those published in our 1930 monograph, but include the present descriptions in order to make comparisons with the several, newly described species of the complex. The insect is closest to, and the gall is identical with, that of C. subfusca from Northern Chihuahua. The insects of plumbea. are not quite as rich and dark a brownish rufous, and average a bit larger. Plumbea insects are also close to C. euconus, from Durango, but euconus is unique in the complex in having conical galls with circular bases. 


description:
  Cynips (plumbea) claripennis, new species
  agamic form

  GALL.-Nearly spherical, with an occasional, short, fine, bluntly pro truding point; gall flattened only on a limited base; dull, slightly shrivel ing, with lead gray scurf rather persistent; older gall reddish tan to dark red brown; up to 7.5 mm. in diameter. Figure 112. 

  HOSTS.—Quercus intricata and Q. Pringlei, two shrubby oaks which were not well separated in the field at the time of our collections. 

  RANGE.-Tamaulipas: Tula, 22 N, 6100' (types). Probably restricted to a more northern area in the Eastern Sierra of Mexico, possibly restricted to the intricata group of dwarf oaks. Figure 102

  LIFE HISTORY..—Adults: February 2

  This insect is close to plumbea, subfusca, and glabrescens, but its wing veins are heavier, and the cubital cell (in our two insects) is practically free of all spots and blotches. The gall, moreover, is so distinctive that we can be certain that there is a distinct species represented here, even if the available insect material is quite inadequate. The gall is not broadly flattened as in the other species; it is dull and more or less persistently Scurfy, and decidedly warty, with an occasional protruding point. The hosts are such distinctive dwarf oaks that host isolation alone might account for the segregation of the species, even if it were not separated from the related species by two or three hundred miles of largely oak-free plateau lying between the Eastern and Western Sierras. 


description:
  Cynips (plumbea) euconus, new species
  agamic form

  GALL.-Unique, a warped cone (quite like a “chocolate bud”) in shape. The base strictly circular, flat or somewhat concave, up to 11.0 mm., averaging near 9.0 mm. in diameter, centrally bearing a very short ridge or single point of tissue by which the gall is attached to the leaf; the conical body of the gall rising directly from the circular base, rising less abruptly basally, prolonged into a long, sharply-pointed tip which is often a bit bent or twisted; the whole gall up to 11.5 mm., averaging under 9.0 mm. in height; in color a peculiar chocolate tan with a more or less abundant bluish gray scurf. Figure 115. 

  HOST.-Quercus sacame [arizonica]

  RANGE.-Durango: Patos, 15 W, 8000' (types). Known only from this one locality in more southern Durango. Figure 102.

  LIFE HISTORY..—Adults: February 10. March 10.

  This species is unique in the complex in having sharply conical galls with circular, flattened bases. This gall represents a mutation from the spherical or hemispherical galls of the com plex which is, in its way, as striking as the short-winged insect mutants which we have found so interesting in Cynips. The gall would never lead one to place the species in the plumbea complex, but the insect is strikingly close to C. plumbea and C. subfusca, the relatives immediately to the north, differing from them only in being more yellow brown in color with darker markings on the thoracic lines and on the sides of the mesothorax. 

  In our collections, euconus was found at only a single locality, Patos, in more southern Durango, but it was fairly common there. On the same trees, in the same locality, we found C. (plumbea) glabrescens. The significant thing is the close identity of euconus with the more northern insects of the com plex, and its more remote relation to glabrescens which occurs in the same locality with euconus. Euconus and glabrescens appear to have been independent mutants from the more northern stock of the complex; but the two species represent such diverse developments that they are apparently (probably physiologically) incapable of interbreeding. Thus they maintain their specific identity even though they occur on the same oaks in the same locality.


description:
  Cynips (plumbea) fucosa, new species
  agamic form

  GALL.-Very close to gall of texcocana. Almost perfectly spherical, or slightly flattened at base; dull, only slightly shriveling; with a conspicuous scurf which is persistent; old gall bluish gray; denuded gall dark purplish tan; up to 8.5 mm. in diameter. Figure 113.

  HOST.-A relative of Quercus chihuahuensis or of the dwarf oak, O. microphylla. The host of our material a small tree apparently not yet described. 

  RANGE.-Querétaro: Querétaro, 28 N, 7300' (types; trees near Q. microphylla). Guanajuato: San Felipe, 20 SW, 8000' (Q. chihuahuensis). Probably restricted to the high mountain areas of Central Mexico, centering in the states of Querétaro and Guanajuato. Figure 102

  LIFE HISTORY..—Adults: March 2, 5, 17

  This is the plumbea representative on the Q. chihuahuensis group of oaks in the high mountains which lie to the east of León and north of Querétaro, in Central Mexico. Both in insect and gall characters it is close to texcocana. Texcocana is confined to the mountains which rim the Valley of Mexico. 


description:
  Cynips (plumbea) fuscipennis, new species
  agamic form

  GALL.-Large, spherical, rarely flattened, almost always drawn out to a conical point basally; dull, slightly shriveling, with a conspicuous and heavy scurf which is quite persistent so old galls are often blue gray; de nuded surface light tan to (less often) dark purplish brown; up to 11.0 mm., averaging over 8.0 mm. in diameter. Figure 116. 

  HOSTS.—Q. nudinervis, Q. conglomerata, Q. texcocana, Q. rhodophlebia, Q. convallata. Apparently on all white oaks of the region except Alpine dwarfs and oaks of macrophylla group

  RANGE.-Morelos: Cuernavaca, 8 N, 8700' (Q. teaccocana; galls on Q. conglomerata). Guerrero: Taxco, 8 NE, 8000' (Q. mudimervis, types). Michoacán: Morelia, 14 E, 7000' (Q. conglomerata). Jalisco: Sayula, 13 SW, 7700' (Q. rhodophlebia, Q. convallata). Probably occurring over a wide area west and southwest of the mountains which rim the Valley of Mexico, from Morelos to Southern Jalisco and Eastern Guerrero. Figure 102.

  LIFE HISTORY.—Adults: February 20. March 2, 3, 12, 15, 22, 29. April 5. Most of the emergence late in March.

  The main road between Mexico City and Cuernavaca climbs Over a mountain pass at 10,200 feet. Although black oaks occur at the highest elevation, white oaks fail to reach the divide, and C. texcocana, occurring on the eastward face of the mountains, looking into the Valley of Mexico, seems effectually isolated from C. fuscipennis which occurs on the westward slopes of that same mountain ridge. Fuscipennis then ranges more than 300 miles to the west, over a very much dissected mountain country with arid, tropic valleys in it. The species extends even to the south ern end of the state of Jalisco. In the same area, C. hirsuta is a segregate of the same complex, but it occurs on the alpine dwarf oak, Q. repanda ; and C. subcostalis and C. scutata are segregates on the Q. macrophylla group of oaks. 


description:
  Cynips (plumbea) glabrescens, new species
  agamic form

  GALL.-Large, subspherical, with a broadly flattened or somewhat con cave base; dull; shallowly but very distinctly shriveling; with some lead gray scurf; older gall rather dark brown or reddish brown in color; up to 9.0 mm. in diameter. Often indistinguishable from galls of C. (arida) saxw lum which occur in parts of the same area. Figure 118. 

  HOSTS.–Quercus undata, Q. chihuahuensis, Q. sacame [arizonica], Q. reticulata [rugosa]; and possibly on the dwarf oak Q. depressipes. 

  RANGE.-Chihuahua: Santa ſsabel, 6 E, 6000' (gall; Q. umdata). Durango: Durango, 2 N, 6500' (Q. undata, types). Canatlán, 7 N, 7400' (Q. undata). Patos, 15 W, 8000' (Q. sacame, Q. chihuahuensis). Otinapa, 8500' (Q. sacame, Q. reticulata; galls on Q. depressipes 2). Probably restricted to a portion of the Western Sierra of Mexico, in southernmost Chihuahua, and in Durango. Figure 102.

  LIFE HISTORY..—Mature larvae: November 8–14. Adults: November 19. February 5, 10, 15, 18, 20, 22, 25, 28, 29. March 2, 4, 8, 11, 22. Most of the emergence late in February, early in March.

  This insect is closer in general appearance to plumbea than it is to subfusca, its immediate neighbor to the north. It differs from plumbea in several details, and in the interesting character of having hairs only on the second abdominal segment. The gall of glabrescens is obviously dull and shriveled on the surface, distinctly different from the smooth and shining galls of plumbea and subfusca. 

  We are under the impression that the mountains of Central Durango are so separated from the rest of the Sierras that the oaks do not range continuously from there into the northern part of the state. We found it impossible to travel the mountain roads between Indé and Santiago Papasquiaro, so we cannot fix the northern boundary of the range of glabrescens. To the south, from Zacatecas through Jalisco, glabrescens is replaced by C. jalisco. 


description:
  Cynips (plumbea) hirsuta, new species
  agamic form

  GALL.-Of moderate size, almost perfectly spherical; dull, slightly shriveling, with only a limited amount of scurf; denuded surface yellow to rosy tan (darker when weathered); up to 9.5 mm., averaging 7.0 mm. in diameter. Figure 117. 

  HOST.-Quercus repanda, a dwarf, alpine oak.

  RANGE.-Michoacán: Morelia, 14 E, 7000'. Probably restricted to alpine dwarf oaks, probably not extending beyond the mountains of Morelia. Figure 120.

  LIFE HISTORY.—Adults: February 25. March 2.

  The galls of this insect were fairly common on the low, dwarf oak, a member of the Q. repanda complex, in the mountains east of Morelia. Since our material represents a single collection, it is uncertain how limited the range of the species may be; but the closely related dwarf oak in the mountains rimming the Val ley of Mexico is host to C. texcocana, another (although unrelated) member of the plumbea complex. In the same mountains east of Morelia, C. (plumbea) fuscipennis occurs on the scrub or tree oak, Q. conglomerata. 

  The gall of hirsuta is similar to that of texcocana, fuscipennis, and fucosa, but hirsuta galls are more strictly spherical and lighter rose tan in color.


description:
  Cynips (plumbea) jalisco, new species
  agamic form

  GALL.-Indistinguishable from that of C. (plumbea) munda. Also in distinguishable from galls of C. (arida) eluta which occur in part of the same area. Of moderate size or small, either with a flattened base (when occurring singly) or drawn out to a conical tip (when occurring in clus ters); dull; very prominently shriveling; bluish gray with a more or less persistent scurf; older gall rose or purplish brown in color; up to 7.5 mm., averaging below 5.0 mm. in diameter. Figure 119. 

  HOSTS.—Quercus macrophylla [magnoliifolia], Q. haematophlebia [magnoliifolia], Q. undata, and Q. sacame [arizonica]. Probably on most of the oaks of the area except the dwarf, alpine oaks and the scrub oaks of the Q. potosima complex.

  RANGE.-Zacatecas: Cantuna, 6 N, 7600' (galls, Q. undata). Zaca tecas, 22 SW (Q. undata). La Mesa, 7000' (Q. saca me, Q. macrophylla). Aguascalientes: Pabellón, 20 W, 7000' (Q. undata, Q. macrophylla; galls on Q. sacame). Guanajuato: San Felipe, 20 SW, 8000' (Q. macrophylla). San Felipe, 8 NE, 7000' (Q. macrophylla). León, 20 NE, 9000' (Q. reticulata). Jalisco: Aguascalientes, 35 E, 7000' (Q. macrophylla; gall on Q. saca me). Atequiza, 5 E, 6500' (Q. haematophlebia). Guadalajara, 25 S, 6000' (Q. haematophlebia). Guadalajara, 10 W, 6000' (Q. macrophylla, types). Magdalena, 4 N, 4300' (Q. macrophylla; galls on Q. haemato phlebia). Probably on most of the oaks except those of the potosima complex, and restricted to an area in the Western Sierra and Western Cordillera of Mexico, from Zacatecas through Jalisco and Guanajuato. Figure 102. 

  LIFE HISTORY.—Adults: November 24. December 15, 18. January 1, 12, 30. February 11, 13, 15, 20, 25, 26. March 2, 4, 6, 20. Most of the emergence late in February, early in March.

  This species is very close to C. glabrescens, differing chiefly in being lighter in color, having usually heavier wing veins, a heavier infuscation on the first abscissa of the radius, darker splotches in the cubital cell, a smaller size, and a gall which is small and very much shriveled, being often drawn out to a point basally. The two galls are never to be confused, although it is not always possible to determine every insect. The geographic isolation of the two types of galls, and their correlation with the several insect structures, make it clear that in the high plateau plains which separate the mountains of Durango from the south ern portion of the Western Sierra, we have factors which have isolated distinct species. 

  Cynips munda is a segregate from the present species, which occurs on the scrub oaks of the potosina group in part of the range of C. jalisco. 


description:
  Cynips (plumbea) munda, new species
  agamic form

  GALL.-Indistinguishable from that of C. (plumbea) jalisco. Also in distinguishable from galls of C. (arida) eluta which occur in parts of the same area. Of moderate size or small, either with a flattened base (when occurring singly) or drawn out to a conical tip (when occurring in clusters); dull; very prominently shriveling; bluish gray with a more or less persistent scurf, the older gall rose or purplish brown in color; up to 7.5 mm., averaging below 5.0 mm. in diameter. Figure 111. 

  HOSTS.—Quercus potosina and Q. jaralensis, related scrub oaks, and probably on other species of the same complex. 

  RANGE.-Zacatecas: Zacatecas, 22 SW (Q. potosima, galls). Aguascalientes: Pabellón, 20 W, 7000' (Q. potosina ; types). Jalisco: Aguascalientes, 35 E, 7000' (Q. potosima). Guanajuato: San Felipe, 20 SW, 8000' (Q. Jaralensis). León, 20 NE, 9000' (Q. Jaralensis). 

  Probably restricted to oaks of the potosina complex wherever they occur, in an area in the Western Sierra of Mexico, from Zacatecas through Jalisco and Guanajuato. Figure 102.

  LIFE HISTORY..—Adults: February 20. March 2. April 15.

  Cynips munda occurs in much the same area as C. (plumbea) jalisco, from which species it is isolated by occurring on the very distinct potosina group of oaks. The galls of the two are identical. The insects are close, but munda is distinctly smaller. It has rather finer wing veins, and a cubital cell which is less heavily marked. The galls of munda are also indistinguishable from the galls of C. (arida) eluta which occur in the same region and on the same oaks occupied by munda. The insects must, therefore, be seen before determinations can be made. 


description:
  Cynips (plumbea) scutata, new species
  agamic form

  GALL.-Similar to gall of C. subcostalis. Rather large, subspherical, prominently drawn out to a conically tipped or truncate base; dull, only slightly shriveling, with light gray scurf prominent, persistent; denuded surface rosy tan or (more often) a peculiar salmon or rose brown; up to 9.0 mm. in diameter. Figure 107. 

  HOSTS.– Quercus macrophylla and Q. nudinervis [both magnoliifolia], which are the prevalent oaks of Guerrero. 

  RANGE.-Guerrero: Taxco, 8 NE, 8000' (types; Q. macrophylla). Possibly restricted to the eastern end of the state of Guerrero, Mexico. Figure 102

  LIFE HISTORY..—Adults: March 10.

  The gall is practically identical with the galls of subcostalis. Scutata and subcostalis are clearly geographic segregates from the texcocama group of species. Scutata occurs in Eastern Guerrero (known from near Taxco) on Q. macrophylla. Fuscipennis is found in the same area on Q. nudinervis. Subcostalis occurs in Western Guerrero (known from west of Chilpancingo) on both Q. macrophylla and Q. nudinervis. 


description:
  Cynips (plumbea) subcostalis, new species
  agamic form

  GALL.-Similar to gall of C. scutata. Very large, spherical, promi nently elongated into a conical or truncate base; dull, only slightly shriveling; light gray scurf prominent, quite persistent; denuded surface of gall rosy tan or (more often) a peculiar salmon or rosy brown to dark brown in color; up to 12.0 mm., averaging nearer 10.0 mm. in diameter. Figure 104. 

  HOSTS.—Quercus macrophylla and Q. nudinervis [both magnoliifolia] two oaks that seem more closely related than the botanists have indicated. These were the only white oaks we found in the area. 

  RANGE.-Guerrero: Chilpancingo, 6 S, 4000' (Q. macrophylla, types; Q. nudinervis). Tierra Colorada, 14 N, 3200' (Q. macrophylla). Probably confined to a more western portion of Guerrero and immediately adjacent states in Southwestern Mexico. Figure 102.

  LIFE HISTORY..—Adults: February 10. March 10, 18, 26. April 9, 10, 15. Emergence mostly late in March and early April.

  This species is known only from the western half of the state of Guerrero. The insect is close to C. fuscipennis which occurs in a territory just north and east. From fuscipennis, subcostalis is, however, perfectly distinct in having a very light, almost straw brown body, in having the full dorsal half of the abdomen naked, and in having a gall which is very large, basically conical, and peculiarly pink or salmon brown in color. The gall of subcostalis is nearly duplicated by the gall of scutata, a species which occurs on Q. macrophylla (not on Q. nudinervis) in the eastern part of Guerrero (known from near Taxco).  

  The galls collected on January 8, 1932, were peculiarly fresh and not yet mature—less mature than the galls of any other species of the complex which we had collected after the middle of November. The galls of subcostalis may be delayed because their hosts occur in Southern Mexico at lower elevations (down to 3200 feet), in regions which are warmer than most of those in which we found oak in that country. 


description:
  Cynips (plumbea) subfusca, new species
  agamic form

  GALL.—Practically identical with galls of C. plumbea. Large, hemi spherical, with a broadly flattened base; surface smooth and shining, not shriveling; bluish gray scurf easily wiped off, exposing the tan brown color of older galls; up to 11.0 mm. in diameter. Figures 105, 108. 

  HOST.-Quercus sacame, an oak closely related to Q. arizonica. 

  RANGE.-Chihuahua: Pacheco, 20 E, 5400'. San Buenaventura, 3 E, 5600'. Occurring on the eastern side of the Western Sierra of Mexico, prob ably restricted to Chihuahua (and northernmost Durango?). Figure 102.

  LIFE HISTORY..—Adults: January 20. February 6, 15.

  The galls of subfusca are practically identical with the galls of plumbea. The insects of subfusca differ only in being richer and darker brown and in averaging rather smaller; but these differences are so evident to the naked eye, and so constant in a series of 40 individuals of subfusca, that it seems evident that here is a population with a distinct genetic make-up. Although lack of adequate maps and records of explorations makes our knowledge of the topography and flora of Northern Chihuahua uncertain, we ourselves were unable to find oak in that state north of a point lying between Casas Grandes and Pacheco. This leaves something like 100 miles of oak-free mountains and very arid deserts isolating subfusca from plumbea. 

  The insects of subfusca are also close to those of C. euconus, from Southern Durango; but euconus galls are conical with circular bases.


description:
  Cynips (plumbea) texcocana, new species
  agamic form

  GALL.-Large, spherical, rarely flattened on a small base, more often drawn to a short point; dull, slightly shriveling, with a conspicuous and often heavy scurf (stellate pubescence) which is so persistent that even old galls are conspicuously blue gray; denuded surface of gall light tan to (more often) dark purplish brown; up to 10.0 mm., averaging near 7.5 mm. Figure 106. 

  HOSTS.—Quercus texcocana [deserticola], Q. conglomerata [rugosa], Q. rhodophlebia [rugosa], Q. repanda. Apparently one species on all oaks, both trees and alpine dwarfs, in the region. 

  RANGE.-Mexico: Mexico City, 12 W, 8500' (Q. tercocama, types; galls on Q. conglomerata, Q. repanda). Mexico City, 17 S, 8200' (Q. tear cocama, Q. conglomerata). Mexico City, 20 S, 9200-9400' (Q. rhodophlebia, Q. repanda). Mexico City, 15 E, 7000' (Q. tercocana). Apparently confined to the mountains bounding the Valley of Mexico; range limited by the divides which rise to elevations above those at which white oaks occur. Figure 102

  LIFE HISTORY..—Adults: March 15, 20, 23, 29, 30. Emergence thus late in March.

  On the inside of the remarkable mountain rim which bounds the Valley of Mexico, this is the only species of the plumbea. complex. Over the divide, to the west and southwest, on the outside of that same mountain rim, this species is replaced by C. fuscipennis, with still other species occurring on other groups of oaks in Central (and more southern ?) Mexico. 

  Texcocana is a very striking insect, in no likelihood of being confused with the plumbea-glabrescens group of species of more northern Mexico and our own Southwest. The galls of texcocana and fuscipennis are very similar, but the ground color of the texcocana gall is more often dark purplish brown, that of fuscipennis light tan.


description:
  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


description:
  Cynips (fulvicollis) gigas (Weld)
  agamic form

  Philonix gigas Weld, 1922, Proc. U.S. Nat. Mus. 61 (18): 12, fig. 2.
  Cynips fulvicollis gigas Kinsey, 1930, Ind. Univ. Studies 84-86:259, figs. 40, 229, 241, 250. Houard, 1935, Marcellia 28:126.

  To the data given in Kinsey 1930, add:

  RANGE.-Missouri: Kansas City (galls, Q. Michauxii). See Figure 120.

  LIFE HISTORY. —Galls immature on August 14.


description:
  Cynips (fulvicollis) insulensis, new species
  agamic form

  GALL.--Typical of the fulvicollis complex; pubescent, up to 9 mm. or more in diameter. Figure 121. 

  HOSTS.–In the Rockies: Q. Gambelii. In the Black Hills: A member of the Q. macrocarpa complex. East of the Rockies: Q. macrocarpa. 

  RANGE.-Illinois: Stockton, 3 W (gall). Wisconsin: Portage, 14 W. Portage, 7 W (gall). Stevens Point, 5 S (gall). Stevens Point, 14 S (gall). Waupun, 8 N. Lake Geneva. Sparta, 7 E (gall). Wausau, 5 S (gall). Minnesota: Badger (galls, R. Voris in Kinsey coll.). Harmony, 6 N. Iowa: Postville, 3 E (galls). Nebraska: Seward (galls). Nebraska City (galls). South Dakota: Black Hills State Park. Deadwood (types). Wyoming: Sundance (galls). Colorado: Happy Canyon (20 SE Denver). Manitou (galls). Palmer Lake (galls). Occurring in the Rocky Mountains east of the Divide, and throughout the northern Middle West north of Missouri and west of Lake Michigan. Figure 120.

  LIFE HISTORY.—Galls: immature mid-August to September 1. Adults: November 4, 13, 20, 25. December 1, 10, 24, 26, 28. Emergence in part in the first year, in part in the second year after formation of the gall. 

  The present species closely resembles the eastern canadensis and the Utah species latigenae; but insulensis has parapsidal grooves which are more sharply defined than in any other species of the fulvicollis complex. 

  It is interesting to find the range of this species discontinuous. It extends into the Rocky Mountains east of the Divide in Colorado (and Northern New Mexico?); into the essentially Rocky Mountain island which is the Black Hills; and into the eastern deciduous forests extending from the eastern part of the Dakotas through Minnesota and Wisconsin to Lake Michigan. While these three areas are well separated by arid plains, there is no good barrier to limit the range toward the south and east, and through Iowa and Northern Missouri we may expect to find the species hybridizing with C. vorisi. We have numer ous insects of the species, but select the types from our limited material from Deadwood, on the north side of the Black Hills. There the complete isolation of the area eliminates all hybridization of this with other species of the complex. 

  Most interesting of the data presented by this species is the evidence that Quercus Gambelii, representing the only complex of broad-leaved white oaks in the Rocky Mountain area, has its closest relative in Q. macrocarpa in the eastern part of the United States. Extending east from its original home in the Rockies, the fulvicollis complex changed from Gambelii to macrocarpa, and only subsequently to Q. alba and the chestnut oaks. Indeed, the change from Gambelii to macrocarpa was so simple that the one species, insulensis, bridged the gap without having to develop mutant forms which would adopt the new host. 


description:
  Cynips (fulvicollis) latigenae, new species
  agamic form

  GALL.--Typical of the fulvicollis complex; always pubescent, not smooth; averaging 7.0 mm. in diameter. Figures 122, 123.

  HOST.-Quercus utahensis [gambelii]

  RANGE.-Utah : Kamas (types). Possibly wide-spread in the mountains of more northern Utah. Figure 120

  LIFE HISTORY..—Adults: second winter after formation of gall

  This species was collected in Northern Utah, very near the extreme, northeastern limit of oak in the Rocky Mountain system, at a point over 800 miles west of the range known for the complex at the time of publication of our Cynips monograph. In this same period we have found the complex in the Black Hills of South Dakota, and in Eastern Colorado. 

  It is remarkable that latigenae very closely resembles the most northeastern member of the group, C. canadensis. The ranges of the two are separated by nearly 1000 miles of plains and mountains in which oak does not occur. I have considered the possibility of canadensis having been carried westward on nursery stock of its eastern host, Q. alba. The alternate genera tions of these insects live in small and inconspicuous cells in otherwise normal buds, and they might easily escape inspections exercised in horticultural quarantines. But we have good herbarium material of the oak host of our Kamas material, and it certainly belongs to the Utah species, Q. utahensis, and not to the Q. alba group of oaks. 

