Andricus archboldi (sexgen)

Andricus archboldi (agamic)

Andricus catalinensis (sexgen)

Andricus chapmanii (agamic)

Andricus chiricahuensis (sexgen)

Andricus coconinoensis (sexgen)

Andricus columbiensis (agamic)

Andricus cooki (sexgen)

Andricus fitzpatricki (sexgen)

Andricus fitzpatricki (agamic)

Andricus highlandensis (sexgen)

Andricus mellificus (sexgen)

Andricus menkei (agamic)

Andricus mogollonensis

Andricus nichollsi (sexgen)

Andricus schickae (agamic)

Andricus schickae (sexgen)

Andricus torreyaensis (sexgen)

Andricus williami (sexgen)

Antron lovellae (agamic)

Antron tomkursari (agamic)

Dryocosmus archboldi (sexgen)

Loxaulus virginianae (agamic)

Neuroterus alexandrae (sexgen)

Neuroterus aliceae (agamic)

Neuroterus aliceae (sexgen)

Neuroterus bussae (sexgen)

Neuroterus oblongifoliae (sexgen)

Neuroterus quaili (agamic)

Neuroterus rosieae (sexgen)

Neuroterus stonei (sexgen)

Zapatella abrahamsoni (agamic)

Zapatella brooksvillei (agamic)

Andricus archboldi Melika & Abrahamson, sp. nov.
Sexual generation

Diagnosis. No other gallers on Q. inopina and other red oaks in Florida induce a similar rounded, integral leaf gall, which is always pale yellow-green (in contrast to the darker green of the leaf) and located on the midrib. The sexual generation gall of Callirhytis quercuspunctata (Bassett, 1863) is similar but occurs on red oak species distributed further north.

Gall (Fig. 29). Integral leaf galls, always on the midrib, ovoid 4–6 mm in diameter and protruding on both sides of the leaf lamina; pale yellow and contrasting with the darker green of the leaf. Usually one larva per gall, rarely two larvae.

Biology. Both generations have been found exclusively on Lake Wales Ridge in Florida, on an endemic oak, Q. inopina. Final instar larvae overwinter in the galls and pupate in March. Asexual females begin to emerge in early April and are fly actively throughout the month, laying eggs that result in integral leaf galls also on Q. inopina. These sexual generation leaf galls grow through the autumn and mature in November. Adults of the sexual generation emerge in December. 

Distribution. USA, Florida only, Highlands Co., Lake Wales Ridge endemic; quite common at the Archbold Biological Station, Lake Placid.

Andricus archboldi Melika & Abrahamson, sp. nov.
Asexual generation

Diagnosis. Galls of the asexual generation very closely resemble the galls of C. quercusclavigera (Ashmead, 1881), and C. quercuspunctata; the former species is known to associate with Q. myrtifolia, a closely related species to Q. inopina. The galls of these three species are difficult if not impossible to distinguish. However, we never found the galls of A. archboldi on Q. myrtifolia. Furthermore, galls induced by C. quercuspunctata and C. quercusclavigera were never found on Q. inopina. Developing galls of C. quercusclavigera appear very similar to asexual galls of A. archboldi, however, at maturity the larval cells of C. quercusclavigera emerge through the bark of the gall and drop to the ground. The larval cells of A. archboldi remain inside the gall at maturity and the adults chew through the gall tissues to emerge. 

Gall (Fig. 30). Large abrupt, rounded and multilocular stem swellings encircling woody stems. The gall has a cheese-like texture when young and actively growing, but becomes woody and hard at maturity. The larval cells never emerge through the surface of the gall. The gall is 2 to 4 cm in diameter and height. 

Biology. Both generations have been found exclusively on Lake Wales Ridge in Florida, on an endemic oak, Q. inopina. Final instar larvae overwinter in the galls and pupate in March. Asexual females begin to emerge in early April and are fly actively throughout the month, laying eggs that result in integral leaf galls also on Q. inopina. These sexual generation leaf galls grow through the autumn and mature in November. Adults of the sexual generation emerge in December. 

Distribution. USA, Florida only, Highlands Co., Lake Wales Ridge endemic; quite common at the Archbold Biological Station, Lake Placid.

Andricus catalinensis Melika, Nicholls & Stone, sp. nov.

Gall (Fig. 45). A spherical succulent unilocular gall on leaves or catkins, usually at the base of either structure, sometimes with small point at end of gall, pale green sometimes with a pinkish tinge.

