Transactions of the Society for British entomology

PART 7

TRANSACTIONS

OF THE

SOCI ETY FOR BRITISH
ENTOMOLOGY

World List abbreviation : Trans. Soc. Brit. Ent.

CONTENTS.

Claridge, M. F.

An Advance Towards a Natural Classification of Eurytomid
Genera (Hym., Chalcidoidea), with Particular Reference to

British Forms.

Date of Publication, 29th April 1961.

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TRANSACTIONS OF THE SOCIETY
FOR BRITISH ENTOMOLOGY

VOL. 14

APRIL 1961

PART VII

An Advance Towards a Natural Classification of
Eurytomid Genera (Hym., Chalcidoidea), with
Particular Reference to British Forms

By M. F. Claridge, M.A., D.Phil.

(Department of Zoology, University College, Cardiff)

Contents

Principles of Natural Classification ... ... ... ... ... 167

Consideration of Eurytomid Genera ... ... ... ... ... 169

Terminology ... ... ... ... ... ... ... ... 170

Key to Genera of Northern Palaearctic Eurytomidae ... ... 170

Classification . 171

Rileyinae ... . . 17*2

Eurytominae ... ... ... ... ... .... 172

Harmolitinae ... ... ... ... ... ... 179

Eudecatominae ... ... ... ... ... ... 181

Phylogenetic Speculation ... ... ... ... ... ... 182

Acknowledgments ... ... ... ... ... ... ... 183

References ... ... ... ... ... ... ... ... 183

Principles of Natural Classification

The most useful form of biological classification is generally
agreed to be that based on the so-called natural* system. How-

*Since preparing the manuscript for this paper Cain and Harrison
(1960, Proc. zool. Soc. Lond., 135: 1-31) have suggested that the term
natural, as applied to biological classifications, should be rejected com¬
pletely since it has been misinterpreted and confused by so many
authors. They propose the new term phenetic for natural classifications
as defined here, i.e. those based on overall similarity. However, since
it is unlikely that taxonomists will cease using such well entrenched
terminology, I incline to the view that it is better to attempt to
clarify and stabilize existing terms rather than promote new ones.

t 5

168

[April

ever, there is probably no term used in taxonomy, the meaning
of which has been so badly confused. An admirable and full
historical discussion has been given recently in a series of papers
by Cain (1958, 1959a, b, c), but it is not out of place to discuss the
matter briefly here, since it is of great importance in entomological
taxonomy.

A natural classification is one which is based on overall
resemblance (affinity) ; that is, on as many characters of all kinds
as possible. It cannot be achieved by single-character, or
virtually single-character, methods, unless the character can be
shown to be correlated with the natural grouping. The taxonomic
importance of any single character can only be judged by its
co- variation with other characters of the group. A necessary
consequence of the natural system is the possibility that any
character confined to members of a particular natural group may
be absent in one or more of its members and thus could not be
used as diagnostic of it ( vide Cain, 1954). The obvious corollary
is that if classificatory groups, particularly genera, are to be
natural, it may not be possible, in many cases, to give exclusive
definitions of them on the basis of a particular character or
character complex: a procedure which is generally considered
essential. I have briefly mentioned this problem before in con¬
nection with the Eurytomidae (Claridge, 1958a).

Most of the misunderstandings concerning the natural system
in recent literature have been concerned with the equation of
natural classifications with phylogeny, in the total absence of
adequate fossil records. In order to produce a phylogenetic
classification, detailed phylogenetic data are required. Such
data can only be obtained from fairly complete fossil records,
which are very rare and almost unknown in insects. The natural
system, based on comparative data of living forms only, may
take no account of parallel and detailed convergent evolution and
may even tend to obscure such phenomena. Undoubtedly it is
probable that in many cases, those species which in a natural
classification are said to be closely allied, are truly phylogenetically
close. However, we can never be sure, and as more and more
groups are hypothetically related the probability of the system
representing phylogeny becomes increasingly less. As Cain
(1959c), in discussing avian taxonomy, has said “ . . . . ; the
theory [of evolution] is now so strong that it will not matter
if we have to say we just don’t know the phylogeny of 80% of
passerine birds”. This applies perhaps with even greater force to
almost all insect groups.

An important consequence of the application of a natural
generic system in practice will usually be a reduction in the
number of genera in a group, since often they have been erected
only for couplets of an artificial key. There are many cases of
such single character genera in the Chalcidoidea. A good example
is A chrysopophagus Girault (Encyrtidae) distinguished from

169

1961]

Cheiloneurus Westwood only by its extruded ovipositor ( vide
Claridge, 1958b). Many species with exserted ovipositors show
greater affinity with some species of Cheiloneurus than with
others with extruded ovipositors. The value of such completely
artificial genera is very slight. However, since unfortunately it
is necessary to place a species generically before it can be
described, a taxonomist without the time and material available
to revise the group completely may have to erect more such
genera.

The great advantage of a natural classification is that it is a
reservoir of the maximum amount of information about a group,
and thus it allows at least limited generalizations to be made
about its components. Also it provides a background against
which phylogenetic speculation can be made, but such speculation
should not be allowed to affect the system.