  Although the characters are few, I believe that latigemae can be separated from canadensis by its more brightly rufous face and legs, its slightly longer wings, and cheeks which are distinctly broadened back of the eyes. This last is the distinctive character. In canadensis the cheeks are not broadened behind the eyes, and the head, in dorsal aspect, appears solid, somewhat chunky. The protruding cheeks in latigemac give a dorsal aspect of a finely formed head that seems shallow in its anterior posterior dimension. Finally, the 15 galls which we have of latigenae are all roughened and entirely pubescent, never smooth and shining as they are in canadensis. 

  From insulensis, of the Black Hills of South Dakota, Eastern Colorado, Minnesota, and Wisconsin, the present species is well distinguished by its dark thorax, more poorly defined parapsidal grooves, and shorter wings. 


description:
  It is unclear whether this should be considered a variety of [Philonix fulvicollis](https://www.gallformers.org/gall/825) or a distinct species in modern taxonomic terms.


description:
  Cynips (arida) arida Kinsey
  agamic form

  Cynips (Acraspis) arida Kinsey, 1930, Ind. Univ. Studies 84-86:313, figs. 57, 290, 291, 333, 354, 403. 

  GALL.-Indistinguishable from the galls of most other species of the complex. Mature gall dome-shaped with a broadly flattened base, shriveling, purplish brown in color, averaging near 5.0 mm. in diameter, a little smaller than saxulum or eluta. Figures 137, 141. 

  HOST.-Quercus grisea. The only evergreen white oak of the area. 

  RANGE.-Texas: Alpine. Fort Davis. Marfa, 22 S. Chisos Mountain Park. - Apparently restricted to West Texas. Figure 131

  LIFE HISTORY.—Adults: November 21. December 14. January 8.

  This species seems to be confined to West Texas, the most nearly adjacent localities from which we have material in New Mexico and Chihuahua yielding another species of the complex, namely C. tenebrica. The galls of arida are indistinguishable from those of tenebrica and, except for their slightly smaller size, from those of C. saxulum and C. eluta.

  There is no species of the C. plumbea complex known from the range of arida, and the most nearly adjacent species of plumbea have smooth, hemispherical galls which are readily distinguished from the dome-shaped galls of arida.


description:
  Cynips (arida) conica Kinsey
  agamic form

  Cynips (Acraspis) conica Kinsey, 1930, Ind. Univ. Studies 84-86:338, figs. 57, 288-289, 336, 361, 404

  GALL.-A smooth, pointed, conical gall with broadened base bearing spiny projections, differing in shape but similar in surface and internal structure to all other galls of the complex. Mature gall regularly or irregularly conical, up to 7.0 mm. in height, sometimes with tip rather sharply pointed and even curved; base sometimes flaring, its edge entire or bearing up to 6 short, blunt, spiny projections, up to 6.0 mm. in diameter; light pinkish brown in color, with a violet tinge due to a puberulence. Figure 133, 134. 

  HOST.-Quercus subturbinella (erroneously ascribed to Q. grisea in the original publication). 

  RANGE.-Arizona: Globe. Utah : Leeds (galls). Zion Park (galls). Apparently extending from the Apache Trail country of Central Ari zona to Southwestern Utah. Figure 131.

  LIFE HISTORY.—Galls: immature first week in September. Adults: mostly emerged before January 20, stray adults after that date.

  This insect is far enough removed from the others in the complex to have been published originally as a representative of another group, and it might still be maintained in that posi tion. In its nearly uniform brownish rufous color, in its rugose foveal groove, and particularly in its absence of infuscations and Splotches in the radial and cubital cells, this insect is distinct from any neighbor in the arida complex—although, curiously, it approaches the two most southern of the Mexican and, even more closely, the Guatemalan species of the complex. But conica is also close to C. arida in West Texas, and occupies a geographic area in which no other member of the complex occurs on the host, Q. subturbinella. Conica appears to be the host or geo graphic isolate, albeit an extreme development, of the arida complex. It is an academic question whether it should be considered in another complex, or a link in the endless chain of species by which one complex is transformed into another


description:
  Found frequently throughout the Western US on Great Basin sagebrush. The shape varies from rounded to lumpy/lobed.


externallink: https://www.biodiversitylibrary.org/page/8635512#page/195/mode/1up
description:
  Gutierrezia sarothrae
  Oval, flower bud gall, length 7 mm. Felt '15i, p. 274
  Itonid. Gutierrezia bud midge, Rhopalomyia gutierreziae


externallink: https://www.biodiversitylibrary.org/item/109257#page/280/mode/1up
description:
  This species produces a pale green, fusiform or suboval swelling in the flower heads of Gutierrezia sarothrae. The gall is about 7 mm long, 3 mm broad and was taken at Las Vegas, N. M., adults appearing October 31st. An examination of the types in the United States National Museum shows that it probably belongs to this genus. 


description:
  All rearing records of this species are of the second generation which produces galls in the top half of the plant stems. The overwintering galls are probably crown or root galls. Two galls on *Bidens frondosa* were found on the node of the plant.

  Stem gall on *Bidens frondosa, Chrysanthemum sp., Helianthus giganteus, H. tuberosus, Vernonia noveboracensis*


description:
  *E. divisa* has been reared from two galls of *Artemisia tridentata var. tridentata*. The galls measure 8 by 9 mm. and 7 by 9 mm. and are very much like the galls of *E. diana*.

  Stem gall on *A. tridentata*


description:
  A crown gall formed (Stoltzfus, 1966) as the larva developed. No stunting of infested plants was observed in the fall of the year but in the spring as the larvae matured rapidly the stems of infested plants were sometimes severely stunted, resulting in lack of flowering.  Galls were first noticed in October when the larvae were in the second instar. By June 15 of the following year the larvae were fully grown. The first puparium was taken June 17 in northeastern Ohio. The galls form at ground level and are nearly round.  They range in size from 7 to 11 mm. The larval cavity in the gall is only slightly larger than the larva. Pupariation occurs within the gall.

  Crown gall on *Aster simplex*


description:
  Galls were found on the lateral branches of *Artemisia cana* at the nodes which produced the last years growth. The size of the branch beyond the gall was generally not affected by the gall development, although in two cases where unusually large galls formed, a branch did not develop beyond the galls.

  Stem gall on *Artemisia cana*


description:
  *E. novaeboracensis* is probably an oligophagous species attacking rhizomes of *Solidago altissima* and occasionally attacking one or more species of *Solidago*. Such a range of larval food follows more closely the typical pattern of the gall-forming Tephritidae.

  Tunnels rhizomes, rhizome gall on *Solidago sp.*


description:
  *E. oregona* produces galls on *Artemisia tridentata tridentata* and *A. tridentata vaseyana*. The galls range in size from 3 by 5 mm. to 4 by 6.5 mm. Eggs laid in the lateral buds during June and July hatch and larvae induce galls from which adults emerge the following June.

  Stem gall on *A. tridentata ssp.* 


description:
  This species has now been reared from upper stem galls in *V. alternifolia* and *V. occidentalis* (L.) Walt. from western Cades Cove with adults emerging in May through September. Amnon Freidberg also reared two specimens (USNM, TAUI) of this species in 1993 from galls on *V. alternifolia* from Washington, DC, Rock Creek Park (A. L. Norrbom, personal communication). These are new host records for E. rotundipennis. *Verbesina alternifolia* appears to be the preferred host with *V. occidentalis* utilized only late in the summer. (p.3)


description:
  This unknown midge induces white haired galls at the terminal end of Ericameria nauseosa stems. The galls are sometimes composed of small hairy bumps and sometimes are only made up of white wool. There are always wide leafy bracts flanking the gall on all sides, sometimes standing erect and other times (usually with the bumpy galls) at a 90 degree angle to the stem. The galls apparently do not stop stem growth, as stems have been seen growing through galls several times. Very common on Ericameria in New Mexico. This species is tentatively synonymized with Russo's "Rhopalomyia sp. B". 


description:
  The gall of *A. lutea* is a simple but abrupt swelling on a stem (Fig. 2). It may occur on older stems as well as the current year's growth, but it is always woody with thickened walls. Among 3 galls dissected, 2 were monothalamous and measured 9.5-14.5 mm in length, 4.5-5.5 mm in width, and 5.0-8.0 mm in depth. The polythalamous gall contained 2 larvae and measured 14.0 x 5.0 x 8.0 mm. The cavity size for each was 5.0 x 2.0 mm. The only noticeable effect *A. lutea* galls had on the host plant was the stunting or prevention of stem growth from axillary buds on the swelled portions of stems.


description:
  The *A. maculata* gall matches an earlier description by Bates (1935). The gall is monothalamous, basically ovoid in shape but usually with an irregular profile due to bracts on the surface (Fig. 3). The entire gall is covered with a thick, white, cottony tomentum. Ten galls ranged from 7.5 to 10.0 mm (Jc = 8.9 mm) at their largest diameter. In cross section there are three distinct tissue layers. The innermost layer is hard surrounded by a thicker, pithy tissue layer which is covered by a thin outer tissue layer. The internal cavity of 10 galls measured 3.0 to 5.0 mm (Jc = 4.35 mm) in length and 1.0 to 3.0 mm (Jc = 1.90 mm) in width. Galls occurring on different subspecies of *C. nauseosus* are not noticeably different in form. Galls first appear in August as shining green, sticky, globular structures at the axillary buds. At this stage they are not distinguishable from young galls of *Aciurina semilucida* (Bates) and *Aciurina trixa* Curran. Within a few weeks they increase in size and a noticeable tomentum forms; although less than half grown, the young galls resemble mature ones. *Aciurina maculata* gall development is unusual in that growth occurs at a time of year when the plant becomes dormant; most commonly, the growth of galls occurs simultaneously with the growth period of the host (Mani, 1964). In such instances it is not always clear whether the formation of the gall is a mechanical reaction associated with the normal plant growth processes or if the gall-former produces growth stimulation substances. In the case of *A. maculata* the occurrence of a growth promoting chemical is a distinct possibility. The apparent stimulus for growth is the feeding of an *A. maculata* larva. Galls grow and develop as long as the gall-former feeds. By fall, when *A. maculata* has reached the third instar and ceases feeding, the gall is fully formed. Galls with dead *A. maculata* larvae were found to be consistently smaller than those containing living larvae. A gall may remain attached up to one or two years following emergence. *Aciurina maculata* galls are found primarily on the lower portions of branches, oftentimes in the adjacent axils of a stem. Also, new galls may form on stems from previous years' growth near the base of the plant. The distribution of galls is often clumped. Galls prevent growth of the affected axillary bud since the tissue of the bud is totally modified in gall formation. 

  [pg. 714 photo 3 is a monochrome photo of the gall]


description:
  *Aciurina opaca* is so rarely collected that little can be stated about its biology. Each of two specimens collected were reared from galls on *C. nauseosus*. A single female emerged from a gall collected on 18 March 1974 and a single male was reared from a gall collected on 21 April 1974. The flies issued from stem swellings that appeared identical to those described for *Valentibulla spp*. (Wangberg, 1978). It is not known how the apparent gall-forming habit of *A. opaca* differs from *Valentibulla spp*.


description:
  (as *Trupanea (Tephritoides) subpura*)

  The larvae feed in the tender growing stems of Baccharis glomeruliflora, tunneling out as much as an inch of stem and usually, but not always, forming slight swellings.


description:
  Palmer (1987) reported this fly to be common on *B. halimifolia* in southeastern Texas and southern Louisiana where it forms stem galls. Wasbauer (1972) lists *B. halimifolia* in reference to an unpublished host list of Foote. Various specimens examined here were reared from flowers and stems of *B. halimifolia* and others were recorded from only the stems of *B. glomeruliflora*.


description:
  A single rearing record was found in material examined, although numerous specimens bore labels that in some way associated them with *Baccharis*. Four light morphs (three males, one female) emerged on 19 May 1972 from *B. pilularis* collected at Walnut Creek (Contra Costa County), Calif. The plants were in a stage of vegetative growth, but the exact structure from which the flies emerged is unknown (J. A. Powell, personal communication). However, these data, when compared with similar data for the three other seasonally varying species, suggests that the specimens were reared from stems. This species also has been reared from *B. pilularis consanguinea* and otherwise associated with *B. emoryi* A. Gray and unidentified *Baccharis* species (Foote & Blanc 1963).


description:
  Adults of *T. palmeri* have been reared from female flowers and terminal stem galls of *B. neglecta* from southcentral and southeastern Texas (Palmer [1987] and specimens examined). Three specimens collected on 25 April 1984 at Victoria, Tex., were reared from *B. halimifolia*.


externallink: https://www.biodiversitylibrary.org/page/28254937#page/780/mode/1up
description:
  Aciurina semilucida (Bates) is a rare, highly localized, tephritid fly forming pubescent, axillary bud galls on branches of Chrysothamnus viscidiflorus (Hooker) Nuttall in southern California. Documentation is provided that the wing patterns of the adults are very strongly sexually dimorphic in southern California, but much less dimorphic in some other parts of its range, e.g. in Idaho, where it previously has been reported only from C. nauseosus [Ericameria nauseosa] (Fallen) Britton.

  The gall of A. semilucida is a stunted axillary branch, reminescent of the beaked galls of A. michaeli (Goeden and Teerink 1996a), but not as strongly attenuated or "beaked" apically (Fig. 6E, 6F). As such, they are dark green and subspheroidal when young (Fig. 6B, 6D), elongating and externally becoming pyriform in outline, light green, and pubescent as they grow and mature (Fig. 6E, 6F). They also bear several basally expanded, sessile, linear, smooth- margined leaves, which are shortest apically (Fig. 6E). The gall body is largely composed of whitish, spongy parenchymatous, expanded nodes, and a harder core surrounding the central locule (Fig. 6D, 6E); this description agrees with that of Wang- berg (1981), which stated that, apparently including the epidermis, the galls are composed "of three distinct layers" (Fig. 6E). The gall is uniloculate (monothalmus [sic]) and characteristically contains an ovoidal to pyriform-shaped, central, open chamber within which the larva feeds by scraping away at the inner wall which becomes shallowly pitted (Fig. 6D, 6E). The photographic illustration in Wangberg (1981) shows the surface of the gall of A. semilucida as dull, not shiny, and thus more commensurate with pubescent, and what he termed "smooth." His photograph also better reflects what he described as its "ovoid," but not its alternative "tear-drop" shape. However, the 12 voucher specimens of galls attached to some of his reared specimens of A. semilucida on C. nauseosus from Idaho were, indeed, pubescent, slightly attenuated apicad, and bore shortened, basally expanded leaves more like the California galls described above (Fig. 6E, 6F). The galls of A. semilucida at our study sites externally also closely resembled the whitish gall of a gall midge (Diptera: Cecidomyiidae) also found on C. viscidiflorus, possibly Asphondylia bigeloviae brassicoides (Townsend) as described and illustrated by Gagne (1989).


description:
  Given that this gall is nearly identical in host, range, and morphology to Kinsey's [Cynips (arida) conica](https://gallformers.org/gall/3397), it is conspicuous that the description of A lovellae makes no mention of that species. It seems likely that the two are conspecific but this cannot be confirmed with current literature. 


externallink: https://www.biodiversitylibrary.org/item/54981#page/627/mode/1up
description:
  Valentibulla dodsoni Foote is a recently described species (Blanc and Foote, 1987) from New Mexico which represents a considerable range extension for the genus. Valentibulla was established by Foote and Blanc (1959) and the other 5 members of the genus are restricted to California, Idaho, Nevada, Oregon, Utah, and Washington (Blanc and Foote, 1987). The host plant reported here, C n. ssp. bigelovii, is added to the only other host plant records for this genus: C parryi and two other subspecies of C nauseosus (Wangberg, 1978). The gall of this species is a rather inconspicuous swelling of the stem (Fig. 1 D). The biology and behavior of V. dodsoni is described elsewhere (Dodson, 1987). [According to this paper all other Valentibulla species are found outside of New Mexico, Arizona, and Colorado, so maybe any Valentibulla found within this region can be presumed to be dodsoni]


externallink: https://www.biodiversitylibrary.org/page/16152770#page/571/mode/1up
description:
  Another ambiguity is the report by Novak et al. (1967) that V. steyskali Foote [as munda (Coquillett)] larvae form "large, polythalmous galls" on stems of C. nauseosus in Idaho.


externallink: https://www.biodiversitylibrary.org/page/16213158#page/501/mode/1up
description:
  A cecidomyiid gall midge, Rhopalomyia bigeloviae (Cockerell), was reared from galls formed on galls of the tephritid, Aciurina trixa Curran, on Chrysothamnus nauseosus (Pallas) Britton in southern California. Only one of the two A. trixa gall types in southern California hosted the gall midge. Rhopalomyia bigeloviae galls were originally discovered in Colorado on the large cottony galls of A. bigeloviae, a species closely related to A. trixa, that forms galls on C. nauseosus outside of California. The midge galls are similar to the host tephritid galls externally, i.e. either smooth or cottony. The number of midge galls per tephritid gall varies, but they can, in some cases, cover the entire surface of the tephritid gall. The gall midge is probably bivoltine, and the tephritid host is univoltine. Gall midge galls first become visible in early spring (March). Adults emerge later in the spring. The fate of these adults remains unknown, but females may oviposit into other parts of the host plant to form a second generation of cecidomyiid galls not dependent on the presence of a tephritid gall.

  On one of the two gall forms in southern California (Fig. lA), we observed that its exterior sometimes was covered by small nodules at different densities among different locations. Upon dissection, each of these nodules was found to contain a small cecidomyiid identified for us as Rhopalomyia bigeloviae (Cockerell) by Raymond J. Gagne (Systematic Entomology Laboratory, Agricultural Research Service, USDA). This species was originally described from a large woolly gall up to 12 mm long, typical of that made by another species of Aciurina (Gagne 1986). Dodson and George (1986) and Headrick et al. (1997) have clarified the status of Aciurina species on C. nauseosus, and this woolly gall is now attributed to the tephritid, A. bigeloviae (Cockerell). We have examined such woolly galls from various collections (Washington State University, Pullman; University of Idaho, Moscow; G. Dodson, personal collection) and they do, indeed, bear R. bigeloviae galls. None of the other gall types, as described by Wangberg (1981), examined from museum collections throughout western U.S., including the smaller resinous gall of A. trixa found in California hosted this gall midge (Headrick et al. 1997).


description:
  [*T. arizonensis* has been reared from *B. sarothroides* in Tucson, AZ, and Otay, CA. These rearing records come from terminal galls, stem tip mines, and flower heads. Swelling from stem-feeding larvae only occurred during pupation.]


description:
  *Tephritis arizonensis* Quisenberry is bivoltine and monophagous, or nearly so, on *Baccharis sarothroides* Gray (Asteraceae). The F1 larvae feed and develop in non-galled, branch-tip mines, the only Nearctic Tephritidae known to do so; whereas, the F2 larvae develop singly in male or female flower heads. (p. 215)

  The lengths of 177 branch-tip mines measured from base to branch apex was 6.6 +- 0.1 (range 3-8.8) mm. The widest width was 1.3 +- 0.1 (Range, 1.0-1.8; n=138) mm at the base of the mine. Mines were scattered over the crowns of host plants. (p. 216)


description:
  [*Tephritis baccharis* was described in Coquillett, 1894, as being from a "irregular oval gall... on tender twigs of *Baccharis viminea*". *T. baccharis* has been reared from San Diego County and Los Angeles County.]


description:
  Tephritis baccharis (Coquillett) is a bivoltine and monophagous on Baccharis salicifolia (Ruiz & Pavon) Persoon in southern California... First instar larvae hatch and tunnel into the pith just basad to the apical bud where they initiate gall formation. Gall and larval growth continue slowly into the fall, where pupation occurs. (p.86)

  *T. baccharis* has been reared only from bud galls on *B. salicifolia*. The young larva settles, molts, and initially excavates a small ellipsoidal cell about 3 mm long and <2 mm wide in the juvenile gall, evidenced externally as only a slight swelling of a branch terminal... Twenty of 23 fully formed F2 galls of the previous year’s overwintering generation sampled at Oriflamme Canyon were fusiform; the remaining galls, subovoidal. The former shape resulted from continued growth of the bud at each gall apex; whereas, the latter shape resulted from death of the terminal bud from larval feeding by *T. baccharis*... The galls of *T. baccharis*, like those of *T. stigmatica*, are shortened, thickened, succulent terminal parts of main or axillary branches. The 23 mature galls incorporated an average of 3.8 +- 0.3 (range, 1-6) nodes, and were smooth surfaced, light green and longitudinally striped or unilaterally colored red-purple when occupied. When empty, the galls turned tan, shriveled, and dehisced, or became woody and persisted as branch swellings. Externally, the 23 galls averaged 13.1 +- 0.5 (range, 8-9) mm in length, and 5.6 +- 0.2 (range, 4-7) mm in width. (p. 94)

  Twenty of 30, F1 galls collected on Santa Cruz Island were fusiform; the remainder, subovoidal. These 30 galls incorporated 3.8 +- 0.1 (range, 3-5) axils and externally measured 20.4 +- 1.0 (range, 12-33) mm long by 8.3 +- 0.3 (range, 6-13) mm wide. (p.95)


description:
  Aceria sp.

  An eriophyid leaf-curling mite (recently identified as an undescribed species of Aceria) that can have severe impacts on the growth of New Mexico olive (Forestiera pubescens Nuttall var. pubescens) in nursery and landscape settings in the southwestern US. Once infected, individual trees generally retain populations of the mite from year to year, suggesting that it overwinters under the bud scales. Symptoms of leaf curl appear about 3 to 4 weeks after bud-break, and the growth of affected trees can be severely impacted. The entire margin of affected leaves curls inward on the underside of the leaf, forming a tight seal against the edge of the petiole; the mites feed and reproduce within this protected habitat, making it hard to reach them with non-systemic pesticides. 


description:
  Tight and profuse bud proliferation on Forestiera pubescens, creating a black-red-brown color and intricately rough-folded texture similar to the galls induced by [Aceria parapopuli](https://www.gallformers.org/gall/676). Observed in New Mexico in January. 


description:
  This undescribed species induces globular galls at the end of twigs, sometimes with active buds on it. The galls are very hard and the surface is mottled with white over the dark gray color of the branch. On the top of the gall there is always a sizeable hole, flanked on each side by tan flaps, that appears to be a natural feature of the gall rather than being chewed by emerging adults. Adults were reared in New Mexico and identified as an undescribed Contarinia species. 


description:
  Identification to genus Rhopalomyia provided by Raymond J. Gagné in a personal communication to Greg Froelich on June 2, 2024, as documented [here](https://www.inaturalist.org/observations/215352655). Dr. Gagné based the ID on examination of the empty gall. 

  A tight, white-hairy, leafy rosette on Hazardia squarrosa. Observed in California in January. 


externallink: https://www.biodiversitylibrary.org/item/54815#page/554/mode/1up
description:
  Hormaphis hamamelidis (Fitch 1851) 

  Brysocrypta hamamelidis Fitch, 1851. 
  Hormaphis hamamelidis (Fitch): Morgan & Shull, 1910. 
  Hormaphis shulliana Bomer, 1952. 
  Hormaphis hamamelidis (Fitch): von Dohlen and Gill (1989) [in part].

  Gall: Conical with slight or no basal constriction, tip often bent to one side, protruding from upper leaf surface and usually on a major vein; length from tip to base on leaf underside 6.3-11.8 mm (mean = 8.1) and maximum width 3.1-5.5 mm (mean = 4.4). Base of gall on leaf undersurface circular and somewhat swollen beyond leaf surface, the circular opening at first obstructed by plant hairs, later forming a small exit hole. Color pale green, turning yellow. 

  Biology: The host is Hamamelis virginiana L., witch-hazel, on which conical leaf galls can be found from May through September. Eggs hatch in mid-April, and first- instar fundatrices crawl to leaf buds. At bud break (early May) first-instar fundatrices start feeding on emerging leaves and galls begin to form. By mid-June fundatrices are mature, and they reproduce sexuparous nymphs through September. Alate sexuparae leave galls from late July to September and produce nymphs of sexuales directly onto the undersides of witch-hazel leaves. From early August until late September, oviparae and males are found on undersides of witch-hazel leaves where they mature (by late August) and mate. Each ovipara lays several eggs in bark crevices on the host plant, usually on older stems with more bark texture but occasionally at the base of leaf buds. Overwintering eggs are found on witch-hazel from September to April. See von Dohlen and Gill (1989) for additional information. 

  Summary: Hormaphis hamamelidis forms small, conical galls usually without basal constriction, is autoecious on witch-hazel, and lives at high elevations in northern regions of the eastern United States. Hormaphis cornu forms larger, conical galls that are usually constricted at the base, is host-alternating between witch-hazel and river birch, and is found at low elevations in the mid-Atlantic and southern regions of the eastern United States. 

  Range: VA, DC, NY


externallink: https://www.biodiversitylibrary.org/item/54815#page/559/mode/1up
description:
  Hormaphis cornu (Shimer, 1867) 

  Hamamelistes cornu Shimer, 1867. 
  Hormaphis hamamelidis Osten-Sacken, 1861 [secondary homonym of Hamamelidis Fitch 1851]. 
  Hormaphis hamamelidis (Fitch): Walsh, 1867; Pergande, 1901; von Dohlen and Gill, 1989 [misidentifications]. 
  Hormaphis hamamelidis Osten-Sacken: Thomas, 1877 [misidentification]. 
  Pemphigus hamamelidis (Fitch): Thomas, 1877 [misidentification]. 