Biology. Only a sexual generation is known, which induces galls on Q. oblongifolia; adults emerge in April.  

Distribution. USA, Arizona, Santa Catalina Mountains

Andricus chapmanii Melika & Abrahamson, sp. nov.

Gall (Fig. 56). The unilocular gall is shaped like a tall cup or wine glass, 5–6 mm long, 2–3 mm in diameter, with delicate longitudinal surface grooves. The gall is slightly broader distal to the point of attachment to the leaf and the apex of the gall is truncate and depressed centrally. The gall is green to brown, develops on the underside of leaves, typically in clusters of 3–10 galls. 

Biology. Only an asexual generation is known, which induces galls on Q. chapmanii. The galls mature in October–November, at which stage the galls fall easily from the leaves. The larvae overwinter in the gall and the adults probably emerge in March–April. Rare. 

Distribution. USA, Florida, Highlands Co., Lake Placid, Archbold Biological Station. 

Andricus chiricahuensis Melika, Nicholls & Stone, sp. nov.

Diagnosis. Type material of this species was obtained from a mixed rearing of galls of this species and Neuroterus stonei. Hence exact details of the gall are unknown, but it is probably a small round bud gall, easily confusable with the gall induced by N. stonei (Fig. 434) and the sexual generation of the closely-related species Andricus coconinoensis (Fig. 88).

Gall. This species definitely induces bud galls on Q. gambelii, but we do not yet know the exact details of gall structure or appearance. It is most likely a globular, smooth gall, either greenish or pale brown.

Biology. Only a sexual generation is known, which induces bud galls on Q. gambelii; adults emerge in April. 

Distribution. USA, Arizona, Chiricahua Mountains.

Andricus coconinoensis Melika, Nicholls & Stone, sp. nov.

Diagnosis. Andricus coconinoensis induces a small bud gall on Q. turbinella and Q. oblongifolia (Fig. 88). Only two other described Andricus species are known to induce bud galls in the southwest of the USA: Andricus cellularius Gillette, 1892 and A. multicostatus Weld, 1952. 

Gall (Fig. 88). A small unilocular bud gall, roughly spherical, about 3 mm in diameter, with a smooth hard surface. Pale brown colour, sometimes with a slight darker brown mottled pattern. Similar to or possibly the same as the undescribed gall in Figure 20 of Weld (1960).

Biology. Only a sexual generation is known, which induces galls on Q. arizonica, Q. turbinella and Q. oblongifolia. Adults emerged in April. 

Distribution. USA, Arizona: Santa Catalina Mountains, Chiricahua Mountains, Mogollon Rim.

Andricus columbiensis Melika, Nicholls & Stone, sp. nov. 

Diagnosis. The galls of A. columbiensis on Q. garryana resemble those of A. occultatus (Weld, 1926), previously known from California, which induces galls on Q. douglasii, Q. dumosa, Q. lobata. Asexual A. opertus (Weld, 1926), also known from California, induces similar cryptic bud galls on Q. douglasii and Q. lobata.

Gall (Fig. 100). A cryptic bud gall, located within a slightly swollen terminal or lateral bud; gall contains a single larval chamber (occasionally two) at base of bud within the meristematic tissue, below the point at which the bud scales are attached.

Biology. Only an asexual generation is known, which induces cryptic bud galls on Q. garryana. Adults emerge in late autumn. 

Distribution. Canada, British Columbia, southern Vancouver Island and adjacent Gulf Islands.

Andricus cooki Melika, Nicholls & Stone, sp. nov.

Diagnosis. Catkin galls of Andricus cooki most closely resemble those of two known species from Florida (C. myrtifoliae (Beutenmueller, 1917) on Q. myrtifolia and C. quercusturnerii (Ashmead, 1881) on Q. nigra) and also Andricus fitzpatricki described herein. In A. cooki the galls are covered with dense whitish pubescence, while in A. fitzpatricki and C. myrtifoliae the clusters of catkin galls are smooth and shiny, without pubescence. 

Gall (Fig. 113). Dark green, sometimes purplish, clusters of densely pubescent, rounded unilocular galls on catkins. Clusters contain 10–20 individual galls and appear to cause shortening of the catkin. Similar to or the same as the undescribed gall in Figure 55 of Weld (1959). 

Biology. Only a sexual generation is known, which induces galls on catkins on Q. myrtifolia and Q. laurifolia. Adults emerge in late April–May. 

Distribution. USA, Florida: Lake Placid; Archbold Biological Station; Lake Manatee State Park; Spanish Ponds Wildlife Reserve, Jacksonville.