Consideration of Eurytomid Genera

In the Eurytomidae, probably since morphological diversity
is relatively slight, there have not been so many genera described
as in some other Chalcidoid families such as the Encyrtidae.
Of workers on Palaearctic forms, Walker (1832, 1846, etc.)
recognized only four major genera, though several others were
also erected, mostly for tropical and oriental species. Thomson
(1875) recognized only the four of Walker and more recently
Ferriere (1950) has included eight in four subfamilies from a
rather wider geographic area within the Palaearctic. Many more
genera were included by Ashmead (1904) in his classification of
world Chalcid genera. Of those of Ashmead described for
Nearctic species, some have been synonymized by Peck (1951),
who recognized only fourteen genera from America north of
Mexico. Burks (1958) recognizes a further four from the same
area. I have not been able to study sufficient Nearctic material
to produce any reliable correlation with Palaearctic forms, but
it seems probable that the genera as recognized here include a
greater diversity of form. My system is based mainly on European
species.

With the key given below it should be possible to identify any
known European species to a genus, though it is hoped that it may
also serve as a basis for eastern Palaearctic and Nearctic forms.
The characters of the occiput have not been used extensively
before. Though not completely diagnostic for the genera as
recognized here, they make possible the construction of such a key
and fail only for odd species groups, particularly of Eurytoma.
The arrangement of the key is entirely artificial and intended as
an aid to identification only.

The figures of antennae were prepared from slide-mounts in
Canada balsam. However, such mounts should not be necessary
for generic identification.

170

[April

Terminology

Richards (1956) has been followed for the terminology of most
morphological features, but unless otherwise stated the term
thorax is taken to include the propodeum.

The terminology of the antennal segments poses a number of
problems which have been discussed elsewhere (Claridge, 1959b).
The term flagellum is here used for that part of the antenna distal
to the anellus: that is it does not include the latter. Though
contrary to most general entomological usage, such a terminology
is very valuable for those groups of Chalcidoidea in which there
is no clear distinction between a funicle and club.

In discussing characters of the occiput I have previously
(1958a) used the terms hypostomal carina and hypostomal
lamella for the distinctive structures of some Eurytoma species
(figs. 1 and 2). Prof. O. W. Richards has pointed out to me
(personal communication) that the carina is unlikely to represent
the boundary of the true hypostoma and that the terms post
genal carina and lamella would probably be preferable. These
terms seem to represent a better interpretation of the structures
and will be used here.

Key to genera of Northern Palaearctic Eurytomidae

1. Antennae (figs. 24 and 25) with anelli indistinct and not
clearly differentiated from flagellar segments .....................

Archirileya Silvestri.
Antennae (figs. 12-18, 26-28) with one distinct anellus, clearly
differentiated from flagellar segments . . . . 2.

2. Occipital carina strongly developed and post genal lamella
present (figs. 1 and 2) ; occiput highly polished, unlike rugose

face . . . . Eurytoma Illiger (partim).

Occipital carina variably developed ; post genal lamella never
present; occiput usually sculptured similarly to face, but
polished in Ahtola and some Eudecatoma . . 3.

3. Female gaster with petiole elongate, at least one and a half

times as long as wide, from above; remainder of gaster
flattened laterally and distinctly globular (figs. 9 and 10).
Forewings with more or less developed, pigmented sub¬
marginal band below marginal vein (figs. 21-23). Male
antenna (fig. 16) little specialised, resembling that of female,
but with only 4 free basal flagellar segments ................. _

Eudecatoma Ashmead
Female gaster with petiole shorter, rarely longer than wide.
Forewings without distinctly pigmented submarginal band.
Male antenna (figs. 13, 18, 26, 28) with flagellar segments
longer and bearing variably arranged long hairs ... _ ..... 4.

171

1961]

4. Occipital carina strongly developed; occiput polished; post
genal lamella not present (fig. 4). Post marginal vein well
developed, almost as long as marginal; longer than stigmal

vein (fig. 20) . . . . . Ahtola gen. nov.

Occipital carina weakly developed or absent (figs. 3 and 5);
occiput not clearly differentiated from face in sculpture. Post
marginal vein less well developed and shorter . 5.

5. 5th segment of female gaster longest, about twice as long as

4th in dorsal view (figs. 7 and 8) ... Eurytoma Illiger ( partim ).
5th segment of female gaster not longest; segments either
subequal (fig. 11) or another segment longest . . 6.

6. Marginal vein short, about as long as or slightly longer than
stigmal vein. Small, short species with gaster shorter than
head and thorax together. Male antenna with only four

clearly separate basal flagellar segments (fig. 18) . . 7.

Marginal vein longer; usually more than twice as long as
stigmal vein. Mostly elongate species. Male flagellum of
seven distinctly separate segments (figs. 26, 28) ; terminating
in a short spine, sometimes very short ... Tetramesa Walker.

