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150
G
EORGE
M
ELIKA
& W
ARREN
G. A
BRAHAMSON
Systematic Parasitoid Laboratory
R
EVIEW OF THE
W
ORLD
G
ENERA OF
O
AK
C
YNIPID
W
ASPS
(H
YMENOPTERA
: C
YNIPIDAE
: C
YNIPINI
)
George M
ELIKA
1
& Warren G. A
BRAHAMSON
2
1
Systematic Parasitoid Laboratory, Plant Protection and Soil Conservation Service of County Vas, Kőszeg,
Hungary (e-mail: chalcini@savaria.hu)
2
Department of Biology, Bucknell University, Lewisburg, PA 17837 USA (e-mail: abrahmsn@bucknell.edu)
Abstract – The current classification of world genera of Cynipini follows Weld (1952a), who divided
Cynipini into 39 genera. Later, Monzen (1954) described a new genus, Neoneuroterus from Japan;
Maisuradze (1961) reported a new genus, Repentinia Belizin & Maisuradze from Ciscaucasus (Azerbaijan);
Kovalev (1965) described two new genera, Belizinella and Ussuraspis from the Far East of Russia; Lyon
(1993) synonymized Xystoteras to Phylloteras and described a new genus, Euxystoteras; and Melika &
Abrahamson (1997b) described a new genus Eumayriella from Florida and synonymized Trisoleniella to
Eumayria. Later, Melika, Ros-Farré & Pujade-Villar (2001) synonymized Fioriella to Plagiotrochus.
Recently 41 genera of Cynipini were known to associate with oaks, generic division of which was based on
the presence or absence of a basal lobe on the tarsal claws. A reclassification of world genera of oak gall-
inducing cynipids of the tribus Cynipini (Hymenoptera: Cynipidae) is given in which 26 genera are proposed
as valid, 15 are synonymized, one Neuroterus subgenus, Latuspina Monzen, 1954 has an uncertain status; 73
comb. nov. and 26 comb. rev. are made.
Key words: Cynipidae, Cynipini, gall-wasps, taxonomy
Introduction
The clear definition of genera is the primary difficulty in the classification of Cynipidae
(Hymenoptera), particularly in the tribus Cynipini (Dailey & Menke 1980). The presence of
alternating asexual and sexual generations in many genera creates considerable morphological
variation among adults that markedly complicates the assessment of generic limits and hence
classification. The assessment of generic limits requires that generic characters be defined that
include the character states of both generations.
Burks (1979) listed 485 species of oak gall-inducing cynipids for North America north of
Mexico and subsequent to this publication many new species have been described. Although the
majority of North American species of oak gall wasps are known from only one generation, there
are nearly 150 species from Central America, mainly Mexico, whose lifecycles are not known.
Accumulation of controlled rearing data to determine lifecycles of cynipids will take decades.
However, we can improve the classification of the Cynipini by using information from the existing
literature on cynipid lifecycles and by detailed analysis of type specimens.
The diagnostic features that are used currently in the keys to the world genera of Cynipini
frequently include morphological characters that are inconsistent. Consequently these keys
frequently are unable to distinguish some genera. For instance, identification of the representatives
of Plagiotrochus Mayr, Bassettia Ashmead, Eumayria Ashmead, Trisoleniella Rohwer and Fagan,
Xanthoteras Ashmead, Sphaeroteras Ashmead, and some species groups of Callirhytis Foerster
are especially complicated. The current systematic arrangement of Bassettia, for instance, includes
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George Melika & Csaba Thuróczy (editors)
several Callirhytis and Andricus Hartig species, Eumayria includes some Callirhytis species,
and Trisoleniella appears to be a synonym of Eumayria (Melika & Abrahamson 1997b). These
systematic difficulties result from the use of inappropriate diagnostic characters for some genera.
Such characters are not useful because they are shared by the representatives of several genera
and, thus, are inappropriate to use at the generic level. In some cases, the structure and location of
a gall or novel host associations were more heavily weighted in the description of new species,
species groups, or even new genera than were morphological peculiarities of adult wasps. For
example, the North American genus Heteroecus Kinsey was established on the basis that its
species associate only with Quercus chrysolepis Liebm. with distribution limited to California.
A historical review of Cynipidae classification was given in Melika & Abrahamson (2000b).
The current classification of world genera of Cynipini follows Weld (1952a). No corrections
to Weld’s Cynipidae classification were made in a manuscript Weld prepared later entitled
“Supplement to Cynipoidea (Hym.) 1905-1950 (1952).” This manuscript, which is dated 1964,
is available at the United States National Museum, Smithsonian Institution. Weld’s Cynipini
classification needs a substantial alteration. It is important to recognize that very little was known
about the alternation of generations in North American cynipids during Weld’s time. This alone
markedly complicated the task of establishing a more natural classification of Cynipini. Excellent
studies of the alternation of generations for USA cynipids have been completed during the past
several decades (Doutt 1959, 1960; Dailey 1969; Dailey & Sprenger 1973a, 1973b; Dailey, Perry
& Sprenger 1974; Evans 1967, 1972; Lyon 1959, 1963, 1964, 1969, 1970; Melika & Buss 2002).
These results have increased our understanding of gall-inducing cynipids and given us a better
background to establish a more natural classification of Cynipini, particularly for those restricted
to North America.
Materials and Methods
Our study analyzed the type species of genera and also many other species types. Specimens
were examined from the following museums and institutions: USNM (United States National
Museum of Natural History, Smithsonian Institution, Washington DC, USA, A. Menke), AMNH
(American Museum of Natural History, NYC, NY, USA, J. Carpenter), CNCI (Canadian National
Collection of Insects, Ottawa, Canada, J.T. Huber), BMNH (British Natural History Museum,
London, England, J. Noyes, D. Notton and N.D.M. Fergusson), OUM (Hope Entomological
Collections, University Museum, Oxford, UK, C. O’Toole and G. McGaven), NHMW (Natur-
historisches Museum, Vienna, Austria, S. Schödl), NHMH (Hungarian Natural History Museum,
Budapest, Hungary, L. Zombori and J. Papp), ZIN (Zoological Institute of Russian Academy of
Sciences, St. Petersburg, Russia, O.V. Kovalev), IZU (Schmalhausen Institute of Zoology of the
Ukrainian Academy of Sciences, Kiev, Ukraine, M.D. Zerova and L.V. Diakontshuk), and MNHN
(Muséum National d’Histoire Naturelle, Paris, France, C. Villemant-Ait-Lemkadem). In addition,
we examined specimens from different genera that were kindly sent to the authors by J. Pujade-
Villar (Universitat de Barcelona, Barcelona, Spain), J.L. Nieves Aldrey (Museo Nacional de
Ciencias Naturales, Madrid, Spain), G.N. Stone (Institute of Cell, Animal and Population Biology,
University of Edinburgh, Edinburgh, Scotland). Authors express their deepest appreciation to
mentioned colleagues for help in obtaining material for research. Also the Cynipidae collection of
Systematic Parasitoid Laboratory (Kőszeg, Hungary) was examined.
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We use the following terminology: for mesosoma (Gibson 1985; Menke 1993), for head,
metasoma, and ovipositor (Fergusson 1988; Ronquist & Nordlander 1989) (Figs 1-4). The surface
sculpturing follows Harris (1979). Abbreviations for the forewing venation follow Ronquist &
Nordlander (1989). The measurements and abbreviations used herein include: F1-F12, 1
st
and
subsequent flagellomeres; POD (post-ocellar distance), the distance between the inner margins of
the posterior ocelli; OOD (ocellar-ocular distance), the distance from the outer edge of a posterior
ocellus to the inner margin of the compound eye; LOD, the distance between lateral and frontal
ocellus; transfacial line, distance between inner margins of compound eyes measured across
antennal sockets (Fig. 1). The width of the radial cell is measured along 2r (Fig. 3). Drawings were
made with the aid of a Leica drawing tube or from stereomicroscope photographs, which were
scanned into a PC and modified in Adobe Photoshop 6.0.
Results and Discussion
Morphology and current arrangement of genera
The tribus Cynipini (Hymenoptera, Cynipidae) includes species that induce galls on members
of Fagaceae, primarily Quercus L. with exceptions for only a few species. For instance,
Dryocosmus kuriphilus (Yasumatsu), the only species of Cynipini known to associate with
Castanea, threatened the chestnut industry of Japan and Korea and was discovered in Georgia,
USA in 1974 (Payne et al. 1975). Another species, Dryocosmus castanopsidis (Beutenmueller),
known from Oregon and California, induces galls on catkins of Castanopsis chrysophylla and
C. sempervirens (Burks 1979).
Three genera, Liebelia Kieffer, 1903 Paraulax Kieffer, 1904, and Poncyia Kieffer, 1903 that
were included in Weld’s (1952a) Cynipini key belong elsewhere. The genera Liebelia and
Diplolepis Geoffroy, 1762 belong in the Diplolepidini (Ronquist 1999). Paraulax possesses some
character states (e.g., the structure of the antenna and forewing venation) that indicate the
genus should be placed in the Charipinae subfamily (Cynipoidea: Figitidae) (Ronquist 1995).
Furthermore, on the basis of its description, Poncyia may be a Plagiotrochus, or if the description
of the pronotum is incorrect, the genus must be placed among inquilines on the basis of its
longitudinally striate petiole and long tergum II. Currently it is placed in the Cynipidae, incertae
sedis (Ronquist 1999). The genus, Synophrus Hartig, 1843, with only three known species from
Western Palaearctic, S. pilulae Giraud in Houard, 1911, S. politus Hartig, 1843 (Europe, Asia
Minor, North Africa), and S. olivieri Kieffer, 1899-1901 (Algeria, Israel, Iran), originally was
described as a gall inducer, however, on the basis of the morphological characters of adults, this
genus belongs to cynipid inquilines (Ronquist 1994). Synophrus has never been a subject to
detailed research and, thus, it is quite possible, that Synophrus species, in spite of their taxonomic
identity with cynipid inquilines, lack gall-inducing capability. This genus is also excluded from the
below analysis.
Around 1000 species of Cynipini are known worldwide, all in the Holarctic region, mainly in
the Western Palaearctic (around 140 valid species) and America North of Mexico (Burks (1979)
listed 485 species of oak gall-inducing cynipids). The taxonomy of Cynipini species have
undergone substantial changes: new names, synonymies, new name combinations, and new alternate
life forms were experimentally obtained for several gall wasps. Currently 41 genera of Cynipini
are known to associate with oaks (Table 1).
R
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Table 1 Current arrangement of Cynipini genera and their distribution
(after Pujade-Villar, Bellido, Segú & Melika 2001, with some changes)
Genus name Geographic distribution Number of species
Acraspis Mayr, 1881 Nearctic probably > 30
Amphibolips Reinhard, 1865 Nearctic around 30
Andricus Hartig, 1840
(= Adleria Rohwer & Fagan, 1917)
Holarctic and Oriental? probably > 300
Antron Kinsey, 1930 Nearctic around 40
Aphelonyx Mayr, 1881 Palaearctic 4
Atrusca Kinsey, 1930 Nearctic uncertain; probably > 40
Bassettia Ashmead, 1887 USA 9
Belizinella Kovalev, 1965 Russia, Far East 2
Belonocnema Mayr, 1881 USA 2
Besbicus Kinsey, 1930 USA 8
Biorhiza Westwood, 1840 Palaearctic 2
Callirhytis Foerster, 1869 Holarctic around 150
Chilaspis Mayr, 1881 Occidental Palaearctic 3
Cynips Linnaeus, 1758 Palaearctic around 25
Disholcaspis Dalla Torre & Kieffer, 1910 Nearctic around 40
Dros Kinsey, 1937 Nearctic 11
Dryocosmus Giraud, 1959 Holarctic Around 25
Erythres Kinsey, 1937 Mexico 2
Eumayria Ashmead, 1887
(=Trisoleniella Rohwer & Fagan, 1917)
USA 5
Eumayriella Melika & Abrahamson, 1997 USA 2
Euxystoteras Lyon, 1993 USA 1
Heteroecus Kinsey, 1922 USA 15
Holocynips Kieffer, 1910 USA 4
Liodora Foerster, 1869 Europe, USA 3
Loxaulus Mayr, 1881 Nearctic 14
Neoneuroterus Monzen, 1954 Russia, Far East and Japan 5
Neuroterus Hartig, 1840 Holarctic About 100
Odontocynips Kieffer, 1910 USA 1
Paracraspis Weld, 1952 USA 3
Parandricus Kieffer, 1906 China 1
Philonix Fitch, 1859 USA 8
Phylloteras Ashmead, 1897
(= Xystoteras Ashmead, 1897)
Nearctic 6
Plagiotrochus Mayr, 1881
(= Fioriella Kieffer, 1903)
Western Palaearctic &
Himalaya
14
Repentinia Belizin & Maisuradze, 1961 Transcaucasus 1
Sphaeroteras Ashmead, 1897 USA 8
Trichagalma Mayr, 1907 Japan and China 2
Trichoteras Ashmead, 1897 USA 8
Trigonaspis Hartig, 1840 Palaearctic Around 10
Ussuraspis Kovalev, 1965 Russia, Far East 1
Xanthoteras Ashmead, 1897 USA 12
Zopheroteras Ashmead, 1897 USA 6
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With the exception of the genus Neuroterus Hartig, 1840 (Fig. 14) the current classification of
the world genera of the Cynipini is based on the presence or absence of a basal lobe (tooth) on
tarsal claws (Figs 5-13) (Weld 1952a). The use of this character dates to earlier treatments
including Ashmead’s (1903) key to 33 genera of Cynipini. Earlier, Mayr (1881), Ashmead (1885),
Dalla Torre (1893), Kieffer (1897-1901) used this character for subgeneric level to separate
Callirhytis and Andricus as subgenera of Andricus. Again with the exception of Neuroterus, Weld
(1952a) divided all the genera into two groups: (a) those that possess a basal lobe on tarsal claws
and (b), those that have simple tarsal claws without a basal lobe (Table 2). Consequently, the
number of oak gall-inducing Cynipini genera distinguished by Weld (1952a) was 39. Later,
Monzen (1954) described a new genus, Neoneuroterus from Japan; Maisuradze (1961) reported a
new genus, Repentinia Belizin & Maisuradze from Ciscaucasus (Azerbaijan); Kovalev (1965)
described two new genera from Far East of Russia, Belizinella and Ussuraspis; Lyon (1993)
synonymized Xystoteras to Phylloteras and described a new genus, Euxystoteras, which differs
from Phylloteras only by having simple tarsal claws; and Melika & Abrahamson (1997b)
described a new genus Eumayriella from Florida and synonymized Trisoleniella to Eumayria.
