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Psyche 10:140-155, 1903.

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1 40
PSYCHE [August
CLASSIFICATION OF THE GALL-WASPS AND THE PARASITIC CYNIPOIDS, OR THE SUPERFAMILY CYNIPOIDEA. 111. BY WILLIAM 13. ASHMEAD, A. M., S. D., ASSISTANT CUKATOR, U. S. NATIONAL MUSEUM, WASHINGTON, D, C.
Subfamily V1.- Xystinae.
1869.
Allotriaidae, Familie 3, Forster, Verb. zoo1.-bot. gesell. Wien. bd. 19, p, 3299 338-
1877.
Allotriina, Tribus, Thornson, Opus. ent., fasc. 8, p. $11. 1890. Allotriina, Subfamily, Cameron, Monogr. Brit. Phyt. Hym., vol. 3, p, 157, 232.
1897.
Allotriinae, 3e Tribu, Kieifer, Monogr. dcs cyiiipides d'Eur., torn., I, p. 54. 1902. Allotriinae, Subfamille 5, Dalla Torre cl KiefEer, Wytsman's Gen. Ins. Fam. Cynipidae, p. I.
1903.
Xystinae, Subfamily VI, Ashmead, Psyche, vol. m, p. 8. The species falling in this subfamily are all smooth, highly polished, and are easily recognized by the short, globose, or subglobose, abdomen, the second segment being usually the largest, by the short thorax, the scutellum being convex and smooth, rarely foveated at base, and by the hind tibiae having only one apical spur. All the species are genuine parasites and destroy various Aphids (&h- didae).
The group seems to form a connecting link between the Figitidae and the C'tzipidm, many of the species closely resembling those in the genera Ceroptres, ATezt~ote?~, Dyopha&a, etc.
Two tribes may be distinguished : -
Pronoturn laterally and the femora and tibiae toward apex, with broad, foliaceons dilations; claws with a tooth beneath , Tribe I. Loboscelidiini Fronoturn and legs normal; claws simple . Tribe 11. Xystini This tribe is based upon the genus Loboscelidia Westwood, described from Sulu Island, and its position is uncertain. Only a single species is known and that
Psvhe 10:HO-155 (1903). http://psyche cnu-luborefJWIO.HO hld



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is a most striking looking wasp on account of the shape of the head and the foliaceous dilations of the thorax and legs. Professor Westwood, uncertain whether it was a cynipid or a proctotrypid, finally placed it doubtfully among the latter in the subfamily Diaprtinae.
Unlortunately, I have not yet had a specimen for examination, but, judging from Westwood's description and figure, and especially of the venation, I have very little doubt of its being a cynipoid and not a proctotrypoid, so for the present treat it as a tribe in the subfamily Xysfinae. Besides the foliaceous dilations on the pronolurii, femora and tibiae, it may be recognized by the following characters : - Wings well developed, with the venation distinct, the marginal cell large, as in Xystvs; head subglobose. with a short, porrect snout; antennae in Q 14-jointecl, filiform (8 unknown) . . Loboscelidia Westwood (Type L. rufescens Westw.)
This tribe is distinguished by the head, thorax, and legs being normal, the pronotnm and legs always w;:'/ho/;/ foliaceous dilations. The antennae in the females are 12- or 13-jointed, in the males 13- or 14-jointed. The species are numerous and attack almost exclusively species belonging to the hon~opterous family Afiiicliilae.
Mesonotum entirely without parapsidal furrows . I Mcsonotuni with the parapsicla1 furrows more or less distinct .
7
I
Wings usually fully developed or if abbreviated always with a, distinctly defined marginal cell 2
2.
Wins"; abbreviated, without a marginal cell. Antmnae i:i 9 1.3-jointed, in (T 14-jointed , I'ezophycta FOrster (Type Xystus brachypterii Hartig.)
3. Xngina? cell ofni along the front margin . 3 Marginal cell completely closed . S
3. Radius extending to the front margin; antennae in 9 13-joinled, in ,7 14-jointed . 4
Radius not. extending to the front margin; antennae in 9 12-jointed, in
1.3-jointed . . Dilyta FOrster
(Typc 1). su?xlavata Forst.)