Andricus fitzpatricki Melika & Abrahamson, sp. nov.
Sexual generation

Diagnosis. In A. fitzpatricki the clusters of catkin galls are smooth and shiny, without pubescence while in A. cooki the galls are covered with dense whitish pubescence.

Gall (Fig. 139). Greenish to yellowish clusters of smooth, rounded unilocular galls, with a nipple on the top, causing reddish tints on affected catkins. 

Host oaks. Q. laurifolia and Q. myrtifolia

Biology. Alternating sexual and asexual generations are known. The sexual generation catkin galls develop on Q. myrtifolia and Q. laurifolia, while the asexual acorn galls have been found on Q. myrtifolia, Q. laurifolia and Q. inopina. Asexual galls develop in October - November, maturing in November. Adults overwintering in galls, emerge in February - March. Sexual generation catkin galls develop in early spring, adults emerge in late spring. 

Distribution. USA, Florida: Highlands Co. (Archbold Biological Station, Lake Placid), Brevard Co. (Malabar Scrub Sanctuary and Coconut Point Sanctuary), Martin Co. (Jonathan Dickinson State Park), Sarasota Co. (Myakka River State Park)

Andricus fitzpatricki Melika & Abrahamson, sp. nov.
Asexual generation

Gall (Fig. 140). Near-spherical ‘pip’ galls growing on one year-old acorns. Galls secrete honeydew when young and green. When mature, the galls became brown and fall from the affected acorn.

Biology. Alternating sexual and asexual generations are known. The sexual generation catkin galls develop on Q. myrtifolia and Q. laurifolia, while the asexual acorn galls have been found on Q. myrtifolia, Q. laurifolia and Q. inopina. Asexual galls develop in October - November, maturing in November. Adults overwintering in galls, emerge in February - March. Sexual generation catkin galls develop in early spring, adults emerge in late spring. 

Distribution. USA, Florida: Highlands Co. (Archbold Biological Station, Lake Placid), Brevard Co. (Malabar Scrub Sanctuary and Coconut Point Sanctuary), Martin Co. (Jonathan Dickinson State Park), Sarasota Co. (Myakka River State Park)

Andricus highlandensis Melika, Nicholls & Stone, sp. nov.

Diagnosis. Galls resemble those of A. quercusutriculus (Bassett, 1881); however, galls of A. highlandensis develop on Q. geminata while A. quercusutriculus is found on Q. alba, Q. chapmani and Q. stellata. Such host specificity at the level of oak section is typical of Nearctic oak gall wasps, with species galling section Virentes oaks (including Q. geminata) never attacking oaks in section Quercus s.s. (Abrahamson et al. 1998a,b, 2003) and vice versa.

Gall (Fig 158). Small rounded integral unilocular leaf gall, blister-like, 2.5–4 mm diameter (n = 50); most often develops on young and incompletely expanded leaves. Gall can be found anywhere on the leaf including the margin, petiole (often stopping leaf development) or end of leaf blade; commonly causes distortion of the leaf; green, covered in slight pale pubescence; very thin walled, with a spacious internal airspace in which the free-feeding larva develops. Old galls can persist on the leaves through the whole year. 

Biology. Only a sexual generation is known, which induces galls on Q. geminata. The galls begin to develop in early April on unfolded young leaves. They mature very quickly and the adults beginning to emerge from late April–May. 

Distribution. USA, Florida, Highlands Co. (Lake Placid, Archbold Biological Station), Martin Co. (Jonathan Dickinson State Park).

Andricus mellificus Nicholls, Stone & Melika, sp. nov.

Diagnosis. The gall is very distinctive; no other species from Arizona are known to induce such a specifically shaped and coloured gall (Fig. 171).

Gall (Fig. 171). A small unilocular bud gall, reaching 4–5 mm in diameter, green at first then becoming red- purple, in an axillary bud. Gall ovoid with a flattened top and a central nipple; nectar-secreting. Also illustrated in Figure 24 of Weld (1960). 

Biology. Only females are known, which induce small bud galls on Q. arizonica; Weld (1960) also lists Q. rugosa as a host. Galls mature in April, and adults emerge soon afterwards. Although only females are known, the spring phenology of this gall strongly implies that this is a sexual generation and males remain to be discovered. This species was included under the species code spAZb7 in the Nicholls et al. (2017) study of nectar secretion in oak gall wasps. This gall was collected at several sites in Arizona, with DNA data used to confirm the morphological assessment that emerged individuals were conspecific. 