  Gall: Conical but usually constricted at the base, tapered tip often bent to one side, protruding from upper leaf surface and usually on a major leaf vein; length from tip to base on leaf underside 8.2-19.4 mm (mean - 13.5) and 4.0-8.1 mm (mean = 5.6) at greatest width. Underside of gall on lower leaf surface circular and somewhat swollen beyond leaf; the opening at first obstructed by plant hairs, later forms a small circular exit hole. Color pale green, turning yellow in early summer. 

  Biology: Fundatrices hatch in late March from overwintering eggs laid on the bark of witch-hazel, form cone-shaped galls on leaves and produce virginoparous emigrants into mid-June. Alate virginoparae depart from galls in June and migrate to river birch to produce the first aleurodiform generation on the undersides of leaves. Two more aleurodiform generations follow; the last produces sexuparous nymphs. Mature sexuparae depart from leaves of river birch and fly back to witch-hazel in mid-September to early October and deposit sexual nymphs. Sexuales mature in late September to late October and mate. Ovipara produces several overwintering eggs which are laid in crevices in the bark or occasionally at the base of leaf buds. See von Dohlen and Gill (1989) for additional information.

  Summary: Hormaphis hamamelidis forms small, conical galls usually without basal constriction, is autoecious on witch-hazel, and lives at high elevations in northern regions of the eastern United States. Hormaphis cornu forms larger, conical galls that are usually constricted at the base, is host-alternating between witch-hazel and river birch, and is found at low elevations in the mid-Atlantic and southern regions of the eastern United States. 

  Range: VA, DC


externallink: https://www.biodiversitylibrary.org/item/35191#page/203/mode/1up
description:
  Hemispherical or subglobose, green, glabrous, cortical, stem swellings, becoming brown and resembling Lecaniums, diameter 3.5 mm, height 2 mm. Fig. 226. Felt 'i6f, p. 188
  Itonid. Lecanoid gall, Rhopalomyia glutinosa Felt


description:
  Hormaphis cornu (Shimer, 1867)

  Hamamelistes cornu Shimer, 1867.
  Hormaphis hamamelidis Osten-Sacken, 1861 [secondary homonym of Hamamelidis Fitch 1851].
  Hormaphis hamamelidis (Fitch): Walsh, 1867; Pergande, 1901; von Dohlen and Gill, 1989 [misidentifications].
  Hormaphis hamamelidis Osten-Sacken: Thomas, 1877 [misidentification].
  Pemphigus hamamelidis (Fitch): Thomas, 1877 [misidentification].

  Alate virginoparae depart from galls in June and migrate to river birch to produce the first aleurodiform generation on the undersides of leaves. Two more aleurodiform generations follow; the last produces sexuparous nymphs. Mature sexuparae depart from leaves of river birch and fly back to witch-hazel in mid-September to early October and deposit sexual nymphs.

  Summary: Hormaphis hamamelidis forms small, conical galls usually without basal constriction, is autoecious on witch-hazel, and lives at high elevations in northern regions of the eastern United States. Hormaphis cornu forms larger, conical galls that are usually constricted at the base, is host-alternating between witch-hazel and river birch, and is found at low elevations in the mid-Atlantic and southern regions of the eastern United States. 

  Range: VA, DC


description:
  Hamamelistes spinosus

  Records:
  Hamamelis japonica, DC
  Betula nigra, UT, WI

  Hosts: Hamamelis virginiana; Betula alleghaniensis, lenta, nigra

  Range: Eastern North America


description:
  Hamamelistes spinosus

  Records:
  Hamamelis japonica, DC
  Betula nigra, UT, WI

  Hosts: Hamamelis virginiana; Betula alleghaniensis, lenta, nigra

  Location: Flower bud

  Range: Eastern North America


description:
  Hamamelistes sp. (undescribed)

  Reports:
  Betula occidentalis, ID
  Betula sp., NH

  North and west of the range of H. virginiana, a parthenogenetic Hamamelistes sp. has been collected on birch (Betula alleghaniensis in northern New Hampshire and Betula occidentalis in northern Idaho) (von Dohlen, 1991; unpublished data).

  Hosts: Betula spp.

  Range: Northern North America


description:
  Hormpahis cornu

  Reports: 
  Hamamelis virginiana, GA

  Hosts: Hamamelis virginiana

  Conical; leaf lamina

  Simple, no secondary host [sic; opposite of other von Dolhen paper and text of this paper]

  Range: Eastern North America


description:
  Hormaphis hamamelidis

  Reports:
  Hamamelis virginiana, CT

  Hosts: Hamamelis virginiana, Betula nigra [sic; opposite of other von Dohlen paper and text of this paper]

  Conical; leaf lamina


externallink: https://babel.hathitrust.org/cgi/pt?id=wu.89041294323&view=2up&seq=77&skin=2021&size=125
description:
  Eriophyes sp.

  Irregular, granulated, woody bud enlargements, diameter 1.5 to 2.5 inches, mite

  Eriophyes populi

  Irregularly globose, tomentose bud masses, diameter .25 to .5 inch on Balm-of-Gilead, mite

  ?Eriophyes populi

  Irregular masses of globose galls at base of leaf stem, mite


externallink: https://babel.hathitrust.org/cgi/pt?id=wu.89041294323&view=2up&seq=77&skin=2021&size=125
description:
  Mecas ornata

  Ovate twig swellings, cracking the second season, on large toothed and quaking aspens, beetle.


externallink: https://babel.hathitrust.org/cgi/pt?id=wu.89041294323&view=2up&seq=77&skin=2021&size=125
description:
  Cecidomyia irregularis Steb

  Irregular one-celled, blister galls, mostly on upper surface, light colored, on large-toothed poplar, midge


externallink: https://babel.hathitrust.org/cgi/pt?id=wu.89041294323&view=2up&seq=77&skin=2021&size=125
description:
  Pemphigus populitransversus

  Oval petiole galls with transverse openings and near middle of leaf stem, aphid, summer


externallink: https://babel.hathitrust.org/cgi/pt?id=wu.89041294323&view=2up&seq=77&skin=2021&size=175
description:
  Pemphigus populicaulis

  Twisted, subglobular galls with oblique openings at base of cottonwood leaves, diameter .25 to .5 inch, aphid


externallink: https://babel.hathitrust.org/cgi/pt?id=wu.89041294323&view=2up&seq=79&skin=2021&size=150
description:
  Morwilkoja vagabunda

  Irregularly folded, convolute deformed leaves, diameter 2 inches, aphid, summer


externallink: https://babel.hathitrust.org/cgi/pt?id=wu.89041294323&view=2up&seq=81&skin=2021&size=150
description:
  Pemphigus rileyi

  Globose, red and yellowish, clustered leaf galls, usually near the base and on underside, diameter 1/5 inch, on quaking aspen, aphid


externallink: https://babel.hathitrust.org/cgi/pt?id=wu.89041294323&view=2up&seq=81&skin=2021&size=150
description:
  Eriophyes sp

  Populus

  There are a number of leaf galls and leaf "piles" or erineums produced by unnamed species of mites


externallink: https://www.biodiversitylibrary.org/item/71437#page/12/mode/1up
description:
  Pemphigus populicaulis

  An irregularly spherical gall at union of blade and petiole. The opening is closed by a twisting of the leaf. Common on several poplars. The insects emerge when the gall dries somewhat and opens, usually about the middle of the summer.


externallink: https://www.biodiversitylibrary.org/item/71437#page/12/mode/1up
description:
  Pemphigus transversus

  Pemphigus populi-transversus

  This oval gall is formed midway of the petiole, on one side, with the opening opposite the petiole and transverse. Not common. On poplar, Populus.


externallink: https://www.biodiversitylibrary.org/item/71437#page/13/mode/1up
description:
  Pemphigus vagabundus

  Byrsocrypta vagabunda

  Populus deltoides

  A deformation of the terminal bud with the leaves so changed as to lose all appearance of leaves, each becoming bag-like, sonic like cock's combs. Yellow-green, becoming nearly black in winter. Persistent. Very common. On poplar, Populus deltoides.


description:
  Mecas inornata

  Saperda inornata

  Populus grandidentata, tremuloides

  Irregularly spherical swelling of the twig, 1-3 cm. in diameter. At first smooth, pithy, in second season becomes cracked without and hollow within. Beetle bores its way out in May. On Populus grandidentata and P. tremuloides. 


externallink: https://www.biodiversitylibrary.org/item/71437#page/13/mode/1up
description:
  Cecidomyia? irregularis, n. s.

  Irregular, monothalamous, blister-galls, between veins. Usually more than one on a leaf. Projecting above mainly, presence indicated below by a light-colored, roughened space. On large-toothed aspen, Populus grandidentata. Not rare.


externallink: https://www.biodiversitylibrary.org/item/71437#page/13/mode/1up
description:
  Pemphigus? rileyi n. s.

  A spherical, red and yellow gall occurring in clusters, usually on the under side of the leaf, near the petiole. Above there is a conical projection at the top of which is the opening. Thin-walled. About 5 mm. in diameter. On aspen, Populus tremuloides. Rather common.


externallink: https://www.biodiversitylibrary.org/item/71437#page/14/mode/1up
description:
  Ectoedemia populella

  This subglobular, one-celled gall is found on the petiole at its union with the blade of the leaf. There are distinct longitudinal ridges. Color like that of the petiole. About 5-7 mm. in diameter. The cell-walls are 1-2 mm. thick. Very abundant on American aspen, Populus tremuloides. In October the larvae go down to pupate in the ground. The moth emerges in May.


description:
  Diplodia tumefaciens
  [Page 58]

  These large, dark brown to black eruptions of outer bark tissue can be 35 cm in diameter and occur one after the other in a vertical row up the trunk.


description:
  Taphrina populisalicis
  [Page 58]

  Hosts: Populus balsamifera subsp trichocarpa, fremontii, Salix laevigata

  Convex, light green bumps correspond to yellow depressions on the opposite side of the leaf.

  Range: worldwide


description:
  Eriophyes parapopuli 
  [Page 59]

  Hosts: Populus tremuloides, balsamifera subsp trichocarpa, fremontii

  The galls usually form on axillary and terminal buds and often involve large sections of stems.

  Eriophyes sp
  Cauliflower gall mite
  [Page 59]

  Hosts: Populus tremuloides, balsamifera subsp trichocarpa

  When fresh in late spring or early summer, galls can be bright red and appear with numerous eruptions and fissures across the surface, but they turn dark brown with age.


description:
  Phyllocoptes didelphis
  [Page 60]

  Hosts: Populus tremuloides

  Some galls have fresh hairs that are hardened lobes and not the typical hairs found in other erineum galls.


description:
  Mordwilkoja vagabunda [Russo's photo seems to depict [Aceria dispar](https://www.gallformers.org/gall/4004), not Mordwilkoja. No observations of M vagabunda have been uploaded to iNaturalist west of Wyoming.]
  [Page 60]

  These convoluted, bladderlike leaf galls occur on nearly all cottonwoods.

  Range: throughout much of the US and especially from the Rocky Mountains to the West Coast.


description:
  Pemphigus populitransversus
  [Page 61]

  Hosts: Populus fremontii, balsamifera subsp trichocarpa, angustifolia, tremuloides

  Rapid growth of the petiole often causes a bend at the site of the gall.


description:
  Pemphigus populicaulis
  [Page 61]

  Hosts: Populus tremuloides, balsamifera subsp trichocarpa, several cottonwoods

  This gall, unlike [the preceding species](https://www.gallformers.org/gall/3460), has a spiral slit along which a series of small holes develop later in the season, through which aphids squeeze out.


description:
  Pemphigus populiramorum
  [Page 62]

  Hosts: Populus fremontii, balsamifera subsp trichocarpa

  The vertical slit, which distinguishes this species' galls, sometimes has slightly protruding lips.


description:
  Pemphigus populivenae [Russo seems to have completely mistaken [Thecabius populimonilis](https://www.gallformers.org/gall/4009) for P populivenae]
  [Page 62]

  Hosts: cottonwoods

  Often, the galls are located in rows along the midrib or along the edges of leaves. 


description:
  Pemphigus bursarius
  [Page 63]

  Hosts: Populus nigra var italica

  This aphid induces tubular, flask-shaped, mid-petiole galls on Lombardy poplar.

  Range: Europe, widespread across North America


description:
  Pemphigus spirothecae
  [Page 63]

  Host: Lombardy poplar, Populus nigra var italica; Populus balsamifera subsp trichocarpa [tristis]

  Feeding activity of the stem mother causes the petiole to twist, forming an ever-tightening spiral.


description:
  Pemphigus sp. B
  Midrib-gall aphid
  [Page 64; plates 43 & 44]

  Hosts: Populus fremontii

  This aphid induces large, globular galls that protrude from the ventral midrib of Fremont cottonwood leaves.


description:
  Ectoedemia populella
  [Page 65]

  Hosts: Populus tremuloides

  Galls are brown, with furrows or shallow cracks over the surface.


description:
  Saperda populnea
  [Page 65]

  Hosts: cottonwoods and willows

  Galls of this beetle are recognized by the hemispherical cuts (scars) made by adult beetles into which eggs are inserted between the phloem and xylem tissues. 


description:
  Hexomyza schineri
  [Page 66]

  Hosts: Cottonwoods, poplars, and especially aspen 

  Young branches can support dozens of these galls, sometimes in chains with one gall after another.


description:
  Contarinia sp.
  Stem-gall midge
  [Page 67]

  Hosts: Populus tremuloides

  This gall midge induces integral polythalamous stem galls that are either straight or bent and twisted on spring terminal shoots of quaking aspen.


description:
  Prodiplosis morrisi
  [page 67]

  This gall midge induces rose-pink to yellow-green roll galls along the leaf margins of aspens.

  Range: from the East Coast to Kansas and Texas. central Sierra Nevada in California


description:
  Harmandia sp. B
  Bead-gall midge
  [Page 68]

  The bead galls of this midge on aspen leaves are easily distinguished from galls of Harmandia sp C by having openings at the top of each gall, even if they are temporarily plugged with fibrous strands.


description:
  Harmandia sp C
  Bead-gall midge
  [Page 68]

  Hosts: Populus tremuloides

  These galls protrude on either side of leaves and show a hairy slit on the ventral side.

  Range: WA


description:
  Unknown
  Petiole-gall midge
  [Page 69]

  Hosts: Populus tremuloides

  Immature galls can be recognized by this abnormal flattening of the petiole and an apical bump that eventually becomes the lip or exit hole.

  Range: WA


externallink: https://www.biodiversitylibrary.org/item/54766#page/358/mode/1up
description:
  Gall. "These galls are corky formations about half an inch in diameter and varying from almost round to long galls, some being an inch or more in length. . . . The general position of the larval cell is, as a rule, in a plane almost perpendicular to the long axis of the gall which is the same as the course of the woody portion of the root. They are arranged in an irregular order close to the wood of the root."


description:
  Aceria dispar

  Phytoptus dispar

  Hosts: Populus tremula, nigra, tomentosa, tremuloides?

  Relationship: Leaf rolling, deformed foliage at end of twigs; leaf rolling

  Range: Europe; China; OR, WV 


description:
  Aceria neoessigi

  Eriophyes neoessigi

  Hosts: Populus fremontii, deltoides subsp wislizeni, tremuloides

  Relationship: The mites form large pendant male catkin galls 20 cm or longer. The catkins remain on the tree until early summer, unlike normal catkins which have dried and dropped.

  Range: CA, NV, UT, WY


description:
  Aceria parapopuli

  Eriophyes parapopuli

  Hosts: Populus fremontii, nigra, tremuloides, angustifolia

  Relationship: The mites cause irregular woody proliferation around buds, eventually hindering growth. In several years, the galls may be 1 inch or more in diameter.

  Range: British Columbia, CA, CO, MT, OR, ND, NV, SD, UT, WY

  Notes: This appears to be a complex of "cryptic" species (sibling species), adapted to each species and selected hybrids of cottonwoods.

  Aceria populi [a distinct European species that may be present in the US but more likely represented by misIDs of parapopuli-complex mites]

  Hosts: Populus tremula, italica, nigra, simonii

  Relationship: Cauliflower Gall on Quaking Aspen; Bud galls on White Poplar; 

  Range: Ontario (probably an error); CO, ID, NY (error)

  Notes: See records by Banks and Chadwick; but their records are probably in error. The first host listed was Populus tremula in 1889; that description was adequate for the code.

  The record in NY was "Lebanon Springs, July 11, 1895, W. H. Harrison."


description:
  Phyllocoptes didelphis

  Hosts: Populus tremuloides, grandidentata

  Relationship: The mites form open, cream-white, lower leaf surface erineum pockets that bulge above.

  Range: CA, MI, SC, SD, WV


description:
  Dasineura sp.

  Enlarged winter bud

  Host: Populus trichocarpa [tristis]

  Infested buds are conspicuous in summer. The red larvae are immersed in a sticky exudate between the bud scales. Larvae form cocoons in fall, diapause as larvae, and pupate in spring. Infested buds burst, but the leaves do not develop.

  Range: WA and OR


description:
  Harmandia stebbinsae

  Host: Populus grandidentata

  Harmandia galls are monothalamous and yellow, green, or reddish. In late spring or summer full-grown larvae drop to the ground to overwinter. Various other cecidomyiids have also been reared from these galls. These include a Clinodiplosis sp., Mycodiplosis thoracica, and Rhizomyia absobrina, all presumably mycophages.

  The gall is a blister, convex and reddish on the upper surface of the leaf, flat and light green below, occurring between or on veins.

  Range: MA and MD


description:
  Prodiplosis morrisi

  Hosts: Populus deltoides, tremuloides, hybrid poplars

  Rolled leaf or curled leaf margin

  Eggs are laid in newly developing leaf buds. Larvae are gregarious within the leaf rolls of marginal curls. Full-grown larvae drop to the ground to pupate. Up to five generations occur per year in Ontario. Infested leaves become deformed and later turn brown. Severe damage can stunt terminal leaders. 

  Range: Eastern North American to Kansas and Texas


description:
  Harmandia helena

  Host: Populus tremuloides

  Harmandia galls are monothalamous and yellow, green, or reddish. In late spring or summer full-grown larvae drop to the ground to overwinter. Various other cecidomyiids have also been reared from these galls. These include a Clinodiplosis sp., Mycodiplosis thoracica, and Rhizomyia absobrina, all presumably mycophages.

  This species was taken from a globular gall reportedly on the underside of the leaf. 

  Range: MA


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_S.htm#Stegophylla
description:
  Stegophylla quercina Quednau  (= quercicola of Blackman & Eastop 1994)  

  Apterae are pale yellowish-green with brown head, covered with bluish-white wax wool; BL 1.6-1.9 mm. On leaves of Quercus spp., usually of the red oak group, in dense flocculent masses which may cause curling and yellowing (Quednau, 1966). Oviparae and apterous males occur in October (the males are true apterae without a sclerotised thorax). In eastern North America. Quednau (2010) distinguished two subspecies, with S. quercina s.str. known from Quebec and New York (BMNH collection, leg. R.L. Blackman), and S. quercina acutirostris Quednau, known from Wisconsin and North Carolina. The latter subspecies makes pseudogalls on the infested Quercus spp. (Quednau 2010). 2n > 30 (for quercina s. str.).


externallink: https://www.biodiversitylibrary.org/item/34032#page/295/mode/1up
description:
  Dicrodiplosis helena, new species

  Gall. — Length 3 mm., subglobular, whitish, on the under surface of the leaf, opening on the upper surface.

  This species was reared April 5, 1912, from subglobular galls on the under side of aspen leaves, Populus tremuloides, collected by Miss Cora H. Clarke at Magnolia, Mass., the preceding August. It is easily separated from the allied D. californica by the shorter distal enlargement and the decidedly longer distal stem of the fifth antennal segment. It is also quite different from D. populi reared earlier from a similar gall. We are not certain that this is the maker of the gall.


externallink: https://www.biodiversitylibrary.org/item/54938#page/826/mode/1up
description:
  Harmandiola stebbinsae NEW NAME

  A North American species, Cecidomyia irregularis Stebbins (1910), is preoccupied by C. irregularis Bremi and is here renamed Cecidoinyia stebbinsae Gagne. NEW NAME.


description:
  Thecabius populi-monilis

  Host: Populus trichocarpha [tristis]

  This is a common gall on black cottonwoods. In spring a single black aphid stem mother produces a red pocket, 5 to 15 mm long and 2 mm wide, that bulges up from the upper leaf surface. The gall lies between the midvein and leaf edge and runs parallel to the midvein. The underside of the leaf shows only the gall's slit opening. Usually only a single gall occurs on a leaf. The galls have a single chamber and, depending on the time of spring, contain one to several aphids. 

  As the stem mother produces young in midsummer, they move out of the spring galls and travel to the new flush of summer leaves, where they produce many of their own summer galls (photo 25). The summer galls look very much like the spring galls, except that they are usually wider (5 to 7 mm) and are arranged end-to-end for the length of the leaf. They look like sausage links and have a single chamber each. Several rows of the summer galls occur on a single leaf, with the result that the leaf is often curled. The summer foliage is often very heavily galled by these aphids.

  The galls are fairly simple in construction and design. The wall of both spring and summer galls contains no layer of thick-walled cells and no food cells line the aphid chamber. Unlike most free-living aphids that feed on phloem sap, these and other gall-forming aphids feed for part of their lives on the contents of non-vascular cells. Unlike other species of gall-forming aphids on cottonwood, T. populi-monilis probably completes its entire life cycle on black cottonwood. 


description:
  Cecidomyiid. Undescribed.
  Big Bud

  Hosts: Populus trichocarpa [tristis]

  These galls (photo 27) are most easily spotted during the fall and winter when the leaves are off the trees. The gall is an enlarged terminal or side bud and will stay attached to the plant for some time, even after the gall has died and turned brown and brittle. In fact, a dead big bud from the previous year can frequently be found on the same twig with a current year's big bud. 

  Attacked buds begin to swell in early May and are often covered with a yellow sticky resin. The bud is enlarged because the bud scales and stipules are thicker than normal. Nestled between the thick scales, you should be able to find several very small orange larvae that are often bathed in yellow plant resin. These are the midge larvae. They use the spaces between scales as their larval chambers. 

  In August, the fully formed galled buds are green and about 2.5 cm long and 1.5 cm wide. All bud parts are enlarged. The outer scales are green, and the inner ones are yellow to cream-colored. By mid-October, the 5 to 15 larvae in each bud are white (because of stored fat) and have a prominent sternal spatula. Most larvae occur near the center of the big bud and are often awash in the sweet-smelling resin. The bud tissue near a larva is sometimes brown. The larvae probably feed by scraping the cells on the surface of the bud parts. Feeding may cause the browning. 


description:
  Eriophyes parapopuli Keifer

  A fairly common gall on black cottonwood caused by eriophyoid mites, Eriophyes parapopuli Keifer, occurs at the base of buds along the stem. When branches are bare in the autumn, the galls are woody, brown, cauliflower-like, and rarely more than 2.5cm in diameter. Several occur along a single stem. The mites live in the cracks and fissures of the gall.


description:
  Saperda populnea

  Long-horned beetles, (Saperda populnea L. or a closely related species) cause spherical or spindle-shaped stem galls (2 cm in diameter) on black cottonwood and on willows. Infestations are often heavy; the shape of the tree may be somewhat disfigured as a result. Adult females lay eggs in late spring in holes that they chew in the bark. When the egg hatches, the larva mines the cambium and then turns into the wood where it excavates a gall chamber. Growth from the wounded cambium causes the swelling. One or two adults emerge from each gall after one to two years of larval development. The adults feed on poplar leaves.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_T.htm#Thecabius
description:
  Thecabius affinis (Kaltenbach) 

  Galls occur on Populus spp., in May-June, mainly on P. nigra and its varieties, and are formed by folding of young leaves in half along mid-ribs; the outer, upper side of the leaf becomes blistered and yellowish or reddish (see [influentialpoints.com/Gallery](https://influentialpoints.com/Gallery/Thecabius_affinis_Poplar-buttercup_gall_aphid.htm)). These are secondary galls containing only the alatiform progeny of the fundatrix, which inhabits a small gall of its own formed by folding the edge of a leaf (fig. 131I). 

  T. affinis occurs on a wide range of Populus spp. throughout Europe and across Asia to east Siberia, Japan and Korea, and also in North America (see below). 

  COI barcode sequences have confirmed that T. affinis is present in North America (Skvarla et al. 2017). DNA data have also confirmed the existence of a separate North American species, Thecabius populiconduplifolius (Cowen), which has 2n=28 (Harper & Macdonald 1966). T. populiconduplifolius is virtually indistinguishable from T. affinis and has an identical biology, but with native poplars as primary hosts (Palmer 1952).