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142
PSYCHE [August
4.
Scutellum with a fovea at base ; third abdominal segment longer than the second . Glyploxysta Thomson
(Type G. heterocera Thorns.)
Scutellurn without a fovea at base ; third abdominal segment much shorter than the second . , Alloxysta FOrster
(Type Xystus macrophadnus Hartig.)
5, Wings abbreviated, not or hardly as long as the abdomen . . 6 Wings much longer than the abdomen; antennae in 9 I~-jointcd, in J I 4-jointed.
Scutellum with one or two foveae at base , Auloxysta Thornson (Type A. nigripes Thomson)
Scutellum not foveate at base . . Xystus Hartig = Allotna Westwood
(Type A. victrix \hTestw.)
6. Antennae in 'i' 13-jointed, in (T 14-jointed . . Nephycta FOrster (Type N. discreta Forst.)
7.
Parapsidal furrows abbreviated ; scutellum without foveae at base; antennae in ? 13-joinled, in J 14-jointed . , Hemicrisis FOrster (Typc H. ruficornis Forst.)
Parapsidal furrows complete, entire ; scutellum with one or two foveac at base; antennae in 9 13-jointed, in (T 14-jointed . . Phaenoglyphis h'orstcr (Type P. xanthochroa Forst.)
1840.
Cynipides, Kamilie (partim), IIartig, Zeitsch. f. enl., bd. 2, p. 187. 1869.
Cynipoidae. Kamilie (partim), FOrster, Verli. zoo1.-bot. gesell. Wien, bd. I 9, P. 329.
1877.
Cynipina, Suh-familia (partim), Thomson, Opus. ent., fasc. 8, p. 778. ~897.
Cynipinae, ae Tribu (partim), Kieffer, Monogr. des cynipicles d' Eur., torn. -1, p. 54.
1902.
Cynipidae, Subfamille (partim), Dalla Torre et Kiefler, \Vytsman's Gen. Insectorum~ p. 42.
To this family, as I have restricted it, belong all the genuine gall-makers, the gall-inhaling species (Synerginaej, and the Ibaliinae, the latter reprcscnting a small group of parasites. The gall-makers and gall-inhaling species are very numerous, ,
closely resemble each other, often living side by side together in the same galls, and are separated with difficulty, the experienced eye alone being able lo detect the difference. They produce galls or live in galls, on various trees and plants,



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the oak, bramble, rose, and various Compositae being especially subject to their attacks. From the FigHidae they are distinguished principally by abdominal peculiarities, the tergites being shorter and not meeting along the venter, not enclosing or hiding the sternites, as in the former, except in some Anacharines. All, however, have a habitus or tout-eiisemfile peculiarly their own, which with practice one soon perceives, and is thus able to recognize the different groups at a glance.
Three subfamilies have been recognized, distinguished by the characters employed in the following table : -
Basal joint of hind tarsi usually shorter than joints 2-5 united or never much longer; abdomen not or very little longer than the head and thorax united .
I
Basal joint of hind tarsi at least twice as long as 2-5 united ; joints 2-4 scarcely longer than thick, the second with a long spined process outwardly . . 2
I. Second segment in female very large, occupying the whole or nearly the whole surface of abdomen, very rarely showing an indistinct dividing suture; it this suture is distinct or complete it is very oblique and the segment dorsally is fully two thirds the length of the abdomen ; males with the second and third segments nearly equal, but here two segments occupy most of the surface of the abdomen; venter more or less covered basally Subfamily I. Synerginae Second segment in female much shorter, but the longest segment; the second and third together not occupying two thirds the whole surface or rarely; venter always visible . . . Subfamily 11. Cynipinae 2. Abdomen very strongly compressed, cultriforin, and much longer than the head and thorax united, the four or five basal segments nearly of an equal length . . Subfamily III. Ibaliinae
Subfamily I. Synerginae.
1896.
Synerginae, Subfamily VII, Ashmead, Trans. Amer. ent. soc., vol. 23, p. 186. 1900.
Synerginae, Subfamily I, Ashmead, Smith's Ins. New Jersey, p. 548, 1901.
Cynipinae, a' Tribu (partim), Kieffer, Monogr. des cynipides d'Eur. tom. I, 1'- 54-
1~02.
Cynipinae, Subfamille (partim), Dalla Torrc et Kieffer, Wytsman's Gen. Ins. p. 2.
This group is classified by Dalla Torre and Kieffer among the Cynipinae; it is evidently an offshoot of the genuine gall-makers but now sufficiently differenti- ated in structural characters and in habits to be kept apart. The SperKinae may