7 . Female basal flagellar segments clearly narrower than suc¬
ceeding segments (fig. 14). Sculpture of head and thorax
regularly alutaceous, rarely with slight traces of shallow

umbilicate punctures . . Systole Walker.

Female basal flagellar segment little narrower than succeed¬
ing segments (fig. 17). Sculpture of head and thorax
irregularly rugose, with scattered shallow umbilicate
punctures . . . Bruchophagus Ashmead.

Classification

The suprageneric classification within the family Eurytomidae
has received little attention, probably because of the extreme
morphological homogeneity of the group. Ashmead (1904), in
revising the world genera, recognised five tribes, viz. — Aximini,
Isosomini, Eurytomini, Riley ini, and Decatomini. These tribes
have all been raised to subfamilies by various more recent
authors. Ferriere (1950) grouped the Palaearctic genera into four
subfamilies, viz. — Rileyinae, Harmolitinae ( = Isosomini Ash¬
mead), Decatominae, and Eurytominae. The only one of
Ashmead’s tribes not included was the Aximini which includes
aberrant forms mostly confined to central America.

Though I only recognize eight Palaearctic genera I retain the
same subfamily grouping as that of Ferriere, since there does
seem to be a wider gulf between the genera so separated than
between those included in the Eurytominae. My classification is
thus as follows : —

Archirileya Silvestri

Rileyinae

172

Eurytominae

[April

Eurytoma Illiger
Ahtola gen. nov.

Systole Walker
Bruchophagus Ashmead

Harmolitinae

Tetramesa Walker

Ailomorphus Walker — probably restricted to the extreme south¬
eastern Palaearctic and the Oriental regions.

Eudecatominae ( = Decatominae Ferriere)

Eudecatoma Ashmead

Rileyinae

The Rileyinae seem to be characterized reliably by the pre¬
sence of two or three antennal anelli, which are not clearly
distinguishable from the basal flagellar segments (figs. 24 and 25),
and a general facies quite unlike the other subfamilies, lacking
the typical umbilicate punctation. The single Palaearctic genus
Archirileya Silv. of southern and central Europe is represented by
two described species. So far as known they are predators of the
eggs of various Orthoptera, like the allied Macrorileya oecanthi
(Ashm.) of North America. The known species of Rileya Ashm.
are parasites in various Cecidomyiid galls.

Archirileya Silvestri

1920, Archirileya Silvestri, Boll. Lah. Zool. Portici , 14: 223-225.
1951, Anarchirileya Boucek, Acta Mus. nat. Prag., 27: 54-56.
1957, Sidonia Erdos, Ann, hist. nat. Mus. hung., 8: 350-351.

The genus is characterized by its elongate body form in which
it resembles Macrorileya Ashm. Rileya differs in its shorter body
form and apparent fusion of the post spiracular sclerite to the
mesepisternum .

Boucek (1958) has synonymized Sidonia podagrica Erdos with
A. inopinata Silv. and has included his own genus and species
Anarchirileya femorata in the genus Archirileya.

Neither A. inopinata nor A. femorata have been recorded from
northern Europe or Britain.

Eurytominae

This is the largest and most difficult to define of the sub¬
families recognized here. There is an extremely wide range of
habits, from endophagous and ectophagous parasitism (most
Eurytoma species) to complete phytophagy (Bruchophagus,
Systole and some Eurytoma species).

173

1961]

Figs. 1-6. — Female occiput in posterior view of: 1, Eurytoma tibialis
Boh.; 2, E. rufipes Walk.; 3, Bruchophagus platypterus
(Walk.); 4, Ahtola atra (Wtalk.); 5, Eurytoma cynipsea
Boh. ; and 6, Tetramesa calamagrostidis (v. Schlecht.) (to
scale a). oc — occipital carina; pf — proboscidial fossa;

pg — post gena; pi — post genal lamella.

Figs. 7-11. — Lateral view of female gaster of: 7, Eurytoma salicvperdae
Mayr; 8, E. cynipsea Boh.; 9, Eudecatoma submutica
(Thoms.); 10, E. mellea (Curtis); and 11, Bruchophagus
platypterus (Walk.) (to scale b).

174

[April

The status of Nikanoria Nikol’skaya ( = Biro-Lajosia Erdos,
vide Boucek, 1958) is uncertain. I have not been able to examine
either of the included species from central Europe and the
U.S.&R., but it is probable that it is best considered as a part of
Eurytoma Ill., as suggested by Boucek ( loc . tit.).

Eurytoma Illiger

1807, Eurytoma Illiger, Mag. Insektenk, 6: 192.

1807, Eurytoma Illiger, in Rossi, Fauna Etrusca, 2nd ed., 2: 128.
1811, Decatoma Spinola, Ann. Mus. Hist. Nat., 17 : 138-152.

Type species — Chalcis abrotani Panzer — designated by West-
wood (1839).