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_T.htm#Thecabius
description:
  Thecabius (Parathecabius) gravicornis (Patch)

  Galls are formed by folding leaves downward along the mid-rib to form a yellowish-green, swollen pocket which becomes filled with the developing emigrant alatae. The fundatrix inhabits a separate small pocket on the upper side of a leaf lamina (Palmer 1952). On Populus balsamifera in North America, widely distributed but apparently rather rare; also collected on P. angustifolia in Utah (BMNH collection, leg. R. Danielsson). Alatae (BL 1.7-2.8 mm, with olive-yellow abdomen) migrate in July-August, presumably to an unknown secondary host.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_T.htm#Thecabius
description:
  Thecabius populimonilis (Riley)   

  The galls on Populus spp. (angustifolia, balsamifera, fremontii, trichocarpa) are rows of ovoid, bead-like cells, pale green or with a reddish tinge, developed from upper side of leaf lamina on both sides of mid-rib (see [influentialpoints.com](https://influentialpoints.com/Gallery/Thecabius_populimonilis_bead-like_cottonwood_gall_aphid.htm)). Each gall in spring contains one developing aptera or alata. The fundatrices inhabit similar, solitary galls. This species is apparently facultatively monoecious; Gillette (1913) suggested that the large alatae (BL 2.7-3.0mm) produced in individual galls in June-July might recolonize cottonwoods, but it seems more likely that they are emigrants and found colonies of apterous exules in wax on Salix roots such as those observed by Lange (1943) in California. Presumably alatae produced in these colonies in October are sexuparae and return to Populus. However, alate sexuparae (BL 1.7-2.0mm) are also produced in galls on Populus, up to 12 per gall, in August-October (Maxson & Knowlton 1929, Palmer 1952, Harper 1959a). A more recent study by Aoki et al. (1996b) has confirmed most of these observations; they found heteroecious and monoecious life cycles at different locations, but their data on sex ratios indicate that some gene flow is probably occurring, rather than that two separate species are involved. Pike et al. (2012) have also now confirmed by DNA barcoding that colonies on willow roots (probably Salix dodgeana) in Washington are T. populimonilis. This species is widely distributed in USA, western Canada and Mexico. (Pemphigus saliciradicis may be a synonym; see note under that species.)


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_T.htm#Thecabius
description:
  Thecabius (Parathecabius) lysimachiae Borner  

  Galls containing developing emigrant alatae are formed by folding and convoluting the leaf of Populus nigra (fig. 131J). The fundatrix inhabits a separate oval pocket-gall on the upper side of the leaf lamina, as in other Parathecabius. Alatae (BL 2.3-2.6 mm, with dark, dirty greenish abdomen) migrate in late June-July to roots and runners of Lysimachia spp., especially L. nummularia. Other species, e.g. Lysimachia (= Glaux) maritima, may also be colonised. Apterae on Lysimachia are brownish or greyish green, with wax; BL 1.1-1.8 mm. Alate sexuparae return to poplars in September, but anholocyclic overwintering on Lysimachia is also common. In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007) and central Asia, and recently (2000) recorded from New York State, USA (Foottit et al. 2006). T. luppovae (Narzikulov), described from Populus densa (= P. talassica) in Tajikistan, may be a synonym. 2n=18.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pachypappa
description:
  Pachypappa pseudobyrsa (Walsh)  

  The fundatrix is wax-dusted, yellowish olive green with a blackish head; BL 4.0-4.7 mm.  Its feeding causes a small blister-like swelling in the centre of the underside of the leaf near the mid-rib (see [influentialpoints.com/Gallery](https://influentialpoints.com/Gallery/Pachypappa_pseudobyrsa_poplar_leaf-purse_aphid.htm)).  Its numerous progeny spread out in May-July along the main veins, developing into dusky-dark alatae, BL c. 3.5 mm.  On Populus spp. (deltoides, fremontii, tremuloides) in eastern North America, where it is widely distributed, and also in Colorado (Palmer 1952, as Asiphum).  The life cycle is unknown; the alatae presumably migrate to an unknown secondary host, although Smith (1974a) thought that it might be monoecious.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pachypappa
description:
  Pachypappa rosettei (Maxson)  

  Feeding by the fundatrix (BL 2.5-2.8 mm, appearance in life unknown) causes bending of leaves and stunting of stems to form a rosette-like nest, as in the closely-related European species P. tremulae.  On Populus tremuloides in Colorado and Utah, USA, and secondary host populations are now recorded from Washington and Alaska (Pike et al. 2012). Alatae (BL 2.2-2.5 mm) are produced in July.  Host alternation to roots of Picea spp. has now been confirmed by DNA bar-coding, and the apterous exule (BL 0.8-1.4 mm) redescribed (Pike et al. 2012). Aphids on roots of Pseudotsuga menziesii have also been tentatively assigned to P. rosettei (Danielsson 1990a). Anholocyclic overwintering may occur on spruce roots (Pike et al. 2012). Aphids infesting the roots of potted conifers in Canada and identified as P. tremulae may also be this species.

  [records of Pachypappa tremulae, a European species are likely attributable to P rosettei:] The numerous records from North America on roots of Picea spp. and Pseudotsuga menziesii (also as Asiphum tremulae and Rhizomaria piceae; Lavallee 1987) may all be due to confusion with P. rosettei (Footit et al. 2006).


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pachypappa
description:
  Pachypappa sacculi (Gillette)  

  The fundatrix is wax-dusted, slate grey with brown head; BL 4.5-5.0 mm.  It forms a large bag-like gall on P. tremuloides, with edges and tip of leaf curled downward (Harper 1966; as Asiphum).  In western USA (Arizona, Colorado, Utah, Washingtom, Alaska) and across Canada (British Columbia, Alberta, Quebec, Ontario and Nova Scotia).  Progeny of the fundatrix are all alate, with dark olive-green abdomen, BL 3-4 mm.  Host alternation to Picea spp. has now been confirmed by DNA bar-coding, and a redescription provided of the apterous exule (BL 0.9-1.8 mm) (Pike et al. 2012). Anholocyclic overwintering may occur on spruce roots (Pike et al. 2012). 2n=10.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus betae Doane (= balsamiferae Williams)  

  Galls are yellowish green sometimes tinged with red, pocket-like, formed usually on the underside of the poplar leaf near its base, on or adjacent to the mid-rib, with little or no deformation of the leaf (Harper 1959a, Whitham 1978).  The opening can be either on the upper side of the leaf or underneath along the mid-rib.  Primary hosts are Populus angustifolia, P. balsamifera, or occasionally P. deltoides ssp. occidentalis or P. trichocarpa.  Alatae (BL 1.5-2.1 mm) leave galls in late June-early August (Harper 1959b), to colonise roots of Beta vulgaris, and also perhaps on other Amaranthaceae (formerly Chenopodiaceae) such as Chenopodium album, and possibly Spinacia.  Apterae on chenopods are yellowish aphids secreting white wax, with BL 1.9-2.4 mm. Anholocyclic populations can persist all year on secondary hosts (see Moran & Whitham 1988), and are widely distributed in North America. The life cycle was studied by Harper (1963) in southern Alberta, and by Moran & Whitham (1988) in Utah.  This aphid's evolutionary ecology has been studied in some detail, including the territorial behaviour of first instar fundatrices in selection of galling sites (Whitham 1979), population dynamics in relation to galling site (Whitham 1978, 1980), genetics of resistance in the host plant (Paige & Capman 1993), induction of sexuparae (Moran et al. 1993a), and discrimination between host trees by returning sexuparae in autumn (Moran & Whitham 1990).  However, interpretation of all this work is problematic as Foottit et al. (2010a) have demonstrated that there is a complex of Pemphigus species in western North America  sharing the same cottonwood primary hosts. Taxonomic affinities with palaearctic populations of Pemphigus on chenopod roots (see P. fuscicornis) are unclear. See also Blackman & Eastop (2000).


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus bursarius (Linnaeus)  Poplar‑Lettuce Aphid,  Lettuce Root Aphid     

  Galls on Populus spp. (most commonly P. nigra) are yellowish or reddish when mature, purse-shaped, always formed on leaf petioles (fig. 131L, and see [influentialpoints.com/Gallery](https://influentialpoints.com/Gallery/Pemphigus_bursarius_Poplar-lettuce_gall_aphid.htm)).  The fundatrix is greyish green, slightly wax-dusted.  Alatae (BL 1.6-2.5 mm) leave the gall through a lateral opening over an extended period from late May to September, with peak emergence in late June-July (northern hemisphere). They found colonies mainly on roots of Compositae/Asteraceae (e.g. Cichorium, Lactuca, Lampsana, Sonchus, Taraxacum, Tussilago). Many of the records of P. bursarius from roots of plants in other families may be referable to other Pemphigus spp. Return migration of sexuparae to poplar is in October-September, but anholocyclic overwintering on roots of secondary hosts is common. P. bursarius occurs in Europe, across Asia to eastern Siberia (Babichev & Kirichenko 2020), northern and southern Africa, North and South America, and (perhaps) Australia and New Zealand. There is some uncertainty about its occurrence in Australia and New Zealand because, although  bursarius-like aphids occur in bursarius-type galls on Populus and are also recorded from Sonchus roots, the secondary host populations are mainly recorded from Amaranthaceae and Polygonaceae. Miller et al.. (2005) found genetically distinct populations on different secondary host plants, indicating a complex of races and/or subspecies. Dunn (1960a) studied gall formation and (1960b) reviewed natural enemies.  Alleyne & Morrison (1978a, b) gave accounts of its biology and natural enemies in Quebec. See also Blackman & Eastop (2000).  2n=20.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus knowltoni Stroyan   

  Galls are green, pouch-like, of variable shape, on undersides of leaves of Populus angustifolia alongside the mid-rib at the base of the lamina (Stroyan 1970).  Fundatrix and immatures in gall are blackish green.  Alatae (BL 1.8-2.1 mm) are mature in the gall in July, and migrate to an unknown secondary host.  Described from Utah, USA. 


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus longicornus Maxson   

  Galls are elongate, thin-walled, on upper sides of leaves of Populus deltoides.  The fundatrix is pale yellowish white.  Alatae (BL 1.7-2.1 mm) emerge in May-June and migrate to an unknown secondary host (Maxson 1923).  In eastern and southern USA.  [According to Smith (1985), there is also an undescribed Pemphigus sp. on P. deltoides in North Carolina that causes elongate to slightly globular galls on the upper sides of the leaves, and has alatae with antennae intermediate in length and antennal sensoriation between longicornus and populivenae.]


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus monophagus Maxson  

  The gall is globular, pale greenish yellow, consisting of a swelling of upper side of base of leaf lamina with petiole broadened and somewhat twisted, and a slit-shaped opening underneath between petiole and leaf (fig. 131N, and Maxson 1934).  Fundatrix honey-yellow to pale olive-green, producing a generation of creamy white apterae in the gall.  Monoecious, alatae (BL 1.4-1.9 mm) emerging from galls in August-September to deposit sexuales on the bark of the trunk.  On Populus angustifolia, P. balsamifera and P.trichocarpa in western North America.  Aoki & Kurosu (1988) discussed the evolution of monoecy in this species.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus nortonii Maxson  

  Galls on Populus deltoides especially var. occidentalis (= monilifera) are pale yellowish green, formed by a swelling at the base of the leaf lamina, twisted on the flattened, broadened petiole; resembling those of P. populicaulis (Palmer 1952), but with a long slit-like opening (Harper 1959a).  Alatae emerge over an extended period (June-Sept; Harper 1959b, Grigorick & Lange 1968), migrating to an unknown secondary host.  Apparently widely distributed in USA, and in Alberta, Canada.  Apart from the greater number of rhinaria on ANT V and VI this species closely resembles P. junctisensoriatus.  Populations from Alberta (Harper 1959a, b) seem to be intermediate in this respect, and the distinction between these two species needs to be verified.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus obesinymphae Aoki & Moran  

  Galls are formed on the petioles of late flushing leaves of Populus fremontii (Aoki & Moran 1994), and P. deltoides (Abbot 2009), usually close to the leaf blade. When mature they are almost spherical, 8-16 mm in diameter, with the lips of the transverse slit sometimes bulging outwards, and a circular ostiole about 1.2 mm in diameter somewhere along the slit. First instar progeny of the fundatrix in the gall are heavily sclerotised with well-developed fore legs and attack predators (Moran 1993). Abbott (2009) demonstrated using microsatellite DNA analysis that there was a high level of clonal admixture within galls, and Grogan et al. (2010) investigated its consequences. Alatae (BL 1.9-2.5 mm) fly in August-October, presumably migrating like P. populitransversus to cruciferous hosts (N. Chen et al. 2009), on which overwintering occurs. Sexuparae return to Populus in May of the following year, and the eggs hatch only a short time after oviposition. Apparently widely distributed in USA (Ohio, Tennessee, Arizona, Utah, Texas), and probably in northern Mexico. N. Chen et al. (2009) recorded molecular and phenological differences between P. obesinymphae and P. populitransversus on cabbage in Texas. This species was confused for many years with P. populitransversus (q.v.), and its distribution and range of both primary and secondary hosts requires further confirmation.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus populicaulis Fitch   

  The gall is yellowish green to creamy, formed by swelling of base of leaf lamina, twisted around broadened petiole, rather like that of P. nortonii and P. populiglobuli but with a small, rounded exit hole on the underside (Palmer 1952, Harper 1959a).  Especially associated with P. deltoides and P. deltoides ssp. occidentalis (= monilifera), but also recorded from other cottonwoods and P. tremuloides.  The fundatrix is pale yellowish to whitish green.  Alatae (BL 1.7-2.0 mm) emerge in July-September.  The secondary host has not definitely been established; Glendenning (1924) reported that it was Oenanthe sarmentosa, but did not provide evidence for this assertion.  Hottes & Frison (1931) suggested that P. brevicornis Hart, described from corn roots, was the sexupara of populicaulis.  Widely distributed in North America.  Setzer (1985) studied gall mortality.  Sokal et al. (1980) and Bird et al. (1982) studied geographical variation in eastern North America using multivariate techniques.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus populiglobuli Fitch   

  The gall is green, often tinged with red and/or yellow when mature; globular, conical or bullet-shaped, formed by swelling of base of leaf lamina and thickening of petiole, the exit hole being a slit on the underside between leaf and petiole (Maxson & Knowlton 1929).  Primary hosts are Populus angustifolia and P. balsamifera, more rarely P. trichocarpa.  The fundatrix is variously reported to be dark green (Palmer 1952) or pale yellowish white (Maxson & Knowlton 1934).  Alatae (BL 1.5-2.3 mm) leave gall in June-early July (Alberta; Harper 1959b) for an unknown secondary host.  Widely distributed in North America.  Differences from P. populicaulis and other closely-related species are discussed by Maxson & Knowlton (1929).


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus populitransversus Riley 

  The gall is formed by swelling and bending of petioles of the leaves of Populus deltoides (incl. ssp. occidentalis) and P. fremontii; it is green, globular to ovoid, with a usually transverse slit (see [influentialpoints.com/Gallery](https://influentialpoints.com/Gallery/Pemphigus_populitransversus_poplar_petiole_gall_aphid.htm)).  Sometimes recorded from other Populus spp. (acuminata, angustifolia, balsamifera, nigra, tremuloides, trichocarpa)  However, some of these records, particularly those from western cottonwoods, may apply to other species.  The fundatrix is whitish to pale dirty greenish yellow (Palmer 1952).  Alatae (BL 1.7-2.0 mm) emerge in July and found colonies of pale yellow aphids in flocculent wax on roots of Brassicaceae (Jones & Gillette 1918), where it can be a serious pest (Royer & Edelson 1991).  Sexuparae return to cottonwood in September-October to produce sexuales which lay overwintering eggs on the bark. Geographic variation in eastern USA has been studied extensively using multivariate methods (Sokal & Riska 1981).  Bingham & Sokal (1986) studied host tree effects on morphology.  P. populitransversus is recorded throughout most of North America east of the Rocky Mountains. However it is still unclear whether the name is being applied to a single species; apart from the confusion with P. obesinymphae (q.v.), Bird et al. (1979) found that two forms with different gall shape and phenology, and corresponding morphological differences in fundatrices and emigrant alatae, occur sympatrically on P. deltoides over a large area of the USA east of the continental divide. It should be noted that the morphology of their aphids from globular galls on late flushing leaves does not correspond with that described for P. obesinymphae.

  Pemphigus identified as P. populitransversus have been found on Populus sp. in Mexico (BMNH collection, leg. R. Peña) and Chile (BMNH collection, leg. G. Monsalve); it has also been introduced to South Africa (Müller & Scholl 1958), where it occurs on Populus nigra (BMNH collection, leg. E McC. Callan) and P. deltoides, and the Azores (on Populus sp.; BMNH collection, leg. A. van Harten).  Presumed anholocyclic populations also occur in New Zealand on Eutrema wasabi roots (BMNH collection, leg. L.M. Emms), and aphids collected on Arabis roots in England (BMNH collection, leg. G. Fox-Wilson) may also be this species.  2n=20.


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus populivenae Fitch   

  Galls are elongate pocket-shaped or cock's comb-shaped, greenish or yellowish or pale red, arising alongside mid-rib on upper side of leaf, with a slit-shaped opening underneath.  On heavily-infested trees, galls may occur on both sides of the leaf, and those on the underside may then be confused with those of P. betae (Maxson & Knowlton 1934; however see also Footit et al. 2010). Whitham (1978) reported that P. populivenae galling Populus fremontii in Utah caused chlorosis of the leaf distal to the gall. As in the case of P. betae, several species of cottonwoods and their hybrids are colonised (angustifolia, balsamifera, deltoides, fremontii and trichocarpa; the latter species in experimental conditions only).  The fundatrix is yellowish green.  Alatae (BL 1.1-1.8 mm) emerge in June-July and migrate to found colonies on roots of Amaranthaceae.  Rumex (Polygonaceae) may also be colonised. The evidence for this host alternation is work in California by Grigorick & Lange (1962), who transferred sexuparae from sugar beet to poplar bark under controlled conditions and obtained fundatrices that produced galls specifically on P. trichocarpa.  The alatae from these galls were identified as P. populivenae, and colonised roots of B. vulgaris and Chenopodium album.  However, the suggested synonymy of both P. betae and P. balsamiferae with populivenae indicates some confusion of identity, and further work seems advisable to confirm that the correct name for populations damaging sugar beet in California (e.g. Summers & Newton 1989) is in fact populivenae rather than betae.  Foottit et al. (2010) have found molecular evidence that at least three taxa with similar galls occur sympatrically on cottonwoods in Alberta. 


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus tartareus Hottes & Frison  

  (= Pemphigus junctisensoriatus Maxson)   

  Described originally from alate sexuparae collected on roots of Bidens sp. in Illinois, USA. Galls on Populus deltoides are pale yellowish green, composed mostly of swollen leaf petiole around broadened base of leaf lamina, with an exit hole in the form of a long slit between the petiole and the leaf (Maxson 1934).  The fundatrix is pale yellowish white, with brownish head and tibiae. Alatae (BL 1.4-1.8 mm) emerge in June-July and migrate to Bidens sp(p). Smith (1985) established the synonymy with P. junctisensoriatus and made successful host transfers from Populus to Dichondra repens (Convolvulaceae) as well as to Bidens.  In USA and eastern Canada.


description:
  The larvae of this species live in numbers between the unexpanded young leaves of the common white or Nettle-leafed Vervain, causing them to become deformed and twisted. Collected July 8th to 10th. The flies emerged from July 15th to 25th. The larvae transform in the ground.


externallink: https://www.biodiversitylibrary.org/item/34032#page/184/mode/1up
description:
  This fly was reared from cylindric enlargements of the flower stalks of Verbena prostrata collected by P. H. Timberlake, November 13, 1910, in the Puente Hills, Whittier, Cal. ... November 30 one gall was full of healthy larvae evidently hibernating in silken tubes attached to the walls of the gall.


description:
  The gall (PI. 2, fig. 9) consists of parts of the abort ed ovary and lower corolla-tube. In the case of those containing larvae the inside of the tube becomes quite pubescent; but this disappears with the formation of the pupa. The length is 5 mm., the width 2 mm. The upper part of the corolla retains the bud-like structure until the adult pushes through it. The galls appear as buds until the fruits mature.


externallink: https://www.biodiversitylibrary.org/item/89742#page/541/mode/1up
description:
  Galls are about 5 mm in diameter, rounded and smooth on the top of the leaf and warty on the bottom of the leaf. Galls are first green, then turn brown after the larvae are fully fed. The larva pupates in the gall.


description:
  Hairy, grayish, ovoid leaf gall

  Galls are about 3mm high, monothalamous, and dehiscent when the larvae are mature.

  Host: U dioica
  Distr: New York


externallink: https://www.biodiversitylibrary.org/item/22105#page/216/mode/1up
description:
  Galls on the upper side of the leaves of Urtica gracilis, either on the midrib, or, more often, on the lateral veins. Urn-shaped (I mean the shape produced by cutting off the
  smaller end of a slender pear) up to 3 m. m. high, subsessile (that is, connected by a very small surface with the leaf), pale green, semi-transparent, succulent gall, bearing a short style or nipple at the upper, truncate end. Inside, the larva of a Cccidomyia. Lake George, July, 1863 ; Trenton Falls, July, 1874 ; not uncommon, but not in large numbers. Each leaf bears one, sometimes two galls, seldom more.


description:
  Tapered stem swelling

  Swellings are 1-2cm long and turn woody in the fall.

  Host: P. pumila
  Distr: Maryland


externallink: http://www.aphidsonworldsplants.info/d_APHIDS_P.htm#Pemphigus
description:
  Pemphigus populiramulorum Riley   

  Galls are formed by swelling of new growth of twigs, or rarely at the bases of petioles; they are quite large (up to 2.5 cm diam.), rounded, green to brownish, with a transverse or vertical slit-shaped opening which often has protruding lips.  The primary host is usually Populus deltoides ssp. occidentalis (= monilifera), but it is also recorded from P. balsamifera and P. fremontii.  The fundatrix is pale creamy yellow (Palmer 1952).  Alatae (BL 2.2-2.5 mm) leave galls in July-September (Harper 1959b), flying to an unknown secondary host.  In western North America (Utah, Colorado, South Dakota).


description:
  Prodiplosis morrisi Gagne

  Hosts: Populus deltoides

  Biology. Larvae curl newly developing leaves of Populus deltoides and Populus tremuloides and can be an important pest in hybrid poplar nurseries in southern Canada and of cottonwood in the southern United States. The affected leaf parts turn brown. Lightly damaged ones dehisce. According to Morris (1981), five generations are possible in southern Ontario between June and September. Each generation affects 4-5 leaves. Long-run damage includes stunting of terminal leaders. Morris (1981) reported typical damage from Illinois, Ontario, and New Brunswick, south to Mississippi and west to Texas and Kansas. Larvae of P. morrisi hibernate in the duff; adults appear in Ontario around 1 June. Another Prodiplosis, a species near myricae (q.v.), has also been reared from leaf rolls of Populus sp. ("wild aspen") in southern Canada. 

  Range: Ontario, MS


description:
  A hitherto undescribed leaf spot of balsam poplar (Populus balsamifera L.) associated with a gall midge (Cecidomyiidae) was first observed in Waterton Lakes National Park, Alberta, in 1963. Subsequent collections indicated its wide distribution in Alberta. Only a minor portion of the foliage of a tree was affected and there was no indication of premature leaf fall. Observations and culture studies were undertaken because, although this spot resembled those caused by pathogenic fungi, it did not fit an existing description

  The first conspicuous symptom of the leaf spot was observed near the end of June as a tiny chlorotic papilla, approximately 1.5 mm in dia and 1 mm high, which protruded adaxially with an abaxial concavity. The concavity was partially filled with a clear sticky fluid within which lay a cecidomyiid larva. Papillae occurred singly or in groups. Three to 4 days later each papilla or group of papillae was surrounded by a chlorotic zone with a brown to black perimeter (Fig. I-B). On dried specimens the papilla was less chlorotic than the surrounding tissue within the spot, thus resulting in a green island (Fig. 3).

  [Several black and white photos of the gall and larva appear in the paper]


description:
  Observed from Ontario to Montana between June and August.

  This species is not listed explicitly in Gagne 1989 but a reference to the Whitney and Baranyay paper is included in the general "Harmandia sp" section; this is the grounds on which we have moved this out of the generic Cecidomyia and into Harmandiola. 

  This gall can only be distinguished from the similar [p-tristis-leaf-spot](https://www.gallformers.org/gall/4027) by host ID. There are places where both hosts occur, and distinguishing the two plants can be difficult. It is possible that the two midges are in fact conspecific. 


description:
  An exposed larva on the lower side of the leaf lamina of Populus tristis causing a circular green spot and a slight raised papilla on the upper side. Observed from southern CA up to Washington. There may be multiple generations per year; fresh galls with larvae have been seen in May and September. 

  This gall can only be distinguished from the similar [p-balsamifera-leaf-gall](https://www.gallformers.org/gall/3987) by host ID. There are places where both hosts occur, and distinguishing the two plants can be difficult. It is possible that the two midges are in fact conspecific. 


description:
  A green to pink-red pea gall on the midrib or major side vein of Populus tremuloides. The gall is nearly entirely found on the upper side of the leaf, and appears detachable, but its exit hole is found on the lower side at the contact point. Observed from Alaska to Nova Scotia south to New Mexico, fresh between June and August.

  This gall is nearly identical to that of [Harmandiola tremulae](https://bladmineerders.nl/parasites/animalia/arthropoda/insecta/diptera/nematocera/cecidomyiidae/cecidomyiinae/cecidomyiidi/cecidomyiidi-unplaced/harmandiola/harmandiola-tremulae/), found on Populus tremula in Europe. It is possible that the same species occurs holarctically on both hosts but no existing literature seems to confirm this. 