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144
PSYCHE [August
be popularly known as "the false gall-makers " or Inquilines, since most of them, if not all of them, are not genuine gall-makers, although often mistaken for them ; nor are they genuine parasites, i. e. they do not destroy the genuine gall-makers; on the contrary all, with possibly two or three exceptions, are inquilines or commen- sals and merely dwell, often side by side, in the galls made by other insects. Most of them are bred from cynipidous galls, found on oak trees, but some are also bred from galls made by other insects, Diptera, etc., occurring on the oak, willow, ctc. The genus Symfhrus is said to be a genuine gall-maker as well as the genus Rhciopkilus, described from Africa 'and bred from a gall on Rhus; but, judging from the structural characters of these wasps I suspect both are really commensals in cynipidous and cecidomyidous galls.
Marginal cell completely closed . . I
Marginal cell open along the front margin . . 6 I. Facc, or at least the cheeks, striated, the striae usually converging towards the mouth ; mesopleura longitudinally striated aciculated, rarely smooth . 3 Face not striated, smooth, coriaceous, or punctate ; mesopleura smooth, highly polished, or at least not striated.
Second abdominal segment occupying nearly the whole surface, without a trace of a dividing suture ; sheaths of ovipositor projecting . a 3 Second abdominal segment divided into two by a delicate, or a distinct, suture, which is either vertical or oblique . 2 2.
Suture dividing the second segment distinct and very oblique: extending towards the base of the petiole, the first division appearing tongue-shaped, dor- sally long, ventrally very short : sheaths ot ovipositor not prominent ; antennae in $ 13-jointed, the third joint longer than the fourth, in J 15-jointed, the
third joint longer than the fourth, excised beneath . Euceroptres Ashmead
(Type E. primus Ashm.)
Suture dividing the second segment very delicate, vertical, the first division not longer dorsally than ventrally ; sheaths of ovipositor prominent, projecting ; antennae in Y 12- or 13-jointed, the third joint not or scarcely longer than the fourth; in {, 14- or 15-jointed, the third joint not longer than the fourth, not excised beneath . . Ceroptres IIartig
(Type C. clavicornis Hartig.)
3.
Mesonotum with the parapsidal furrows complete, distinct; face entirely stri- ated, withoz~t a smooth median elevation ; sheallis of ovipositor usually, but not always prominent . 5