Eurytoma is the oldest described genus in the family and a
large number of species have been described, including a number
from all the major geographic regions. The generic limits are
very difficult and I have found it impossible to give completely
diagnostic key characters without recognizing artificial assem¬
blages of species. In most of the species groups the occiput is
strongly margined and polished with the post genal lamella well
developed (figs. 1 and 2). This complex of characters provides
the best means of identifying the genus. However, it is not
present in the cynipsea Boh. group (fig. 5), saliciperdae Mayr
group, and a few odd species such as E. setigera Mayr. The
lengthened fifth tergite of the gaster distinguishes these groups
from species of Bruchophagus and Systole. It would be possible
to restrict the genus to those species with the occipital characters,
but the species groups so included have no more relationship
with each other than with the excluded groups. Thus an
extremely artificial system would result.

I have not examined specimens of Ipideurytoma Boucek &
Novitsky (1954), but it seems probable that it is best considered
as a very distinct species group of Eurytoma, characterized by
the very curious flattened head. The single European species,
I. spessivtsevi Bek. & Nov., has not been recorded from Britain.

Ahtola gen. nov.

Type species — Isosoma atrum Walker (1832).

The species of Ahtola may be separated from Eurytoma by
the following combination of characters : —

Occiput (fig. 4) strongly margined; polished; sculptured
differentially to face; post genal carina clearly differentiated, but
not formed into a lamella. Forewings (fig. 20) with post marginal
vein well developed, as long as or longer than marginal vein;

175

1961]

thickly haired with small submarginal speculum. Gaster (fig. 19)
clearly petiolated; tergites subequal, fifth not or little longer than
fourth.

Ahtola resembles superficially some species of Tetramesa, but
is separable by means of the occipital characters. The latter
together with the structure of the male antenna (fig. 13) place the
genus in the Eurytominae.

Isosoma atrum together with Eurytoma globiceps Boucek
(1954) and E. cylindrica Thomson (1875) represents a distinct
natural group which I believe warrants generic status. I had
supposed that Eurytomocharis Ashmead might be congeneric
with atrum. However, Dr. B. D. Burks, Washington, informed me
(in litt.) that in his opinion it was not congeneric with the type
species of Eurytomocharis, E. minuta Ashm., and that it probably
represented an undescribed genus. Thus, at present it seems best
to give a new generic name even though it may have to be
synonymized later.

Ahtola atra (Walker) comb. nov.

1832, Isosoma atrum Walker, Ent. Mag., 1 : 14.

The following is a redescription based on modern material : —

Female. Predominantly black species. Tarsi and extreme
apex of femora and base of tibiae rufo-fuscous ; venation fuscous ;
disc of wing variably infumate, particularly in large specimens.
Maximum length about 4-2 mm.

Head, in frontal view, distinctly transverse, width to height
as about 11: 8 (cf. fig. 4); cheeks evenly narrowed to mouth;
antennae inserted at or slightly above lower level of eyes ;
antennal scrobes strongly margined; surface sculpture consisting
of distinct umbilicate punctation, with short white hairs arising
from each puncture. Head, from above, distinctly transverse,
varying from about twice as wide as long to distinctly less;
rather gibbous between eyes; POL: OOL, measured from edges
of ocelli, as slightly more than 4:3. Malar space to height of
eye as about 3 : 4. Antennae (fig. 12) with scape simple, reaching
to about level of ocelli; pedicel short globular; anellus distinctly
transverse: seven clearly distinct flagellar segments, sixth and
seventh closely associated ; proportions of flagellar segments
rather variable; rhinaria well developed, tending to form two
rows on each segment.

Thorax variable, from about twice as long as wide to
distinctly less — length to width as about 8:5; pronotum similarly
variable, from about twice as wide as long to distinctly less —
width to length as about 15:9; surface sculpture heavily
alutaceous with densely distributed umbilicate punctures ;
mesepisternum sloping evenly to mid coxa. Propodeum heavily
and variably rugulose, usually with distinct median furrow ; often

176

[April

with tendency to develop a dorsal face; angle of slope shallower
in narrow specimens. Forewings (fig. 20) densely clothed with
short hairs; stigmal vein slightly shorter than marginal vein and
making an angle of about 30° with post marginal vein; post
marginal vein, though fading apically, about as long as or slightly
shorter than marginal vein; submarginal vein broken into a long
basal and a short apical section.

Gaster varying from about as long as thorax, in large specimens
(fig. 19) to clearly longer, usually in smaller narrower specimens;
petiole variable, at least half as long as wide in dorsal view ; basal
segments, after petiole, subequal; tergites highly polished; short
hairs and faint alutaceous sculpture developed dorsally on 5th
to 8 th tergites.

Male. Resembles female closely. Antennae (fig. 13) longer;
scape distinctly swollen medially; seven clearly separate flagellar
segments, flattened laterally and bearing dense long hairs with
tendency to be arranged in whorls. Abdominal petiole elongate,
about one and a half times as long as wide.

A. atra is subject to variation in body width like many other
species whose larvae inhabit stems. It is a distinct species, but
may really consist of a number of very closely allied forms which
it has not yet been possible to separate on adult morphology.
Detailed breeding experiments are required to elucidate the
problem. The generic characters given in the key, together with
its size, should immediately separate the morphospecies from all
other British Eurytomids.