Later, Melika, Ros-Farré & Pujade-Villar (2001) synonymized Fioriella to Plagiotrochus. We will
discuss these new genera and synonymizations below.
Table 2 Division of the Cynipini based on the presence or absence of a basal lobe
on the tarsal claws (* – genera which include species in both groups)
Genera with toothed claw Genera with simple claw
Acraspis Aphelonyx
Amphibolips Bassettia
Andricus Belizinella *
Antron Belonocnema
Atrusca Biorhiza
Belizinella * Callirhytis *
Besbicus Chilaspis
Callirhytis * Dryocosmus
Cynips Erythres
Disholcaspis Eumayria
Dros Eumayriella
Liodora Euxystoteras
Neoneuroterus Heteroecus
Paracraspis Holocynips
Parandricus * Loxaulus
Philonix Odontocynips
Phylloteras Parandricus *
Repentinia Plagiotrochus
Trichoteras Sphaeroteras
Trigonaspis * Trichagalma
Xanthoteras Trigonaspis *
Ussuraspis
Zopheroteras
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Figures 1–14 Cynipidae: Cynipini. 1, Head, front view: cl, clypeus; msl, malar sulcus; msp, malar
space; POD, post-ocellar distance; OOD, ocellar-ocular distance; LOD, the distance between
lateral and frontal ocellus; TFL, transfacial line. 2, Thorax, dorsal view: apl, anterior parallel line
(anteroadmedian line); mms, median mesoscutal sulcus (median mesoscutal line); n, notaulus; pl,
parapsidal line; sc, scutum (mesoscutum); scf, scutellar fovea; scl, scutellum; sss, scutoscutellar
sulcus; tsa, transscutal articulation. 3, Forewing: length, length of radial cell; w, width of radial cell.
4, Gaster: II, 2
nd
tergite; hyp, hypopygium; sas, subapical setae; ov, ovipositor; vsp, ventral spine of
hypopygium. 5-13, Tarsal claw: 5, Amphibolips quercuscinerea. 6, Andricus quercusfoliatus.
7, Cynips quercusfolii. 8, Cynips divisa. 9, Neuroterus quercusbaccarum. 10, Neuroterus tricolor,
asexual female. 11, N. tricolor, sexual female. 12, Belonocnema quercusvirens. 13, Sphaeroteras
ocala. 14, N. tricolor (scutellar foveae absent)
156
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Thus, classification of the Cynipini has relied on the presence or absence of the tarsal claw
(Figs 5-13). Lyon (1993), for example, stated that in cynipid taxonomy “the presence or absence of
a tooth on the tarsal claw is considered to be of fundamental importance in separating major
genera of the group.” However, this morphological criterion, dividing the Cynipini genera into two
major groups, is insufficient for all taxonomic distinctions. There are a number of exceptions to
this criterion, including those discussed by Weld (1952a). For instance, the majority of Neuroterus
species have tarsal claws without a basal lobe, however, quercusbaccarum L., numismalis Olivier,
and petioliventris Hartig, have toothed tarsal claws with a basal lobe in the asexual generations but
simple claws in the alternate sexual generations. The same is true for Callirhytis, in which both
generations of the European species glandium (Giraud) have toothed tarsal claws as the asexual
generation of bella (Dettmer) (Nieves Aldrey 1992). One species from Trigonaspis genus,
T. megaptera (Panzer) has a weak tooth on claws, while other species, T. synaspis have simple tarsal
claws. The sexual generations of the North American genus Xystoteras have a very weak tooth on
claws, and in Parandricus mairei Kieffer, the females have toothed tarsal claws while the males
have simple claws without a basal lobe. In the genus Belizinella, one species, B. gibbera Kovalev,
has claws with a basal lobe while another species, B. vicina Kovalev, have simple claws (Kovalev
1965). The presence or absence of a basal lobe is a likely homoplasy and probably evolved
separately in different cynipine genera. The ancestral condition is a simple tarsal claw without a
basal lobe, based on its appearance in the majority of genera from the Aylacini. However, even in
this tribe, Xestophanes Foerster, Diastrophus Hartig and Gonaspis Ashmead have a basal lobe. An
additional complication is the use of the term “tooth” Given that the term basal lobe would be
more precise. In some cases the claws have a strong basal lobe, for example as in Amphibolips and
many Andricus species. However in numerous species, including those that Weld treated as
possessing a tooth, no distinct tooth occurs on the claws.
This character has been used for generic separation in other Hymenoptera groups as well, as for
example in the Chalcidoidea. However, later works reconsidered the diagnostic value of this
character in these groups and subsequently dropped it. Anura Wijesekara (Sri Lanka, pers. comm)
eliminated the use of this character for the separation genera of Chalcididae and Leucospidae; Eric
Grissell (pers. comm.) attempted to use this character to separate species (but not genera of
Torymidae) and found that this character does not work even at the species level.
The main diagnostic characters for genera separation used by Weld (1952a) and characters that
we use in genera separation were discussed earlier (Melika & Abrahamson 2000b) and they are
provided below.
Acraspis Mayr, 1881
Acraspis Mayr 1881: 2, 29. Type species: Acraspis pezomachoides (Osten Sacken, 1862). Designated by
Rohwer & Fagan 1917. Types examined. Ashmead 1903 and Beutenmueller 1909 (synonym of Philonix).
Weld 1922 (Acraspis and Philonix distinct genera).
Paracraspis Weld 1952b: 324. Type species: Callirhytis guadaloupensis Fullaway, 1911. Designated by
Weld 1952b. Types examined. New synonym.
Diagnosis. Similar to Philonix, however, the scutellum of asexual Acraspis females is
elongated and pointed posteriorly (Figs 25, 26), while in Philonix it is rounded (Fig. 27); the
projecting part of the ventral spine of the hypopygium in Acraspis is equally broad through entire
length or slightly tapering to a point at the apex, with dense subapical setae (Fig. 18); in Philonix
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the projecting part of the ventral spine is flattened, broadest at the apex, with long dense subapical
setae reaching far beyond the apex of the spine to form a broad truncate tuft (Fig. 17). The asexual
females of Acraspis also resemble Biorhiza but differ in having distally elongated and flattened
scutellum, which overhangs the metascutellum and hides the propodeum; the latter declines more
strongly than in Biorhiza.
In sexual Acraspis females, the scutum is coriaceous or microreticulate, bare or with very
sparse short setae; notauli are incomplete in the anterior one-third or very indistinct. Closely
resembling Disholcaspis but F1 of the antenna is equal or very slightly shorter than the
scapus+pedicellum; the head is more broad from above; the scutum and mesopleuron are
coriaceous, while in Disholcaspis F1 is 2.0 times as long as scapus+pedicellum; the head is more
narrow from above; the scutum and mesopleuron are more densely reticulate. In Acraspis males,
F1 is straight or only slightly incised, less than 2.0 times as long as scapus+pedicellum; the scutum
is shorter; the scutellum is coriaceous, elongated, gradually and slightly depressed toward the
transscutal articulation, without scutellar foveae or transversal groove, with white setae. In
Disholcaspis males, F1 is 2.0 times as long or longer than scapus+pedicellum; the scutum is
longer. We are unable to find appreciable diagnostic characters for the precise separation of
Acraspis males from Cynips. It is, however, possible that Acraspis forms a brachypterous species-
group within Cynips as Kinsey (1930, 1936) proposed. Additional biological experimental data on
alternation of generations and/or using of molecular techniques are required to solve this problem.
See also Diagnosis to Cynips.
Comments. The only diagnostic character given by Weld (1952b) for differentiating his newly
described Paracraspis from Acraspis is that it has “less reduced wings, a more robust thorax with
a normally rounded scutellum.” However, the scutellum in Paracraspis is longer than broad,
slightly overhanging the metascutellum; not tapering posteriorly to a point as in a typical Acraspis
but distinctly elongated, not rounded as in Philonix; the scutum is flattened as in Acraspis (Fig.
25). For instance, A. inflata (Kinsey) and A. gemula var. suspecta (Kinsey) have an elongated
scutellum, which do not taper posteriorly to a point. Some Acraspis species are also densely
pubescent. Thus, we consider Paracraspis as a syn. nov. of Acraspis. Three Paracraspis species
must be transferred to Acraspis: guadaloupensis (Fullaway, 1911), comb. nov., insolens Weld,
1926, comb. rev., and patelloides Weld, 1926, comb. rev.
Biology. The majority of species are known from the asexual generations only and the galls
produced by asexual species are very similar, known commonly as “hedgehog” galls. They are
usually globular, hard and slightly elongated, detachable leaf galls, with surface reticulate or
rough, with points or spines. The sexual generation induces tiny bud galls. The sexual form was
described by Bassett (1881) for the first time as Cynips gemula. Triggerson (1914) established
alternation of generations experimentally for A. erinacei (Beutenmueller, 1909).
Distribution. Seventeen species of Acraspis were listed for the United States and Canada
(Burks 1979), however, two species must be transferred into Cynips: arida Kinsey, 1930, comb.
rev. and conica Kinsey, 1930, comb. rev. Currently the number of known species is 18. Several
species were also described from Mexico (Kinsey 1930, 1936), however, the validity of some
species is very dubious and must be revised. Several of them are likely synonyms and several other
species described in the Acraspis genus must be transferred to other genera.
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Figures 15–38 15–24, Ventral spine of hypopygium: 15, Cynips quercusfolii, asexual female;
16, Sphaeroteras ocala; 17, Philonix fulvicollis; 18, Acraspis echini; 19, Cynips longiventris, sexual
female; 20, Neuroterus laeviusculus, sexual female; 21, N. quercusbaccarum, asexual female;
22, N. numismalis, sexual female; 23, Andricus solitarius; 24, A. quercusramuli. 25–29, Thorax,
dorsal view: 25, Acraspis insolens; 26, A. echini; 27, Philonix fulvicollis; 28, Trigonaspis
quercusforticorne; 29, Andricus askewi. 30, Andricus kollari, asexual female, mesosoma in lateral
view. 31, Atrusca quercuscentricola, forewing. 32–33, Bassettia pallida, head: 32, from above;
33, front view. 34–35, Plagiotrochus quercusilicis, head: 34, from above; 35, front view.
36, B. pallida, forewing. 37–38, propodeum: 37, B. pallida; 38, P. quercusilicis
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Amphibolips Reinhard, 1865
Amphibolips Reinhard 1865: 10. Type species: Cynips spongifica Osten Sacken, 1862. Designated by Rohwer
& Fagan 1917. Types examined.
Trissandricus Kieffer 1910: 115. Type species: T. maculipennis Kieffer, 1910. Original designation. Monotypic.
Weld 1931 (synonym of Amphibolips quercusspongifica).
Diagnosis. The head and mesosoma are dull, coarsely rugose. The mesosoma is robust, highly
arched, broader than the head; the scutum is broader than long; the scutellum is subquadrate or
cushion-shaped, slightly broader than long, with large, deep and sometimes wrinkled scutellar
foveae. The projecting part of the ventral spine of the hypopygium is narrow, needle-like, long,
usually more robust and broader than in closely related Andricus. The forewing has distinct or less
distinct smoky spot(s). Tarsal claws have strong basal lobes. The genus is very uniform in the
characteristics of adults and the galls that they induce. The morphology of the asexual and sexual
females is identical.
Comments. On the basis of diagnostic characters given above, 6 species from the genus Andricus
herein are transferred to Amphibolips: A. ellipsoidalis Weld, 1926, comb. rev., A. femoratus
Ashmead, 1887, comb. nov., A. quercusostensackenii (Bassett, 1863), comb. nov. (= Andricus
quercussingularis (Bassett, 1863), syn. nov.), A. ruginosus (Bassett, 1890), comb. nov., A. vernus
Bassett, 1900, comb. rev. Galls of these six species structurally are also typical Amphibolips galls.
Méhes (1953) described Amphibolips mernyensis on the basis of one gall, collected from
Quercus cerris in Hungary, however it appears to be a nomen nudum (Melika, Csóka & Pujade-
Villar 2000).
Biology. Both asexual and sexual generations induce stem, bud and leaf galls, the majority of
which are uniform in their structure: the larval chamber occurs in the center of the gall and
supported by radiating filaments.
Distribution. North and Central America. With the species transferred herein, 40 species are
known: 30 from America north of Mexico (Burks 1979) and 10 from Mexico (Beutenmueller 1911;
Kinsey 1937b).