Distribution. USA, Arizona: Chiricahua Mountains, Sonoita

Andricus menkei Melika & Abrahamson, sp. nov.

Diagnosis. The galls resemble those of Callirhytis quercusgemmaria (Ashmead, 1885). However, the small ribbed galls of C. quercusgemmaria are in clusters surrounding twigs, green and nectar-secreting when young, later turning grey-green and dehiscing. In contrast, galls of Andricus menkei are solitary and are not known to secrete nectar.

Galls. Unilocular stem galls, usually solitary or rarely in pairs (very rarely threes) that burst through the bark as they develop. The gall surface is marked with parallel longitudinal ribs. When attacked by parasitoids and inquilines the galls do not dehisce but enlarge, become woody and remain on tree over the winter. 

Biology. Only the asexual generation is known, which induces dehiscent bud galls on Q. pumila. The larva overwinters in the gall; adults emerge in March. 

Distribution. Florida, Dade-Metropolitan Co., Deering Estate Pinelands, Coral Gables.

Andricus mogollonensis Melika, Nicholls & Stone, sp. nov.

Diagnosis. The gall resembles sexual generation pip galls induced by the sexual generations of Disholcaspis Dalla Torre & Kieffer, 1910 and Acraspis Mayr, 1881. No other Andricus species from Arizona is known to induce such a gall. In A. mogollonensis the tarsal claws are simple, thus it is Callirhytis sensu Burks (1979). No species of Callirhytis are known from AZ, NM which induce bud galls on Q. gambelii or other white oaks.

Gall. (Fig. 193). A tiny chestnut brown, unilocular pip gall in axillary buds. The gall is a smooth thin-walled ‘pip’ gall, comprising only the larval chamber, reaching 3–4 mm long by 2 mm in diameter, with a blunt rounded apex.

Biology. Only females are known, whose morphology most closely resembles asexual generation females (genae are broadened behind the eye, visible in front view). However, this character alone is inadequate to confirm that this is an asexual generation. Mature galls were collected in April from Q. gambelii; adults emerged soon afterwards. This spring developmental phenology is more compatible with a sexual generation. In addition, the gall morphology of a bare larval chamber on a bud is more typical of a sexual generation gall. 

Distribution. USA, Arizona: Chiricahua Mountains, Mogollon Rim

Andricus nichollsi Melika & Stone, sp. nov.

Diagnosis. The gall somewhat resembles the sexual gall of Andricus kingi Bassett, 1900 but it is larger and greenish. 

Gall. (Fig. 204). A small pointed unilocular leaf gall, 4–5 mm tall, 2 mm across, on edge of leaf, which causes the leaf to curve inwards at the point of gall attachment. Gall pale green with scattered brown flecks and covered with a dense coating of short hairs. Some of the hairs and the gall tip become red and then brown as the gall matures.

Biology. Only females are known. Their morphology strongly suggests that they belong to the sexual generation rather than the asexual generation. This is also suggested by the gall’s rapid development in spring. In addition, very similar galls with comparable morphology (small, unilocular, roughly conical, developing on the edge of leaves) are known from the sexual generations of some Nearctic Andricus species (see Doutt 1960, Dailey & Sprenger 1973a,b). Galls develop on Q. oblongifolia, mature in April; adults emerge soon afterwards. 

Distribution. USA, Arizona, Santa Catalina Mountains

Andricus schickae Nicholls, Melika & Stone, sp. nov.
Asexual generation

Diagnosis. Galls closely resemble those of Andricus occultatus, but the adults are different. Also closely resembles A. columbiensis. 

Gall. (Fig. 232). A cryptic unilocular gall within a swollen terminal or lateral bud, whose scales remain intact and surround the gall. The gall is ca. 2–2.5 mm long and 1–1.5 mm in diameter, and consists only of the larval chamber, with a thin woody wall. 

Biology. Alternate asexual and sexual generations are known. The asexual generation bud galls develop on Q. garryana, mature in November, adults emerge the following spring. The sexual generation galls (Figs. 233–234) were found on Q. lobata leaves in late March-April; adults emerge soon afterwards. We expect both generations can be found on both Q. lobata and Q. garryana. 

Distribution. USA, California (Los Molinos, Stockton, Cosumnes River Preserve) and Canada, British Columbia (Victoria).

Andricus schickae Nicholls, Melika & Stone, sp. nov.
Sexual generation

Diagnosis. No other sexual generations of Andricus species from California are known to induce unilocular integral leaf galls that are simply swellings of the petiole/midrib.