In the British Museum (Nat. Hist.) type collection there is a
single Walker specimen (B.M. Type Hym. 5.569) bearing a green
type label. This specimen, a female, fits well the original descrip¬
tion and is here designated as lectotype.

Biology

A. atra may be bred from stems of Alopecurus species together
with Eurytoma tapio Claridge (1959b). During the winter I have
removed large numbers of last instar larvae from the stems where
they appeared to have been feeding phytophagously. However,
the exact host relationships are uncertain. I have also reared a
series of specimens from a collection of stems, thought to be of
Anthoxanthum odoratum L., but this record requires confirmation
and the specimens were not used in making the above redescrip¬
tion.

Material Studied

ENGLAND. Berkshire, Cothill, Nat. Grid Ref. SU4699 : 1 9 ,
26.V.1957. Buckinghamshire, Oakley Wood, Nat. Grid Ref.
SP6111: 12 9 9, 2 6 d, emerged 29.iv.-2.v.l958 from larvae in
stems of Alopecurus pratensis L.; 1 9, 26.vi.1957. Oxfordshire,
Otmoor, Nat. Grid Ref. SP5514-5614: 4 9 9, 2 dd, em. 1-2.V.1958,
larvae in stems A. pratensis; 1 9, 3 6 d, 20.V.1958; 1 9,1 d,
21.V.1957 ; 2 9 9, 1 d, 4.vi.l958; 1 d, 14.vi.1958 (all coll. M. F.
Claridge).

1961]

177

Figs. 12-19 —Left antenna in lateral view of : 12, Ahtola atra (Walk.),
female; 13, A. atra, male; 14, Systole albipennis Walk.,
female; 15, Eudecatoma mellea (Ciirtis), female; 16, E.
mellea, male; 17, Bruchophagus platypterus (Walk.),
female; 18, B. platypterus , male (figs. 12, 14-17 to scale a;
figs. 13, 18 to scale b).

Fig. 19. Lateral view of female gaster of Ahtola atra (scale c).

Figs. 20-23. Part of leading edge of forewing of: 20, Ahtola atra ;

21, Eudecatoma submutica (Thoms.); 22, E. flavicollis
(Walk.); and 23, E. concinna (Boh.) (figs. 20, 22 and 23
to scale b; fig. 21 to scale c).

178

[April

Ahtola cylindrica (Thomson) comb. nov.

1875, Eurytoma cylindrica Thomson, Hym. Scand., 4 : 52.

I have only been able to examine a single specimen, the type,
of this species, and thus cannot evaluate it adequately. It seems
to differ from A. atra in its smaller size and extremely narrow
body. It is possible that it is a very small specimen of A. atra
though I am inclined to think that it represents a good species.

I have seen the five specimens from the Thomson collection
which stand as Eurytoma cylindrica. One of these has been
provisionally selected as lectotype by Dr. A. Jansson. I agree
with his designation and here validate it. The specimen has been
remounted and is staged on a pin which also carries a male
Tetramesa species and a printed label “Wit”. The other specimens
are not conspecific.

Ahtola glohiceps (Boucek) comb. nov.

1954, Eurytoma glohiceps Boucek, Acta Mus. nat. Prag., 29:

72-74.

Dr. Z. Boucek has kindly sent me two specimens of his species
Eurytoma glohiceps. It is closely allied to A. atra, but differs
in its smaller size, more gibbous head and longer abdominal
petiole. Its biology is unknown, but Boucek ( loc . cit.) believes
that it is probably associated with grasses.

Bruchophagus Ashmead

1888, Bruchophagus Ashmead, Ent. Amer., 4: 42.

1950, Bruchophagus Ferriere ( partim ), Mitt. Schweiz, ent. Ges.,
23: 379.

Type species — Bruchophagus horealis Ashm. — designated by
Ashmead (1894).

The group of species represented in the Palaearetic region by
Eurytoma platyptera (Walk.) and closely allied forms, is here
separated from Eurytoma and recognized as the genus Brucho¬
phagus. It differs from Eurytoma in its shorter gaster without
an elongated fifth segment (fig. 11), and in the lack of the
occipital carina and post genal lamella (fig. 3). The only western
Palaearetic species that have been studied intensively are un¬
doubtedly phytophagous in the fruits of various Papilionaceae —
B. platypterus (Walk.) ( = gihhus Boh.) in Trifolium species
(Claridge, 1959c), and B. ononis (Mayr) in Ononis species
(Crevecoeur, 1951). In this habit they resemble the known
Nearctic species (vide Burks, 1957). Nikol’skaya (1952a, b, and
1955) has described several phytophagous species from the
U.S.S.R. including one, B. mutahilis, from Primrose seeds, Primula

179

1961]

species. Burks (loc. cit.) has included a new species bred from
seeds of an Aloe species (Liliaceae), imported from South Africa.
It seems likely that all the species attributable to Bruchophagus
as recognized here are similarly phytophagous. Species differ¬
entiation within the genus is very difficult and it is probable that
careful breeding experiments will show there to be more species
involved than the adult morphology suggests.

