Neuroterus archboldi (sexgen)

Neuroterus chapmanii (sexgen)

Neuroterus christi (sexgen)

Neuroterus weldi (agamic)

Neuroterus quercusbatatus (agamic)

Neuroterus quercusirregularis (sexgen)

Neuroterus quercusminutissimus (agamic)

Neuroterus quercusverrucarum (agamic)

Neuroterus quercusbatatus (sexgen)

Neuroterus pallidus (sexgen)

Neuroterus floccosus (agamic)

Neuroterus archboldi Melika and Abrahamson, NEW SPECIES

Distribution.-Florida (Archbold Biological  Station, Lake Wales Ridge, Highlands Co.; Jonathan Dickinson State Park, Martin Co.). 

Biology.-Only the bisexual  generation is known, and  the  only  known host for N. archboldi is Quercus chapmanii. This species induces  blister-like  parenchyma  thickenings  which  are  visible on both  sides of the leaf but protrude  more on the underside of the leaf. The gall is rounded, monothalamous, 1.5-2.0 mm in diameter, 1.0-1.5 mm high,  green or whitish green,  very slightly lighter than the leaf,  and drier than the succulent polythalamous N. quercusirregularis galls. The larval cell is situated on the  underside of the leaf, not separated from the outer layers of the gall. There are 18 to 25 or more galls per leaf. After adults emerge,  the galls shrink and  wrinkle  and dry out  and  drop. Consequently,  rounded holes remain  in the leaf. We observed these galls  actively  growing for the  first time on 9  March 1995. Adults emerged the second and third week of March into April.  They were very  common leaf  galls on Q. chapmanii at the Archbold Biological Station. Simultaneous with this  species, N. quercusirregularis galls could develop on the same leaf,   but   they are easily   distinguishable from  those of the former on the  basis of galls. Our  efforts to rear the  alternate generation in  cages containing emerged wasps on Chapman oaks were  unsuccessful. 

Neuroterus chapmanii Melika and Abrahamson, NEW SPECIES
Distribution.-Florida    (Archbold Biological  Station,  Lake Placid,  Highlands Co.; Jonathan  Dickinson State  Park, Martin Co.). 
Biology.-Only the bisexual  generation is  known. Galls are on catkins of Quercus chapmanii, are ovoid, thin walled, up to 1.5 mm long,  and with a  groove  running from one  side to the  other  across  the  upper  surface thus resembling a  closed  purse. The  gall is covered  with  white  pubescence, especially on the top, and is monothalamous.  Galls are randomly scattered along the staminate axis and perpendicular to it. Sometimes two galls develop together. The galls  are surrounded by anthers. As the gall  and catkins mature,  the  color changes from  green  to  pale  brown.  They  remain  on the catkins until emergence of the adult or longer, dropping along  with  the  catkins.  In Florida, on the Lake Wales Ridge, Chapman oak begins to flower the  end of February to early March. We found the gall for the first time on 3 March 1995 when some of the  galls had emergence holes, likely made by the  gall-inducing wasps; also, pupae and adult wasps were cut from the galls on this date. 

Neuroterus  christi Melika and Abrahamson, NEW SPECIES

Diagnosis.--The galls are also  quite  similar  in  appearance  to  those of N. quercusirregularis, but those of N. christi are  strictly  associated with Q. geminata and Q. virginiana Mill., not  nearly so succulent,  and  usually of a  regular  cylindrical shape and not so irregularly  shaped as those of N. quercusirregularis.

Distribution.-FL

Biology.-Only the  bisexual  generation is known. The host oaks are Quercus geminata and Q. virginiana. The galls are leaf parenchyma thickenings,  equally  protruding on both sides of the leaf along one side of the main  vein, green when  young but turning brownish green when  mature,  polythalamous, generally with four to six larval  cells   per gall,  about 12.0-15.0  mm long,  3.0-4.5 mm wide, and 3.0-4.5 mm high, and  usually of a  regular  cylindrical form. The walls of the inner cells are whitish green and  the  surface of the gall is smooth, without hairs. There  are usually one or  two  galls  per  leaf.  Adults usually emerge from  the upper  side. The first  galls appear in mid-March, and the adults  emerge the last week of May and beginning of June. 