Andricus Hartig, 1840
Andricus Hartig 1840a: 185. Type species: A. noduli Hartig, 1840 (= A. trilineatus Hartig). Designated by
Foerster 1869.
Aphilotrix Foerster 1869: 331, 336. Type species: Cynips corticis Linnaeus, 1758. Original designation. Mayr
1881 (synonym of Andricus).
Manderstjernia Radoskowsky 1866: 304. Type species: M. paradoxa Radoszkowski. Original designation.
Later, the name was determined to be the elder synonym of Cynips albopunctatus Schlechtendal.
Oncaspis Dettmer 1925: 123. Type species: O. filigranata Dettmer. Original designation. Dettmer 1928
(O. filigranata is the sexual generation of Andricus solitarius (Fonsc.) and, thus synonym of Andricus.
Weld 1930 (synonym of Andricus).
Euschmitzia Dettmer 1925: 122. Type species: E. rara Dettmer. Original designation. Later, the author
thought it was the sexual generation of Andricus nudus Adler and asked Weld to publish it. Weld 1930
(synonym of Andricus).
Femuros Kinsey 1937a: 65. Type species: F. repandae Kinsey. Original designation. Weld 1952a (synonym
of Andricus).
Feron Kinsey 1937a: 69. Type species: F. verutum Kinsey. Original designation. Weld 1952a (synonym
of Andricus).
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Druon Kinsey 1937a: 56. Type species: D. protagion Kinsey. Original designation. Weld 1952a (synonym
of Andricus).
Conobius Kinsey 1938: 262. Type species: C. strues Kinsey. Original designation. Weld 1952a (synonym
of Andricus).
Adleria Rohwer and Fagan 1917: 359. Type species: Cynips kollari Hartig, 1843. Original designation.
Benson in Marsden-Jones 1953 (synonym of Andricus).
Dros Kinsey 1937a: 49. Type species: D. petasum Kinsey. Original designation. Types examined. New
synonym.
Erythres Kinsey 1937b: 461. Type species: E. hastata Kinsey. Original designation. Types examined. New
synonym.
Liodora Foerster 1869: 331, 334. Type species: L. sulcata Foerster. Original designation. Mayr 1881
(synonym of Dryophanta). Dalla Torre 1893 (treated as a separate genus). Dalla Torre & Kieffer 1910
(synonym of their “Diplolepis L. Geoffr.”). Mayr 1903 (restored its generic status). Types of North
American species examined. New synonym.
Parandricus Kieffer 1906: 102. Type species: P. mairei Kieffer. Original designation. Monotypic. Types
examined. New synonym.
Trichoteras Ashmead 1897a: 67. Type species: T. coquilletti Ashmead. Original designation. Types
examined. New synonym.
Diagnosis. Projecting portion of the ventral spine of the hypopygium is needle-like, long, with
subapical setae that do not reach beyond the apex of the spine; if short then slender and thin,
tapering to point at the apex, at least 2.0 times as long as broad, with short sparse subapical setae
which if reaching beyond the apex of the spine then never dense and they do not form a truncate
tuft (Figs 23, 24). Mesosoma is arched, notauli usually complete, sometimes absent or indistinct in
the anterior 1/3-1/4; scutum is usually as long as broad or subequal, subquadrate; scutellar foveae
usually distinct, separated by a median carina (Figs 29, 30).
Comments. Mayr (1881) treated Callirhytis as a subgenus of Andricus. Dalla Torre & Kieffer
(1910) erroneously synonymized Trisoleniella Rohwer & Fagan to Andricus.
Adleria. Rohwer & Fagan (1917) proposed Adleria instead of Cynips sensu Authors, which
they did not accept along with the type designations. The European species of the former Cynips
sensu Authors [not Cynips Linnaeus!] were transferred to Andricus after the sexual generations
were discovered and found to have the diagnostic characteristics typical of the sexual Andricus
(Benson in Marsden-Jones 1953). Zerova, Diakontshuk & Ermolenko (1988), in their review of
the gall-inducing insects of the European portion of the former Soviet Union, removed 11 species
from Andricus and transferred them again to reestablished Adleria. Their decision was based
solely on the analysis of the adults of the asexual generations. The diagnostic characteristics given
by these authors to separate Adleria from Andricus are the same as those in Weld (1952a).
Kovalev (1965) described two other Adleria species from south of Far East of Russia, which were
synonymized later to Andricus (Abe 1986). Currently the genus Adleria with 6 species is listed as
a valid genus in Burks (1979) for America north of Mexico. Five of these species must be
transferred to Andricus: A. dimorphus (Beutenmueller), comb. nov., A. flavicollis (Ashmead),
comb. nov., A. nigricens (Gillette), comb. nov., A. quercusstrobilanus (Osten Sacken), comb.
nov., A. vacciniiformis (Beutenmueller), comb. nov., and A. weldi (Beutenmueller), comb. nov.,
while one species, A. arizonica (Cockerell) must be transferred to Disholcaspis: D. arizonicus
(Cockerell), comb. nov. (the projecting part of the ventral spine of the hypopygium is shorter with
very long setae reaching far beyond the apex of the spine, see Disholcaspis).
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Weld (1951) synonymized Andricus ashmeadii Bassett to Adleria nigricens (Gillette). However,
examination of the types showed an appreciable difference: in A. ashmeadii the ratio of antennal
segments 1:3:4 is 20:20:11; the gena are only slightly broadened behind the eye; anterior tentorial
pits are much deeper; the frons punctate; the scutum is bare with fine sculpture; scutellar foveae
are larger with sculptured bottom; the transscutal articulation continuous, while in A. nigricens the
ratio of antennal segments 1:3:4 is 15:20:13; the gena are much broader behind the eye; the frons
is dull, transversely rugose; the scutum is densely pubescent, with more dull sculpture; scutellar
foveae are smaller with smooth shiny bottom; the transscutal articulation is interrupted medially;
the areolet of the forewing is twice shorter than in A. ashmeadii. We believe that these are distinct
species and consequently we restore species status for Andricus ashmeadii Bassett, comb. rev.,
status nova.
Erythres. The only character separating this genus from Andricus is the simple tarsal claw.
Both known species, E. hastata Kinsey and E. jaculi Kinsey occur on red oaks and induce cone-
like galls inside enlarged bud scales and form terminal clusters of aborted leaves and bracts that
enclose a small, seed-like cell. Galls resemble those of Dryocosmus floridensis (Beutenmueller),
Andricus stropus Ashmead, A. quercusfoliatus (Ashmead), and A. fecundator (Hartig). Thus,
A. hastata (Kinsey) and A. jaculi (Kinsey) are comb. nov.
Liodora and Dros. Dalla Torre & Kieffer (1910) treated Liodora sulcata Foerster as the sexual
generation of Cynips quercusfolii (Linnaeus), however, Weld (1930, 1951, 1952a) considered
Liodora as a distinct genus. Kinsey (1937a) stated that Dros “includes some species which have
previously been assigned to “Andricus”, “Driophanta” [sic], etc. Its further limitation must await a
monographic revision of the whole group”. He also mentioned that the genus may be related to
Feron and Druon which were synonymized by Weld (1952a) to Andricus. Dros, according to
Kinsey (1937a), is most readily distinguished from these “other genera” by a head which is
narrowed behind eyes, by a 14- or 15-segmented antenna, entirely smooth scutum and
mesopleuron, widely separated, pear-shaped scutellar foveae, and by distinctive vase or urn-
shaped galls. The only difference between Liodora and Dros is that the head in Dros is narrower
than the thorax while in Liodora it is equal to the thorax (Weld 1952a). However, this character is
insufficient to distinguish Liodora from Dros. Dros viscidum (Weld), for example, differs from
Liodora only by having a polished scutum, while D. sessile (Weld) has an alutaceous and polished
scutum. The two genera were also confusing for Weld. Weld (1952a) considered caepula (Weld)
and discale (Weld) as Dros species, but later moved them to Andricus; viscida was described by
Weld (1944) in Liodora, but later was transferred to Dros. Dros atrimentum (Kinsey) and
D. pedicellatum (Kinsey) originally were described in Andricus. Dailey & Sprenger (1973a)
moved atrimentum (Kinsey) from Dros to Andricus. The same character states found in Dros and
Liodora are encountered in many European Andricus species as in for instance, inflator Hartig,
and gallaeurnaeformis (Fonscolombe). The latter species induces urn-shape galls as well.
Evans (1972) determined that Liodora dumosae Weld is the sexual generation of Andricus
pattersonae Fullaway. Rosenthal & Koehler (1971) were incorrect when they concluded that this
species was the bisexual generation of Andricus kingi Bassett, the latter was described by Dailey &
Menke (1980). Probably Rosenthal and Koehler had the sexual generation of A. kingi.
Liodora and Dros are distinguished easily from the sexual Cynips by the presence of a long
ventral spine of the hypopygium, with short sparse setae, which never reach beyond the apex of
the spine.
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Consequently, three Liodora and 11 Dros species known from North America must be
transferred to Andricus: apiarum (Weld), comb. nov., clarkei (Bassett), comb. nov., comata
(Weld), comb. nov., amphorus (Weld), comb. nov., atrimentus Kinsey, comb. rev., moreliensis
(Kinsey) (Mexico), comb. nov., pedicellatus Kinsey, comb. rev., periscellus (Kinsey) (Mexico),
comb. nov., perlentus (Kinsey) (Mexico), comb. nov., petasus (Kinsey) (Mexico), comb. nov.,
picatus (Kinsey) (Mexico), comb. nov., repicatus (Kinsey) (Mexico), comb. nov., sessilum
(Weld), comb. nov., and viscidus (Weld), comb. nov.
Parandricus. The genus is known from one species, P. mairei Kieffer, 1906, described from
China, and is known to induce catkin galls on oaks (Kieffer 1906). The location of types is
unknown. We examined one male and one female labelled as “Hankow” which are deposited at
the USNM, Washington, DC and are originated from the type series. Weld (1952a) differentiated
Parandricus from Andricus in that females have an “abdomen in side view longer than high, ...
and hind femur stout”. In many sexual Andricus species, which induce catkin galls, both females
and males have “stout” hind femur. In males, except that the hind femur is very short and stout,
metatarsus is equal 2+3+4 tarsomeres; 5
th
is equal 1
st
in length (Weld 1952a). However, several
Andricus males, e.g., A. quercuscalicis (Burgsdorf), A. burgundus Giraud, Callirhytis blastophaga
(Ashmead) and others, have the same proportions of hind tarsomeres and are “stout.”
Consequently, the proportions of hind tarsomeres and length of hind femur vary markedly in the
sexual Andricus. Thus, the diagnostic characters given for Parandricus are insufficient to treat it as
a separate genus and, thus Parandricus is a syn. nova of Andricus, and Andricus mairei (Kieffer)
is a comb. nov.
Trichoteras. One of the diagnostic characters offered to separate Trichoteras (for the type
species T. coquilletti Ashmead) from Andricus is a 12-segmented antenna, F2 is equal or slightly
longer than F1. Andricus formosalis Weld, for example, has F2 longer than F1, has enlarged gaster
as in coquilletti; in A. pattoni (Bassett) F2=F1; the shape and proportions of the head, scutum,
scutellum, central area of propodeum, the forewing venation, the ventral spine of the hypopygium
are the same as in coquilletti. The females of the sexual generation of the European
A. gallaeurnaeformis (Fonscolombe) have F2=F1, the head is slightly broader than the mesosoma,
the scutum is same as in Trichoteras. Three species are known as brachypterous: coquilletti (fully
winged females known also), frondeum (Weld), and tubifaciens (Weld). Consequently, all 8
known species must be transferred to Andricus: burnetti Dailey & Sprenger (1983), comb. nov.,
californicum (Beutenmueller), comb. nov., coquilletti (Ashmead), comb. nov., frondeum (Weld),
comb. nov., perfulvum (Weld), comb. nov., rotundula (Weld), comb. nov., tubifaciens (Weld),
comb. nov., and vacciniifoliae (Ashmead), comb. nov.
Biology. Alternation of generations is known. These species induce galls on all organs of oak
trees, and the shapes and sizes of galls vary markedly. Life-cycles are known for numerous
European species, however, we know little about the life-cycles of the American and Asian
species.
Distribution. Holarctic.
Aphelonyx Mayr, 1881
Aphelonyx Mayr 1881: 29. Type species: Cynips cerricola Giraud, 1859. Type designated by Mayr (1881).
Diagnosis. Very closely resembles asexual Andricus, differs in antennae 2.0 times as long as
head+mesosoma, notauli incomplete in the anterior 1/3, tarsal claws are simple, while in asexual
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Andricus notauli usually complete, antennae are less than 2.0 times as long as head+mesosoma,
tarsal claws are usually toothed.
Biology. Only the asexual generation is known.
Distribution. Four species are known: Aphelonyx cerricola (Giraud, 1859) from Western
Palaearctic, A. crispulae Matsumura, 1920, A. glanduliferae Matsumura, 1920, and A. acutissimae
Monzen, 1953 from Japan (Monzen 1953), however, the last three Japanese species must be revised.
Atrusca Kinsey, 1929
Cynips (Atrusca) Kinsey 1930: 276. Type species: Cynips (Atrusca) dugesi var. simulatrix Kinsey. Original
designation. Weld (1952a) gave it genus status.