Gall. (Figs. 233–234). A unilocular integral gall in the leaf midrib or petiole, 3–4 mm long by ca. 2 mm across, visible as a slight longitudinal swelling that is a paler yellow-green than the rest of the leaf, sometimes also reddish. The gall causes distortion of the leaf lamina, in some cases bending it back through almost 180°.

Biology. Alternate asexual and sexual generations are known. The asexual generation bud galls develop on Q. garryana, mature in November, adults emerge the following spring. The sexual generation galls (Figs. 233–234) were found on Q. lobata leaves in late March-April; adults emerge soon afterwards. We expect both generations can be found on both Q. lobata and Q. garryana. 

Distribution. USA, California (Los Molinos, Stockton, Cosumnes River Preserve) and Canada, British Columbia (Victoria).

Andricus torreyaensis Melika & Abrahamson, sp. nov.

Diagnosis. The gall resembles that of Zapatella quercusmedullae (Ashmead, 1885), but the morphology of adults is very different from that of Zapatella species. This species was also compared with species of the newly erected genus Melikaiella, and again the morphology of adults differs from that of Melikaiella species. Three Callirhytis species, C. quercusscitula (Bassett, 1864), sexual generation only; C. seminosa (Bassett, 1890), asexual generation only; C. quercuspunctata (Bassett, 1863), asexual generation only are known to induce similar stem swelling galls, however all three species are found further north and are unknown in Florida.

Gall. (Figs. 252–255). A shoot swelling just behind the bud. The galls are multilocular, usually terminal, cylindrical swellings, 5– 9 mm in diameter and 10–22 mm in length, hard and woody when mature. The leaf buds on the gall can continue to grow after the gall has begun to develop, and ultimately cover the gall with leaves. Usually two to three neighboring twigs or buds are attacked, creating small clusters of two or three, sometimes four galls. The shape of the swelling differs slightly on different oak hosts. 

Biology. Only a sexual generation is known, which induces stem swelling galls on Q. incana, Q. inopina, Q. laevis, Q. laurifolia, Q. myrtifolia, and Q. nigra. Galls mature in April-May, and adults emerge soon afterwards.  

Distribution. USA, Florida: Torreya State Park, Archbold Biological Station, Jacksonville, Lake Manatee State Park.

Andricus williami Melika, Nicholls & Stone, sp. nov.

Gall. (Figs. 273–274). An acorn gall, developing as a swelling external to the cotyledons around the base of the acorn inside the acorn cup (see also Weld 1960, Fig. 18). The larval cell intrudes into the body of the acorn; adults emerge through holes they create along the rim of the acorn cup. The gall is unilocular and consists only of the larval chamber ca 3 mm long and 1.5–2 mm in diameter, surrounded by a hard woody wall. Several galls may develop around the same acorn. 

Biology. Only a sexual generation is known, which induces acorn galls on Q. arizonica. Weld (1960) also records this gall type from Q. gambelii, Q. rugosa and Q. turbinella. Galls mature in April, adults emerge soon afterwards.

Distribution. USA, Arizona, Chiricahua Mountains. 

Comments. All known Callirhytis and Andricus species that induce acorn galls, whether pip galls (external to the acorn cup) or stone galls (within the acorn cup and/or also inside the cotyledons of the acorn), have their asexual generations making such galls. The only exception, other than A. williami, is the sexual generation of Andricus chrysobalani Weld, 1926 which develops on Q. chrysolepis and is known only from California.

Antron lovellae Melika, Nicholls & Stone, sp. nov.

Diagnosis. The gall is somewhat similar to the asexual generation gall of the California species Cynips (Antron) douglasii (Ashmead, 1896) but rounder and with fewer spines.

Gall. (Fig. 286). A small (up to 8 mm in height and diameter) unilocular leaf gall on underside of leaves, with a reddish-brown surface covered with grey pubescence. Roughly conical with short stubby spines protruding from near the base of the gall. Also pictured in Fig. 163 of Weld (1960).

Biology. Only an asexual generation is known, which induces leaf galls on Q. turbinella. Galls mature in October-November, adults emerge soon afterwards. 

Distribution. USA, Arizona, Flagstaff

Antron tomkursari Melika, Nicholls & Stone, sp. nov.

Diagnosis. Molecular data cluster this species with A. acraspiformis (Weld, 1926), A. plumbeum (Weld, 1926) and A. quercusnubila (Bassett, 1881), all three of which occur in the US southwest and induce leaf galls in their asexual generations. 