Ferriere (1950) introduced the genus Bruchophagus to the
Palaearctic literature, but defined it almost completely on the
single character of a four segmented male funicle. As I have
pointed out previously (1959b), such a character may be difficult
to appreciate and, more important, usually breaks down within
any natural genus, at least in the Eurytomidae. Ferriere included
such diverse species as Eurytoma cynipsea Boh. and E. setigera
Mayr, together with those species here included. Boucek (1951)
and I (1958a) have criticized the genus as recognised by Ferriere.
I believe that it can be recognized as a natural unit only when
some of the species which resemble the platypterus group in the
possession of a four segmented male funicle alone, are excluded.
This view is enhanced by the biological information.

Systole Walker
1832, Systole Walker, Ent. Mag., 1 : 22.

1959, Systole Walker, Claridge, Ent. mon. Mag., 95: 39.

Type species — Systole alhipennis Walk. — only originally in¬
cluded species.

I have previously (1959c) discussed the genus and given
characters by which it is distinguished from Eurytoma,

The species of Systole, like those of Bruchophagus, form a
small and very compact group, all the known species of which
are phytophagous exclusively in the seeds of Umbelliferae. The
genus is recognized on rather slight characters, but again like
Bruchophagus, the biological generalizations which can be made
about it justify its recognition.

Harmolitinae

In the northern Palaearctic region I recognize only one genus
in this subfamily, viz., Tetramesa Walker. It may be divided
into a number of species groups which grade into each other and
which I do not consider to warrant generic rank. The larvae of
all known species are phytophagous in grasses.

Ailomorphus rhopaloides Walk. (1871), described from Hong
Kong, is closely allied to Tetramesa. It differs in having the
gaster more laterally compressed with the fifth tergite distinctly
the longest. Of species known to me, it resembles most closely

180

[April

T. rornana (Walk.) of southern Europe. A. rhopaloides, together
with an undescribed species represented in the British Museum
(Nat. Hist.), is probably best kept separate from Tetramesa at
least until the south-eastern Palaearctic and Oriental forms are
better known. It is interesting that A. rhopaloides has been
recorded as phytophagous in Bamboo, Phyllostachys species
(Gahan, 1924).

0 2 mm

Figs. 24-28. — Left antenna in lateral view of : 24, Archirileya i/nopi-
nata Silv., female; 23, A. inopinata, male; 26, Tetramesa
brevicornis (Walk.), male; 27, T. hyalipennis (Walk.),
female ; 28, T. hyalipennis , male (figs. 24, 25, 27 to scale
a; figs. 26, 28 to scale c).

Fig. 29. — Part of leading edge of forewing of Systole tuonela Clar.
(to scale b).

1961]

181

Tetramesa Walker

1832, Isosoma , Walker, Ent. Mag., 1 : 14 (preoccupied).

1848, Tetramesa Walker, List spec. Hym. coll. Brit. Mus., 2,
Chalcidites, appendix: 154, London.

1863, Harmolita Motschulsky, Bull. Soc. Nat. Moscow, 36 (3): 58.
1871, Philachyra Walker, Notes on Chalcidiae, 1 : 7-8, London.
1920, Isthmosoma Hedicke, Archiv. Naturgesch., 86 (11): 165.
1958, Tetramesa Walker, Claridge, Ent. mon. Mag., 94: 84.

Type species — Tetramesa iarbas Walker ( = T. crassicornis
(Walk.) (vide Claridge, 1958a)) — only originally included species.

I have previously ( loc . cit.) given a description of the genus
and no further information need be added here. A complete
diagnosis will be given in a revision of the European species now7
in preparation.

Eudecatominae

The Eudecatominae is the Decatominae of Ferriere. I have
previously discussed the genus Decatoma Spinola ( nec auctt .)
and concluded that it should be taken as a synonym of Eurytoma
(Claridge, 1959a). Thus the group name based on it cannot be
used to refer to a group not including it. Since the only genus
attributable to the subfamily is Eudecatoma Ashm., I suggest the
name Eudecatominae.

Eudecatoma Ashmead

1888, Eudecatoma Ashmead, Ent. Amer., 4: 42.

Type species — Decatoma batatoides Ashm. — only species in¬
cluded by Ashmead (1894).

The genus is characterized as follows : —

Often extensively pale marked species; exact marking vari¬
able. Female flagellum (fig. 15) with five clearly separate basal
segments. Male flagellum (fig. 16) resembles that of female but
with only four distinctly separate basal segments. Occiput
variable — clearly margined and polished in some larger species
(cf. E. biguttata (Swed.) — not margined and sculptured more
or less as face in most smaller species (cf. E. mellea (Curtis)).
Forewings with distinct pigmented submarginal band, often ex¬
tensive (figs. 21 and 22), but sometimes small (fig. 23). Gaster
(figs. 9 and 10) distinctly petiolated and laterally flattened; in
female, rather circular in outline, with ovipositor clearly directed
dorsally when not in use.

The species of Eudecatoma form a clearly distinct natural
group. I have previously (1959a) discussed the British species.