Diagnosis. The malar space lacks sulcus; the scutum is reticulate; the forewing possesses dark
spots and/or dark stripes along veins, the radial cell is 2.0-2.5 times as long as broad, the 4
th
abscissa of Rs is strongly angulate (Fig. 31). The scutum and the pronotum laterally have dense
setae; the projecting part of the ventral spine of the hypopygium is at least 3-4 times as long as
broad, subapical setae always long and dense, reaching far beyond the apex of the spine. Most
closely resembles Cynips (= Antron and Besbicus), however, differs in diagnostic characters given
above. See also Diagnosis to Cynips.
Comments. On the basis of a short radial cell and the absence of a malar sulcus, five species
from the Sphaeroteras genus herein are transferred into Atrusca: A. carolina (Ashmead), comb.
nov., A. rydbergiana (Cockerell), comb. nov., A. texana (Ashmead), comb. nov., A. trimaculosa
(McCracken and Egbert), comb. nov., and A. unica (Weld), comb. nov.; one Xanthoteras species,
A. pulchripenne (Ashmead), comb. nov. Dailey and Menke (1980) indicated that Antron
clavuloides Kinsey must be placed in Xanthoteras. In our opinion, it must be transferred to
Atrusca: A. clavuloides (Kinsey), comb. nov. because the ventral spine of the hypopygium is long
and the radial cell is short, the 2nd abscissa of Rs is strongly angulate.
Biology. Only asexual generations are known currently that induce rounded detachable leaf galls.
Distribution. North and Central America.
Bassettia Ashmead, 1887
Bassettia Ashmead 1887: 146. Type species: Bassettia floridana Ashmead, designated by Ashmead (1903).
Types examined.
Diagnosis. Closely resembles Plagiotrochus, however, the head is 2.0-2.5 times as broad as
long from above (Fig. 32), higher or equal to width in front view; the malar space lacks radiating
striae (Fig. 33); the female forewing margin lacks cilia (Fig. 36); the median propodeal area is
narrow, limited by nearly straight or slightly outward bent lateral propodeal carina, without or with
fragmented median longitudinal carina (Fig. 37); F1 of the male antenna is nearly straight, not
incised. In Plagiotrochus, the head is less than 2.0 times as broad as long from above (Fig. 34); as
broad as high, or broader than high (transverse), but it is never higher than broad in front view; the
malar space has radiating striae (Fig. 35); the female forewing margin has cilia; the central
propodeal area is broad, limited by a strongly outward bent lateral propodeal carina, with complete
or partially complete median longitudinal carina (Fig. 38); F1 of the male antenna is strongly
incised and broadened distally. The asexual females of two Bassettia species (B. ligni Kinsey and
B. pallida Ashmead) and one Plagiotrochus species (Plagiotrochus australis (Mayr)) share a
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unique character within the Cynipini: the vertex has a deep longitudinal depression from the
median ocellus to the antennal sockets, with or without a carina at the bottom.
Another closely related genus is Loxaulus Mayr, which can be easily distinguished from
Bassettia by the structure of the propodeum similar to Plagiotrochus and the scutum is very finely
transversely sculptured (Melika & Abrahamson 2000a).
Figures 39–60 39–40, Belonocnema quercusvirens: 39, fore leg, 40, forewing. 41–42, antenna,
asexual female: 41, Biorhiza pallida; 42, Trigonaspis quercusforticorne. 43, Cynips quercusfolii,
asexual female, head from above. 44–45, B. pallida, asexual female, head: 44, from above;
45, front view, asexual female. 46–49, Trigonaspis megaptera, sexual generation, head:
46, female, from above; 47, female, front view; 48, male, from above; 49, male, front view.
50–52, thorax, dorsal view: 50, Callirhytis glandium; 51, Bassettia pallida, 52, Plagiotrochus
quercusilicis. 53, C. glandium, head, front view. 54–55, Hind tarsus: 54, C. quercusfolii, asexual
female; 55, B. pallida, asexual female. 56–57, Antenna: 56, C. quercusfolii; 57, B. pallida.
58–59, first four segments of antenna: 58, Disholcaspis quercusmamma, female
59, D. quercusmamma, male; 60, Acraspis gemula
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Bassettia also resembles some Callirhytis species, however, the head is greater or equal to its
width; the mesosoma is compressed dorso-laterally, usually 1.3-1.5 times as long as high in lateral
view; the scutum 1.5-2.0 times as long as broad (Figs 32-33), while in Callirhytis the head is
transverse, broader than high (Fig. 53); the mesosoma is arched, if slightly compressed dorso-
laterally, then less than 1.2-1.3 times as long as high in lateral view; the scutum equal or only
slightly longer than broad.
Comments. Four species from Bassettia must be transferred to Callirhytis: C. aquaticae
(Ashmead), comb. nov., C. ceropteroides (Bassett), comb. rev., herberti (Weld), comb. nov., and
C. quercuscatesbaei (Ashmead), comb. nov.
Biology. Alternate asexual and sexual generations are known. Rosenthal & Koehler (1971) and
Evans (1972) described the sexual generation of B. ligni Kinsey. The present authors also found
sexual Bassettia in Florida. The asexual generation induces stem galls with cells hidden under the
bark of twigs. The sexual generation induces small oval swellings on leaves, which protrude both
sides.
Distribution. Currently five Bassettia species are known, all from North America north of
Mexico.
Belonocnema Mayr, 1881
Belonocnema Mayr 1881: 16. Type species: B. Treatae Mayr. Original designation. Types examined. Ashmead
1885 (synonym of Dryorhizoxenus).
Dryorhizoxenus Ashmead 1881: xxv. Type species: D. floridanus Ashmead. Original designation. Monotypic.
Types examined. Ashmead 1886 (synonym of Belonocnema).
Diagnosis. Readily distinguished from all other Cynipini genera by the apex of the foretibia
extending far beyond the base of the foretarsomere I (Fig. 39); 4th abscissa of Rs is strongly
angulate, the forewing possess a short radial cell and narrow dark stripes prolong veins (Fig. 40).
Comments. It appears that B. kinseyi Weld, known to induce detachable pea-like leaf galls on
Q. virginiana and distributed in Texas only, is the alternate generation of B. treatae, known to
induce root galls on Q. virginiana and distributed from Florida and Georgia to Texas. Thus,
B. kinseyi is a synonym of B. treatae (Lund, Ott & Lyon 1998). A congeneric species,
B. quercusvirens (Osten Sacken) is known from Florida and Georgia only. Both species,
B. quercusvirens and B. treatae (asexual generations) induce pea-like leaf galls on Q. virginiana,
which are indistinguishable. However, in B. quercusvirens females, the spur on the foretibia is as
long as the metatarsus, 2.0 times as long as the furcula; the midtibia possess a distinct spur, while
in B. treatae the spur on the foretibia is 0.25 times as long as the metatarsus and is not longer than
the furcula; the midtibia lacks a spur. So, it is questionable, if B. quercusvirens and B. treatae are
distinct species or only different geographical races of the same species.
Biology. Recently alternation of generations for B. treatae Mayr was established experimentally
(Lund, Ott & Lyon 1998).
Distribution. USA only (Florida, Georgia and Texas). Only two species are known.
Biorhiza Westwood, 1840
Biorhiza Westwood 1840: 56. Type species: Cynips aptera Fabricius, 1793. Designated by Westwood
(1840).
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Apophyllus Hartig 1840a: 185, 193. Type species: Cynips aptera Fabricius. Designated by Hartig (1840a).
Mayr 1881 (synonym of Biorhiza). Monotypic.
Teras Hartig 1840a: 185, 193. Type species: Cynips quercus
terminalis Fabricius, 1798. Designated by
Hartig (1840a). Monotypic. Mayr 1881 (synonym of Biorhiza).
Heterobius Guerin-Meneville 1865: 138. Dalla Torre (1893), Dalla Torre & Kieffer (1910), Rohwer & Fagan
(1917) (synonym of Biorhiza).
Dryoteras Foerster 1869: 331, 334. Type species: Cynips quercus
terminalis Fabricius, 1798. Designated by
Foerster (1869). Mayr 1881 (synonym of Biorhiza).
Sphaeroteras Ashmead 1897a: 67. Type species: Biorhiza mellea Ashmead, 1887. Designated by Ashmead
(1897a). Mayr (1902), Beutenmueller (1909), Dalla Torre & Kieffer (1910) treated Sphaeroteras as a
synonym of Biorhiza. Weld (1951) reestablished the validity of this genus. Types examined. New
synonym.
Beutenmueller (1909) erroneously treated Xanthoteras and Phylloteras also as synonyms of Biorhiza, while
Dalla Torre (1893) and later Dalla Torre & Kieffer (1910) erroneously treated also Philonix Fitch as a
synonym of Biorhiza.
Diagnosis. The asexual generation of Biorhiza most closely resembles Trigonaspis. Differs in
having a more dorso-ventrally flattened mesosoma, the frons has longitudinal carina or elevation
which reach between antennal sockets; F1 is only 1.2-1.5 times as long as F2 (Fig. 41); the
scutellum does not or slightly overhangs the metascutellum; the dorsal median part of the
pronotum is longer and more pronounced; the propodeum is nearly in the same plane as the rest of
the mesosoma or only slightly declined; the projecting part of the ventral spine of the hypopygium
is 1.0-1.5 times as long as broad, while in Trigonaspis the mesosoma is strongly arched anteriorly;
the frons lacks elevation; F1 is 1.6-2.0 times as long as F2 (Fig. 42); the scutellum strongly
overhangs the metascutellum; the pronotum is shorter in dorsal median line, the propodeum
declines strongly, the projecting part of the ventral spine of the hypopygium is 2.0-2.5 times as
long as broad.
The sexual generation of Biorhiza differs from Trigonaspis in that the head is straight between
eyes from above, not lunate, ocelli smaller, the malar space without sulcus (Figs 43–45); scutellar
foveae is absent, only a transverse groove is present anteriorly, while in Trigonaspis the head is
lunate between eyes from above; ocelli larger; the malar space with deep sulcus (Figs 46–49); an
indistinct ridge separates scutellar foveae, the bottom of which is smooth and shiny.
Some sexual Andricus females are very similar to Biorhiza but differ in that the hind tarsomer
II is equal or longer than tarsomer V; the scutellum is smooth or delicately reticulate; antennae are
filiform, antenna nearly equally broad through entire length, while in Biorhiza the hind tarsomere
II is nearly 2.0 times as short as tarsomere V; the scutellum is dull rugose; antennae shorter,
subsequent flagellomeres broadento the apex.
Comments. Sphaeroteras. Ashmead (1897a) distinguished his newly described Sphaeroteras
genus from “the true Biorhiza in having no carina on the frons between the antennae, in having
only 13-jointed antennae, by the scutellum being rounded, and finally by the hind tarsi being much
shorter than the tibiae.” In the asexual generation of Biorhiza pallida Olivier, the first character
varies extraordinarily from a distinct strong carina to a delicate elevation only. Biorhiza nawai
(Ashmead, 1904) (= B. weldi Yasumatsu & Matsuda, 1955, synonym in Pujade-Villar, Ros-Farré
& Melika 2002, in press) described from Japan also has a weak elevation only. Biorhiza pallida
and B. nawai have 14-segmented antenna. However, even the type specimen of S. melleum has an
indistinct suture that separates F11 from F12; antenna of one of the syntypes of Ashmead’s
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melleum has a distinct suture that separates F12. Sphaeroteras caepuliforme (Beutenmueller) has
14-segmented antenna. In Biorhiza, the shape of the scutellum varies from rounded to slightly
elongated posteriorly; in some specimens the scutellum is more rounded than in the type specimen
of melleum. In the majority of examined S. caepuliforme specimens, scutellum is slightly elongated
posteriorly. The hind tarsus in the asexual B. pallida females is shorter than tibia; in the sexual
females, which may be also apterous, the length of hind tarsus is nearly equal to the tibia. Weld
(1951, 1952a) and later Burks (1979) placed 8 species into the genus Sphaeroteras. Only two of
them belong to Biorhiza: B. mellea Ashmead, comb. rev. and B. caepuliformis Beutenmueller,
comb. rev., all other species must be transferred into Atrusca, based on the absence of the malar
sulcus, short radial cell and strongly angulate Rs (see Atrusca above).
Sphaeroteras ocala (Weld) is known from Florida only and from the bisexual generation
known to induce root galls on white oaks. It closely resembles asexual Atrusca because the
forewing possesses dark stripes along veins, the radial cell is only 2.0-2.5 times as long as broad,
the 2nd abscissa of Rs is strongly angulate. However, the ventral spine of the hypopygium is very
short, less than 2.5 times as long as broad, which resembles the sexual Cynips. Thus, the status of
this species is currently uncertain.
Also two species from Xanthoteras must be transferred to Biorhiza: eburnea (Bassett), comb.
nov. and emoryi (Ashmead), comb. nov. (see also in Trigonaspis below).
Biology. Alternation of asexual and sexual generations is known.
Distribution. Holarctic. Six species are currently known: 4 from North America north of
Mexico (herein transferred Sphaeroteras and Xanthoteras species), one species from Western
Palaearctic and one species from Japan and Far East of Russia.
Callirhytis Foerster, 1869
Callirhytis Foerster 1869: 331, 335. Type species: C. hartigi Foerster. Original designation. Monotypic.
Eusymphagus Dettmer 1930: 54, 55, 56, 57. Type species: E. bellus Dettmer. Original designation. Dettmer
1933 (synonym of Callirhytis). Weld 1952a (synonym of Andricus, because tarsal claws with basal lobe).
Nieves Aldrey 1992 (revision of the European species).