Gall. (Fig. 288). Small, round, dehiscent and unilocular leaf gall, on the underside of the leaf, to 4 mm in diameter, pale green with widely scattered pink spots. The inner larval chamber is surrounded by solid parenchyma tissue and not suspended on fine filaments.

Biology. Only an asexual generation is known, which induces leaf galls on Q. arizonica. Galls mature in April- May, adults emerge soon afterwards. 

Distribution. USA, Arizona: Strawberry (on Mogollon Rim), Chiricahua Mountains

Dryocosmus archboldi Melika & Abrahamson, sp. nov.

Diagnosis. The gall most closely resembles that of D. quercuslaurifoliae (Ashmead, 1881), D. quercusnotha (Osten Sacken, 1870) and D. quercuspalustris (Osten Sacken, 1861). In D. archboldi, the gall narrows to a pointed tip (Fig. 314), while in D. quercuslaurifoliae, D. quercusnotha and D. quercuspalustris the galls are always rounded, never with a pointed tip (Figs. 315–316).

Gall. (Fig. 314). An integral, succulent unilocular leaf gall, reaching 5 mm in diameter and height. Conical in shape, narrowing to a point; on underside of leaf; green with paler longitudinal stripes. The larval cell becomes free-rolling once the gall matures. Very rarely the galls develop on catkins.

Biology. Only a sexual generation is known, which induces integral leaf galls on Q. myrtifolia and Q. laurifolia. The galls develop very quickly, appearing in the second half of March and producing adults by the end of March into April. 

Distribution. USA, Florida:Archbold Biological Station, Lake Placid, Josephine Creek, Highlands Co.; Coconut Point Sanctuary, Brevard Co.; Gratin Beach Island, Walton Co.; Spanish Ponds Wildlife Reserve, Jacksonville.

Loxaulus virginianae Melika & buss, sp. nov.

Diagnosis. A key to all 14 Loxaulus species known from America north of Mexico was published by Melika & Abrahamson (2000). There are two species of Loxaulus, L. pattersoni (Kinsey, 1922) and L. virginianae, which are associated with Q. virginiana. Loxaulus pattersoni, known from Texas, induces galls on roots of Q. virginiana.

Gall. (Fig. 328). Galls are tiny cryptic cells hidden under the bark of twigs, with little or no visible swelling of the twig. The galls are 2 mm long and consist only of the larval chamber. The galls are commonly induced in a longitudinal cluster and are most easily detected by the emergence holes in the bark. 

Biology. Only an asexual generation is known, which induces stem galls in twigs of Q. virginiana. Galls mature in May, adults emerge soon afterwards. 

Distribution. USA, Alabama, Grand Bay

Neuroterus alexandrae Nicholls & Melika, sp. nov.

Diagnosis. Neuroterus alexandrae belongs to Kinsey’s subgenus Neospathegaster. It is one of only two Neuroterus known from the southwestern US that induce round bud galls. Closely resembles N. stonei.

Gall. (Figs. 342–343). Spherical, unilocular bud galls, 3–4 mm in diameter, on terminal or axillary buds. Gall surface smooth and shiny, initially green, sometimes developing mottled or striped patterning in red or purple. Possibly the same as the undescribed gall pictured in Fig. 25 of Weld (1960).

Biology. Only a sexual generation is known, which induces bud galls on Q. turbinella. Galls mature in April– May, adults emerge soon afterwards. 

Distribution. USA, Arizona, Mogollon Rim area

Neuroterus aliceae Melika, Nicholls & Stone, sp. nov. 
Asexual generation

[Photo on page 100 of the pdf]

Diagnosis. Neuroterus alicae belongs to Kinsey’s subgenus Diplobius. The asexual generation of Neuroterus aliceae induces dehiscent leaf galls on Q. oblongifolia. The only previously known Neuroterus species from the US southwest to induce detachable leaf galls, N. argentatus Weld, 1944 asexual generation, induces small, flat, rounded spangle galls on the underside of the leaves of Q. gambelii, Q. oblongifolia and Q. turbinella, which are obviously different from the aggregated brown fluffy leaf galls induced by N. aliceae. 

Gall. (Fig. 367). Brown fluffy leaf galls, consisting of an aggregated group of larval chambers ≥ 5 mm across, on midrib on the underside of leaf. Covered in a dense, somewhat woolly, coat of reddish-brown hairs. Similar in structure to Andricus pattoni (Bassett, 1881) from Florida.