182 [April

Evidence suggests that at least some of the species are
endoparasitic, a habit rare in the family.

Phylogenetic Speculation

Several authors have speculated on the phylogenetic relation¬
ships of the Eurytomidae and of the genera within the family
(vide Bugbee, 1936; and Nikol’skaya, 1956). Particular attention
has been paid to the importance of the phytophagous habit in
the evolution of the group. Bugbee concluded that “Whether the
primitive Eurytomid stock was parasitic, phytophagous or both
is uncertain, but the evidence favours a plant feeding origin”.
Nikol’skaya on the other hand favours a parasitic ancestry.

Bugbee, after giving a very useful summary of the comparative
anatomy of the Nearctic genera, wTent on to suggest that certain
features were phylogenetically primitive and others more recent
from the results of his preceding comparative account alone.
With regard to the structure of the antennae for example, he
uses six definite, but unjustifiable, a priori criteria for determining
phylogenetic relationships. These include : —

“ . . . 2. The more alike the male and female antennae,
the more primitive are the insects.

3. Reductions in the number of segments is a sign
of recent development. ...”

Such criteria are entirely subjective and without fossil evidence
mean nothing. Bugbee’s conclusions on such grounds are that
the Harmolitinae include structurally the most primitive forms.
This may be so, but all known species are specialized plant feeders
associated with Gramineae. Probably they represent a branch or
branches derived from the ancestral Eurytomids which early took
up the phytophagous habit and radiated following their host
plants. Modern species seem to be restricted mainly to the grass¬
lands of the Holarctic region.

The genera of Eurytominae exhibit the widest range of habit,
structure and geographical range known within the family. It
seems most likely that the subfamily includes forms nearer to
the ancestral Eurytomid than do the others. Within it a number
of forms have independently taken up the phytophagous habit.
Some such as Systole and Bruchophagus species specialized as
seed-eaters. Many species of Eurytoma are only parasitic for a
short period and complete their development phytophagously,
e.g. E. flavimana Boh., a parasite of Tetramesa linearis (Walk.),
a gall-former in stems of Agropyron species. Larvae of the allied
E. suecica von Rosen are completely phytophagous in stems of
Wheat, Triticum species (v. Rosen, 1956). Such a habit probably
evolved from a condition such as that found to-day in E. flavi¬
mana, Thus there is little doubt that the phytophagous habit has
arisen many times within the subfamily.

183

1961 ]

The Eudecatominae and Rileyinae show a number of possibly
specialized features and probably represent rather early divergents
from the ancestral stock. It seems most likely that the ancestral
Eurytomids were parasitic forms, probably associated with insect
galls. The modern forms then specialized in various directions,
thus retaining only some of the ancestral characters. Without
the help of a good fossil record such characters cannot be differ¬
entiated from the specialized and more recent features. However,
since the phytophagous habit can be shown almost certainly to
have arisen several times within the Eurytominae, there is no
reason why it should not have done so in the Harmolitinae. In
fact, any of the subfamilies or genera could be polyphyletic in
origin and any supposed phylogeny can be read in either direction.
We simply have to admit that in the final analysis we just do
not know.

Acknowledgments

Much of the work for this paper was carried out in the Hope
Department of Entomology, University Museum, Oxford, during
the tenure of a research studentship of the Department of
Scientific and Industrial Research. I should like to express my
thanks to Dr. M. W. R. de V. Graham, Dr. B. M. Hobby, Prof.
O. W. Richards, and Prof. G. C. Varley for their criticisms of parts
of the manuscript at various stages in its preparation. I should
like also to thank Dr. A. J. Cain, Dr. B. W. Staddon, and Dr. H.
H. Williams for their valuable criticism and discussion, particu¬
larly concerning the first section. I wish also to express my
thanks to the following who have given or loaned much valuable
material : — Dr. B. D. Burks, Washington ; Prof. P. Brinck and
Prof. C. II. Lindroth, Lund; and Dr. Z. Boucek, Prague. Mr. G.
J. Kerrich and Mr. R. D. Eady of the Commonwealth Institute
of Entomology and Mr. J. F. Perkins of the British Museum (Nat.
Hist.) have always given me every assistance in examining the
collections in their care, for which I am very grateful.

References

Ashmead, W. H. 1894. Descriptions of new parasitic Hymenoptera.
Trans, amer. ent. Soc., 21 : 318-344.

Ashmead, W. H. 1904. Classification of the Chalcid flies or the super¬
family Chalcidoidea, with descriptions of new species in the
Carnegie Museum, collected by Herbert H. Smith. Mem.
Carnegie Mus., 1 (4) : i-xi, 225-555, 9 Plates.

Boucek, Z. 1951. Results of the zoological scientific expedition of
the National Museum in Praha to Turkey 7 Hymenoptera
1 Chalcidoidea (first part). Acta Mus. nat. Prag., 27 : 47-57.

Boucek, Z. 1954. Chalcidologicke poznamky I, Pteromalidae, Tory-
midae, Eurytomidae, Chalcididae (Hymenoptera). Acta Mus.
nat. Prag.. 29 : 49-80.