Diagnosis. A transversely rugose scutum characterizes only three Cynipini genera, Bassettia,
Callirhytis, and Plagiotrochus (Figs 50–52). The diagnostic characters that separate Callirhytis
from Bassettia and Plagiotrochus are discussed above (see Diagnosis to Bassettia). Additionally
Callirhytis differs from Plagiotrochus in having a transverse head (front view), the malar space
with a sulcus (Fig. 53); and forewing margins lack cilia. In Plagiotrochus, the head in front view is
higher than broad, the malar space with radiating striae but without distinct malar sulcus (Fig. 35),
and forewing margins are ciliate. The central propodeal area in Callirhytis is usually narrower than
in Plagiotrochus, the latter in both generations has a very broad central propodeal area with
strongly bent lateral longitudinal carinae, the area is usually smooth and delicately striate, except
P. marianii, in which the central median area has strong parallel striae, but the area is large, broad.
Comments. Mayr (1881), Ashmead (1885), Cameron (1893), Dalla Torre (1893), Kieffer
(1897-1901) and many others treated Callirhytis as a subgenus of Andricus, which differed in
possessing simple tarsal claws, while the subgenus Andricus (Andricus) had tarsal claws with basal
lobe. Mayr (1902) restored the generic status of Callirhytis and Dalla Torre & Kieffer (1910)
followed this convention. The main diagnostic character for the reestablished Callirhytis genus
was the simple tarsal claw. Weld (1930) wrote “in the original description of the genotype,
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Callirhytis hartigi Foerster, it is not stated whether the tarsal claws are toothed.” It is puzzling why
no attention was given to the transversely rugose scutum; a diagnostic character of Callirhytis
described by Foerster as the main diagnostic feature of the genus. As a consequence of
misunderstanding the character’s importance, the North American Callirhytis genus, in our
opinion, currently includes more Andricus than Callirhytis species. Presence or absence of the
basal lobes on tarsal claws is a specific nor a generic character. Nieves Aldrey (1992) in his
revision of the European Callirhytis genus showed that different species vary in the presence or
absence of toothed tarsal claws; for instance, both generations of C. glandium (Giraud) and the
asexual C. bella (Dettmer) have tarsal claws with basal lobes, while other species have simple
tarsal claws.
Of the 115 species placed into Callirhytis in North America north of Mexico (Burks 1979),
only 15 species have the scutum transversely sculptured as in the typical Western Palaearctic
Callirhytis species: cedrosensis Dailey & Sprenger, corrugis (Bassett) (= defecta Kinsey),
eldoradensis (Beutenmueller), electrea Weld, flora Weld (= C. milleri, the asexual generation
(Dailey, Perry and Sprenger 1974), fructicola Ashmead, fructuosa Weld, intersita Weld, lapillula
Weld, medularis Weld, morrisoni (Ashmead), perrugosa Weld, petrina Weld, petrosa Weld,
quercusmedullae (Ashmead).
Two Callirhytis species, quercuspomiformis (Bassett) and quercusrugosa (Ashmead), herein
are transferred to the Amphibolips genus (see Amphibolips above); all other species in our opinion
belong to the genus Andricus.
Seven species of Andricus known from North America north of Mexico have the scutum
transversely sculptured and, thus must be transferred into Callirhytis genus: albobalani
(Weld), comb. nov., chrysobalani (Weld), comb. nov., coortus (Weld), comb. nov., coronus
(Beutenmueller), comb. nov., montezuma (Beutenmueller), comb. nov., rhizoxenus Ashmead,
comd. rev., and wheeleri (Beutenmueller), comb. nov.
Two species of Bassettia are transferred to Callirhytis: ceropteroides Bassett and herberti
(Weld) (see Bassettia above).
Biology. Alternate asexual and sexual generations are known. The asexual generations are
known to induce galls on/in acorns, while the sexual generations develop in stem swelling-like
galls in young twigs or in cells hidden under the bark in twigs.
Distribution. Holarctic.
Chilaspis Mayr, 1881
Chilaspis Mayr 1881: 6, 32. Type species: Andricus nitida Giraud, 1882. Designated by Mayr (1881). Pujade-
Villar, Ros-Farré & Melika (2002, in press) (revision of the genus).
Diagnosis. The genera Cynips and Biorhiza resemble Chilaspis in having a smooth
mesoscutum and mesopleuron, but they possess malar sulcus, and the malar space lacks striae at
the base of the clypeus. Chilaspis also resembles Plagiotrochus, however, it differs in possessing a
smooth scutum and mesopleuron. Chilaspis is very closely related to Dryocosmus. In Chilaspis,
striae on the frons are indistinct or weak, radiating from the clypeus in the malar space and in the
lower face only; the vertex and occiput are smooth or very weakly coriaceous; the scutellum
usually is uniformly smooth or weakly sculptured in the central part and occasionally with some
wrinkles that prolong the marginal carina; scutellar foveae are separated by a distinct carina; the
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pronotum is smooth in females, sometimes with indistinct striae in males, while in Dryocosmus
striae radiate from the clypeus to one half of the eye height, some reaching the antennal foramina;
the vertex and occiput are sculptured, sometimes strongly coriaceous or rugose; the scutellum is
uniformly wrinkled; scutellar foveae are separated or not by a weak carina; the pronotum,
especially in females, has long and distinct striae in the posterior lateral portion.
Closely related also to the genus Cynips, Clilaspis differs in the head, the scutum and scutellum
are smooth and shiny, without reticulation, while in Cynips the scutellum and/or the head are
reticulate or coriaceous.
Biology. Alternate asexual and sexual generations are known. The asexual generation emerges
from rounded detachable leaf galls, while the bisexual generation develops in catkin galls.
Distribution. Three species are known from the Western Palaearctic only (Pujade-Villar, Ros-
Farré & Melika 2002, in press). Known from Europe, North Africa, Israel, and Iran.
Cynips Linnaeus, 1758
Cynips Linnaeus 1758: 553. Type species: Cynips quercus-folii Linnaeus, 1758. Designated by Westwood
(1840).
Dryophanta Foerster 1869: 331, 335. Type species: Cynips quercus-folii L. Designated by Foerster (1869).
Monotypic. Rohwer & Fagan 1917 (synonym of Cynips).
Antron Kinsey 1930: 180. Type species: Cynips (Antron) echinus var. schulthessae form schulthessae Kinsey.
Original designation. Weld (1952a) gave the genus status. Kinsey (1930) proposed as a subgenus of
Cynips. Types examined. New synonym.
Besbicus Kinsey 1930: 222. Type species: Cynips (Besbicus) multipunctata var. conspicua Kinsey. Original
designation. Weld (1952a) gave the genus status. Kinsey (1930) proposed as a subgenus of Cynips. Types
examined. New synonym.
Diagnosis. Asexual females are fully winged; the ventral spine of the hypopygium is short,
broadened at the apex, with dense, long subapical setae that reach far beyond the apex of the spine
and form a dense truncate tuft (Fig. 15). Sexual females resemble those of Biorhiza, but differs in
that the hind tarsomere II is equal or only slightly shorter than tarsomere V (Fig. 54); the head is
narrower from above (Fig. 43), antennae filiform, subsequent flagellomeres of the same width
(Fig. 56); the transscutal articulation is bent; the scutellum is longer, while in Biorhiza females the
hind tarsomere II is shorter than V (Fig. 55); the head is broader from above (Fig. 44), antennae
are shorter, subsequent flagellomeres broadento the apex (Fig. 57); the transscutal articulation is
straight; the scutellum is shorter. Males differ from Biorhiza in that they have F1 straight, not
incised, while in Biorhiza F1 is strongly incised and enlarged distally. Males are also very similar
to Acraspis and we are unable to find characters to distinguish them. See also Diagnosis to
Acraspis and Biorhiza.
Comments. Mayr (1881) synonymized Liodora Foerster to Dryophanta. Dalla Torre (1893)
treated it as a separate genus with one species, L. sulcata Foerster. Dalla Torre & Kieffer (1910)
synonymized it with their “Diplolepis L. Geoffr.” After revising Foerster’s types of L. sulcata,
Weld (1930) concluded that they are “not congeneric with the sexual generation of Diplolepis folii
(L.)” and he restored it to generic status. See Liodora also above in Andricus.
Kinsey (1936) in his revision of the genus Cynips divided it into six subgenera: Cynips
(European species), Antron, Besbicus, Atrusca, Philonix, and Acraspis. Under the “mellea”
species-complex in the Acraspis subgenus, he included also all known Sphaeroteras species. It is
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possible that Acraspis and Philonix might be synonyms of Cynips, however, more information on
the alternation of generations is necessary. See also Diagnosis to Acraspis and Atrusca.
Antron and Besbicus. Weld’s (1952a) key is incorrect in separating Besbicus from Cynips on
the basis of structural differences in the ventral spine of the hypopygium (couplet 59). The
structure of the ventral spine of both genera is identical (Fig. 19). The only diagnostic character
given by Weld (1952a) for separation asexual generations of Antron and Cynips is the presence of
smoky spots on forewing in Antron and their absence in Cynips. Kinsey (1936) described that the
darker spotting on the forewings of Cynips (in his understanding of the genus) can vary strongly
from no spots to numerous fused spots. Dark stripes along veins are typical for the European
Cynips species. The peculiarities of Rs configuration, carinae on propodeum, width of genae,
notauli completeness, and pubescence of the scutum, given by Weld (1952a) are specific rather
than generic -level characters.
Alternation of generations occurs in Antron and Besbicus, however, the only Besbicus species
known to have an alternate sexual generation is B. mirabilis (Kinsey) (Evans 1967). The adults of
sexual B. mirabilis are entirely congeneric with the sexual generations of Cynips. Two species of
Antron were listed among those with alternating asexual and sexual generations, A. douglasii
(Ashmead) and A. quercusechinus (Osten Sacken). Andricus ribes Kinsey was listed as the sexual
generation of A. quercusechinus (Weld 1951; Burks 1979). Weld (1952a) described for this
species that “on circumstantial evidence this is considered the alternating generation of echinus”.
There is no evidence for this. Kinsey (1922a) did not mention that A. ribes might be the sexual
generation of any other cynipid wasp in the original description. Kinsey (1930) gave no acceptable
evidence for the synonymization of A. ribes to A. quercusechinus. However, A. ribes is entirely
congeneric with sexual Cynips and, of course, is the sexual generation of one of the Californian
Cynips (Antron) species. However, it is too early to state that it is the sexual generation of
A. quercusechinus; this must be proved.
Weld (1951) and Burks (1979) listed Dryophanta lobata as the sexual generation of Antron
douglasii (Ashmead), although, it was not mentioned in the original description (McCracken &
Egbert 1922). Kinsey (1930) synonymized A. lobata to A. douglasii on circumstantial grounds. An
examination of paratypes of females and a male [Paratype No. 53990, USNM] as well as the
original description (McCracken & Egbert 1922) indicates that it is not a bisexual Cynips. There
are no other bisexual cynipids known that have such broad, short ventral spines of the
hypopygium. Consequently, the status of D. lobata is uncertain.
The following species must be transferred from Besbicus to Cynips: conspicuus Kinsey, comb.
rev., heldae Fullaway, comb. rev., indictus Kinsey, comb. rev., leachii Kinsey, comb. rev.,
maculosus Weld, comb. rev., mirabilis Kinsey, comb. rev., multipunctatus (Beutenmueller),
comb. nov., tritior Kinsey, comb. rev., as well as the following Antron species: acraspiformis
(Weld), comb. nov., douglasii (Ashmead), comb. nov., dumosae Kinsey, comb. rev., magdalenae
(Weld), comb. nov., plumbeum Weld, comb. rev., quercusechinus Osten Sacken, comb. rev.,
quercusnubila Bassett, comb. rev., russus Kinsey, comb. rev. These species are known from
North America north of Mexico. We have not analyzed Besbicus and Antron species mainly
described from Mexico by Kinsey. They require a detailed revision.
We also transfer one species from the genus Xanthoteras to Cynips: C. pulchellum
(Beutenmueller), comb. nov.
Biology. Alternation of asexual and sexual generations is known. The bisexual generation
induces detachable leaf galls, while the asexual generation induces tiny small bud galls.
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Distribution. Holarctic. One species, Cynips staminobia Kovalev, 1965, was described on the
basis of males reared from catkin galls on Quercus mongolica from Far East of Russia (Kovalev
1965).
Disholcaspis Dalla Torre & Kieffer, 1910
Disholcaspis Dalla Torre & Kieffer 1910: 371. Type species: Callaspidia quercus-globulus Fitch, 1858.
Designated by Ashmead (1903). Types examined.
Holcaspis Mayr 1881: 35. Name preoccupied by Chaudoir in Coleoptera.
Diagnosis. Asexual females. Robust specimens, the ventral spine of the hypopygium is short,
the projecting part is a maximum of 2.0-3.5 times as long as broad or much shorter, subapical setae
are long, dense, reaching far beyond the apex, but never forming a dense truncate tuft; notauli
incomplete; usually scutellum has a transverse impression anteriorly, scutellar foveae absent, if
present then indistinct. See also Diagnosis to asexual Andricus. Sexual generation
. The mesosoma
is bare, with setae on the pronotum laterally only; the head is narrower than the mesosoma; the
scutum anteriorly is microreticulate or coriaceous; the scutellum is elongated, gradually and very
slightly depressed toward the transscutal articulation, without foveae or transversal groove; in
females, F1 is 1.6-1.8 times as long as scapus+pedicellum, in males, F1 is 2.0 times as long as
scapus+pedicellum (Figs 58-60). See also Diagnosis to sexual Acraspis and Cynips.