Biology. Alternate asexual and sexual generations are known, both of which induce galls on Q. oblongifolia. Sexual catkin galls mature in April, adults emerge soon afterwards. The asexual fluffy leaf galls mature in October- November; adults overwinter in the galls and emerge the following spring. Distribution. USA, Arizona, Santa Catalina Mountains

Neuroterus aliceae Melika, Nicholls & Stone, sp. nov. 
Sexual generation

Diagnosis. The sexual generation of N. aliceae induces catkin galls on Q. oblongifolia. The only known Neuroterus species from the US southwest which induces flower galls (also on white oaks) is N. floricomus Weld, 1957, sexual generation. The galls of N. floricomus are dense wooly enlargements of the catkin axis (Weld 1957b) and are clearly different from those of N. aliceae (Fig. 366).

Gall. (Fig. 366). A whitish swelling surrounding individual flowers, with sparse very short pubescence. A swollen flower is 1–2 mm in diameter; multiple flowers per catkin are galled.

Biology. Alternate asexual and sexual generations are known, both of which induce galls on Q. oblongifolia. Sexual catkin galls mature in April, adults emerge soon afterwards. The asexual fluffy leaf galls mature in October- November; adults overwinter in the galls and emerge the following spring. Distribution. USA, Arizona, Santa Catalina Mountains

Neuroterus bussae Melika & Nicholls, sp. nov. 

Diagnosis. Neuroterus bussae belongs to Kinsey’s subgenus Neuroterus. Only one Neuroterus species north of Mexico, the sexual generation of N. christi Melika & Abrahamson, 1997, is known to induce integral leaf galls on Q. virginiana and Q. geminata. The galls of N. christi are multilocular and integral thickenings of the leaf parenchyma that protrude equally on both sides of the leaf along one side of the midrib, generally with four to six larval cells per gall (Melika & Abrahamson 1997a). In contrast, galls of N. bussae are small, unilocular blister-like leaf galls, that commonly develop all over the leaf lamina.

A taxon with a similar gall was described from Texas on Q. virginiana, N. niger var. alimas Kinsey, 1923, which is one of the five varieties of N. niger Gillette, 1888 described by Kinsey (1923) that were still considered forms of that species by Burks (1979). When the galls of this new species were first encountered, we initially considered that it could be N. niger var. alimas. The holotype of that species was deposited at the USNM and labelled as “Acc.24856”, pink ”N. alimas, female, Holo- Paratype’, ”Q. virginiana”, ”Austin, Tex. 4.16.21., Patterson”; however, when inspected the insect was absent and only the gall was found. One paratype is present, but in a very poor condition, and being male is hard to determine to species. According to Kinsey (1923) this variety induces a sexual generation spring gall, while the other four varieties induce autumn asexual generation galls. Moreover, N. niger var. alimas is the only variety associated with section Virentes oaks while the other four are on Quercus oaks, and as mentioned above no Nearctic cynipid species are shared across the oak sections (Abrahamson et al. 1998a, 2003; Stone et al. 2009). Hence biological information suggests that N. niger var. alimas is a species distinct from N. niger and the other four of Kinsey’s varieties. Kinsey (1923) described N. niger var. alimas based on one female (holotype—lost) and 13 males and gave no detailed morphological description for this variety. Thus, there is no morphological evidence for synonymisation of N. niger var. alimas to N. niger and at the same time it cannot be treated as a valid species so we move it to nomen dubium

Gall. (Figs. 379–380). Blister-like small integral parenchyma galls, visible on both sides of the leaf; on new leaves. Emergence holes typically on upper surface of leaf. 

Biology. Only a sexual generation is known, which induces integral leaf galls on Q. virginiana. Galls mature in late March-April, adults emerged immediately after the galls were collected, in March-April. 

 Distribution. USA, Florida: Morriston, Gainesville, Jacksonville.

Neuroterus oblongifoliae Nicholls, Stone & Melika, sp. nov.

Diagnosis. Neuroterus oblongifoliae belongs to Kinsey’s subgenus Diplobius. One other species, Neuroterus lamellae Weld, 1957, is known from Arizona that induces galls in the form of integral leaf parenchyma thickenings (Weld 1957b).

Gall. (Fig. 393). An integral leaf gall visible as a 2–3 mm thickening of the leaf lamina, sometimes spanning the whole width of the leaf, more usually on one side of midrib, green underneath, pink or red on upper surface. Multilocular. 