Neuroterus weldi Melika and Abrahamson, NEW SPECIES
Comments.-Weld (1959)  wrote of a stem  swelling found on Q. chapmanii, but he never reared the adults and consequently did not describe the species. The galls induced by N. weldi are probably those Weld described.
Distribution.-Central and coastal sand ridges  of  south  central Florida (Archbold Biological  Station, Lake Placid,  Highlands Co.;  Avon  Park  Air  Force  Range, Highlands Co.;  Jonathan Dickinson State  Park, Martin Co.). 
Biology.-Only the unisexual generation is known. The gall is cylindrical,  mostly  a terminal stem swelling, covered  with normal bark, on second year or older twigs and branches  only of Quercus chapmanii. It is 25.0-40.0 mm long, 6.0-9.0 mm in diameter, and polythalamous. Galls are not  woody,  rather  they are easily cut like cheese. The leaf petioles are never involved in gall formation. Galls persist on stems for several years.  Fully  developed  adults  over- winter in galls and emerge from the second half  of February  into March of the following spring. The first galls observed are formed by the end  of May.

Neuroterus quercusbatatus (Fitch) 

Cynips Quercus-batatus Fitch 1859: 810. Females and males (type examined). 
Neuroterus batata Basset: Ashmead 1885a: 296, 303. 
Neuroterus  quercusbatatus: Burks 1979: 1074. 
Cynips noxiosus Bassett 1881: 108. Females,  males, galls of both  generations (types examined). New synonymy. 

Distribution.-Ontario, Rhode  Island, Connecticut  west to Illinois, Colorado, south to  Florida. 

Biology.-Alternate bisexual  and unisexual generations  are known. Both  generations  induce  stem-swelling  galls  on Quercus alba L. (Ashmead 1885a, Burks 1979). The  gall is a  woody,  elongate stem swelling. It is polythalamous,  with an irregular  shape and a surface that is covered by normal  bark,  twisted,  but  in  large  part cylindrical, tapering  gradually  to  the  stem at  both ends and up to 60 mm long by 20 mm wide in the unisexual form. Sometimes several  galls  can  more or less fuse, drying brown in bisexual forms, or with a  whitish or purplish bloom in unisexual forms [an apparent mistake, copied from Kinsey; these should the other way around]. Internally hard and woody, the tissue is  little modified except by the larval cells which  are  densely  packed,  each  with a distinct but wholly inseparable lining. The unisexual form develops on  young stems, involving  petioles  and  leaf midveins. The unisexual form begins to develop in mid-summer, forming woody stem galls. The adults overwinter in the galls and emerge in spring after the oaks are  actively  growing. 

Neuroterus quercusirregularis (Osten Sacken) 

Cynips q. irregularis Osten Sacken 1861: 65. Species  described from one damaged male (sex not  certain) and one gall. Type lost. 
Neuroterus   irregularis: Ashmead 1885a: 296,  304. 
Neuroterus quercusirregularis: Burks  1979: 1074. 
Cynips quercus-majalis Bassett  1864: 683. Females,  males, galls (types examined). New synonymy. 
Neuroterus  majalis: Mayr  188 1 : 37. 
Neuroterus (Dolichostrophus) irregularis var. majalis: Kinsey 1923: 100

Distribution.-Ontario, New  York, Massachusetts, Connecticut, New Jersey, Maryland, Virginia,  west to Illinois,  Missouri, Oklahoma, Texas, and southward (Beutenmueller   1910). We found this  species  in North Carolina (on Q. stellata), and in Florida (common,  locally  abundant, occurs  everywhere  the host plants  grow).