  This gall morphotype can be distinguished from other midge galls on Populus leaves by the following traits:

  - Larva concealed within galled tissue, protruding only on the upper side of the leaf
  - Thick walled, nearly spherical, large


description:
  The literature on Harmandiola helena is inconsistent. Felt's original description places the opening on the upper side, while Gagne 1989's drawing (though only listed as "possible helena") shows the opening on the lower side. Russo suggests that his Harmandia sp. A, which seems to match our [p-tremuloides-like-populnea](https://www.gallformers.org/gall/4030) and does open on the upper side, might be related to H helena, suggesting the name may belong to that morphotype. Our current interpretation follows Gagne. An additional complication is that in iNaturalist observations, galls closely resembling Gagne's drawing are largely observed on Populus grandidentata, a host not listed in any existing literature. It is added tentatively as a host based on those unconfirmed observations [(eg)](https://www.inaturalist.org/observations/93855751) or [(eg)](https://www.inaturalist.org/observations/54278543). 

  Regardless of its correct name, the gall pictured in this entry closely matches the description of the European species [Harmandiola pustulans](https://bladmineerders.nl/parasites/animalia/arthropoda/insecta/diptera/nematocera/cecidomyiidae/cecidomyiinae/cecidomyiidi/cecidomyiidi-unplaced/harmandiola/harmandiola-pustulans/) on Populus tremula. 

  The gall pictured in this entry can be distinguished from other midge gall morphotypes on Populus leaves by the following traits: 

  - Larva concealed within galled tissue
  - Gall evenly protruding from both sides
  - Opening in linear slits on the lower side of the leaf


description:
  A rough, hairless, light-green to pink-red gall on the lamina of Populus tremuloides, protroding on both sides, with a globular mass below and a puckered pair of protruding lips on either side of a slit-like opening above. It varies from nearly globular to flattened against the leaf. Either single or in scattered groups. Observed from Alaska to Quebec and south to Colorado between May and September.

  This gall resembles that of [Harmandiola cavernosa](https://bladmineerders.nl/parasites/animalia/arthropoda/insecta/diptera/nematocera/cecidomyiidae/cecidomyiinae/cecidomyiidi/cecidomyiidi-unplaced/harmandiola/harmandiola-cavernosa/) known in Europe from Populus alba and tremula. 

  This gall morphotype can be distinguished from other midge galls on Populus leaves by the following traits:

  - Larva concealed within galled tissue, protruding primarily on the lower side of the leaf 
  - Noticeably protruding lips on the upper side of the leaf surrounding a linear slit exit hole


description:
  This morphotype may be the best fit for the name [Harmandiola helena](https://www.gallformers.org/gall/4005); see ID Notes on that page for more info.

  Green to pink-red globular galls on Populus tremuloides showing equally on both sides of the leaf, often confluent where several grow nearby and sometimes sprawling onto adjacent veins, frequently found at the base of the leaf (but apparently never galling the petiole), succulent and thick-walled. They are unique among Populus leaf galls in having a circular rather than linear exit hole, which is typically but not always found on the upper side of the leaf. Circles are often visible before midges emerge, and these incipient holes are often seen on the opposite side of the one where midges eventually emerge as well. They are glossier and more succulent in texture than those of most Harmandiola galls. Variation in shape and density is greater for this morphotype than in many others, including some galls which present as a single large sphere rather than a flat cluster. Some of these variations may end up representing distinct species of midge inducers. 

  These galls resemble those Bladmineerders attributes to either [Lasioptera populnea](https://bladmineerders.nl/parasites/animalia/arthropoda/insecta/diptera/nematocera/cecidomyiidae/cecidomyiinae/lasiopteridi/lasiopterini/lasioptera/lasioptera-populnea/) or [Contarinia populi](https://bladmineerders.nl/parasites/animalia/arthropoda/insecta/diptera/nematocera/cecidomyiidae/cecidomyiinae/cecidomyiidi/cecidomyiini/contarinia/contarinia-populi/). It may not belong in the genus Harmandiola.

  This gall morphotype can be distinguished from other midge galls on Populus leaves by the following traits:

  - Larva concealed within galled tissue protruding equally on both sides of the leaf
  - Circular rather than linear exit hole on the upper side of the leaf, though occasionally rings suggesting incipient openings are seen on the lower side as well


description:
  Diplolepis nebulosa

  Mature galls are spherical, averaging 5.0–7.0 mm in diameter, spineless and on the abaxial leaf surface. They occur singly or in rows. They are most common on short R woodsii on dry exposed prairie. Immature galls are green to light brown with scaly, thick walls and a prominent depression distal to the point of attachment. Mature galls are pale brown and the depression is less prominent. Walls are thin and easily crushed between fingers. Larvae feed over the entire chamber surface. Adults removed from cold storage take an average of 26.3 days to emerge and are present in the field from mid-May to mid-June. Immature galls are found from early July to mid-August. Both immature and maturing galls appear together suggesting a lengthy period of gall initiation. Periclistus are found in 66.6% of the galls by mid-season and are major source of inducer mortality. 


description:
  The larvae normally feed in galls on the roots of goldenrod (*Solidago*), growing in damp or wet soils. Occasionally the root galls project above the surface of the ground. Sometimes galls are formed on the stems, especially when the roots of the host are covered with water, during oviposition. A patch of infested goldenrod will frequently yield many galls, often several on the roots of a single plant, but with only a single larva or puparium in each gall. The infestations are usually in the form of local colonies. There are many species of closely related goldenrods in Florida, and possibly more than a single species serves as host. One species was identified, through the courtesy of authorities at the United States National Museum, as *Solidago fistulosa*. Daecke recorded the host on Long Island, N.Y., as *S. juncea*. p.28

  [NOTE: *S. juncea* is for *E. comma*]


description:
  The larvae feed in root galls on goldenrod (*Solidago sempervirens*). From 1 to 4 galls were found on a single host plant. Those having galls were growing in soil subject to standing brackish water during spring and summer. These galls, which may attain a length of over 45 mm and a diameter greater than 25 mm, are normally just below the surface of the ground. Usually each gall contains only a single larvae or puparium, but very occasionally the galls are compound, containing two larvae or puparia separated by a thin partition. (p. 29, as *E. reticulata*)


description:
  Evidently, the gall consists of an aborted and transformed growing shoot. At maturity the cylindrical gall is about 2.0 x 0.5 cm (Fig. 43). External traces of leaves appear on the gall in the form or imbricated scales that are 1-3 cm long. (p. 214, as *E. reticulata*)


description:
  Diplolepis polita

  Galls of D. polita (fig. 3) are monothalamous, spherical, 3.5 to 5.0 millimeters in diameter, clothed with weak spines and formed on the adaxial surface of leaflets. Immature galls are yellowish-green, but mature galls are often bright red. They usually occur in clusters with several galls per leaflet. They are found only on Rosa acicularis Lindl. in forested regions of Alta. and Sask. growing in partly shaded habitats. Walls of mature galls are thin (about 1 millimeter) and easily crushed with fingers. The larvae feed over the entire chamber surface. Adults removed from cold storage take an average of 26.8 days to emerge (fig. 9) and are present in the field in early May with the first galls present by about May 20. Periclistus take an average of 23.4 days to emerge after cold storage. Periclistus are found in about 71 percent of the galls by mid-season (table 1) and are the major source of inducer mortality. Galls are mature and on the ground by mid-August. 


description:
  Diplolepis polita

  Diplolepis polita was first described from galls collected in the United States by Ashmead (1890). This species is now known to occur from the coast of British Columbia north to central Alaska and the Yukon, east to western Quebec, and from central Ontario north to James Bay. It is common throughout the Aspen Parkland Ecoregion from Alberta to Manitoba on roses in partially shaded areas at the edge of forests (Fig. 4). 

  Mature galls are spherical, averaging 3.5–4.5 mm in diameter, clothed with weak spines, and found on the adaxial (upper) surface of leaves (Fig. 16). They are single-chambered (Fig. 17) with thin walls and usually grow in clusters, several galls per leaflet. Galls are found both on the sucker shoot and on multi-year roses. Galls always abscise in late summer. Immature galls in shaded areas are usually greenish-yellow, whereas those growing in areas open to the sun are often bright red. All turn light brown when dry. 

  Galls of D. polita are found only on R. acicularis east of the Rockies, never on R. woodsii growing in open areas of the southern prairies. However, they are found on R. woodsii in more shaded areas in the Okanagan in British Columbia. A large population was found on R. arkansana near the top of the Cypress Hills in Alberta (Shorthouse 1991). 

  A large population of galls from within Prince Albert National Park in Saskatchewan illustrates the composition of a typical component community (Fig. 44A). Only 4% of the inhabitants were inducers by fall, 29% were Periclistus, and the rest (67%) were parasitoids. Periclistus (average of 5.4 per gall; Table 4) were commonly found in about 70% of all maturing galls and 12% of mature galls (Table 5). The most common parasitoids were Eurytoma sp. and Pteromalus sp. (Fig. 44A). About half of the Eurytoma sp. and half of the Torymus sp. matured and exited galls by the end of the season (Table 3).


description:
  The larvae feed in small round galls on the stems of goldenrod (Solidago sp.) (p. 27, as *E. nicholsoni*)


description:
  Numerous small pouch galls along the midrib or major lateral veins of Populus tremuloides. They are pale green to pink-red, hairless, thin-walled, sometimes spotted with white inclusions, and typically seem to lay flat against the leaf more than protruding downwards. When growing densely they become confluent. They are similar to [Harmandiola helena](https://www.gallformers.org/gall/4005), which also occurs in numbers along the midrib of P tremuloides in similar patterns, but differ in having the majority of the gall on the lower side of the leaf and a slightly protruding slit opening on the upper side of the leaf. Further, H helena galls are typically oriented more or less perpendicularly to the leaf, while the galls of this species seem to be oriented at an angle. Observed from Nova Scotia to Manitoba and south to Michigan from July to September. 

  This gall is similar to the galls induced by [Harmandiola populi](https://bladmineerders.nl/parasites/animalia/arthropoda/insecta/diptera/nematocera/cecidomyiidae/cecidomyiinae/cecidomyiidi/cecidomyiidi-unplaced/harmandiola/harmandiola-populi/) on Populus alba and tremula in Europe.

  This gall morphotype can be distinguished from other midge galls on Populus leaves by the following traits:

  - Larva concealed within galled tissue, protruding entirely on the lower side of the leaf
  - Only a slight protrusion on the upper side around a linear slit opening


description:
  A small, green to pink-red, hairless, globular to capsule-shaped gall on the upper side of leaves of Populus tremuloides. Only a small flat surface, not protruding at all, is visible on the lower side. Observed from Alaska to Prince Edward Island and south to New Mexico, fresh specimens reported between May and August.

  This gall closely resembles the gall induced by [Harmandiola globuli](https://bladmineerders.nl/parasites/animalia/arthropoda/insecta/diptera/nematocera/cecidomyiidae/cecidomyiinae/cecidomyiidi/cecidomyiidi-unplaced/harmandiola/harmandiola-globuli/), known from Populus alba and tremula in Europe.

  This gall morphotype can be distinguished from other midge galls on Populus leaves by the following traits:

  - Larva concealed within galled tissue, protruding only on the upper side of the leaf
  - Thin walled, capsule shaped, small


description:
  Harmandia sp. A [Russo suggests this might be related to the eastern [H helena](https://www.gallformers.org/gall/4005), see ID Notes on that page]
  Leaf-gall midge
  [Page 68]

  Hosts: Populus tremuloides

  Galls occur along the midrib and occasionally lateral veins of leaves in early summer.


description:
  Schismatodiplosis lantanae (Rubsaamen)

  S. lantanae induced uni-chambered pouch galls inhabited by one larvae or pupa. This gall consisted predominantly of hypertrophied spongy parenchyma.

  Schismatodiplosis lantanae induced spherical, green pocket galls, mostly prominent on the abaxial leaf side (Fig. 15), 7.4 ± 0.3 mm in diameter and with a regular, hairy surface. The unique larval chamber bore one larval inducer or its pupa (Fig. 16) and was bound by a nutritive tissue. This tissue was differentiated from the hyperplasia of the adaxial epidermis (Fig. 17). From the inside of the chamber, the ostiole channel was observed facing the adaxial surface (Fig. 18). 

  Schismatodiplosis lantanae galls had oval-shaped ordinary cells, and trichomes similar to those of the healthy leaves on the adaxial epidermal layer (Fig. 19). Stomata were absent. On the abaxial epidermal layer, the ordinary cells presented sinuous anticlinal walls and stomata and trichomes similar to those observed in healthy leaves (Fig. 20). 

  The nutritive tissue was one-layered, with thin-walled and oval-shaped cells (Fig. 21). Some necrotic areas indicated cicatrisation spots (Fig. 22). 

  Gall parenchyma could be divided into the following three zones: (1) hypertrophied cells with conspicuous nuclei surrounding the nutritive tissue (Figs 21 and 22), (2) hypertrophied spongy parenchyma cells with few chloroplasts and conspicuous intercellular spaces occupying the majority of gall cortex (Fig. 23) and (3) a one-layered palisade parenchyma just below the adaxial epidermis (Fig. 23). The vascular system consisted of small collateral bundles concentrically aligned between the fourth and the fifth parenchyma layers (Fig. 23).


description:
  Aceria lantanae (Cook)

  A. lantanae crinckle galls were uni- or multi-chambered and contained several mites, caused by several leaf foldings, and consisted of hyperplasic epidermis and parenchyma.

  Aceria lantanae induced crinkle-leaf light green galls with an irregular and hairy surface, more prominent to the adaxial side of the leaf. They occurred along the secondary veins and were 4.5 ± 0.4 mm in length. These were covering galls that could eventually form a pocket, when the adaxial portion invaginated, provoking an intumescence. As a result, an adaxial folding, centrally opened, was formed (Fig. 8). 

  On the adaxial epidermal layer of the gall, glandular and non-glandular trichomes were similar to those found on healthy leaves. However, around gall ostiole trichomes presented a higher degree of alteration (Fig. 9) and emergences were formed at trichome basis (Fig. 10). The nymphal chamber sheltered numerous A. lantanae individuals (Fig. 11). 

  The nymphal chamber was bound by 1–3 layers of quadrangular cells differentiated from the hyperplasia of the adaxial epidermis, forming an atypical nutritive tissue and several emergences alternating with mite-feeding sites (Fig. 12). Some suberised spots were observed (Fig. 13). 

  Gall cortex consisted of hyperplasic small parenchyma cells with conspicuous nuclei. These cells were irregular in shape, with few chloroplasts. Intercellular spaces were not conspicuous (Fig. 13). The vascular system of the galls was composed of sparse small bundles with well developed vascular parenchyma (Fig. 14)


description:
  This gall morphotype can be distinguished from other midge galls on Populus leaves by the following traits:

  - Larva concealed within galled tissue
  - Gall evenly protruding from both sides
  - Opening in linear slits on the lower side of the leaf
  - Covered in pits, bumps, and small hairs


description:
  Andricus sp. E
  Russet-midrib-gall wasp
  [Page 191]

  Hosts: Quercus oblongifolia, turbinella

  Which fresh, the woolly hairs covering the individual galls are deep rose or brick red, which fades to dark brown with age.


description:
  Descriptions of the gall induced by Harmandiola stebbinsae are contradictory. The original Stebbins description states the gall is found "above mainly, presence indicated below by a light-colored, roughened space", suggested something most similar to our [p-tremuloides-like-globuli](https://www.gallformers.org/gall/4033). However, Gagne 1989's drawing shows the species to induce a concentric circular gall similar to [p-balsamifera-leaf-gall](https://www.gallformers.org/gall/3987), though it seems unlikely that Gagne would not note that this was an exposed-larva gall if it were. Adding to the confusion is the fact that no observations on iNat that match either description have been reported from the type host, Populus grandidentata. 


description:
  The image Stebbins provides for this species resembles a Harmandiola midge gall more than any other Pemphigus species. The undescribed midge that most closely conforms to Stebbins' description is [p-tremuloides-pouch-gall.](https://www.gallformers.org/gall/4031) Since this aphid is not listed in any later summaries of the genus Pemphigini, this may be a mistaken ID. Aphids are known to colonize midge galls on poplar after the midges depart, so there is a plausible reason this error could have been made.


externallink: https://www.biodiversitylibrary.org/page/61038784#page/158/mode/1up
description:
  Dicrodiplosis helena Felt

  The midge was reared from subglobular galls on the underside of Aspen leaves, Populus tremuloides, collected by Miss Cora H. Clarke at Magnolia, Mass. It is by no means certain that in this species we have the gall maker.


description:
  Mordwilkoja vagabunda

  The stem mother attacks the terminal bud of shoots to form a hollow, convoluted, bladder-shaped structure (Figs. 4 and 5). This structure apparently represents a cluster of galls, with each gall induced from a single leaf (Gillette 1914). Large clusters may reach diameters of 80-95 mm and attain a thickness of about 50-60 mm (Gillette 1914). Galls in southern Alberta produce an average of 637 (range: 98-2,040) alatae, with emergence beginning in mid- to late July and being 90% complete by August 5 (Harper 1959b).

  The primary host is P deltoides, on which empty galls are often retained during winter months (Palmer 1952; Harper 1959a). Secondary hosts have not been reported. The author has seen one case in Montana of this aphid on what appeared to be a hybrid between P deltoides and P angustifolia. The aphid can be common on individual trees, but its distribution seems to be highly localized (Gillette 1914; KF, pers. obs.).


description:
  Pachypappa pseudobyrsa (Walsh, 1863) 

  Feeding by the stem mother induces a pocket-like pseudogall adjacent to the midrib of the leaf blade (Fig. 6). The gall projects up from the upper leaf surface, opening on the lower leaf surface. The gall is too small to contain the apterae, which crawl out to feed on the secondary leaf veins on the underside of the leaf. This feeding results in the lower leaf surface becoming coated with a powdery white wax formed by the apterae. Feeding also causes the two sides of the lower leaf surface to gradually bend toward one another to become almost parallel and form a purse-like pseudogall (Fig. 7). Galls are light green but occasionally have a tinge of red along the midvein of the leaf. Each gall may produce an average of 625 alatae (Alleyne and Morrison 1977). The species has also been referenced in the literature as Asiphum pseudobyrsum (Palmer 1952; Alleyne and Morrison 1977) and Pachypappa pseudobursa (Floate and Whitham 1993).

  The primary host is P deltoides (Palmer 1952; Alleyne and Morrison 1977; Floate and Whitham 1993). Alatae are found in galls in mid-summer and presumably colonize unknown secondary hosts. This aphid has wide geographical distribution, with populations locally concentrated on individual trees.


description:
  Pemphigus betae Doane, 1900

  (= P balsamiferae Williams, 1911) 

  Galls are initiated on the upper leaf surface, usually along the midrib. Fully formed galls are pocket-like structures about 10 mm in length with a depth of 5 mm extending below the leaf surface. Developing galls are green, with mature galls frequently turning reddish. There is typically one gall per leaf that most often occurs near the base of the leaf blade. This location appears to be optimal for gall initiation and offspring reproduction to the extent that stem mothers will fight to exclude competitors from these sites (Whitham 1979). Gall densities can be highly variable, with particularly susceptible trees having five or more galls per leaf (Fig. 8). Sexuparae appear to have a limited ability to detect this susceptibility and will preferentially colonize those hosts on which stem mothers are most likely to be able to induce galls in the following spring (Moran and Whitham 1990). Galls in southern Alberta have been reported to produce an average of 41 (range: 2-180) alatae, with emergence beginning in late June to early July and being 90% complete by July 23 (Harper 1959b).

  Primary hosts include P angustifolia, P balsamifera, and hybrid intermediates (Palmer 1952; Harper 1959a; Floate and Whitham 1993). Secondary hosts include sugar beet (Chenopodiaceae: Beta vulgaris L.), for which Pemphigus betae has been reported to vector curly top virus and virus yellows (refs. in Harper 1959a). Other secondary hosts in the family Chenopodiaceae include lamb's quarters (Chenopodium album L.), spinach (Spinacia oleraceae L.), Swiss chard (Beta vulgaris var. cicla), and dock (Rumex spp.). Other secondary hosts include lettuce (Asteraceae: Lactuca sativa L.) and pigweed (Amaranthaceae: Amaranthus sp.) (Harper 1959a). This aphid is common and has wide geographical distribution wherever its primary host species occurs.

  Molecular analyses identify three co-occurring genetic "types" of P betae (Foottit et al. 2008). Further studies are needed to determine if these types represent sibling species and to identify mechanisms for their reproductive isolation.


description:
  Pemphigus bursarius (Linnaeus, 1758) 

  Stem mothers form flask-shaped galls on the leaf petiole or occasionally on young shoots (Harper 1966). As many as three or four galls may occur on one leaf, each producing up to 78 alatae (Alleyne and Morrison 1977). First instars of this species include a "soldier" caste distinguished by their aggressive behaviour in attacking natural enemies (Rhoden and Foster 2002). Soldier aphids attack using sustained thrusts of their stylets to pierce the predator's cuticle. Alternatively, soldier aphids may use their legs to squeeze predators to rupture the cuticle and damage internal organs (Rhoden and Foster 2002). Such attacks are costly to the aphids but can kill predaceous maggots (Diptera: Syrphidae) to protect other aphids remaining in the gall.

  Populus nigra L. var. italica is the primary host (Harper 1959a; Alleyne and Morrison 1977). Secondary hosts include lettuce, on which P bursarius is considered to be an economically important pest in Europe and in the United States (refs. in Harper 1966; Alleyne and Morrison 1977). Other secondary hosts include carrot (Apiaceae: Daucus sp.), lamb's quarters, and nipplewort Lampsana communis L. and Sonchus spp. (Asteraceae) (Palmer 1952). The aphid apparently has wide distribution in Canada, being previously reported from Alberta, Ontario, and Quebec (Harper 1966).


description:
  Pemphigus junctisensoriatus Maxson, 1934 

  Galls resemble those of P populicaulis (see section 8), being more or less globular and incorporating tissues at the base of the leaf blade and the petiole. However, galls of P junctisensoriatus are primarily composed of petiole and have a long slit extending more than halfway around the gall. Galls of P. populicaulis are primarily composed of leaf blade tissues and have a small round opening (Maxson 1934). Palmer (1952) reported the species to be "rare." Alleyne and Morrison (1977) reported collections of P. junctisensoriatus from only one tree at Macdonald College Campus in Quebec. There have been no reports of this aphid from the grassland region, but its occurrence seems probable.

  The primary host is P deltoides (Maxson 1934; Palmer 1952; Alleyne and Morrison 1977). Secondary hosts have not been reported.


description:
  Pemphigus monophagus Maxson, 1934. 

  Stem mothers feed near the base of the leaf blade to form a roughly globular gall mainly comprising the swelling of the upper surface of the leaf blade and partially comprising the swelling and twisting of the leaf petiole (Palmer 1952). The gall has a circular opening on the underside of the leaf and is frequently bisected by a shallow groove, suggesting a double gall (Maxson 1934). This species does not have a secondary host, completing its entire life cycle on the Populus host. The only apparent generation of alatae are the sexuparae, which exit the gall in late August and early September. First instars of this species include a morphologically distinct subset of instars that are probably members of a soldier caste, although this has not been confirmed either experimentally or by direct observation (Aoki and Kurosu 1988).

  Primary hosts include P angustifolia and P balsamifera (Palmer 1952; Alleyne and Morrison 1977) (P balsamifera = P tacamahaca in Palmer 1952). Palmer (1952) reports P monophagus to be "rather rare," although high densities of galls were reported on leaves of P. balsamifera at Pigeon Lake, Alberta (Alleyne and Morrison 1977).


description:
  Pemphigus nortonii Maxson, 1934. 

  Descriptions of this gall vary. Maxson (1934) describes galls of this species as "similar" to those of P. populicaulis. Palmer (1952) states that galls of the two species are "indistinguishable." Harper (1959a) reports that galls of the two species are "somewhat similar" but can be distinguished. Whereas galls of P nortonii are formed solely from the swelling and twisting of the petiole and open via a long, semi-spiral slit, galls of P populicaulis include tissues of both the petiole and leaf blade and open via a small round exit hole (Harper 1959a). In southern Alberta, each gall produces an average of 315 (range: 20-812) alatae, with exiting beginning in late June to mid-July and being 90% complete by August 6 (Harper 1959b).

  The primary host is P deltoides (Maxson 1934; Palmer 1952; Harper 1959a). Secondary hosts have not been reported. The aphid is reported to be uncommon (Palmer 1952).


description:
  Pemphigus populicaulis Fitch 1859 [Poplar leaf-petiole gall aphid]. 

  Galls are initiated on the upper leaf surface, with blade and petiole becoming twisted to form a more or less globular structure at the juncture of the leaf petiole and the leaf blade. The structure is composed of petiole tissue but also includes tissues of the leaf blade that usually extend some distance down onto the gall (Figs. 9 and 10). Twisting of the leaf tissues forms a semi-spiral groove that extends two-thirds of the diameter of the gall. One or a few small round openings in the groove of mature galls provide an exit for alatae. An average of 410 alatae per gall was reported for Quebec (Alleyne and Morrison 1977). An average of 123 alatae per gall (range: 8-426) was reported for southern Alberta, with their exiting from galls beginning in late June to mid-July and being 90% complete by September 11 (Harper 1959b).