Biology. Only a sexual generation is known, which induces leaf galls on Q. oblongifolia. Galls mature in April, adults emerge soon afterwards. 

Distribution. USA, Arizona, Santa Catalina Mountains.

Neuroterus quaili Melika, Nicholls & Stone, sp. nov.

Diagnosis. Neuroterus quaili belongs to Kinsey’s subgenus Neospathegaster. Two other Neuroterus species are known to induce catkin galls on section Quercus oaks in California: N. floricola Kinsey, 1922 and N. florulentus Weld, 1957. Both induce different shaped and coloured galls to N. quaili: golden yellow-brown with an apical point in N. floricola, thin-walled, tan and ovate in N. florulentus. In contrast the gall of N. quaili is spherical, purplish, with quite a robust wall. 

Gall. (Fig. 405). A monolocular catkin gall, 2–3 mm diameter, spherical, purple with paler mottling, on flowers rather than on the catkin stem, relatively robust wall around larval cell. Probably the same as the undescribed gall in Fig. 40 of Weld (1957a). 

Biology. Galls develop on Q. berberidifolia, mature in April; adults emerge soon afterwards. Only females were reared, with males as yet unknown; however, the spring phenology of this gall suggests that it is a sexual generation.

Distribution. USA, California, Quail Ridge Reserve.

Neuroterus rosieae Melika, Nicholls & Stone, sp. nov.

Gall. (Figs. 420–421). A small monolocular bud gall, pear-shaped with a spherical body that is drawn out to an apical point, greenish to purple, 3 mm across, 5 mm tall. The gall bears tiny bracts on its surface and is covered in dense, short, pale pubescence. 

Biology. Only a sexual generation is known, which induces bud galls on Q. arizonica and Q. turbinella. Galls mature in April, adults emerge soon afterwards. 

Distribution. USA, Arizona, along Mogollon Rim in area south of Flagstaff

Neuroterus stonei Melika & Nicholls, sp. nov.

Diagnosis. Neuroterus stonei belongs to Kinsey’s subgenus Diplobius. The only other Neuroterus species that is known from the southwestern USA that induces bud galls in its sexual generation is N. alexandrae, although galls of that species have only been found to date on Q. tubinella while N. stonei galls the oaks Q. gambelii and Q. arizonica.

Gall. (Fig. 434). A small unilocular bud gall, spherical, reaching 4–5 mm diameter, that develop in terminal and lateral buds. The lower quarter to third of gall has a few bud scales around it. Gall is initially pale green with some darker green markings, becoming pinkish with darker red or purple lines as it matures. Similar in structure to the sexual generation of N. alexandrae on Q. turbinella. Possibly the same as the undescribed gall pictured in Fig. 22 of Weld (1960).

Biology. Only a sexual generation is known, which induces bud galls on Q. arizonica and Q. gambelii. 

 Distribution. USA, Arizona: Chiricahua Mountains, Mogollon Rim area south of Flagstaff.

Zapatella abrahamsoni Melika, sp. nov.

Gall. (Fig. 445). The galls are rounded, irregular woody expansions of lateral buds, 5.0–7.5 mm in diameter, 4–6 mm high. The tissue of the gall is hard, typically with 2–4 larval cells (very rarely only one) embedded in the surrounding woody tissues. The walls of the larval chambers are distinct, slightly paler than the surrounding woody tissue. The galls persist on the trees for years

Biology. Only an asexual generation is known, which induces galls on Q. inopina, an oak species endemic to the Lake Wales Ridge. The galls become visible in late September and develop through November. The larvae (sometimes pupae) overwinter in the galls. Adult wasps were dissected from galls from the end of January through March; emerge in late April-May. 

Distribution. USA, Florida, Highlands Co., Lake Placid, Archbold Biological Station. This species is likely endemic to south-central Florida and particularly to the Lake Wales Ridge.

Zapatella brooksvillei Melika & Abrahamson, sp. nov.

Gall. Tiny enlargements at the base of young twigs and branches, multilocular with up to 7–8 cells in one gall; sometimes only a barely visible swelling indicates the presence of the gall. 

Biology. Only an asexual generation is known, which induces twig galls on Q. laevis. Mature galls, containing adult wasps, were collected in January-February; adults emerge in March. 

Distribution. USA, Florida, Hernando Co. (Brooksville Ridge) and Highlands Co., Lake Placid (Archbold Biological Station).

New species of Nearctic oak gall wasps (Hymenoptera: Cynipidae, Cynipini)