Comments. There are seven known species of Disholcaspis from the western United States:
D. chrysolepidis (Beutenmueller), conalis Weld, corallina (Bassett), plumbella Kinsey, sulcata
(Ashmead), truckeensis (Ashmead), and washingtonensis (Gillette), which differs from other
species of this genus by the ventral spine of the hypopygium that is broad throughout its entire
length, not tapering to a point at the apex; length of the projecting part of the spine is less or equal
to its width; Rs is slightly curved in the apical one third and slightly expanded, median propodeal
carinae bent, lyre-shaped, while in all other Disholcaspis species, the ventral spine is longer,
needle-like, the projecting part at least 2.0-3.5 times as long as broad, Rs is straight, the radial cell
is slightly longer; propodeal carinae are fragmented or lacking entirely.
Examination of types of some North American asexual Andricus species showed that there are
three species: lasius (Ashmead), reniformis McCracken & Egbert, and spectabilis Kinsey; and also
one Adleria species known from Mexico, lapiei (Kieffer) (= Holcaspis weldi Beutenmueller, 1911,
not Cynips weldi Beutenmueller, 1918), which differs from “typical” Disholcaspis, Andricus, and
Adleria. They agree in their morphology with the mentioned seven Disholcaspis species and form
the same species-group.
This group of 11 species has caused problems in their placement. Weld (1952a) “provisionally
placed in Disholcaspis” the seven Disholcaspis species mentioned above and wrote that “these
species might well form a new genus if the life cycle of any one of them were known.” Three
Andricus species: lasius, reniformis, and spectabilis were “placed” in Andricus. Burnett (1977)
placed 5 species (conalis, corallina, plumbella, sulcata, washingtonensis) in his new genus Weldia
and described a new species, californicus (the name is not valid, since it was not published and the
name Weldia was preoccupied by Yoshimoto (1962) for Eucoilidae). On the basis of the presence
of complete notauli, Dailey & Menke (1980) moved truckeensis to Andricus. Examination of a
series of this species at the USNM indicated that notauli vary, notauli can reach the pronotum but
also can disappear in anterior one third of scutum. Andricus truckeensis is similar to lasius,
reniformis, and spectabilis in possessing a very short ventral spine of the hypopygium and all the
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species occur on oaks of the subgenus Protobalanus. McCracken & Egbert (1922) suggested that
reniformis might be a variety of truckeensis. Dailey & Menke (1980) stated “additional studies
may indicate that these four species should be placed in a new genus.” Burnett (1977) considered
chrysolepidis and truckeensis as a distinct monophyletic group that forms a separate genus
differing from the above mentioned seven Disholcaspis species. Adleria lapiei (Kieffer, 1911)
originally was described in Disholcaspis (Kieffer 1911), but later Weld (1952a) “placed” it in
Adleria. Kinsey (1937a) suggested that D. lapiei Kieffer was “probably a synonym of D. weldi
(Beutenmueller)”. A Weld note in the general collection of USNM included in a box containing
lapiei Kieffer (= Holcaspis weldi Beut.), offers “Not a true Disholcaspis. Test Weld 1936” and
later Weld (1952a) treated them as synonyms. Burnett (1977) provisionally treated lapiei as
Adleria.
Consequently, the following species are transferred to Disholcaspis: lasius (Ashmead), comb.
nov., reniformis McCracken & Egbert, comb. nov., spectabilis Kinsey; comb. nov., and lapiei
(Kieffer), comb. nov.
One species from the current North American Adleria is transferred herein to Disholcaspis:
arizonicus (Cockerell), comb. nov. (see in Andricus).
Biology. Alternation of asexual and sexual generations is known. The sexual generation
usually produces detachable, woody bullet-like stem and subterranean or root galls; the asexual
generation induces small thin-walled bud galls of the same shape and structure as those of Cynips,
Acraspis and many Andricus. For a long time only the asexual generation was known. Evans
(1972) experimentally discovered the sexual generation of D. eldoradensis (Beutenmueller). We
reared females and males from tiny pale grayish-white bud galls on Quercus bicolor Willd.
collected in Pennsylvania (Bucknell Natural Area, Northumberland County, spring, 1996). Trees
were heavily infested with galls of D. quercusmamma (Walsh) while other Disholcaspis species
were not found in this area. The morphologies of these reared females and males agree with those
of sexual D. eldoradensis. We believe these individuals represent the sexual generation of
D. quercusmamma.
Distribution. North and Central America.
Dryocosmus Giraud, 1859
Dryocosmus Giraud 1859: 353. Type species: D. cerriphillus Giraud. Original designation. Monotypic.
Pujade 1985 (revision of the Western Palaearctic species).
Entropa Foerster 1869: 330, 334. Type species: E. lissonota Foerster. Original designation. Monotypic. Mayr
1881 (synonym of Dryocosmus).
Diagnosis. Most closely resembles Chilaspis. Diagnostic characters for separation of
Dryocosmus from Chilaspis are given in Diagnosis to Chilaspis above.
Comments. This genus needs a revision, especially the Nearctic and Eastern Palaearctic
species; more precise and strict diagnostic generic limits must be established (Pujade-Villar, Ros-
Farré & Melika 2002, in press).
Biology. The North American Dryocosmus species are morphologically uniform with the
two known European species and with species described from Asia: D. japonica (Ashmead),
D. hakonensis (Ashmead), D. nawaii (Ashmead), D. mitsukurii (Ashmead), and D. kuriphilus
(Yasumatsu). The latter species threatened the chestnut industry of Japan and Korea and was
discovered in Georgia, USA in 1974 (Payne 1978; Payne et al. 1975). This is the only species of
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Cynipini known to associate with Castanea. Another species, known from Oregon and California,
Dryocosmus castanopsidis (Beutenmueller) induces galls on catkins of Castanopsis chrysophylla
and C. sempervirens. Several Eastern Palaearctic species are known only from either asexual or
sexual generation and probably some could be paired in alternate generations (Pujade-Villar 1985).
Life cycles of these species must be identified in order to know the exact number of species and
their limits.
Distribution. Holarctic distribution with 16 known Nearctic (Burks 1979), 2 Western Palaearctic
and 6 Eastern Palaearctic species.
Eumayria Ashmead, 1887
Eumayria Ashmead 1887: 147. Type species: E. floridana Ashmead, 1887. Designated by Ashmead (1903).
Trisolenia Ashmead 1887: 142. Type species: T. saltata Ashmead, 1887. Designated by Ashmead (1903).
Rohwer & Fagan 1917 (name preoccupied by Ehrenberg in 1861 for Protozoa). Dalla Torre & Kieffer
1910 (synonym of Andricus).
Trisoleniella Rohwer & Fagan 1917: 377. New name for Trisolenia. Melika & Abrahamson 1997b (synonym
of Eumayria).
Diagnosis. Head is 1.5-1.7 times broader than long from above in the sexual generation and is
2.5-2.8 times as broad as long in asexual females; malar sulcus absent (Figs 61-64); thorax is
flattened dorso-ventrally, not arching in the anterior part; scutum is slightly longer than broad (Fig.
65), finely coriaceous or macroscopically punctate, never transversely sculptured; base of tergum
II has pale felt-like ring of dense short setae, interrupted dorsally (in males dense short pale setae
present only ventrally and ventro-laterally). See also the Diagnosis to Eumayriella Melika &
Abrahamson below.
Comments. In Burks (1979), 4 species of Eumayria were listed, from which Eumayria
eldoradensis was transferred into Callirhytis (Dailey et al. 1974) and E. longipennis to Andricus,
as well as four species of Trisoleniella were transferred to Eumayria (Melika & Abrahamson
1997b).
Biology. Five species are known. Eumayria floridana Ashmead is known only from a bisexual
generation and induces stem swelling-like galls, while the four other species have only asexual
generations.
Distribution. Eastern and midwestern United States.
Eumayriella Melika & Abrahamson, 1997
Eumayriella Melika & Abrahamson 1997b: 672. Type species: Eumayria invisa Weld, 1952. Original
designation.
Diagnosis. Closely resembles the brachypterous species of the former genus Trichoteras
(Andricus, part, after authors, see Andricus). In Eumayriella, thorax is flattened dorso-ventrally;
pronotum is dorsally much longer, placed in the same plane as scutum; scutum and scutellum are
pubescent, each longer than broad, scutoscutellar suture distinct, scutellum without foveae; head is
broader than thorax from above, 2.3-3.0 times broader than high; antenna filiform, F2 shorter than
F1; while in Trichoteras thorax is arched in anterior one-third; pronotum is much shorter dorsally
and placed in a different plane than scutum; each scutum and scutellum is as long as broad; head is
narrower or equal to thorax; antennae are short, F2 nearly equal F1. Eumayriella also closely
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resembles Eumayria, but the head is more transverse in front view and from above (2.3-3.0 times
broader than high, while in bisexual Eumayria it is only 1.5-1.7 times and in unisexual Eumayria –
2.5-2.8 times; malar space lacks radiating striae (Figs. 61-64, 67); antenna is filiform, long, 14-
segmented, F1 substantially longer than pedicel and scape together; all flagellomeres are much
longer than broad (Figs. 69-71).
Figures 61–72 61–62, Eumayria floridana, head: 61, from above; 62, dorsal view.
63–64, E. enigma, head: 63, from above; 64, dorsal view. 65–66, Scutum, dorsal view:
65, E. floridana; 66, Eumayriella archboldi. 67, E. archboldi, head, front view. 68, E. floridana,
gaster. 69–71, Antenna: 69, E. floridana, male; 70, E. enigma, female; 71, E. archboldi.
72, Loxaulus huberi, head, front view
Scutum and scutellum are longer than broad, with dense white setae, without median and
anterior parallel lines (Fig. 66); distinctly emarginated laterally and partially posteriorly as well;
scutellum lacks foveae, with transverse depression along scutoscutellar suture; apical one third of
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scutellum is gradually depressed toward apex and narrowed to a point that joins scutum along
median dorsal part; posterior one third of scutellar disk is the highest part, strongly convex;
sculpture of scutellar disk is very finely punctate, posterior one-fourth rugose. In Eumayria, head
is nearly as high as broad, malar space and partially frons with radiating striae; antennae are much
shorter, F1 as long as pedicel, scape as broad as long; F2 to F4 is slightly longer than broad,
subsequent flagellomeres, except last one, subequal, nearly as broad as long; scutellum with
foveae, posterior part of disk never convex; posteriorly rounded and has a dull rugose sculpture;
scutum and scutellum are bare, without dense white setae. Second abdominal segment in
Eumayriella lacks pale felt-like ring of dense short setae at base, while Eumayria has such a ring
(Fig. 68). The latter character is used also to separate several genera of Eucoilidae (Quinlan 1986)
and Figitinae (Figitidae) (Fergusson 1986).
Biology. Only asexual females are known. Galls are cell clusters, hidden under the bark of
twigs. Two species, E. archboldi Melika & Abrahamson and E. invisa (Weld, 1952) represent this
genus.
Distribution. Florida, USA.
Heteroecus Kinsey, 1922
Heteroecus Kinsey 1922b: 81. Type species: Andricus dasydactyli Ashmead, 1896. Designated by Kinsey
(1922b).
Diagnosis. The only diagnostic character provided by Kinsey (1922b), to differentiate this
genus from Andricus is the tarsal claws without lobe. Discovery of the alternation of generations in
this genus (Lyon 1963, 1984) furnished more evidence to synonymize Heteroecus to Andricus or
North American “Callirhytis” (sensu Weld 1952a).
Comments. Kinsey (1922b) wrote “Whether to interpret this group as a species or a genus is
largely a matter of individual opinion and convenience.” The validity of this genus is very dubious
and might be synonym of Andricus, if the presence or absence of tooth on the tarsal claw is
regarded as a non-generic character.
Biology. The other peculiarity of this genus is that the representatives of this genus are
distributed in California only and are associated only with Quercus chrysolepidicola.
Distribution. California, USA. Twelve species were listed by Burks (1979). Later few new
species were described, all from California (Dailey & Sprenger 1983; Lyon 1984). Thus, currently
16 species are known.
Holocynips Kieffer, 1910
Holocynips Kieffer 1910: 114. Type species: H. emarginata Kieffer. Original designation.
Diagnosis. The only distinction of Holocynips from many species of Andricus is the simple
tarsal claw. All other diagnostic characters are found in many Andricus species.
Comments. The validity of this genus is very dubious and might be a synonym of Andricus, if
the presence or absence of tooth on the tarsal claw is regarded as a non-generic character.
Biology. The asexual generation only is known. Galls are on roots or at the base of young
sprouts on white oaks only.
Distribution. Four species are known: two from the Eastern and 2 from Western Coast of the
USA.
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Loxaulus Mayr, 1881
Loxaulus Mayr 1881: 9, 12, 33. Type species: Cynips quercus-mammula Bassett, 1881. Designated by
Ashmead (1903). Monotypic. Melika & Abrahamson 2000a (revision of the genus).
Solenozopheria Ashmead 1887: 149. Type species: S. vaccinii Ashmead. Original designation. Monotypic.
Weld 1951 (synonym of Loxaulus).
Compsodryoxenus Ashmead 1896: 128. Type species: C. maculipennis Ashmead, 1896. Designated by
Ashmead (1903). Weld 1951 (synonym of Loxaulus).