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Mesonotinn with the parapsidal furrows
incomplete, wanting, or evanescent
anteriorly; face with a smooth median elevation, the striae confined to the cheeks and on the space next to the eyes ; sheaths of ovipositor prominent 4
4.
Antennae in 9 12-jointed, the third joint not longer than the fourth, in 8 15- jointed, the third joint usually strongly excised . . Periclistus Forster (Type Aulax cananae Hartig.)
5. Claws simple; petiole of abdomen striated; antennae in $ IS-, 14-, or 15- jointed, in if 15-jointed, the third joint scarcely longer than the fourth, excised outwardly towards base . Synergus Hartig (Type S. nigripes Hartig.)
Claws with a more or less distinct tooth at base beneath ; petiole of abdomen not striated ; antennae in 9 13-jointed, in J1 14-jointcd, the third joint longer than the fourth . . Rhoophilus Mayr
(Type R. l6wii Mayr.)
6.
Scutellum normal, distinctly bifoveated at base ; mesonotuni with more or less distinct parapsidal furrows , 7
Scutellum broad, not foveated at base ; mesonotun without parapsidal furrows. Antennae in 9 I~-jointed, in if 14-jointed, the third joint much longer than the fourth, strongly excised outwardly . . Sapholytus Forster (Type Synergus apicalis Hartig.)
7.
Areolet in front wings rather large distinct ; the two foveae at base of scutellum very large transverse, separated by a carina; mesothorax transversely rugulose ; antennae in 9 13-jointed, in J 15-jointed, filiform, the third joint hardly as long as the fourth or no longer . . Synophrus Hartig (Type S. politus Hartig.)
Areolet in front wings incomplete, the outer side alone present; the two foveae at base of scutellu~n not large oblique; mesothorax coriaceous, not transversely rugulose; antennae in 9 13-jointed, in f 15-jointed, filiform, the third joint much longer than the fourth . . Synophromorpha Ashm., g. nov. (Type S. salicis Ashm.)
Subfamily 11. Cynipinae.
1900. Cynipinae, Subfamily 11, Aslimead, Smith's Ins. New Jersey, p. 548. 1901. Cynipinae, 2' Tribu (partim), Kicffer, Monogr. des cynipides d'Europe tom. I, p. 54.
1902. Cynipinae, Subfamille (partim), Ualla Torre et Kieffer, Wytsman's Gen. Ins. p. 2.
This subfamily, as I have restricted it, comprises only ge-imine gall-makers, or



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I 46 PSYCHE [AU~US~
cynipoids producing galls or deformations on various trees and plants. The genera and species are numerous, much more numerous than most people imagine, and undoubtedly many genera and species yet remain unknown to us. The National collection contains many undescribed species. The vast majority of the described species belonging to this subfamily produce galls on oak trees, and on the rose and bramble (blackberry and raspberry), but this is due probably to the fact that the galls made on these trees and plants are much more conspicuous, or the trees and plants themselves are more thoroughly studied, than those on other trees and plants, and when the galls on other trees and plants are more extensively collected and studied, we may expect a wonderful increase in our knowledge of the gall-making cynipoids. The subfamily Cynipi~/ae is dividable into five minor groups or tribes, which appear to be natural, since the species falling in each tribe confine their attacks to trees and plants of the same order or family, or closely allied orders or families. The species falling in the tribe C'ifini, for example, produce galls only on trees of the order CUPULIFERAE, those of the tribe Rhodiiini attack the ROSACEACKAE, those of the tribe Aulaciniattack the COMPOSITACEAE, etc. These tribes may be recognized by the use of the following table: - 1
Antennae inserted abnormally high up on the face on- an imaginary line drawn across from the apex of the eyes ; face with two short, distinct antennal fur- rows .
5
Antennae inscrtcd normally on or near the middle of the face, or far below an imaginary line drawn across from the apex of the eyes ; face without distinct antenna! furrows . 2
2. Winged forms . 3
Wingless or subaptcrous forms.
These are all dimorphic or agarnous forms, represented only in the female sex ; they produced the fully winged sexual form represented by both sexes (8 9 ) and are easily recognized by the family characteristics, and produce galls on oak trees, or the C7<*diferae in late fall and winter. The sexual form appears in early spring and smiuner , . Tribe I. - Cynipini 3.
Cubitus in front wings wanting or if present originating distinctly below the mid- dle of the basal nervure; areolet often entirely absent; abdomen variable. Cubitus in front wings rarely entirely absent and originating at or near the middle of the basal nervure, never much below the middle; areolet usually present and lying directly beneath the origin of the radius ; abdomen in $ sub-