Biology.-Only the  bisexual  generation is known  to induce integral  leaf  galls  on Quercus alba and Q. montana Willd. (Ashmead 1885a, Beutenmueller 19 10, Kinsey 1923, Weld 1959), and Q. chapmanii and Q. stellata (Burks 1979). We frequently found this species on Q. margaretta as well. According to Kinsey (1923), N. quercusirregularis var. albipleurae Kinsey induces galls  on Q. breviloba (in  Texas only). 

The gall is a leaf parenchyma thickening, with the larval cells mostly deeply embedded, and is polythalamous. It  is usually elongate, oval, as  thick as wide,  up  to 5.0 mm wide by 15.0 mm long, several galls often fusing, green, very succulent,  shriveling considerably  upon drying, and  the larval cell has a distinct  but inseparable layer. The galls are smooth and large  when on Q. alba and the pubescence is denser and the galls are smaller when on Q. stellata, Q. margaretta, or Q. chapmanii. Galls usually  appear on very  young, unfolding leaves, very  quickly mature,  and develop from April until  early June, earlier farther south. Adults emerge from May through July.

Neuroterus  quercusminutissimus (Ashmead)

Cynips q. minutissima Ashmead 1885a: 7. 
Neuroterus minutissimus: Ashmead 1885b: 296. 
Neuroterus (Diplobius) verrucarum var. minutissimus: Kinsey  1923: 72.

Distribution.-Florida. Common everywhere Quercus geminata, Q. minima, and or Q. virginiana grows. 

Biology.--Only the unisexual generation is known. This species induces small, 1.25-2.50 mm, detachable galls,  covered with coarse, light  brown mossy pubescence, on the underside of leaves in numbers  in  the fall on  Quercus  geminata, Q. virginiana, and Q. minima. In Florida, adults  emerge early in the spring, but  the galls do not  develop  until  mid- to late summer. The fully grown adults overwinter in the galls

Neuroterus quercusverrucarum (Osten Sacken) 

Cynips quercus  verrucarum Osten Sacken 1861: 62. Females  and  galls  (types examined). 
Cynips verrucarum: Osten Sacken 1865: 354. 
Neuroterus  verrucarum: Ashmead 1885a: 296,  304. 
Neuroterus   quercusverrucarum: Burks 1979:  1075. 
Neuroterus exiguissimus Bassett  1900: 332. Females and galls (types examined). New synonymy. 
Cynips flocossa Bassett 1881 : 111. Females and galls  (types examined). New synonymy. 
Neuroterus  verrucarum var. pernotus Kinsey  1923: 74. 
Neuroterus flocossus:  Burks  1979:  1073

Biology.--Only  the unisexual  generation is  known. This species induces detachable leaf galls which are small, smooth, seed- like larval cells, hard, thin-walled, and covered with a dense flattened yellowish brown mass of wool  that dries brown. The gall is monothalamous, attached to the underside of the leaf by a small point to the leaf vein, separable,  but the leaf is depressed at the point of attachment. The average diameter of a gall is 2.0 mm by 1.0 mm high. The galls are usually numerous, covering the whole underside of the leaf and  are found on Quercus alba, Q. bicolor Willd., Q. chapmanii, Q. margaretta, and Q. stellata. Viereck (1916) also recorded Q. macrocarpa Michx. as a  host, but this  must be confirmed. Galls begin to develop in September (Weld 1959). According to Kinsey (1923), the galls  appear  after  mid-summer in August and the larvae do not mature  until late in the fall. Adults emerge the next spring in February to March.

Distribution.-Ontario, New York, New Jersey,  Connecticut,  west to Illinois, Oklahoma, Texas,  south to Florida. 