Diagnosis. The head from above is broader than the mesosoma; the gena possess a malar
sulcus (Fig. 72). The scutum is usually finely transversely coriaceous; the scutellum lacks foveae,
with a transverse shallow depression (Fig. 74). The central portion of the propodeum is narrow,
limited by parallel or slightly outward bent lateral carinae and with a median longitudinal carina
and/or longitudinal striae; the median longitudinal carina in some species is indistinct, fragmented
but always present at least in anterior half (Fig. 75). The radial cell of the forewing is short and
broad, not more than 2.5 times as long as broad (except L. quercusmammula with the radial cell
2.6–2.7 times as long as broad), the forewing margin of female with or without cilia, with brown,
smoky spots (or stripes) along the areolet, 2r, Rs, and M. (Fig. 76). Tarsal claws lack tooth. The
ventral spine of the hypopygium is short, slender or needle-like; subapical setae are short and
sparse, do not reach beyond the apex of the spine and the prominent part is never more than 3.0–
3.5 times as long as broad (Fig. 73). The propodeum is similar to that of the Mediterranean genus
Plagiotrochus Mayr. However, in the latter, the central portion of the propodeum is much broader,
and is limited by the strongly outward-bending lateral carinae (Fig. 38).
Biology. The genus includes 14 species (Melika & Abrahamson 2000a). Alternate asexual and
bisexual generations are known only for L. trizonalis Weld.
Distribution. North America only.
Neuroterus Hartig, 1840
Neuroterus Hartig 1840a: 185, 192. Type species: Neuroterus politus Hartig, 1840. Designated by Ashmead
(1903).
Spathegaster Hartig 1840a: 186. Type species: S. petioliventris Hartig. Original designation. Monotypic.
Mayr 1881 (synonym of Neuroterus).
Ameristus Foerster 1869: 333. Type species: Neuroterus politus Hartig. Designated by Rohwer & Fagan
(1917). Mayr 1881 (synonym of Neuroterus).
Dolichostrophus Ashmead 1887: 129. Type species: Cynips quercusirregularis Osten Sacken, 1861.
Designated by Ashmead (1887). Dalla Torre 1893 (synonym to Neuroterus).
Neuroterus subgenus Diplobius Kinsey 1923: 27, 28-31, 35. Type species: Cynips floccosa Bassett, 1881.
Original designation.
Neuroterus subgenus Dolichostrophus Kinsey 1923: 25, 28, 32, 78. Type species: Cynips irregularis Osten
Sacken Original designation.
Neuroterus subgenus Neospathegaster Kinsey 1923: 28, 35, 121. Type species: Cynips vesicula Bassett,
1881. Original designation.
Neuroterus subgenus Neuroterus Kinsey 1923: 27, 128. The nominal subgenus of Neuroterus.
Neuroterus subgenus Pseudoneuroterus Kinsey 1923: 130. Type species: Cynips macroptera Hartig, 1843.
Original designation.
Neuroterus subgenus Spathegaster Kinsey 1923: 28, 131. Type species: Spathegaster petioliventris Hartig,
1840. Original designation.
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Neoneuroterus Monzen 1954: 33. Type species: N. kashiyamai Monzen. Original designation. New
synonym.
Repentinia Belizin & Maisuradze, 1961. In Maisuradze 1961: 28. Type species: R. lencoranica Belizin &
Maisuradze. Monotypic. Original designation. New synonym.
Diagnosis. The scutum is smooth, microreticulate, or alutaceous, shiny, emarginated
posteriorly along dorso-axillar surface; transscutal articulation is strongly emarginated laterally, if
uninterrupted then strongly curved, never straight (Fig. 14); the scutoscutellar suture is deep, the
scutellum lacks separated foveae; fully winged, the radial cell of forewing is elongated, narrow.
Usually notauli are absent or only barely traceable, however, some European species have distinct
notauli, at least in the posterior half, and one species, N. lanuginosus Giraud, has deep notauli
reaching the pronotum. The main diagnostic character that separates Neuroterus from all other
genera of Cynipini is the absence of the scutoscutellar suture. See also in Melika, Stone & Csóka
(1999).
Comments. Kinsey (1923) divided the genus into 6 subgenera. Subdivision is appropriate
since the genus includes morphologically diverse species, however, some subgenera are hardly
distinguishable. The genus Neuroterus requires revision, particularly of the North American and
Eastern Palaearctic species (particularly described from Japan). A number of new Neuroterus
species and synonymizations were made recently on the North American species (Melika &
Abrahamson 1997a).
Neoneuroterus. The only diagnostic characters given by Monzen (1954) for separation
Neoneuroterus from Neuroterus were: antennae are 14-segmented in females and 15-segmented in
males; notauli distinct and complete; head and mesosoma glabrous and smooth; the ventral spine
of the hypopygium is long and pointed. However, all these characters occur in Western Palaearctic
Neuroterus species as well (see the recent key to the Western Palaearctic Neuroterus species in
Melika, Stone & Csóka 1999). We also examined the types of Neoneuroterus boni-henrici
(Dettmer, 1934) and found no substantial characters to separate this species from other Neuroterus
species. Thus, two Neoneuroterus species are transferred to Neuroterus: N. kashiyamai (Monzen,
1954), comb. nov. and N. bonihenrici Dettmer, 1934, comb. rev. Later, Kovalev (1965) described
other Neoneuroterus species from Far East of Russia: N. spumeus Kovalev, 1965 (sexual gen.,
galls on leaves), N. nephroideus Kovalev, 1965 (sexual gen., galls in buds), and N. vernicosus
(asexual gen., galls unknown). The latter species must be transferred into Trigonaspis:
T. vernicosus, comb. nov. Two other species, N. spumeus and N. nephroideus must be revised,
their placement in Neoneuroterus, and thus in Neuroterus, is questionable.
Repentinia. Maisuradze (1961) described this species from Azerbaijan and placed it near
Neuroterus. The asexual females are known to induce stem swelling-like galls on twigs of
Quercus castaneifolia CAM. Maisuradze (1961) mentioned that the new genus closely resembles
Neuroterus and differs in that the head and mesosoma have a dense white pubescence, F1 is very
short, nearly equal F2; lateral propodeal carinae are absent. Weld (1964, ms.) wrote, “Genus can
not be placed in key – perhaps near Trichagalma.” We examined the types of this species at the
Zoological Institute in St. Petersburg (Russia) and found no substantial characters to separate it
from Neuroterus. Moreover, all the diagnostic characters given by Maisuradze (1961) for
Repentinia occur in some Neuroterus species, particularly in N. macropterus (Hartig, 1843).
Analysis of type series of Repentinia lencoranica showed that it is a syn. nova of N. macropterus.
This species is widespread in Europe on Q. cerris only, but it also has been found in Israel on
Q. ithaburensis (Sternlicht 1968), and is known from Bulgaria (Vassileva-Samnalieva 1974),
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Romania (Ionescu 1973), and we identified large series collected from Turkey and Iran (Lorestan).
Thus, it is apparent that the distribution of this species includes a large distribution within the
Western Palaearctic.
Neuroterus (Latuspina) Monzen, 1954: 35. Type species: Neuroterus (Latuspina) stirps
Monzen, 1954: 35. Original designation. The only character given by Monzen (1954) to
differentiate this subgenus and species from Neuroterus is that the projecting part of the ventral
spine of the hypopygium is not normally pointed at the apex, but dilated “like a spatula”. No
Neuroterus species are known to have distally broadened ventral spine. Consequently, Latuspina
cannot be placed into Neuroterus. Weld (1964, ms.) stated that he was unable to obtain the types
and that Dr. Yasumatsu, whom he asked to look for types, didn’t find them in Monzen collection.
Until the types are revised, the status of Neuroterus (Latuspina) Monzen will remain uncertain.
Biology. Nearly 100 described species, many are known to have alternate generations. Usually
induce tiny integral or detachable leaf galls, mono- or polythalamous, or integral stem and catkin
galls. Some species have polymorphic galls.
Distribution. Holarctic.
Odontocynips Kieffer, 1910
Odontocynips Kieffer 1910: 112. Type species: O. nebulosa Kieffer. Original designation. Monotypic.
Diagnosis. Easily distinguished from all other Cynipini by the presence of a strong apical lobe
on the hind tibia (Fig. 77). Otherwise, it is closely allied to Andricus and Holocynips.
Biology. One species, O. nebulosa Kieffer, 1910 is known from the asexual generation only. It
induces root galls on white oaks. Recently details of the biology were described (Wilson, Lester &
Edmonson 2000).
Distribution. USA.
Philonix Fitch, 1859
Philonix Fitch 1859: 783. Type species: P. fulvicollis Fitch, 1859. Designated by Ashmead (1903). Types
examined. Ashmead 1903 and Beutenmueller 1909 (Acraspis is a synonym of Philonix). Dalla Torre &
Kieffer 1910 (synonym of Biorhiza). Weld 1922 (Acraspis and Philonix distinct genera).
Diagnosis. Closely resembles Acraspis, see Diagnosis to Acraspis.
Comments. Kinsey (1930) transferred Dryophanta pallipes Bassett into Philonix and
considered it as the sexual generation of P. fulvicollis. However, Kinsey provided no justification
for this synonymy. Weld (1959) wrote that P. pallipes was “perhaps a synonym of Acraspis
gemula (Bassett)”. We compared the three specimens of P. pallipes (from Beutenmueller
collection, USNM, Washington, DC) with the paratypes of the sexual generation of A. gemula and
found no differences. Philonix pallipes is a syn. nov. of A. gemula. If P. pallipes (Bass.) is treated
as an Acraspis species, there is no evidence that species of Philonix have alternate sexual
generations. Several species of Philonix, described by Kinsey (1930, 1936) from Mexico must be
revised.
Biology. Philonix species are associated with white oaks only. Galls induced by Philonix
species differ from those of Acraspis. They are fleshy, soft, and globular, attached to the leaves,
usually the undersides, with a thick wall covered with a short very dense, felt-like pubescence.
Distribution. North America and Mexico.
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Phylloteras Ashmead, 1897
Phylloteras Ashmead 1897a: 67. Type species: Biorhiza rubinus Gillette, 1888. Designated by Ashmead
(1897a). Monotypic. Dalla Torre & Kieffer 1910 (synonym of Trigonaspis). Beutenmueller 1909
(synonym of Biorhiza). Types examined.
Xystoteras Ashmead 1897b: 260. Type species: X. volutellae Ashmead, 1897. Original designation. Monotypic.
Lyon 1993 (synonym of Phylloteras). Types examined.
Euxystoteras Lyon 1993: 138. Type species: E. campanulatum Lyon. Original designation. Types examined.
New synonym.
Diagnosis. Ant-like, apterous or brachypterous but the forewing is not longer than the
mesosoma. Similar to Zopheroteras, however, the scutellum is not knobbed; the head is equal or
even higher than broad in front view. Phylloteras poculum (Osten Sacken) has a slightly elevated
scutellum similar to Zopheroteras, but not nearly as strongly elevated. See also Diagnosis to
Zopheroteras. Closely related to Trigonaspis but has a longer, needle-like ventral spine of the
hypopygium, the propodeum is placed in the same plane as the rest of thorax or only slightly
declined, while in Trigonaspis the projecting part of the ventral spine is nearly 3.0 times as long as
broad, the propodeum declines strongly.
Comments. Lyon (1993) separated Euxystoteras from Phylloteras “by its simple tarsal claws”.
The inadequacy of this generic character argues that Euxystoteras must be a synonym of
Phylloteras. Thus, Euxystoteras campanulatum (Lyon 1993) must be transferred to Phylloteras,
P. campanulatum (Lyon), comb. nov. If data are obtained on the alternate bisexual generations of
Phylloteras, it is possible that it will join Trigonaspis, forming only a group of species rather than
a separate genus.
Biology. Asexual generation is only known. Induces small detachable leaf galls.
Distribution. Eight species are known from North America.
Plagiotrochus Mayr, 1881
Plagiotrochus Mayr 1881: 32. Type species: Cynips quercus-ilicis Fabricius, 1798. Designated by Ashmead
(1903).
Fioria Kieffer 1903a: 31. Type species: Callirhytis marianii Kieffer, 1903. Original designation. Kieffer
1903b (name preoccupied by Silvestri in 1869 for Myriapoda).
Fioriella Kieffer 1903b: 95. Melika, Ros-Farré & Pujade-Villar 2001 (synonym of Plagiotrochus).
Diagnosis. The gena in the asexual female is broadened behind the eye; the clypeus with
radiating striae, do not project as a distinct lamella between mandibles, the malar sulcus is absent;
the mesopleuron is shiny, flat in the postero-dorsal margin and ventral area; the metasoma is
compressed laterally; the scutellum is equal or only slightly longer than the metascutellum; the
propodeum forms an obtuse angle with the scutellum; the lateral propodeal carinae are strongly
bent outwards, with a more or less impressed median carina; the ventral spine of the hypopygium
is thin, with short sparse white setae not forming an apical tuft. See also Diagnosis to Bassettia,
Callirhytis, and Loxaulus. Plagiotrochus also resembles Chilaspis, however, the latter differs from
the former by a smooth scutum and mesopleuron.
Comments. All the mentioned characters are not exclusive to the genus Plagiotrochus but the
above combination does define this genus. Homoplasies are very common in Cynipini and it is
very difficult to find an exclusive diagnostic character for a given genus of Cynipidae, which is not
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shared with another genus or other genera. For example, the main diagnostic character for
recognizing Plagiotrochus in the Palaearctic area is the presence of a median propodeal carina,
which, however, is also present in some species of the Nearctic Loxaulus and Bassettia (Melika &
Abrahamson 2000a). Another character, the longitudinal depression on the vertex with a median
longitudinal carina, is present in some asexual females of Plagiotrochus but also in some North
American species, e.g., Bassettia ligni Kinsey and B. pallida Ashmead.