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compressed, with the second segment always large, occup\ing usually about two thirds the whole surface of the abdomen, the hypopygium ending in a blunt hairy process of variable length; scape of antennae obconical, about thrice as long as thick at apex, the third joint always distinctly longer than the fourth. (Producing galls on oaks.) . , . Tribe I.- Cynipini 3. Abdomen in 9 not, or only slightly, compressed at apex, the hypopygium neither piominent nor acutely pointed at apex; scape of antennae about thrice as long as thick at apex , 4
Abdomen in ? much con~pressed towards apex, the second segment occupying about two thirds the whole surface, the hypopygium prominent, acutely pointed at apex or plowshare shaped ; front wings with the areolet distinct, its base directly beneath the origin of the cubitus ; scape of antennae subglobosc, hardly twice as long as thick. (Producing galls on rose-bushes, etc., Rosa and Rz;I,z;s.) Tribe 11.- Rhoditini
4.
Abdomen with the second segment large, occupying much more than half the whole surface ; front wings with the areolet distinct, lying directly beneath the origin of the cubitus; third joint of antennae longer than the fourth. (Pro- ducing galls on maple worts, Sapi7zdzcear, maple, ace^.) Tribe 111.- Pediaspiclini
Abdomen with the second segment shortcr, occupying scarcely half the whole surface ; front wings with the areolet often wanting, or if present not lying directly beneath the origin of the radius, usually small ; third joint of antennae not or rarely longer than the fourth, usually shorter. (Producing galls on KOSACEAE : .fibus, &a-gdria, Potentih ; COMPOSITACEAE : Lygodesmia, Hiera- &lm, &Cfu~a, &&/di?l1?L, SoIZ~hus._ flab&, firaxam, etc. ; PAPAVERACEAE: Pafaverus, G/azicitmt, etc.) . . Tribe 1V.- Aulacini 5.
Front ring's without an areolet.
(Producing galls on LEGL'MIKOSAE : Acacia.) . . Tribe V.- Eschatocerini This is the largest and most extensive tribe in the subfamily, and contains many genera and species, and all the species, without a single exception, produce galls on the mastworts (CUPULIFERAE), the oaks (QUERCUS) especially being most frequently subject to their attacks; it is extremely rare for them to attack the chestnut (CASTAREA), or the beech (FAGUS), although their galls are sometin~cs found on these trees, but the wasps producing them are seldom reared, and are still iindescribecl.




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PSYCHE [August
The numerous genera into which these wasps are now divided, may be recognized by the aid of the following table : - Apterous or subapterous forms . I
Wings fully developed . . II
I. Mesonot~~m viitho7d, or with indistinct, or incomplete parapsidal furrows, never deep or sharply defined 2
Mesonotum with deep, sharply defined parapsidal furrows . 6 2.
Mesonotum with traces of furrows, the furrows, however, never complete 3
Mesonotum smooth, polished, without a trace of the furrows; face smooth, highly polished.
Antennae 14-jointed, the third joint not quite so long as the two following united, joints 10-13 a little longer than thick ; scutellum small, rounded, convex, with a slight transverse grooved line at base ; claws of hind tarsi simple ; abdomen bare (agarnous $ ) . Xystoteras Ashmead (Type X. vollutelae Ashin.)
3. Antennae 14-jointed . 4
Antennae 13-jointed . 5
4. Scutellum triangular or conical, as viewed from above, the apex obtuse, or ending in an obtuse thorn, the base not separated from the mesonotum by a grooved line ; body very hairy ; face shagreened, opaque ; antennae very long; the joints cylindrical, 3 to 5, or to 6, very long, the following gradually shortening; claws of hind tarsi with a tooth towards base beneath ; sides of abdomen usually densely pubescent (agarnous 9 ) . . Philonix Fitch =: Acraspis Mayr
(Type P. fulvicoliis Fitch.)
Scutellum rounded or semicircular, always rounded off posteriorly, with a slight arcuate furrow or depression at base, the base separated from the mesonotum by a delicate grooved line and carha; face and mesonotui~~ alutaceous or shagreened ; antennae long, the third joint as long or nearly as long as joints 4-5 united, joints 6-13 a little more than twice as long as thick: claws of hind tarsi simple, without a tooth beneath (agamous $ ) Zopheroteras Ashmead
(Type Acraspis vaccinii Ashin,)
5.
Scutellum rounded, with indications of foveae on either side at base, or at least depressions, the base separated from the mesonoturn by a delicate tiansverse grooved line; face shagreened or coriaceous, the n~esonotum subopaque or