  The primary host is P deltoides (Palmer 1952; Harper 1959; Alleyne and Morrison 1977; Floate and Whitham 1993). Rare occurrences of P populicaulis have been reported on intersectional hybrids (Gillette and Palmer 1934; Floate and Whitham 1993). Secondary hosts include water parsley (Apiaceae: Oenanthe sarmentosa C. Presl ex DC.) and possibly fleabane (Erigeron spp.) and goldenrod (Solidago spp.) (Asteraceae) (refs. in Harper 1959a). Harper (1959a) cited references that raised the possibility that P populicaulis might be synonymous with P brevicornis. This synonymy is not supported in more recent work (Eastop and Hille Ris Lambers 1976).


description:
  Pemphigus populiglobuli

  Galls are semi-globular structures formed by the swelling of the leaf blade at the junction with the petiole and extend above the plane of the upper leaf surface (Fig. 11). Although morphologically similar to galls of P. populicaulis, galls of P. populiglobuli typically have a long, narrow slit between the modified tissues of the leaf blade and the petiole, whereas galls of P. populicaulis have a small, pore-like opening. Mature galls are green, often with a reddish tinge. In southern Alberta, galls produce an average of 56 (range: 8–174) alatae, with their exiting from galls beginning in late June to early July and being 90% complete by July 20 (Harper 1959 b). 

  Primary hosts include P. angustifolia, P. balsamifera, and hybrid intermediates (Maxson and Knowlton 1929; Gillette and Palmer 1934; Harper 1959 a; Floate and Whitham 1993). However, the key in Alleyne and Morrison (1977) associates P. populiglobuli with P. deltoides .


description:
  Pemphigus populiramulorum Riley 1879 [Poplar twig gall aphid*]. 

  Whereas galls of other Pemphigus species are formed from tissues of the leaf blade and/or petiole, galls of P. populiramulorum occur on twigs of the current year. Feeding on the young twigs causes swelling of the tissues that expand into a hollow globular structure that surrounds the stem mother and her offspring (Fig. 12). The exit opening is a narrow slit with slightly protruding lips. The location of galls on the woody tissues of the twig results in vacant galls frequently being retained on the tree during winter months. The point of attachment left by abscised galls is a characteristic flattened “button” region on the twig (Fig. 13). Galls in southern Alberta produced an average of 74 (range: 9–190) alatae, with gall exiting beginning in early to mid-July and being 90% complete by August 23 (Harper 1959 b). The primary host is P. deltoides (Palmer 1952; Harper 1959 a ; KF, pers. obs) . This aphid is described by Maxson and Knowlton (1929) as occurring on P. balsamifera , but this report has been questioned by Palmer (1952).


description:
  Pemphigus populitransversus Riley 1879 [Poplar petiolegall aphid*]. 

  The stem mother feeds on petioles to cause the swelling of tissues that form a hollow, globular structure that surrounds both the stem mother and her offspring (Figs. 12 and 14). Mature galls have a transverse exit slit with margins that may be thickened and slightly protruding, reminiscent of lips. An average of 150 alatae per gall was found in Quebec (Alleyne and Morrison 1977). An average of 192 alatae per gall (range: 25–962) was found in southern Alberta, with gall exiting beginning in late June to mid-July and being 90% complete by August 23 (Harper 1959 b ). Jones and Gillette (1918) provide detailed taxonomic and life history information of this species on both primary and secondary hosts. 

  The primary host is P. deltoides (Maxson and Knowlton 1929; Gillette and Palmer 1934; Harper 1959 a ; Alleyne and Morrison 1977; Floate and Whitham 1993). Secondary hosts include sugar beet, dock, cabbage, turnip, and lettuce (refs. in Alleyne and Morrison 1977). 

  Pemphigus obesinymphae Aoki is a close relative of P. populitransversus and induces a gall that may be easily mistaken for that of the latter species (Aoki and Moran 1994). Pemphigus obesinymphae is not known to occur in Canada, but is reported to co-occur with P. populitransversus on P. deltoides in the United States (Abbott and Withgott 2004). The species is mentioned here to increase awareness of its possible presence in Canada.


description:
  Pemphigus populivenae Fitch 1859 [Poplar vein gall aphid, also known as sugarbeet root aphid*]. 

  Feeding by the stem mother on the underside of the leaf blade causes the leaf tissue to swell and form a pocket-shaped gall. Galls are oval to semi-oblong, open on the underside of the leaf, and project above the upper plane of the leaf blade. Galls are usually closely associated with the midrib or a primary vein. They attain a length of about 8– 12 mm, a depth of about 5 mm, and are green. 

  Primary hosts include P. angustifolia and P. balsamifera (Maxson and Knowlton 1929; Palmer 1952; Harper 1959 a). Whitham (1978) reports P. populivenae on Fremont cottonwood, P. fremontii S. Watson ( Aigeiros ), in the United States. Quinoa, Chenopodium quinoa (Chenopodiaceae), is a secondary host (Cranshaw et al. 1990).


description:
  Pemphigus spyrothecae (= spirothecae ) Passerini 1856. 

  Feeding by the stem mother causes the leaf petiole to twist into spirals (usually three) that surround the aphid. The spirals swell, with their seams becoming tightly pressed together to form a hollow, spiral- shaped gall that completely surrounds the stem mother and her offspring (Fig. 15). Galls mature in late August or early September and are initially green, but often become reddish. Alatae exit the mature gall via a series of small pores that form along the seams of the spirals. Pemphigus spyrothecae does not have a secondary host and, as reported for P. bursarius and suspected for P. monophagus , has a morphologically distinct subset of first instars that form a soldier caste. Experimental studies show that galls containing 50 soldier and 50 non-soldier aphids versus 100 non-soldier aphids can be 10 times less likely to be attacked by predators under field conditions (Foster and Rhoden 1998). 

  Populus nigra L. var. italica is the primary host (Alleyne and Morrison 1977). This aphid was first reported in North America from material collected in 1972–1973 in Quebec, where it was extremely common (Alleyne and Morrison 1974). Blackman and Eastop (1994) report its introduction into western Canada. In addition, high densities were observed by the author in 2003 at Kelowna, British Columbia.


description:
  Thecabius gravicornis (Patch, 1913) (Pemphigus) [Folded-leaf balsam aphid]. 

  Feeding by the stem mother adjacent to the midrib on the lower surface of the leaf blade initiates a small cavity in which apterae are produced. The apterae leave the cavity to feed on expanding terminal leaves. Their feeding causes the leaves to fold ventrally along the midrib to form a purse-shaped pseudogall similar to that of P. pseudobyrsa . In Quebec, alatae appear in the pseudogalls from mid-June to early July (Alleyne and Morrison 1977). 

  Primary hosts include P. balsamifera (Gillette and Palmer 1934) and P. angustifolia (Blackman and Eastop 1994). There has been one report of T. gravicornis on P. nigra L. var. italica (Alleyne and Morrison 1977). This aphid is apparently widespread but uncommon (Blackman and Eastop 1994). 


description:
  Thecabius populimonilis (Riley, 1879) (Pemphigus) [Bead-like cottonwood gall aphid]. 

  The life cycle of this aphid is reviewed by Gillette (1913) and by Maxson and Knowlton (1929). The stem mother feeds on the underside of the leaf blade, causing formation of a blister-like pseudogall that projects above the upper plane of the leaf surface. Her offspring (the second generation) are apterae that leave the gall shortly after birth to each induce their own similarly shaped gall, either on the same leaf or on adjacent leaves of the same shoot. This results in one or more rows of galls forming parallel to the midrib, with each gall containing one aphid (Fig. 17). Most of these aphids become alatae that migrate to other trees. However, some individuals stay in the galls and remain as apterae to produce apterous offspring (the third generation). These apterae leave the gall to move onto adjacent leaves to induce new galls and produce offspring, a portion of which develop into winged sexuparae. Gillette (1913) reports that 10–12 sexuparae may occur in one gall with the parent. The sexuparae exit these galls in late summer to produce male and female sexuales, which mate and produce the overwintering egg from which the stem mother develops in the following spring. 

  Primary hosts include P. angustifolia and P. balsamifera (Maxson and Knowlton 1929; Gillette and Palmer 1934; Harper 1959 a ; Floate and Whitham 1993). There is likely no secondary host.


description:
  Aceria neoessigi (Keifer, 1940) (Eriophyes). 

  Mites overwintering in the buds of Populus emerge in the spring to attack the flowers of the catkin. Attacked flowers become malformed and enlarged and group tightly together such that the catkin forms a large pendant gall that may exceed 20 cm in length (Fig. 18). Galled catkins remain on the tree until early summer, long after unattacked catkins have dehisced. The species does not have a secondary host. Galled catkins may also contain Aceria populinquis Keifer, which occurs as an inquiline (Wilson and Oldfield 1966). In the United States, P. fremontii is the primary host (Wilson and Oldfield 1966). In Canada, the primary host is P. deltoides (KF, pers. obs.), which is closely related to P. fremontii (see Fig. 6 in Eckenwalder 1996).


description:
  Aceria parapopuli (Keifer, 1940) (Eriophyes). 

  Mites overwinter in galls to exit in the spring and attack newly expanding vegetative and reproductive buds. Attacked buds form a woody, cauliflower-like gall that may exceed 3 cm in diameter (Fig. 19). Active galls normally persist for 1 year but can remain active for up to 15 years. New generations are produced every two to three weeks during summer months. Active galls are red and green, whereas dead galls are brown. Aceria parapopuli does not have a secondary host but can disperse between trees using wind currents (ref. on life history cited in McIntyre and Whitham 2003). 

  Primary hosts include P. angustifolia , P. balsamifera , and hybrid intermediates (Floate and Whitham 1993).


description:
  Thecabius populiconduplifolius (Cowen, 1895) (Pemphigus) [Folded-leaf poplar aphid].

  Feeding by the stem mother at the leaf margin causes the edge of the leaf blade to fold under and form a small gall (Fig. 16). The apterae produced in this gall exit to feed on the underside of expanding leaves, which produces galls similar to those of T. gravicornis.

  Reported primary hosts include P. deltoides (Palmer 1952), P. balsamifera (Harper 1966), and P. nigra L. var. italica (Alleyne and Morrison 1977). Secondary hosts include species of Ranunculus (Swain 1919; Palmer 1952). Thecabius populiconduplifolius has an identical biology and is virtually indistinguishable from the Eurasian species T. affinis (Kaltenbach), although the two species can be separated by chromosome number (Blackman and Eastop 1994).


description:
  Pemphigus betae

  The aphid forms pouch-shaped galls on the leaf blades of its host; among its primary hosts are the balsam poplars, Populus section Tacamahaca (e.g., P. angustifolia James, P. balsamifera L.), and hybrids involving these species, and cottonwoods in Populus section Aigeiros (e.g., P. deltoides Marsh, P. fremontii S. Watson) (Fig. 1) (Palmer 1952; Harper 1959; Floate et al. 1997). 

  The taxonomic status of P. betae remains unclear. Pemphigus betae was originally described from specimens on sugarbeets, the gall form being unknown. Pemphigus balsamiferae Williams, described from the gall form, is currently considered a synonym of P. betae (Maxson 1916; Remaudie`re and Remaudie`re 1997). Grigarick and Lange (1962) considered P. betae to be synonymous with Pemphigus populivenae Fitch, which also forms pouch-shaped galls on the leaf blades of species in Populus section Tacamahaca. Summers and Newton (1989) used the name P. populivenae for aphids on sugarbeets in California. Harper (1959) designated aphids that formed galls projecting below the plane of the leaf blade on Populus hosts as P. betae (Fig. 1) and used the name P. populivenae for aphids forming galls that projected above the plane of the leaf blade. However, according to Maxson and Knowlton (1929), in heavy infestations the latter can occur in the position typical of P. betae. In addition, Pemphigus knowltoni Stroyan has been reported from Utah, where it forms galls similar to those of P. betae on P. angustifolia (Stroyan 1970). In Europe the species on beets and other Chenopodiaceae is named Pemphigus fuscicornis (Koch) and has no known associated primary host form (Blackman and Eastop 2000).

  Collectively, these results identify at least three co-occurring populations forming similar galls on the same host trees.The concordance of the mitochondrial and nuclear data suggests that the three groups are reproductively isolated. Specimens from group A (Fig. 2) key to P. betae. Specimens from group C key to P. populivenae. Specimens from group B are intermediate. No specimens matched the state reported for P. knowltoni. The available evidence suggests that group B is an undescribed species. Other than P. betae, P. populivenae, and P. knowltoni, no North American Pemphigus species are known to regularly form elongate galls along the leaf midrib of species in Populus section Tacamahaca. Although group C was identified as P. populivenae, there are several inconsistencies. Galls of this species are reported to project above the plane of the leaf blade (Harper 1959), but galls sampled in the present study resembled those of P. betae, i.e., they project below the leaf blade. Galls of P. populivenae can apparently resemble those of P. betae when populations of the former are high (Maxson and Knowlton 1929). However, galls in the position normally ascribed to P. populivenae are uncommon in southern Alberta (Harper 1959; K.D.F., personal observations), whereas group C samples were common in our collections. These discrepancies are resolved if P. populivenae forms galls that can project either above or below the leaf blade regardless of population density. Alternatively, group C may represent an undescribed species morphologically similar to P. populivenae. Groups B and C co-occur in morphologically pure zones of P. balsamifera (samples from Cypress Hills and Lundbreck Falls, Alberta). Group B was not detected at sites where P. deltoides (section Aigeiros) or hybrids involving it occur.

  It is apparent that several taxa form similar galls on the leaf blades of species and hybrids of Populus section Tacamahaca. The form of a gall in a given position (petiole, petiole2blade junction, along midvein), may depend as much on host physiology or morphology as on the inducing aphid species. Anecdotal observations that we made during recent collections indicate that midvein galls containing morphologically similar aphids tend to be more globular when close the leaf base than those located more distally, especially on thicker leaves, sometimes approaching the state characteristic of Pemphigus populiglobuli Fitch. Maxson and Kowlton (1929) noted galls similar to those of P. populivenae produced by aphids they identified as Pemphigus populicaulis Fitch and P. populiglobuli. Reliance on gall position and form as indicators of species identity is clearly not a viable approach for use with this group of Pemphigus species, given the current state of knowledge.


description:
  Pemphigus populivenae

  Grigarick and Lange (1962) considered P. betae to be synonymous with Pemphigus populivenae Fitch, which also forms pouch-shaped galls on the leaf blades of species in Populus section Tacamahaca. Summers and Newton (1989) used the name P. populivenae for aphids on sugarbeets in California. Harper (1959) designated aphids that formed galls projecting below the plane of the leaf blade on Populus hosts as P. betae (Fig. 1) and used the name P. populivenae for aphids forming galls that projected above the plane of the leaf blade. However, according to Maxson and Knowlton (1929), in heavy infestations the latter can occur in the position typical of P. betae. In addition, Pemphigus knowltoni Stroyan has been reported from Utah, where it forms galls similar to those of P. betae on P. angustifolia (Stroyan 1970). In Europe the species on beets and other Chenopodiaceae is named Pemphigus fuscicornis (Koch) and has no known associated primary host form (Blackman and Eastop 2000).

  Collectively, these results identify at least three co-occurring populations forming similar galls on the same host trees.The concordance of the mitochondrial and nuclear data suggests that the three groups are reproductively isolated. Specimens from group A (Fig. 2) key to P. betae. Specimens from group C key to P. populivenae. Specimens from group B are intermediate. No specimens matched the state reported for P. knowltoni. The available evidence suggests that group B is an undescribed species. Other than P. betae, P. populivenae, and P. knowltoni, no North American Pemphigus species are known to regularly form elongate galls along the leaf midrib of species in Populus section Tacamahaca. Although group C was identified as P. populivenae, there are several inconsistencies. Galls of this species are reported to project above the plane of the leaf blade (Harper 1959), but galls sampled in the present study resembled those of P. betae, i.e., they project below the leaf blade. Galls of P. populivenae can apparently resemble those of P. betae when populations of the former are high (Maxson and Knowlton 1929). However, galls in the position normally ascribed to P. populivenae are uncommon in southern Alberta (Harper 1959; K.D.F., personal observations), whereas group C samples were common in our collections. These discrepancies are resolved if P. populivenae forms galls that can project either above or below the leaf blade regardless of population density. Alternatively, group C may represent an undescribed species morphologically similar to P. populivenae. Groups B and C co-occur in morphologically pure zones of P. balsamifera (samples from Cypress Hills and Lundbreck Falls, Alberta). Group B was not detected at sites where P. deltoides (section Aigeiros) or hybrids involving it occur.

  It is apparent that several taxa form similar galls on the leaf blades of species and hybrids of Populus section Tacamahaca. The form of a gall in a given position (petiole, petiole2blade junction, along midvein), may depend as much on host physiology or morphology as on the inducing aphid species. Anecdotal observations that we made during recent collections indicate that midvein galls containing morphologically similar aphids tend to be more globular when close the leaf base than those located more distally, especially on thicker leaves, sometimes approaching the state characteristic of Pemphigus populiglobuli Fitch. Maxson and Kowlton (1929) noted galls similar to those of P. populivenae produced by aphids they identified as Pemphigus populicaulis Fitch and P. populiglobuli. Reliance on gall position and form as indicators of species identity is clearly not a viable approach for use with this group of Pemphigus species, given the current state of knowledge.


description:
  Pemphigus betae

  As with most host-alternating aphid species, the sugarbeet root aphid is restricted in its primary host range, but the secondary host range is more diverse. Pemphigus spp. prefer poplars in the genus Populus (Fig. 2) as their primary hosts (Blackman and Eastop 2006). Over the years, P. betae primary hosts have been identified as species within the Populus section Tacamahaca, including narrowleaf cottonwood (P. angustifolia James), balsam poplar (P. basamifera L.), and black cottonwood (P. trichocarpa Torr. & A. Gray) in North America (Parker 1915, Harper 1963, Whitham 1978, Floate et al. 1997, Dewar and Cooke 2006, Hein et al. 2009). Although there is some overlap in their geographical distribution, these primary hosts occupy relatively distinct regions in North America. For example, narrowleaf cottonwood is found primarily in the central and southern Rocky Mountain regions, the balsam poplar is found in the northern Rockies into Canada and extending east to the Great Lakes region, and the black cottonwood has a range stretching from Alaska, through western Canada, and primarily into the northwestern United States, but also sporadically down as far south as Baja, CA (U.S. Department of Agriculture [USDA] 2014). Additional ecological complexity of these aphid–primary host relationships was demonstrated by Floate et al. (1997) who found a dramatic increase in P. betae host acceptability for hybrids between the aphid’s primary hosts and other Populus spp.


description:
  Thecabius populi-monilis (Riley). 

  This aphid forms characteristic rows of bead-like cells, particularly on the upper sides of the leaves of narrow-leaved cottonwood, Populus angustifolia, and to a less extent on some of the wider-leaved species of poplars. The stem mother produces a gall similar to that formed by her young. These galls occur singly, usually between the midrib and the margin of the leaf. The first bead-like rows of galls are produced by the young of the fundatrix. Only a single individual is found in each cell. 

  According to Gillette, most of the second generation lice become winged and leave the galls, thus serving to scatter the species from tree to tree. A portion, especially of the later lice that are born, remain apterous, remain in the galls, and give rise to the third generation. These young, like the young from the stem mother, also migrate to the new leaves to continue the production of galls each of which harbors but one louse at first, but a portion of the young of this brood remain with the mother in the gall and become winged sexupara, of which 10 or 12 may be found in a single gall with the parent. Emergence of these sexupara begins about the first week of August and continues until the last of September; soon after emergence the sexupara give birth to the se.xual forms. 

  On the narrow-leaved cottonwood, the galls are most commonly in rows, on one or both sides, and rather paralleling the midrib. On P. balsamifera, the rows may follow the margin, midrib. parallel the midvein, or run at different angles on different parts of the leaf. Sometimes three or four rows of galls will occur on the broad part of the leaf. 

  The stem mother usually has 5-jointed antennæ and is wingless. The winged individuals of the second generation emerge from the galls within a period of a few weeks. The mouth of the gall opens slowly, finally permitting the winged female to emerge. Predators frequently enter the opening galls and kill the pupal nymphs. Several species of syrphid larvæ and a small hemipterous insect were the forms most commonly observed in Utah. 

  Wingless females were numerous in the galls at St. George, on July 9, 1925. 

  This species has been collected in Utah from Blacksmith Fork Canyon, Cedar City, Eden, Farmington, Hyde Park, Murray, Logan, St. George, Salt Lake City, Santa Clara, and Smithfield.  


description:
  Thecabius populi-conduplifolius (Cowen). 

  This aphid folds the leaves of balsam poplar, Populus balsamifera and P. deltoides, producing a pseudogall along the midvein or parallel to it. The stem mothers may be found in galls near the margin of the leaf. The young of the stem mother do not remain in the gall with her, but migrate to the new leaves at the tips of the branches. All of the second generation become winged. They migrate to Ranunculus sp. where the third and several subsequent generations of apterous viviparous females are produced. These summer colonies give a cottony appearance to the base of the Ranunculus leaves and stems. The fall migrants (sexupara) appear during late summer and fall and migrate back to the cottonwoods where the sexual forms are produced. 

  Galls of the second generation are always on terminal leaves. However, late in the season, after the stem mother ceases producing, new terminal leaves are produced which do not become infested. 

  At Fruita, and along the stream below Capital Bridge, nearly all of the leaves on the young poplars were folded on June 27, 1926. By this time most of the aphids had migrated. Many of the aphids had been parasitized, but a few living specimens were obtained. These aphids were quite numerous in the galls at Randlett on July 14, 1927, and less so at the same time in the few collections made at Fort Duchesne and Vernal. Ordinarily, quite a number of aphids occur in each gall. When they were examined at St. George on July 9, 1925, only a few specimens remained in galls. Winged females were becoming abundant at Plain City, by June 15, 1928. 


description:
  Pemphigus populi-caulis Fitch.

  [Two drawings of the gall appear on page 9 of the pdf]

  The Gall.—The gall is semi-globular. Large ones attain a diameter of 1/2 inch or more. Normally located galls are at the base of the leaf where the blade and petiole unite. The gall is composed of the thickened and twisted petiole and a very small portion of the base of the leaf blade which frequently extends a short distance down onto the gall. The greater part of the gall extends below the plane of the leaf blade. Viewed from above the mouth is normally not visible and the midrib and the first diagonal veins on either side are seen to merge into the body of the gall. 

  When viewed from below the midrib is seen to bend abruptly and to merge with a ridge formed by the edge of the petiole which follows the curve of the gall and forms a part of one lip of the opening. The other lip of the opening is formed by a fold in the base of the leaf blade for a part of its length and the balance by the edge of the petiole. The petiole is twisted so that the flattened portion lies in nearly the same plane as the leaf blade instead of at right angles to it as in normal leaves. It is clear that almost the entire gall is made up of the much enlarged and twisted petiole. The mouth is usually full-lipped and extends more or less in a spiral fully 2/3 the distance around the gall. Many times there is a small round opening at some point along the lips through which the winged lice escape. The senior author has collected aphids which correspond in every respect to P. p. -caulis, from galls similar to those produced by P p.-venae at Oak Park MN, and in the Bitter Root Valley in MT. These were from Populus balsamifera trees heavily infested with P p-globuli and P. p-venae. It seems than under certain conditions at least P. p-caulis produces galls on the leaf similar to those produced by P. p.-venae. 

  This aphid was very abundant in Logan during the spring of 1925. The infestation was so severe that by July 1st many leaves had fallen from the infested poplars, Populus balsamifera and P deltoides. About 3 percent of the leaves remaining on the trees were infested with galls.

  The aphids are ordinarily very numerous in these galls, and sometimes a hundred or more pupal nymphs and winged females in addition to the stem-mother are present in a single cell. Ordinarily, there is a large drop of liquid in the bottom of the gall. The waxy material secreted by the wax glands makes the inside of the gall grayish so that when it is opened, the moving aphids appear to be mixed up in a growth of fungus.

  In addition to being present in Logan, this aphid was also found in Big Cottonwood Canyon, Utah, on June 29, 1925, where it was producing its characteristic galls. it has also been collected at Hyde Park during early July, 1928.

  Notes:

  Pemphigus populi-caulis and Pemphigus populi-globuli were described by Dr. Asa Fitch in the " Report of the Noxious and Other Insects of the State of New York," 1859. 

  Fitch's descriptions of the insects are too meager to make a separation of the species certain without the galls. 

  The gall produced by the aphis Fitch described as P. p.-caulis is described as a "bullet-like gall " on the middle of a leaf stalk from a Lombardy poplar. This gall was found on the 27th of June, three years before the description of P. p.-caulis was publishéd. 

  Discussing the gall further, Fitch says: "When the leaves fell from this tree in autumn, a few were found among them, more faded than the others and having these same excrescences, but placed at the base of the leaf instead of on the middle of its stem"

  Farther on in his discussion, Fitch states: "Galls analagous to those here described grow upon the leaf stalks of the Lombardy and Black poplar in Europe, from attacks of the Pemphigus bursarius of Linnaeus. " 

  After comparing it with specimens of that species and its galls, Fitch considered his aphid different from P. bursarius, the fly being paler in its color, and its gall because of spirally coiled somewhat like the shell of a snail."