Plesiomorphic traits of P. semicarpifoliae (Cameron) known from the Himalayan area suggest
a Southeast Asian origin for Plagiotrochus and all the species known from the Mediterranean
region are derived forms (Bellido, Ros-Farré, Kovalev & Pujade-Villar 2000).
Weld (1926) described one species, P. suberi from California, which induces stem galls on the
introduced European cork oak, Quercus suber. Later, Esquivel & De Santis (1953) described
another asexual species, P. abdominalis, from Argentina, which is also known to induce stem
galls. Nieves-Aldrey (2001) in his Fauna Iberica suggested that both, P. suberi and P. abdominalis
are synonyms of P. amenti Kieffer (asexual generation, described as P. pardoi Nieves-Aldrey,
1985). However, Pujade-Villar (1998) regarded P. pardoi as a distinct species from P. amenti and,
thus decided that P. suberi Weld, 1926 is a valid name, while P. abdominalis Esquivel & De
Santis, 1953 and P. pardoi Nieves-Aldrey, 1985 are synonyms. Bailey & Stange (1966) wrote of
P. suberi, “the insect has been found in Switzerland and Portugal (where it probably originated)
only in the past few years.” This species life cycle has changed and its population occurs as
asexual females only. A failure in a genetic regulatory switch from thelyotoky to bisexuality may
have resulted in the deletion of the sexual generation in geographic isolation. Geographic
parthenogenesis, for example, is present in Andricus mukaigawae (Mukaigawa) and A. targionii
Kieffer in Japan (Abe 1986).
Biology. Alternation of generations is known. The asexual generation usually induces stem
galls, the cells of which are hidden under the bark of twigs, while the sexual generation usually
induces catkin galls. Some species induce leaf galls.
Distribution. Mediterranean region (Southern Europe and North Africa) – 7 sexual and 7
asexual Plagiotrochus forms are known (Bellido, Ros-Farré, Kovalev & Pujade-Villar 2000).
Nieves Aldrey (2001) listed 10 Plagiotrochus species for the Iberian Peninsula, one species,
P. vilageliui Pujade-Villar, 2000 was described from Corsica (France) (Pujade-Villar, Villemant &
Andrei-Ruiz 2000). One species, P. marianii (Kieffer) was recorded from Slovakia and Hungary
(Ambrus 1974; Melika, Ros-Farré & Pujade-Villar 2001), one species, P. semicarpifoliae is
known from the Himalayan area. Introduced to California and Argentina.
Trichagalma Mayr, 1907
Trichagalma Mayr 1907: 3. Type species: T. drouardi Mayr. Original designation. Monotypic.
Diagnosis. The ventral spine of the hypopygium is slender and very short, the projecting part
broader than long, with sparse subapical setae, reaching beyond the apex of the spine; the
metasoma is strongly compressed laterally to a knife-edge dorsally; the forewing is 1.7 times as
long as the body, with dark spots, Rs straight, the radial cell very long and narrow, (Fig. 78);
the mesosoma is densely pubescent; the propodeum is very short, without carinae. No other
Cynipini is known to have such strongly compressed metasoma laterally. In general, the head
and mesosoma resemble asexual Cynips, however, the absence of notauli, scutellar foveae
and propodeal carinae and the laterally strongly compressed metasoma clearly differentiate
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Trichagalma from Cynips. Absence of notauli and scutellar foveae; smooth mesosoma, strongly
laterally compressed gaster, the short ventral spine of the hypopygium also resemble Neuroterus.
However, the robust size, strongly arched and densely pubescent mesosoma differentiates the
genus from Neuroterus.
Comments. Ashmead (1904) described Dryophanta serratae from Japan (Sapporo) on the
basis of one female, reared by Dr. Matsumura from the gall collected from Quercus serrata. Later
Mayr (1907) described his T. drouardi, which appeared to be a synonym of D. serratae Ashmead
(Monzen 1954). Thus, the genus is known from only one robust species, T. serratae (Ashmead,
1904).
Biology. Asexual females only are known to induce detachable stem galls, similar to those of
the European Andricus serotinus (Giraud).
Distribution. Known from Japan only.
Trigonaspis Hartig, 1840
Trigonaspis Hartig 1840a: 186. Type species: T. crustalis Hartig, 1840. Original designation. Monotypic.
Xanthoteras Ashmead 1897b: 262. Type species: Biorhiza forticornis Walsh, 1864. Designated by Ashmead
(1897b). Specimens from the original Walsh’s series examined. New synonym.
Belizinella Kovalev 1965: 49. Type species: B. gibbera Kovalev. Original designation. Types examined. New
synonym.
Ussuraspis Kovalev, 1965: 53. Type species: U. nervosus Kovalev. Original designation. Types examined.
New synonym.
Diagnosis. Trigonaspis closely resembles Biorhiza, however, it differs in the presence of
arched mesosoma and shorter pronotum. Phylloteras and Zopheroteras are also very similar to
Trigonaspis, however they differ in having longer ventral spines in both genera, knobbed
scutellum in Zopheroteras, and less arched mesosoma in Phylloteras. See also Diagnosis to
Biorhiza, Phylloteras and Zopheroteras.
Comments. Dalla Torre & Kieffer (1910) treated Zopheroteras as a synonym to Trigonaspis.
Beutenmueller (1909) placed Xanthoteras into Biorhiza, while Mayr (1902) considered
Xanthoteras as a separate genus from Biorhiza (= Sphaeroteras) because of the strongly arched
mesosoma and toothed tarsal claws. The original description of Xanthoteras (Ashmead 1897b)
does not include substantial diagnostic characters to separate it from Trigonaspis. Xanthoteras
fumosum (Weld), X. pumiliventre (Bassett), X. radicola (Ashmead) were described and are known
from the sexual generations only. Although placed into Xanthoteras, they are congeneric with the
European species, T. megaptera (Panzer). The sexual generation of Trigonaspis is quite different
from that of Biorhiza, which earlier workers had synonymized as Xanthoteras (see Diagnosis to
Biorhiza). Weld (1921) treated these three Xanthoteras species as Trigonaspis and only later
transferred them to Xanthoteras.
Weld (1951, 1952a) and Burks (1979) placed 13 North American species into Xanthoteras.
Dailey & Menke (1980) synonymized X. obconicum (Weld) to X. pulchellum (Beutenmueller)
Seven Xanthoteras species here are transferred to Trigonaspis: fumosa Weld, comb. rev.,
mediocre (Weld), comb. nov., ornata Kinsey, comb. rev., pumiliventre (Bassett), comb. nov.,
quercusforticorne (Walsh), comb. nov., radicola (Ashmead), comb. nov., and teres (Weld),
comb. nov. Two species, X. eburneum (Bassett) and X. emoryi (Ashmead) herein transferred to
Biorhiza; X. pulchellum (Beutenmueller) is transferred to Cynips; X. pulchripenne (Ashmead)
transferred to Atrusca.
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Belizinella. Kovalev (1965) mentioned that his new genus is closely related to Xanthoteras,
Xystoteras and Trigonaspis and from the latter, it differs in having a 14-segmented antennae, while
in Trigonaspis antennae are 13-segmented (however, in T. synaspis antennae are 14-segmented).
He mentioned also that in Belizinella notauli are absent, however, in B. gibbera the notauli are
distinct and complete, reaching the pronotum. The galls of two newly described species B. gibbera
and B. vicina are identical in their shape, location and inner structure as the gall of T. synaspis.
Without doubt, it is a synonym of Trigonaspis and, thus, Trigonaspis gibbera Kovalev, comb.
nov. and T. vicina Kovalev, comb. nov.
Figures 73–83 73, Loxaulus huberi, female, lateral view; 74, L. masneri, female, thorax, dorsal
view; 75–76, L. huberi, female: 75, propodeum; 76, forewing. 77, Odontocynips nebulosa, hind
tibia. 78, Trichagalma serratae, forewing. 79–80, Mesosoma, lateral view: 79, Zopheroteras
guttatum; 80, Z. sphaerula. 81–82, Z. guttatum: 81, Head, front view; 82, mesosoma, dorsal view.
83, Z. sphaerula, ventral spine of hypopygium
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Ussuraspis. The diagnostic characters given by Kovalev (1965) for the separation of his newly
described genus from Trigonaspis and Xanthoteras are not satisfactory – they are identical with
those for Trigonaspis. So, Ussuraspis nervosa Kovalev, 1965 must be transferred to Trigonaspis:
T. nervosus, comb. nova.
If data are obtained on the alternate sexual generations of Phylloteras and Zopheroteras, it may
be that both genera will join Trigonaspis and will form only a distinct species groups rather than
separate genera.
Biology. Alternation of asexual and sexual generations is known. Induces detachable leaf and
root galls.
Distribution. Holarctic. We found undescribed Trigonaspis species from the material collected
in China and Malaysia.
Zopheroteras Ashmead, 1897
Zopheroteras Ashmead 1897b: 261. Type species: Acraspis vaccinii Ashmead, 1887. Designated by Ashmead
(1903). Dalla Torre & Kieffer 1910 (synonym of Trigonaspis).
Parateras Ashmead 1897b: 262. Type species: P. hubbardi Ashmead. Original designation. Monotypic. Weld
1922 (synonym of Zopheroteras).
Diagnosis. Ant-like, apterous or brachypterous species. Closely resembles Phylloteras, differs
in that the scutellum is knobbed (Figs 79-80) and the head is transverse in front view (Fig. 81); the
scutellum is elevated, with knobbed disk; notauli complete, strongly curving inward posteriorly
(Fig. 82); the ventral spine of the hypopygium is narrow, needle-like, the projecting part is never
less than 4-5 times as long as broad, subapical setae never reach beyond the apex of the spine (Fig.
83). Species are difficult to distinguish on the basis of the galls only. The adults are required for
precise identification of the species.
Comments. If data are obtained on the alternate bisexual generations of Zopheroteras, it may
be that it will join Trigonaspis and will form only a distinct group of species rather than a separate
genus.
Biology. Only the asexual generation is known. It induces small, usually rounded or elliptic
detachable leaf galls on both red and white oaks.
Distribution. Six species are known from North America (eastern and midwestern United
States only).
Conclusions
From 41 currently valid genera 15 are synonymized. Thus, 26 genera in Cynipini are valid
(Table 3). As the result, 73 comb. nov. and 26 comb. rev. are made.
Acknowledgments
We express our deepest appreciation to A. Menke for his valuable suggestions during our work
in the USNM, Washington, DC. We thank J. Abrahamson, C. Abrahamson, R. Bowman, G. Csóka,
R. Hammer, A. Johnson, I. Kralick, R. Peet, R. Roberts, P. Schmaltzer, A. Schotz, C. Winegarner,
and M. Winegarner for field and technical assistance. We thank D. Notton, N. Fergusson,
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G. McGaven, S. Schödl, L. Zombori, J. Papp, O. Kovalev, L. Diakontshuk, J. L. Nieves-Aldrey,
J. Pujade-Villar for loans and/or gifts of research material. We also thanks to F. Ronquist,
G.N. Stone, J. Pujade-Villar for their valuable suggestions and discussions on the Cynipini generic
classification. Support was provided to GM and WGA by Bucknell University’s David Burpee
endowment, the Archbold Biological Station, and NSF grant BSR-9107150 to WGA; and to GM a
Short-Term Visitor Grant, USNM/Smithsonian Institution, Office of Fellowships and Grants,
February, 1996; The CanaColl Foundation, Canadian National Collection of Insects, Ottawa, June,
1998; American Museum of Natural History, Office of Grants and Fellowships (New York City,
NY), July 1998; Theodore Roosvelt Memorial Fund, NYC, July 1998).
Table 3 Current arrangement of world genera of oak cynipid wasps (Cynipidae: Cynipini)
Proposed valid genera New synonymic genera
1.
Acraspis
Mayr, 1881 Paracraspis Weld, 1952
2.
Amphibolips
Reinhard, 1865
3.
Andricus
Hartig, 1840
Dros Kinsey, 1937, Erythres Kinsey, 1937, Liodora
Foerster, 1869, Parandricus Kieffer, 1906, Trichoteras
Ashmead, 1897
4.
Aphelonyx
Mayr, 1881
5.
Atrusca
Kinsey, 1929
6.
Bassettia
Ashmead, 1887
7.
Belonocnema
Mayr, 1881
8.
Biorhiza
Westwood, 1840
Sphaeroteras Ashmead, 1897
9.
Callirhytis
Foerster, 1869
10.
Chilaspis
Mayr, 1881
11.
Cynips
Linnaeus, 1758
Antron Kinsey, 1930, Besbicus Kinsey, 1930
12.
Disholcaspis
Dalla Torre &
Kieffer, 1910
13.
Dryocosmus
Giraud, 1859
14.
Eumayria
Ashmead, 1887
15.
Eumayriella
Melika &
Abrahamson, 1997
16.
Heteroecus
Kinsey, 1922
17.
Holocynips
Kieffer, 1910
18.
Loxaulus
Mayr, 1881
19.
Neuroterus
Hartig, 1840
Neoneuroterus Monzen, 1954, Repentinia Belizin &
Maisuradze, 1961
20.
Odontocynips
Kieffer, 1910
21.
Philonix
Fitch, 1859
22.
Phylloteras
Ashmead, 1897
Euxystoteras Lyon, 1993
23.
Plagiotrochus
Mayr, 1881
24.
Trichagalma
Mayr, 1907
25.
Trigonaspis
Hartig, 1840
Belizinella Kovalev, 1965, Ussuraspis Kovalev, 1965,
Xanthoteras Ashmead, 1897
26.
Zopheroteras
Ashmead, 1897
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ENERA OF
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