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alutaceous; antennae rather short, the third joint a little longer than the fourth; joints 9-12 twice as long as thick; claws of hind tarsi with a tooth at base beneath (agarnous ?) . . Phylloteras Ashmead (Type Biorhiza rubinus Gillette.)
Scutellum semicircular, bounded by a delicate rim posteriorly, without foveae at base; face smooth shining, or at the most feebly alutaceous, the mesonotum polished, with traces of farrows anteriorly; antennae somewhat short, the third joint nearly twice as long as the fourth, joints 7-12 scarcely longer 1.han thick; claws of hind tarsi simple . Trigonaspis Hartig (Type Cynips megapterus Panz.)
6.
Front tibiae outwardly at apex normal, never prolonged into a large spined process, at most only slightly dilated . 7 Front tibiae outwardly at apex prolonged into a large, spined process . 10 7. Scutelli~m laterally imrnarginal, without a frenum; face sometimes with a ridge or carina between the antennae . 8 Scutelluin laterally margined, or with a distinct frenum ; face ivithwt a ridge or carina between the antennae.
Antennae 14-jointed, the third joint scarcely longer than the fourth, joints I 1-14 hardly longer than wide ; head and thorax shining, but the former more or less coriaceous or alutaccous ; scutellu~n without distinct foveae at base, although there is a slight transverse grooved line; tarsi shorter than tibiae; claws with a tooth within (agamous 9 ) Xanthoteras Ashmead
(Type Biorhiza forticornis Walsh.)
8.
Face with a distinct median ridge or carina between the antennae; scutelluin large, lunate, or semicircular; antennae 14-jointed; hind tarsi as long as their tibiae, the claws simple without a tooth at base beneath (agamous $ ) Biorhiza Westwood
(Type Cynips aptera Linn6.)
Face zuithozil a distinct median ridge or carilia between the antennae ; scutelluin rounded, convex, or a little longer than wide, and separated from the niesonotuin I V a delicate grooved line
Antennae 12- or 13-jointed . .. . 9
Antennae 14-jointed.
Third joint of antennae long but much shorter than 4 and 5 united, joints 11-13 scarcely twice as long' as thick, the last joint hardly as long as the two preceding uniled; head and thorax alutaceous or shagreened, the pleura finely striated; scutellnm small, highly con- vex, with a distinct transverse fovea at base; hind tarsi longer than



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I jo rs YCHE [~ugust
their tibiiie, tlie claws with a distinct tooth at base beneath (agamous 9 j . , Parateras Ashmead
(Type P. liubbardi Ashm.)
9.
Antennae 1.3-jointed, the third joint not quite so long as 4-5 united, joints 9-12 scarcely longer than thick, the last joint as long as tlie two preceding united; body bare or nearly; from alutaceous; mesunotum smooth, shining; scutellum without a fovea at base; hind tarsi much shorter than their tibiae, the claws with a blunt tooth at base beneath (agarnous 9) Spliaeroteras Ashmead
(Type Biorhiza mellea Ashm.)
Antennae I 2-jointed, the third joint a little shorter and thicker than the fourth, but equal to the fifth, joints 6-8 gradually shortening, joints 9-1 I very little longer than thick, the last joint oblong-, fully as long as 10 ancl n combined; head and thorax very closely punctate, hairy, the disk of inesopleura alone polished, but densely pubescent below ; scutellum cushion-shaped, a little longer than wide, with two distinct, smooth, lunate foveae at base; hind tarsi not
longer than their tibiae, the claws with a tooth at Inse beneath (againous 9 ) Trichotei-as Aslimead
(Type T. coquilletti Ashm.)
10.
Antennae 13-jointed, somewhat thickened, the joints after the third short; claws simple (agamous ? ) . . Belonocnema May (Type B. treatae Mayr,)
I I.