  It is very evident from Fitch's discussion that he had in mind, when he described P. p.-caulis, a gall located on the petiole of the leaf at some distance from the leafblade, and one produced by a twisting of the petiole into a spiral. 

  It is quite probable that the petiole gall described by Fitch is not typical of P. p.-caulis, but that those at the the base of leaf which he mentions as being taken in the fall are the typically located galls. 

  The aphid herein figured and described as Pemphigus populi-caulis is from galls located at the base of the leaf and produced by a twisting or spiralling of the petiole. The species described as P. p.-globuli, is considered to be the globuli of Fitch because of the likeness of its gall to the one described by Fitch in his discussion of that species. 

  Since the original descriptions of P. P.-caulis and P. P.-globuli were published these species have received much attention, if we may judge from the frequent references to both. However, in spite of this, several aphidologists appear to have failed to distinguish between them. 

  P. p.-caulis and P. p.-globuli are very similar structurally and their galls are, in a general way, much alike, although quite different when studied in detail. This general similarity of both the aphids and their galls appears to have caused considerable confusion in the literature. This was probably increased by a lack of detail in the earlier descriptions. 

  In studying the two species under discussion, the senior author has collected material from several states and in many localities in some of these states. When collections of both species have been made in the same locality, they rarely have been found on the same host species. In Colorado and Utah P. p.-caulis was taken on Populus deltoides and P. p.-globuli on Populus angustifolia and Populus balsamifera. In California P. p.-caulis was taken on Populus Fremontii and P. p.-globuli on Populus trichocarpa. P. p. -caulis was taken on Populus deltoides at St. Cloud, Minnesota, and both species on Populus balsamifera at Oak Park, Minnesota. East of the Rocky Mountains in Montana P. p.-caulis was taken on Populus deltoides and P. p.-globuli on Populus angustifolia. P. p.-caulis was taken on Populus balsamifera in the Bitter Root Valley of Western Montana and P. p.-globuli on the same species of Populus near the Flat Head Lake. The junior author reports both species on Populus balsamifera. These observations appear to indicate that Populus balsamifera is the only common host of P. p.-caulis and P. p.-globuli east of the Rocky Mountains. Populus trichocarpa may serve as a host for both in the range of this tree. 

  These collections have been carefully separated according to the type of gall and a large series of the aphids from each gall type critically studied. These studies have brought to light certain structural differences by which P. p.-caulis and P. p.-globuli alate forms from the galls may be separated. These differences are constant in relative magnitude but not in actual magnitude. 

  Both species under discussion are quite variable in size and in some other characters. This necessitates studying several individuals and comparing the means of characters in order successfully to separate them. When a sufficient number of individuals to make a representative population is studied, separation becomes easy and certain. 

  Thomas, in the "Third Annual Report on Noxious and Beneficial Insects of Illinois," 1879, pictures a typical Pemp. populi-globuli gall for the gall of Pem. populi-caulis. Dr. O. W. Oestlund described Pem. populi-caulis in his synopsis of the Aphididæ of Minnesota. His description is not sufficiently detailed to determine which of the two species under discussion he really had. The host is given as Populus monilifera. 

  Essig, in Pomona College Journal of Entomology, 1912 describes and illustrates Pem. populi-caulis and several galls. These galls are of 3 quite distinct types. One is typical galls. of Pem. p.-caulis. Several are typical Pem. p.-globuli and one is typical of what the senior author is calling Pem. p-venae. The senior author has collected all three types of galls on Populus trichocarpa in California. The aphids from the typical Pem. p.-globuli and Pem. p.-caulis were the ones used in the studies already referred to. The ones from the other type of gall do not fit with either Pem. p.-caulis or Pem. p.-globuli. Several collections of these galls have been made in Minnesota. The aphids from Minnesota differ from the species being discussed in the same way as those from California. This species is described later as Penz. P.-venae. The descriptions of Essig are not sufficiently detailed to enable one to determine his species with certainty. However, his figures of the antenna, of the alate viviporous female and the stem mother are typical of Pem. p.-caulis.

  It seems quite probable that Essig failed to separate the 3 species in his material. 

  Dr. Edith M. Patch described Pem. populi-caulis from Populus balsamifera and pictures 1 typical Pem. populi-globuli gall and 2 not typical of this species or of Pem. populi-caulis. The latter are similar to galls taken by the senior author at Oak Park, Minnesota, produced by what he calls Pem. populi- venæ.


description:
  Pemphigus populi-globuli Fitch

  The Galls.—The galls are usually semi-globular. Not infrequently they are more or less cone-shaped. Normally located galls are situated at the base of the leaf, on its upper side. The color is green, or when exposed to the tinged with reddish varied with yellow. The gall is composed of a much enlarged and thickened portion of the leaf blade. On the under side there is a somewhat curved mouth-like orifice bounded on one side by the much thickened midrib and on the other by a thickened fold in the leaf blade. A more or less broken portion of the blade usually extends along this fold nearly the entire length of the mouth. 

  Many times the blade of the leaf stands at an angle to the petiole. The opening in the gall is rarely equal to one-half the circumference of the gall and is not visible from above. The lip-like borders of the opening are frequently quite rough. Galls located at some distance from the base of the leaf and resembling those produced by P. P.-venæ sometimes occur, especially on heavily infested trees. 

  This aphid is frequently encountered in Utah, on balsam poplar, Populus balsamifera. It has been collected in this state from Big Cottonwood Canyon, Blacksmith Fork Canyon, Farmington, Hyde Park, Providence, and Salt Lake City. The material described was collected at Salt Lake City. 


description:
  Pemphigus populi-transversus Riley

  This aphid has, as its summer host, plants belonging to the family Cruciferæ. Jones states that Populus balsamifera, P. monilifera, P. trichocarpa and P. fremontii have been mentioned as hosts. He also gives cabbage, turnip, mustard (Brassica nigra), cauliflower, broccoli, (B. oleracca botrytis), Brussels sprouts, rape, and several weeds belonging to the family cruciferæ as hosts of the subterranean forms. While this aphid has been reported as damaging certain cruciferous crops in the south, no damage in the northern portions of its range has been reported. In Utah this aphid was collected at Farmington, on June 28, 1923. At this time spring migrants and pupal nymphs were present in galls formed on the petiole near the leaf of cottonwood, Populus deltoides. Galls and stem-mothers apparently of this species, were collected at Santa Clara by Dr. H. J. Pack, during June, 1928. 


description:
  Pemphigus populi-venæ Fitch

  Pemphigus Populi-venæ was described by Dr. Asa Fitch in 1859. Since the original description was published, this species does not appear to have been recognized but appears to have been confused by some aphidologists with P. p.-caulis and P. p.-globuli. 

  In his studies of this species Dr. Fitch appears to have failed to recognize the real nature of the aphids in the galls. It seems altogether probable that his male and female larvæ and pupæ were the early stages of the alate females (fundatrigenia).

  The winged individuals were so briefly described by Fitch that the description is valueless for purposes of identification. Identifying aphids by the plant deformities produced is very apt to lead to confusion. Easily distinguished species produce very similar deformities and a single species may produce more than one form of deformity. 

  The Galls.—Dr. Fitch describes the galls of P. P.-venæ as "an oblong compressed excrescence like a cock's comb, of a light red color varied with pale yellow growing from the midrib of balsam poplar leaves on their upper side with an orifice on the opposite side. They are nearly semi- circular and half as high as long, being usually over a half inch in length, with uneven surface, their walls thick, brittle and succulent. They are commonly placed near the base of the leaf but sometimes in its middle." 

  Galls similar to those described by Fitch were collected by the senior author at Oak Park, Minnesota, on balsam poplar and near Spreckles, California, on Populus trichocarpa. 

  The aphids from these galls have some of the characters of P. p.-caulis and P. P.-globuli but are not wholly like either. The most noticeable difference between P. P.-venæ and P. p.-caulis and P. P.-globuli is its smaller size. The senior author has taken all three species on a single balsam poplar tree in Minnesota. Because of the similarity of the size of these species to those described by Dr. Fitch and the likeness of the galls of P. p.-venæ as described by Dr. Fitch to those collected in Minnesota and California, the aphid here described is considered the P. p.-venæ of Fitch. 

  The Gall.—Dr. Fitch's description of the gall should be enlarged by stating that the galls vary in form from more or less oblong to semicircular and even almost triangular in outline. On heavily infested trees galls occur on both sides of the leaf. When on the underside they may be confused with those of P. beta, being similar in form and location. 

  Aphids collected at Hyrum, Utah, on June 20, 1926, seem to be the species described by Fitch. These aphids were producing galls on the leaves of the narrow-leaved cottonwood, Populus angustifolia. 


description:
  Pemphigus populi-ramulorum Riley.

  [Drawings of this gall appear on page 17 of the pdf]

  While this species was described from material collected on Populus balsamifera it occurs commonly on Populus deltoides in localities outside the range of the balsam poplar. This species resembles quite closely both P. p.-caulis and P. p.- globuli. 

  This aphid is frequently encountered in Utah and is particularly interesting because of the gall it produces on the young twig bark of poplars, especially Populus balsamifera. Collections made at Leeds, both from twig galls and from galls below the bases of the leaves on the petiole, indicate that this one species produces two types of galls. This is a rather uncommon occurrence among gall-forming aphids. The openings to the leaf galls are often slightly beaked, and the mouth does not always open along the same axis, which gives variation in appearance to the openings of different galls. 

  In Utah this species has been collected from Benjamin, Benson, Farmington, Holliday, Hooper, Leeds, Provo, St. George, Santa Clara, and Zion National Park. 


description:
  Pemphigus betæ Doane.

  A pemphigid from beet roots grown at Pullman, Washington, was named "Pemphigus betae" and described by Doane in 1900. In 1910 Williams described Pemphigus balsamiferae. This aphid was taken at Squaw Creek, Nebraska, on Populus balsamifera. In 1914 the senior author discovered that the sugar-beet root-louse in Colorado is the summer and fall generations of P. balsamiferae from the narrow-leaved cottonwood, Populus angustifolia. Unless the beet is the host of two Pemphigus species and the one in Washington is different from the one feeding on beets in Colorado, P. balsamiferæ Williams, becomes a synonym of P. beta Doane. In this discussion P. balsamiferæ is considered the spring form of P. betae. 

  P. betae occurs on the balsam poplar within the western range of this tree and on Populus angustifolia where this tree occurs naturally. Its galls are located on the underside of the leaf. They may easily be confused with the galls of P. p.-venæ. 

  The following unpublished chapter in the life history of P. betae was worked out by the senior author in 1921. 

  Hibernating aphids were collected from the soil November 2d. These were confined in salve boxes in a greenhouse under ordinary greenhouse temperatures. The aphids collected November 2d. are designated the first generation. The second generation (young of the hibernating lice) appeared December 31st after nearly 2 months' inactivity on the part of the hibernating individuals. The third generation appeared January 21st; the fourth generation, February 24th ; the 5th, March 12th ; the 6th, March 24th; the 7th, April 8th, and the 8th, April 29th. 

  The age of the various generations when reproduction began was: second generation, 21 days old; third, 34 days; fourth, 16 days; fifth, 12 days; sixth, 15 days; and seventh, 21 days. The variation in ages is probably due to variations in temperature and the condition of the food supply. Low temperatures retard development and high temperatures quicken it. 

  The first, second, third, fourth, and fifth generations were all apterous viviparous females which began reproducing immediately upon becoming mature. The first alate forms (sexuparæ) appeared in the sixth generation. The sixth and seventh generations were composed of about equal numbers of apterous viviparous females and alate sexuparæ. The apterous females of the sixth and seventh generation began to reproduce as soon as mature. The offspring of the sixth generation were of four kinds. Those of the apterous forms were apterous viviparous females and sexuparæ; those of the sexuparæ, true males and females. The offspring of the seventh generation (the eighth generation) were of four kinds also. Those of the apterous viviparous females were apterous forms and sexuparæ. The fortner ceased feeding and became inactive upon reaching mature size. They sought out crevices in the soil where they became completely covered with the wax secreted by their wax glands. Those of the sexuparæ were true males and females. The hibernating forms of the eighth generation differ from the apterous forms of the sixth and seventh generations, since upon reaching sexual maturity they produce only apterous forms. 

  This change into hibernating forms appears to be biological function and not a temperature reaction. 

  This aphid has been collected in Utah from the following localities: Amalga, Benson, Brigham, Collinston, Cornish, Delta, Garland, Hoytsville, Hyde Park, Lehi, Lewiston, Logan, Madison, Millville, Richfield, Richmond, Salt Lake City, Smithfield, and Trenton. 


description:
  Prociphilus fraxinifolii (Riley).

  Prociphilus fraxinifolii differs from typical Prociphilus species in the nature of the individuals composing the various generations. The following observation was made by the senior author at Longmont, Colorado, in 1925.

  A young white ash tree in one of the city parks was very heavily infested with Proc. fraxinifolii on June 18. At this time many first instar lice were travelling from the infested tips of the branches toward the trunk of the tree. Upon reaching the trunk, these young lice travelled up or down until another branch was reached, when they started toward its tip. It was thought that the migrating lice might be in search of fresh leaves upon which to locate. 

  At the time of observation some curled leaves contained stem mothers and many large mature apterous females which were giving birth to the young already mentioned. Others contained only a limited number of these young. 

  A cluster of curled leaves containing the stem mothers and apterous females was removed and fastened to a branch of a young ash tree that had no curled leaves. Immediately . the young were observed travelling toward the trunk of the tree. The second day after the curled leaves were placed on the young tree, the leaves of nearly every branch below the point where the curled leaves were attached were badly curled. 

  On the fourth day no leaves above the point where the curled leaves were attached to the branch were curled. The migration of young, therefore, appears to be mainly downward. The third and fourth day after the curled leaves were placed on the young tree they were completely dry and the mature lice in them were dead. Apparently the stem mothers and second generation apterous females do not migrate. 

  This observation indicates that the sequence of generations of Proc. fraxinifolii is somewhat similar to that of Eriosoma species. No winged individuals were secured. It is believed that the winged lice are produced by the migrating generation (the young of the second generation), and, therefore, belong to the fourth generation. 


description:
  The juniper mistletoe (Phoradendron juniperinum) is common on western juniper and causes noticeable stem swellings.


externallink: https://www.biodiversitylibrary.org/page/11256375#page/77/mode/1up
description:
  Asiphum sacculi n. sp.

  I first saw the gall of this louse about eighteen years ago when on a mountain trip some twenty miles or more northwest of Fort Collins. I did not meet with it again until the present summer, when, on July 13th, I found two of the leaf pockets characteristic of this species in Estes Park on twigs of Populus tremuloides about six miles apart and at an altitude of about 7500 feet. On August 9th, four more of the galls were taken near the Half-way House on Pike's Peak, at an altitude of about 9000 feet. In two of these galls there were no living lice. In each of the occupied galls taken (four), there was one large stem-female present, with a large number of her offspring in all stages of development up to the adult alate lice. There were no adult apterous lice and all that were half grown or more gave evidence that they were to get wings. In the breeding cages the alate lice began at once to deposit young with long beaks. In every case the lice were accompanied by a species of large black ant. 

  The Gall. An infested leaf becomes very much enlarged, and somewhat thickened with the edges turned in so as to make a heart shaped pocket, and the apex of the leaf is extended and turned back as shown in Plate XV, Figure 10. The color of the infested leaf is yellowish green, and lighter than the healthy foliage surrounding it.


externallink: https://www.biodiversitylibrary.org/page/11256375#page/79/mode/1up
description:
  Mordwilkoja vagabunda Walsh.

  Byrsocrypta vagabunda
  Pemphigus vagabundus
  Pemphigus oestlundi n. sp., Cockerell, Ent. News, p. 34, 1906.
  Mordivilkoja oestlundi

  The galls of this louse at the terminal buds of cotton- wood twigs have occurred in greater or less abundance in at least one limited locality near Fort Collins, Colorado, for the past fifteen or more years. The section referred to is mostly rather low, moist land, along the course of an irrigating ditch and near the river. It seems strange that the galls should not have become more generally distributed unless the alternate host is largely limited to the area mentioned. I am assuming that there is an alternate host for the reason that the lice all become winged and leave the galls rather early in the summer. Most of them are gone by August 1st here. The Galls, Figures IS, 19 and 20. When growing, the galls are as green in color as the Cottonwood leaves, and are, in fact, a transformed leaf in each case. On the inside of the green gall the main veins of the leaf are very prominent. Apparently these galls differ from others produced by Aphids by not having any opening to the exterior during their growth, but Mr. L. C. Bragg has discovered a small brown scale (Figure IS A, and 20, b) at the base of the gall, which seems to be the apex of a folded leaf, beneath which is an opening to the interior and through which the blade of a pen- knife may be passed without cutting any tissue. This opening is so narrow that the lice do not escape by it. About the time that fully matured winged lice are developed in a gall (about July 10th to loth, at Fort Collins), little star-shaped mouths (Figures IS and 19, o) appear at the apices of the more prominent lobes thru which the alate lice escape. Many of them appear on a single gall as shown in Figure 18. The galls nearly always are from terminal buds, and whether one or more of the leaves form a single gall I have no certain knowledge, but apparently it is one in each case. Large galls measure as much as SO to 95 mm. in greatest diameter, and about 50 to 60 in the greatest thickness.

  *Professor Oestlund, in his Aphididae of Minnesota, p. 22, states that Walsh's vagabundus is evidently something different from the louse that has since been known to be associated with the coxcomb gall. To be sure, September is late to take the migrants from these galls, and the measurements given by Walsh are too large for this species. But he evidently had a louse belonging to the Genus Pemphigus, as then understood, and in the Walsh-Riley paper published in Vol. I, of American Etomologist, page 107, the vagabond gall was figured, and both the winged lice and the apterous stem mothers from the galls mentioned. As Walsh at that time considered the winged lice from these galls the same as what he had described as B. vagabunda, it seems to me best to abide by his identification of his own species, and especially as we do not know any other species to which to refer his original description, which is quite inadequate for its identification anyway. I am therefore retaining the name vagabunda. 


externallink: https://www.biodiversitylibrary.org/page/11256375#page/73/mode/1up
description:
  Thecabius populiconduplifolius Cowen

  Pemphigus populiconduplifolius, Cowen, Hemiptera of Colorado, Bull. 31, Colo. Exp. Sta., p. 115, 1895. Hunter, Aphidida; of North America, p. 79, 1901. Gillette,* Jour. Economic Ent. p. 355. 1909. Jackson, Cols. Hort. Soc. Vol. 22, p. 217, 1908. Contributions No. 29, Dep. Zool. and Ent., O. S. U., p. 217, 1908. 
  Pemphigus ranunculi n. sp., Davidson, Jour. Economic Ent. p. 372, 1910. 
  Pemphigus populiconduplifolius, Davidson, Jour. Economic Ent. p. 374, 1910. 
  Pemphigus californicus, Davidson, Jour, of Economic Ent. p. 414, 1911. 
  Pemphigus populiconduplifolius . Essig, Pom. Jour. Ent. p. 699, 1912. 
  Pemphigus californicus, Essig, Pom. Jour. Ent. pp. 699, and 700, 1912. Essig, Pom. Jour. Ent. p. 827, 1912. 
  Pemphigus populiconduplifolius. Patch,* Bull. 213, Maine Exp. Sta. p. 76, 1913. 

  *These can hardly be populiconduplifolius, as the stem females were reported in both cases as being present in the colonies of developing lice, a condition which we have never found in Colorado where the types of the species were taken. Furthermore, I have a stem female from Massachusetts that was taken by Mr. L. C. Bragg, and it is readily distinguishable from any of the stem females that I have seen from Colorado by having remarkably thickened femora for all legs. The femora are very nearly twice as great in diameter as they are in the Colorado form and are of about the same length. Four winged migrants taken from Populus balsamifera (Ace. No. 47-10) in Maine by Dr. Edith M. Patch are before me, mounted in balsam. These seem to differ from Colorado examples principally by having weaker sensoria, which are also fewer in number, on the sixth joint of the antennae. I will suggest that this eastern form be known as Thecabius patchii, though it does not have the typical habit of most known examples of this genus of having the stem mother in a gall by herself.

  The above literature may be briefly summarized as follows: 

  The original description by Mr. Cowen dealt with the alate fundatrigenia in the folded leaves of the cottonwoods with the mere mention of yellow apterous individuals, all from Colorado. 

  Hunter lists this species only. 

  The writer, in 1909, recorded the species from Massachusetts. Davidson, 1910, described the alate and apterous forms taken in California from the buttercup (Ranunculus Californicus) to which he gave the name ranunculi, but which is probably populiconduplifolius, as Mr. Bragg and Mr. Asa C. Maxson have repeatedly traced this species to the buttercup in Colorado, where it seems to be perfectly at home. On page 374 of the same paper Davidson records populiconduplifolius in the folded leaves of Populus trichocarpa and mentions seeing the stem mother.

  Jackson merely quotes Cowen's original description. Davidson, finding the name ranunculi preoccupied, suggests californicus instead. Essig lists this species both as populiconduplifoliiis and californicus. Dr. Patch records several captures of this species in Maine on the leaves of Populus balsamifera and gives a figure showing the distribution of the wax glands of what she took to be the stem mother, and also an excellent figure of the antenna of the alate fundatrigenia, or summer migrant. This is a rather common but not abundant species in northern Colorado and the writer has also taken it at Wheatland, Wyoming, upon a broad-leaved cottonwood.

  The fundatrix is never found in the leaves folded along the midrib in which the other lice occur, but is always found in a narrow fold on the margin of one of the first leaves to open and upon the under side, see figures 1 and 5a. The second generation, almost as soon as born, leave the pseudo-gall of the fundatrix and travel to the tenderest little opening leaves at the tip of the twig, where they locate, several to a leaf, upon the lower or ventral surfaces where they begin to feed, causing the leaves to fold along the midrib as shown at figure 5, b. Mr. Maxson in a letter makes the following statement in regard to his observations upon the early occurrence of the second brood: 

  "The first larvae of the second generation were observed June 18th. These were traced to the young leaves at the tips of the branches where they located on the underside. These leaves began to fold along the midrib and in a few days typical P. conduplifolius galls were formed." 

  This second brood all acquire wings and leave the cottonwoods and go to the buttercups. Ranunculus sp., so far as our observations go. At the present writing, December 24th, there are several thrifty colonies in the laboratory on buttercups where they have been since the migrants were put upon the plants in July. They attack, not the roots, but the crown and leaves and stems near the ground. The buttercup seems to be a permanent food plant for this species, upon which it seems to be able to live continuously throughout the year.


externallink: https://www.biodiversitylibrary.org/page/45256502#page/113/mode/1up
description:
  Pemphigus betae Doane

  This insect is a native of the Rocky Mountain region, and is a rather general feeder upon the roots of plants during the summer season. The goosefoot family, especially sugar beets, garden beets and mangel wurtzels, are favorite food plants. Among our native plants and weeds, we have found this louse common upon the roots of Aster, goldenrod (Solidago), Iva xanthifolia, and lambsquarter, Chenopodium. While it does not seem necessary for this insect to take on a fall migratory habit, still, as near as we have been able to estimate, about one-half of the lice acquire wings during September and October and leave their summer hosts and fly to the cottonwoods. Apparently, only a small percentage ever find the desired host. In beet-growing sections it is common for these lice to fly in countless millions, like great swarms of ants, for- more than a month during the warmer portion of the day. Since the fall of 1910 we have known that this louse migrates to the cottonwoods, in large numbers in the fall to deposit the sexual males and females, but not until the past summer has it been definitely determined in what form this species appears upon the cottonwoods early in the summer. Mr. Asa C. Maxson, in charge of the experimental work in the Longmont district for the Great Western Sugar Company, who has been a collaborator with us in some plant louse studies, has succeeded the past summer in determining the form of gall produced by this louse upon the cottonwood leaves and in successfully colonizing the lice in large numbers upon the beets and in rearing from these colonies, the sexuparae. We extract from a report by Mr. Maxson upon his work as follows: "On May 4th . . . the larvae were located upon the upper side of the leaf which was light in color where the larvae were feeding, and showed a depression which, as it grew larger, entirely inclosed the insect and formed a gall on the under side of the leaf. This form of gall is produced by Pemphigus balsamiferae Williams." 

  Many observations by the writers tend to confirm the conclusions reached by Mr. Maxson whom we congratulate on the excellent life history work that he has done on this insect, which is one of the worst pests that the sugar beets have to contend against in the Rocky Mountain States. 

  It seems probable that Parker, in Journal of Economic Entomology, 1914, page 139, hit upon the correct gall for this louse, but referred to it on page 141 as by populicaulis Fitch, which produces a globular gall upon the petiole of the leaf. Furthermore, this gall occurs in the Middle and Eastern States, while P. betae seems to be strictly a Western species. 


externallink: https://www.biodiversitylibrary.org/page/56598181#page/227/mode/1up
description:
  Asiphum sacculi Gillette

  Asiphum sacculi Gillette, 1914b :65; Gillette and Palmer, 1934:217; Palmer, 1952:356.
  Pemphiglachnus kaibabensis Knowlton, 1938a :264.