Neuroterus quercusbatatus (Fitch) 

Cynips Quercus-batatus Fitch 1859: 810. Females and males (type examined). 
Neuroterus batata Basset: Ashmead 1885a: 296, 303. 
Neuroterus batatus form bisexualis  Kinsey 1920:  334. 
Neuroterus (Dolichostrophus) batatus var. batatus form bisexualis  Kinsey  1923: 4. 
Neuroterus  quercusbatatus: Burks 1979: 1074. 
Cynips noxiosus Bassett 1881: 108. Females,  males, galls of both  generations (types examined). New synonymy. 
Neuroterus noxiosus form vernalis Kinsey 1920: 337. Bisexual generation.

Distribution.-Ontario, Rhode  Island, Connecticut  west to Illinois, Colorado, south to  Florida. 

Biology.-Alternate bisexual  and unisexual generations  are known. Both  generations  induce  stem-swelling  galls  on Quercus alba L. (Ashmead 1885a, Burks 1979). In  Florida, the galls of the bisexual  generation also develop on Q. chapmanii and Q. margaretta. The  gall is a  woody,  elongate stem swelling. It is polythalamous,  with an irregular  shape and a surface that is covered by normal  bark,  twisted,  but  in  large  part cylindrical, tapering  gradually  to  the  stem at  both ends and up to  20 mm long  and 8 mm wide in  the bisexual form. Sometimes several  galls  can  more or less fuse, drying brown in bisexual forms, or with a  whitish or purplish bloom in unisexual forms [an apparent mistake, copied from Kinsey; these should the other way around]. Internally hard and woody, the tissue is  little modified except by the larval cells which  are  densely  packed,  each  with a distinct but wholly inseparable lining. The bisexual form is on older stems involving the bases of the petioles. The galls of the bisexual  generation are less  woody, usually on younger twigs, and develop in late spring to early summer. The adults  emerge in June and July.

[no entry for this species, only mentioned in passing in the description of Neuroterus chapmanii]

N pallidus Bassett (galls are in clusters, usually on the end of catkins)

Neuroterus quercusverrucarum (Osten Sacken) [partial, synonymy reversed by Nicholls et al 2022]

Cynips quercus  verrucarum Osten Sacken 1861: 62. Females  and  galls  (types examined). 
Cynips verrucarum: Osten Sacken 1865: 354. 
Neuroterus  verrucarum: Ashmead 1885a: 296,  304. 
Neuroterus   quercusverrucarum: Burks 1979:  1075. 
Neuroterus exiguissimus Bassett  1900: 332. Females and galls (types examined). New synonymy. 
Cynips flocossa Bassett 188 1 : 1 1 1. Females and galls  (types examined). New synonymy. 
Neuroterus  verrucarum var. pernotus Kinsey  1923: 74. 
Neuroterus flocossus:  Burks  1979:  1073

Biology.--Only  the unisexual  generation is  known. This species induces detachable leaf galls which are small, smooth, seed- like larval cells, hard, thin-walled, and covered with a dense flattened yellowish brown mass of wool  that dries brown. The gall is monothalamous, attached to the underside of the leaf by a small point to the leaf vein, separable,  but the leaf is depressed at the point of attachment. The average diameter of a gall is 2.0 mm by 1.0 mm high. The galls are usually numerous, covering the whole underside of the leaf and  are found on Quercus alba, Q. bicolor Willd., Q. chapmanii, Q. margaretta, and Q. stellata. Viereck (1916) also recorded Q. macrocarpa Michx. as a  host, but this  must be confirmed. Galls begin to develop in September (Weld 1959). According to Kinsey (1923), the galls  appear  after  mid-summer in August and the larvae do not mature  until late in the fall. Adults emerge the next spring in February to March.

Distribution.-Ontario, New York, New Jersey,  Connecticut,  west to Illinois, Oklahoma, Texas,  south to Florida. 

Descriptions of four new species of cynipid gall wasps of the genus Neuroterus Hartig (Hymenoptera: Cynipidae) with redescriptions of some known species from the eastern United States