  This gall-forming aphid is relatively rare in the counties of Nova Scotia studied, found only twice in two different locations in Antigonish County on aspen poplar (Fig. 56). In the Fairmont Road region one small tree was studied over a period of three summers. Each year in late June or early July this same tree was infested with this leaf-pocket gall. None of the adjacent poplars observed in the same region were so infested. Unfortunately when the region was revisited on the fourth summer, the plot had been cleared with the resultant end of the observations on this tree. Gillette (1914b:66), the original describer of this species, reports that “all the young of the fundatrix, the second generation, become winged and leave the galls”. He also reports finding the two empty galls near the half-way house, Pike’s Peak. If this species has a summer alternate host which is unknown, and which is implied in Gillette’s comments, how is it possible to find this same leaf-pocket gall on the same host in three consecutive summers, and not find it on any nearby aspens? 

  From the facts at hand it appears that the species is monophagous, and if so, would account for the observations given. The fact that Gillette found some empty galls, does not necessarily imply that they have migrated. Syrphid larvae predators have been observed in some galls opened, which by this means could dispose of the contained aphids.

  Specimens of this species were at first verified by M. E. MacGillivray as A. rosettei , her determination being based solely on alatae in balsam. Upon further comparison with Gillette’s A. sacculi (1914b:65, figs. 10-14) and figures 56 and 57 in this paper, there is little doubt that the species is A. sacculi Gillette. His description of the fundatrix and nymphs fits these specimens as does also the peculiar leaf-pocket gall. 

  The fundatrix is very large, oval and dull gray and pulverulent. The nymphs have a dark olive green abdomen and gray head and thorax, the abdomen being white tipped on five lateral tufts along the abdominal margins, and with intersegmental darker bands transversely placed across the dorsum. The wings of the alate second generation are slightly smoky; head and thorax blackish, abdomen dark olive-green. 

  Collections: In leaf-pocket pseudo-galls on Populus tremuloides Michx. (trembling aspen). ANTIGONISH COUNTY, Fairmont Road, July 4, 1950 (429); June 28, 1951 (501); June 26, 1952 (501); West River Road, July 30, 1952 (652).


externallink: https://www.biodiversitylibrary.org/item/109950#page/329/mode/1up
description:
  Pemphigus longicornis Maxson, n. sp.

  The galls are quite similar in form to those produced by Tetraneura graminis only less notched at top and less ribbed on sides. Walls thin. Length of large galls about five-eighths inch, height about one-half inch. From side to side they measure about three-sixths inch. Slightly constricted where attached to leaf. Placed on upper side of leaves of Populus deltoides.

  Type locality: College Station, TX


description:
  Pemphigus knowltoni sp. n. 

  Biology: Pemphigus knowltoni causes variably shaped pouch galls on the underside of the leaves of Populus angustifolia, typically alongside the mid-rib at the base of the lamina. The galls collected in the type locality were green in colour. The progeny of the fundatrix are all alate, and contain rostrate, i.e. exultype, embryones. From this it appears that the species is heteroecious, but the secondary host is unknown. 

  P. knowltoni lies closest morphologically to P. populitransversus Riley, to which it will key if referred to Palmer (1952). P. populitransversus causes petiolar galls on the broad-leaved Populus sargentii and P. deltoides (see “Biology” above). It is quite possible that P. knowltoni is the primary host form of an already described Pemphigus from roots of a secondary host, but it seems desirable to describe it pending opportunities for further investigation of its biology. 

  Distribution and type material The type material consists of the contents of three galls collected from Populus angustifolia at Garden City, Utah, 7.vii. 1966 (G. F. Knowlton and H.L.G.S.). The holotype is an alate migrans from this collection, and the paratypes three fundatrices and 23 alate migrants with the same data. No other collections of the species are at present known to me. The species is named for Professor George F. Knowlton of Logan, Utah, in recognition of his kindness in showing me the aphid fauna of Utah during two visits paid to Logan in 1964 and 1966. 


description:
  Pemphigus obesinymphae, new species

  [A drawing of the gall appears on page 8 of the pdf]

  Gall (Fig. 3d). The galls are formed on the petioles of late flushing leaves; most are adjacent to the leaf blade. The shape is roughly spherical, with the lips of the gall slit sometimes bulging outward. Diameters of mature galls ranged from 8 to 16 (mean 13.8). Walls of mature galls are approximately 1.5 thick. The gall slit is irregular in shape and extends along the equator of the gall for about 1/3 of the circumference, with the petiole and leaf blade at the poles. The lips remain closed along most of their length until the emergence of the adult fundatrigeniae. However, each gall has a single, round ostiole, 1.2 in diameter; this may occur anywhere along the slit. 

  Material examined: All collections of P. obesinymphae were from petiole galls on Populus fremontii Watson in Arizona and Utah, USA, and were taken between 15 July 1991 and 31 July 1992. Arizona locations include: Santa Cruz County: Sonoita, Canelo (Audubon Research Ranch), Patagonia; Cochise County: Portal (Southwest Field Station); Graham County: Thatcher; Pima County: Tucson (Molino Basin, Santa Catalina Mtns.). The Utah localities include Salt Lake County: Salt Lake City (City Creek Canyon) and Davis County: Uintah (Weber River). Some alates that emerged from galls were confined in vials, and their 1 st instar offspring were obtained. 

  The holotype is an alate fundatrigenia, containing embryos, collected from Populus fremontii by N. A. Moran at Patagonia on 13 October 1991. The paratypes are two adult fundatrices collected at Patagonia on 13 October 1991, two fundatrices collected at Patagonia on 8 September 1991 and 19 October 1991, one fundatrix collected at Portal on 16 September 1991, five emigrants collected at Patagonia on 25 August 1991, six emigrants collected at Patagonia on 13 October 1991, ten 1st instar fun datrigeniae, 14 2nd instar fundatrigeniae, and one 3rd instar fundatrigenia collected at Patagonia on 13 October 1991, and 12 1st instar exules deposited by emigrants collected at Patagonia on 25 August 1991. All the paratypes were collected from P. fremontii by N. A. Moran. The holotype and eight paratypes will be deposited in the U. S. National Museum, Beltsville, Maryland, U.S.A. An additional 16 paratypes will be deposited at the British Museum of Natural History, London and the Canadian National Museum. 

  Pemphigus obesinymphae, n. sp. is closely related to Pemphigus knowltoni Stroyan and, especially, to Pemphigus populitransversus Riley. We hypothesize that the three constitute a monophyletic species group within the genus Pemphigus. Both P. obesinymphae and P. populitransversus form galls only on Populus fremontii, whereas P. knowltoni uses only Populus angustifolia. When adult fundatrigeniae are not present, the presence of a gall ostiole and morphological features of the 1 st and 2nd instar fundatrigeniae can be used in order to distinguish P. obesinymphae and P. populitransversus in galls on Populus fremontii (Table 1). 

  [Table 1 shows that obesinymphae is known from Populus fremontii in AZ and UT and likely in NM, CO and northern Mexico, is found on petioles typically near the leaf blade, has an ostiole as well as a slit, colonizes Populus fremontii in the spring and galls late flushing leaves; that populitransversus is known from Populus fremontii throughout North America, is found near the center or elsewhere on petioles of first flush leaves, colonizes its host in fall, has no ostiole; that knowltoni is known from Populus angustifolia in UT, possibly also in ID, CO, WY, MT, galls leaf midribs, colonizes first flush leaves of its host in the fall]

  Biology: The galls of P. obesinymphae contained aphids of only two generations: the fundatrix and fundatrigeniae, as is typical of Pemphigus. No apterous fundatrigeniae were found in galls of obesinymphae, although such forms are usual in both Pemphigus spirothecae Passerini and Pemphigus monophagus Maxson, other species that are known to have long-lived galls and defenders or suspected defenders (Aoki and Kurosu, 1986, 1988). The adult fundatrigeniae possessed embryos resembling typical Pemphigus exules. Therefore, this species probably alternates between two different host-plant taxa. Despite considerable searching and attempted transfers to several suspected plants (Chenopodium and Poinsettia species), its secondary hosts are yet unknown. The host alternation shows peculiar seasonal timing: fundatrigeniae fly from galls much later in the season than in other Pemphigus, overwintering occurs on secondary hosts, and sexuparae return to P. fremontii in May of the following year, after the first leaves have expanded. Sexuparae were collected from 5-20 May at Patagonia in 1992; they produced sexual offspring of typical appearance for the genus. Hatching fundatrices were collected from the same clusters of sexuales, suggesting that the eggs hatch within a short period rather than overwintering. Hatching fundatrices then establish galls on late flushing leaves; gall establishment was observed at the Patagonia site from 15 May-l June 1992. The first fundatrigeniae began to be deposited in some galls on 10 June. This life cycle recalls that of Schlechtendalia chinensis (Bell), which also produces sexuparae in spring and which also has eggs that hatch without prolonged dormancy or overwintering (Takada, 1991). 

  P. obesinymphae exhibits four peculiar features: First, the life cycle is dramatically altered (Moran, 1993), with the result that galls occur only on the late leaves that expand later in the season as the shoots elongate. Most Pemphigus initiate galls on the early leaves; however, Bird et al. (1979) and Faith (1979) reported a dimorphism in the timing of gall initiation within P. populitransversus, with one gall type restricted to first flush leaves and a second restricted to late leaves. Second, 1 st instar nymphs of the 2nd generation attack predators (Moran, 1993) and have the legs well developed and the tergites heavily sclerotized. Defensive 1 st instar nymphs have been reported in Pemphigus dorocola (Aoki, 1978) and P. spirothecae (Aoki and Kurosu, 1986; Foster, 1990), but their tergites are not so strongly sclerotized as in P. obesinymphae. Third, galls have an ostiole, through which defenders enter and exit the gall (Moran, 1993). Ostioles are characteristic of galls of Pemphigus dorocola (Aoki, 1978) and P. spirothecae (Foster, 1990), but are not known from other American Pemphigus. Fourth, 2nd and 3rd instar nymphs of the 2nd generation are usually much swollen, and look as if they were wingless adults. Such swollen nymphs are not known from galls of any other Pemphigus species. Why they are so swollen remains to be investigated.


description:
  Pemphigus populitransversus

  Pemphigus obesinymphae, n. sp. is closely related to Pemphigus knowltoni Stroyan and, especially, to Pemphigus populitransversus Riley. We hypothesize that the three constitute a monophyletic species group within the genus Pemphigus. Both P. obesinymphae and P. populitransversus form galls only on Populus fremontii, whereas P. knowltoni uses only Populus angustifolia. When adult fundatrigeniae are not present, the presence of a gall ostiole and morphological features of the 1 st and 2nd instar fundatrigeniae can be used in order to distinguish P. obesinymphae and P. populitransversus in galls on Populus fremontii (Table 1).

  [Table 1 shows that obesinymphae is known from Populus fremontii in AZ and UT and likely in NM, CO and northern Mexico, is found on petioles typically near the leaf blade, has an ostiole as well as a slit, colonizes Populus fremontii in the spring and galls late flushing leaves; that populitransversus is known from Populus fremontii throughout North America, is found near the center or elsewhere on petioles of first flush leaves, colonizes its host in fall, has no ostiole; that knowltoni is known from Populus angustifolia in UT, possibly also in ID, CO, WY, MT, galls leaf midribs, colonizes first flush leaves of its host in the fall]


description:
  Asiphum sacculi Gillette 

  [A photo of this gall appears on page 2 of the pdf]

  This species was described from Colorado by Gillette (1914) and recorded from Utah by Knowlton and Bischoff (1939). Palmer (1952) considered Pemphiglacnus kaibabensis described from Arizona by Knowlton (1928) to be a synonym. 

  A. sacculi has been found in the foothills and mountain areas of southern Alberta since 1959. It forms a pseudogall on Populus tremuloides Michaux by causing a leaf to increase greatly in area and its sides and end to fold over (Fig. 1). In the gall are the fundatrix and her offspring with large numbers of ants. A. sacculi has also been found in Ontario on P. tremuloides (G. A. Bradley, personal communication). 


description:
  Pemphigus bursarius L. 

  [A photo of this gall appears on page 2 of the pdf]

  This species, commonly known as the lettuce root aphid, has been found in the United States (Palmer 1952), Europe (Stroyan 1964), New Zealand (Cottier 1953). North Africa, and the Middle East (Bodenheimer and Swirski 1957). In Canada it has been found on Lombardy poplars in Ontario and Quebec. P. bursarius has also been reported as an important pest of lettuce in England and the United States (Dunn 1959; Lange et al. 1957). 

  In Alberta P. bursarius was first found in 1963 on a few specimens of Populus nigra L. var. italica Muenchhausen in a garden at Lethbridge, and could become a pest of lettuce in southern Alberta. The galls, which are generally on the petiole but are occasionally found on young shoots, vary in size and shape but are purseor flask-shaped (Fig. 2). In the gall are the fundatrix and her offspring. 

  Dunn (1959) considered that synonyms of P. bursarius are Aphis bursarius L., Pemphigus glandifomis Rudow, P. lactucarius Passerini, and Eriosoma populi Mosley.


description:
  Thecabius populiconduplifolius Cowen 

  This species was first described from Populus monilifera Aiton (= P. deltoides Marsh) in Colorado (Cowen 1895). Palmer (1952) also reported it from Colorado on Populus sargentii Dode. In Canada T. populiconduplifolius has been found in New Brunswick on Populus balsamifera (M. E. MacGillivray, personal communication).

  This species has been found on Populus balsamifera L. in a farm shelterbelt and in the St. Mary's River Valley near Lethbridge, Alberta. The fundatrix causes the leaf margin to fold under to form a gall (Fig. 3). Her offspring leave the gall and feed on the underside of newly emerging leaves causing the outer edges to fold inwards toward the centre (Fig. 4). These pseudogalls vary considerably in shape. 

  Palmer (1952) reported the summer host as Ranunculus L. in Colorado and California and considered Pemphigus ranunculi Davidson and P. californicus Davidson to be synonyms. 

  Palmer indicated that T. populiconduplifolius is similar to T. affinis (Kaltenbach) and that Mordvilko considered them to be inseparable. The species are separated in her key by the number of secondary sensoria on the sixth segment of the antennae of the fundatrigenia but she indicates that affinis is a European species. In a collection of aphids initially considered to be T. populiconduplifolius, from a few trees in one small area of southern Alberta, the author found specimens with characteristics of each species. In this collection some of the specimens were not bilaterally symmetrical and would have been placed in affinis on the basis of one antenna and populiconduplifolius on the basis of the other antenna. 

  Bodenheimer and Swirski (1957) reported the secondary host of affinis to be Ranunculus. As the secondary hosts of the two species are the same and some specimens have characteristics of both species, it appears that afinis and populiconduplifolius may be synonymous or perhaps that there is hybridization between the species. 


description:
  Pemphigus obesinymphae

  The North American aphid Pemphigus populi-transversus Riley (Hemiptera: Aphididae: Pemphiginae) forms galls on the leaves of the eastern cottonwood (Populus deltoides; Salicaceae) throughout the eastern and central United States. From the 1940s to the 1990s, Dr. Robert Sokal and his associates published extensively on character and geographical variation in P. populi-transversus in the central and eastern United States (citations summarized in Sokal et al. 1991). One of their more intriguing observations was that most populations surveyed exhibited two distinct gall types, which they named ‘‘elongate’’ and ‘‘globular,’’ based on their shapes (Bird et al. 1979; Sokal and Riska 1981). The aphids forming the two types differed at several allozyme loci and in the seasonal production of sexuals, with the sexuals of the elongate-type appearing in the fall and those of the globular-type appearing in the spring (Faith 1979a,b; Rhomberg 1980; Setzer 1980). Faith (1979a,b) hypothesized that the unusual gall polymorphism could reflect these underlying differences in life cycles, suggesting a rare case of incipient allochronic speciation. 

  The relationship between P. populi-transversus and a closely related aphid species, P. obesinymphae Aoki, has been of the most confusing among North American Pemphigus species (Aoki and Moran 1994; Blackmon and Eastop 1994). Pemphigus obesinymphae was originally described in the southwestern United States on Populus fremontii (a cottonwood that is likely the sister taxon of P. deltoides; Eckenwalder 1996). Pemphigus obesinymphae was found to share many morphological and life-history characters with the globular-gall morph of P. populi-transversus, including the globular gall shape and the spring sexuals noted by Faith (1979a,c). Then, in groundwork for this study, we found that the globular morph is in actuality a distinct population of P. obesinymphae, and not a morph of P. populi-transversus. We clarify this relationship here to coordinate our study with the existing literature on P. populi-transversus. Hereafter, to limit further confusion, we will recognize the species that forms the elongate gall alone as P. populi-transversus. The species that forms the globular gall is P. obesinymphae, and is distributed across the United States on two hosts: the eastern cottonwood P. deltoides (on which it is sympatric with P. populi-transversus) and the Fremont cottonwood (on which P. populi-transversus is absent).

  This analysis is necessary because of the complex distribution of P. obesinymphae: as we have described, the species consists of two lineages, one sympatric with P. populi-transversus on P. deltoides, and one restricted to southwestern North America on Populus fremontii (Aoki and Moran 1994; Abbot and Moran 2002).

  In early spring, a first burst of growth occurs from embryonic leaves that have overwintered in the buds. These ‘‘early’’ leaves are followed by a ‘‘late’’ flush of summer leaves, most of which are newly initiated on the growing shoots. Pemphigus populi-transversus foundresses, having overwintered as eggs, form galls on the early, spring leaves. Pemphigus obesinymphae, by contrast, returns to P. deltoides after the first flush of spring leaves, and forms galls on the late flush of summer leaves. Having appeared later on trees than P. populi-transversus, it also remains longer, well into the fall after P. populi-transversus has departed for its secondary hosts (Bird et al. 1979; Fig. 1).

  As expected, there was no evidence that P. populi-transversus forms galls on Fremont cottonwoods, despite a westerly distribution on P. deltoides that overlaps with the eastern boundary of P. fremontii’s range (Fig. 2). Overall, the distribution of P. populi-transversus is in complete accord with the surveys of Sokal and associates.


description:
  Pemphigus populitransversus

  The North American aphid Pemphigus populi-transversus Riley (Hemiptera: Aphididae: Pemphiginae) forms galls on the leaves of the eastern cottonwood (Populus deltoides; Salicaceae) throughout the eastern and central United States. From the 1940s to the 1990s, Dr. Robert Sokal and his associates published extensively on character and geographical variation in P. populi-transversus in the central and eastern United States (citations summarized in Sokal et al. 1991). One of their more intriguing observations was that most populations surveyed exhibited two distinct gall types, which they named ‘‘elongate’’ and ‘‘globular,’’ based on their shapes (Bird et al. 1979; Sokal and Riska 1981). The aphids forming the two types differed at several allozyme loci and in the seasonal production of sexuals, with the sexuals of the elongate-type appearing in the fall and those of the globular-type appearing in the spring (Faith 1979a,b; Rhomberg 1980; Setzer 1980). Faith (1979a,b) hypothesized that the unusual gall polymorphism could reflect these underlying differences in life cycles, suggesting a rare case of incipient allochronic speciation. 

  The relationship between P. populi-transversus and a closely related aphid species, P. obesinymphae Aoki, has been of the most confusing among North American Pemphigus species (Aoki and Moran 1994; Blackmon and Eastop 1994). Pemphigus obesinymphae was originally described in the southwestern United States on Populus fremontii (a cottonwood that is likely the sister taxon of P. deltoides; Eckenwalder 1996). Pemphigus obesinymphae was found to share many morphological and life-history characters with the globular-gall morph of P. populi-transversus, including the globular gall shape and the spring sexuals noted by Faith (1979a,c). Then, in groundwork for this study, we found that the globular morph is in actuality a distinct population of P. obesinymphae, and not a morph of P. populi-transversus. We clarify this relationship here to coordinate our study with the existing literature on P. populi-transversus. Hereafter, to limit further confusion, we will recognize the species that forms the elongate gall alone as P. populi-transversus. The species that forms the globular gall is P. obesinymphae, and is distributed across the United States on two hosts: the eastern cottonwood P. deltoides (on which it is sympatric with P. populi-transversus) and the Fremont cottonwood (on which P. populi-transversus is absent).

  This analysis is necessary because of the complex distribution of P. obesinymphae: as we have described, the species consists of two lineages, one sympatric with P. populi-transversus on P. deltoides, and one restricted to southwestern North America on Populus fremontii (Aoki and Moran 1994; Abbot and Moran 2002).

  In early spring, a first burst of growth occurs from embryonic leaves that have overwintered in the buds. These ‘‘early’’ leaves are followed by a ‘‘late’’ flush of summer leaves, most of which are newly initiated on the growing shoots. Pemphigus populi-transversus foundresses, having overwintered as eggs, form galls on the early, spring leaves. Pemphigus obesinymphae, by contrast, returns to P. deltoides after the first flush of spring leaves, and forms galls on the late flush of summer leaves. Having appeared later on trees than P. populi-transversus, it also remains longer, well into the fall after P. populi-transversus has departed for its secondary hosts (Bird et al. 1979; Fig. 1).

  As expected, there was no evidence that P. populi-transversus forms galls on Fremont cottonwoods, despite a westerly distribution on P. deltoides that overlaps with the eastern boundary of P. fremontii’s range (Fig. 2). Overall, the distribution of P. populi-transversus is in complete accord with the surveys of Sokal and associates.


description:
  Pemphigus populitransversus [Felt's image shows the "globular" morph of P populitransversus now considered part of P obesinymphae]

  Oval petiole galls with transverse openings and near middle of leaf stem, aphid, summer


description:
  Pemphigus populitransversus

  [Drawings of the two "morphs" of P populitransversus appear on page 2 of the pdf]

  On a single, isolated tree in West Point, Georgia, Sokal discovered the simultaneous occurrence of two apparently distinct types of galls, either "globular" or "elongate" in form (see Fig. 1). The latter were more cylindrical than the former and generally were smaller. Field examination of contents of some of the galls suggested differences in the biology of the inhabitants. The elongate galls contained many alates (in addition to nymphs and one stem mother each), while the globular galls held few if any alates, but possessed a stem mother and nymphs.


description:
  Pemphigus spirothecae

  Galls on Lombardy poplar (Populus nigra var italica) formed by both twisting and swelling of the petiole

  This aphid produces galls in early spring on the petioles of Lombardy poplars and has long been known to occur on poplars in Europe (Connold 1901; Theobald 1929; Stroyan, pers. comm.). No previous record of this aphid exists in North America. The feeding activity of the stem mother causes the petiole to twist into a spiral. The spiralling continues until the stem mother is completely surrounded by petiole tissue (Fig. 1). Within this spiral the stem mother reproduces and the gall swells, producing a large cavity in which the aphid progeny develop. Unlike P. bursarius galls which mature in early summer, galls of P. spirothecae mature in late August and early September. This is primarily attributable to the monoecious life cycle of P. spirothecae which produces one migrant form, the sexupara, within the mature gall. This form corresponds to that of P. bursarius produced on the roots of alternate hosts. 

  As galls of P. spirothecae mature they change from green to pale red and a variable number of tiny pores appear along the seams (Fig. 3). Much larger openings may develop on this region of the gall, facilitating mass release. At leaf-fall, large numbers of apterae and alates are still found within the galls. Galls of P. spirothecae aphids were extremely abundant on Lombardy poplars in regions surveyed. 


description:
  Pemphigus bursarius

  Galls on Lombardy poplar (Populus nigra L. var. italica Muenchhausen) formed by petiole swelling only, no twisting

  This aphid is regarded as a major pest of lettuce in Europe (Dunn 1959; Judge 1967; Stroyan 1964) and in the United States (Lange et al. 1957; MacDaniel 1958). Little is known concerning this species in Canada although Harper (1966) found them on Lombardy poplar in Alberta and indicated that it has the potential of becoming a rather important lettuce pest in North America. Reports of attacks by this aphid on lettuce in Nova Scotia (Creelman 1969) and in Quebec (Lange and Cartier, pers. comm.) are further indicators. The primary author is currently engaged in a study of this species. 

  The galls of P. bursarius are flask-shaped (Fig. 4) and are formed on the petioles of leaves of Lombardy poplars. Quite often as many as three or four galls may be present on a single leaf petiole. In such instances the shape of the gall is modified depending on the degree of overlap (Fig. 5). Surveys of poplars conducted during 1972 and 1973 in regions on the west end of Montreal Island and in Pincourt, Ile Perrot, Que., indicate that each gall is the product of a single stem mother or fundatrix hatched in early spring from an overwintering egg. An average of 78 alates per gall were produced by 20 galls examined, and these alates escape from the gall cavity during mid-June and mid-July (by means of tiny pores on the lip). Present studies indicate that this aphid colonizes many alternate hosts, but lettuce is the only one of economic importance. 


description:
  Pemphigus populitransversus

  Galls on Populus deltoides, formed by swelling of petiole. Galls bear a transverse slit-like opening

  This aphid, which is indigenous to North America, forms galls on cottonwood poplars. The gall resembles that of P. bursarius but migrant release is by means of a transverse slit on the lip (Fig. 6). An average of 150 migrants of this aphid were produced by 20 galls. P. populitransversus has been the subject of many investigations in the United States. Jones and Gillette (1918) showed that this aphid attacked turnips in summer. More recently Wene and White (1953), Wolfenbarger (1965), Chalfant (1969), and Harding (1971) have found this aphid to be a serious pest on cabbage. MacNay (1955) recorded this species in Alberta on Populus deltoides, P. tremuloides, and P. angustifolia.