184 [April

Boucek, Z. 1958. To the taxonomy of the European species of Schizo-
notus and Caenocrepis — parasites of economic importance —
with notes, and some new synonymy in the Pteromalidae and
Eurytomidae (Hym.). Acta Mus. nat. Brag., 32 : 395-404.

Boucek, Z., & Novicky [Novitzsky], S. 1954. Ipideurytoma
spessivtsevi n.g. n.sp., ein neuer Borkenkafer-Parasit. Ent.
Tidskr ., 75: 266-271.

Bugbee, R. E. 1936. Phylogeny of some Eurytomid genera. Ent.
Amer., 16: 169-223.

Burks, B. D. 1957. A new Bruchophagus from a Liliaceous plant
with a host plant list for the genus (Hymenoptera, Euryto¬
midae). Proc. ent. Soc. Wash., 59 : 273-277.

Burks, B. D. 1958. Chalcidoidea, in Krombein, K.V., Hymenoptera
of America North of Mexico Synoptic Catalog (. Agriculture
Monograph No. 2), First Supplement. Washington.

Cain, A. J. 1954. Subdivisions of the genus Ptilinopus (Aves,
Columbae). Bull. Brit. Mus. (Nat. Hist.), Zool., 2 (8) :
267-284.

Cain, A. J. 1958. Logic and memory in Linnaeus’s system of taxo¬
nomy. Proc. Linn. Soc. Bond., 169: 144-163.

Cain, A. J. 1959a. The post-Linnaean development of taxonomy.
Proc. Linn. Soc. Load., 170: 234-244.

Cain, A. J. 1959b. Deductive and inductive methods in post-Linnaean
taxonomy. Proc. Linn. Soc. Lond., 170: 185-217.

Cain, A. J. 1959c. Taxonomic concepts. Ibis, 101 : 302-318.

Claridge, M. F. 1958a. Tetramesa Walker 1848, a valid name for
Iso soma Walker 1832 in place of Harmolita Motschulsky 1863,
with a short discussion on some Eurytomid genera (Hym.,
Eurytomidae). Ent. mon. Mag., 94: 81-85.

Claridge, M. F. 1958b. The British and Scandinavian species of the
genus Clieiloneurus Westwood (Hym., Encyrtidae). Ent. mon.
Mag., 94: 156-161.

Claridge, M. F. 1959a. A contribution to the biology and taxonomy
of the British species of the genus Eudecatoma Ashmead
( — Hecatoma Auctt. nec Spinola) (Hym., Eurytomidae).
Trans. Soc. Brit. Ent., 13: 149-168.

Claridge, M. F. 1959b. The identity of Eurytoma appendigaster
(Swederus, 1795) (Hym., Eurytomidae), together with descrip¬
tions of some closely allied species bred from Gramineae. Ent.
mon. Mag., 95 : 2-13.

Claridge, M. F. 1959c. Notes on the genus Systole Walker, includ¬
ing a previously undescribed species (Hym., Eurytomidae).
Ent. mon. Mag., 95 : 38-43.

Crevecoeur, A. 1951. Note sur la biologie d’Euryfoma (. Bruchophagus )
ononis Mayr (Hym., Chalcidoidea). Bull. Ann. Soc. ent. Belg.,
87: 148-150.

Ferriere, C. 1950. Notes sur les Eurytoma (Hym., Chalcidoidea) I. —
Les types de Thomson et de Mayr. Mitt, schweiz. ent. Ges.,
23: 377-410.

1961] 185

Gahan, A. B. 1924. The systematic position of the genus Harmolita
Motschulsky with additional notes (Hymenoptera). Proc. ent.
Soc. Wash., 28: 224-229.

Nikol’skaya, M. N. 1952a. Chalcid fauna of the U.S.S.R. (Chalci-
doidea). Monograph on the fauna of the U.S.S.B., Zoological
Institute of the Academy, Nauk, 44: 1-574 [in Russian].

Nikol’skaya, M. N. 1952b. Two new species of seed-eaters from the
family Eurytomidae (Hymenoptera, Ohalcidoidea). Bev. Ent.
U.B.S.S. , 32: 304-306 [in Russian].

Nikol’skaya, M. N. 1955. New genera and species of Chalcids of the
families Eurytomidae and Callimomidae in Middle Asia
(Hymenoptera, Ohalcidoidea). Trav . Inst. zool. Ao.ad. Sci.
U .B.S.S., 21 : 335-341 [in Russian].

Nikol’skaya, M. N. 1956. Seed-eating chalcids of the U.S.S.R. and
the importance of the phytophagous habits in the evolution
of the group (Hymenoptera, Ohalcidoidea). Bev. Ent.
U.B.S.S., 35 (3): 570-581 [in Russian, with English summary].

Peck, 0. 1951. Ohalcidoidea, in Muesebeck, 0. F. W., Krombein,

K. V., and Townes, H. K., Hymenoptera of America north of
Mexico Synoptic Catalog, U.S. Dept. Agric., agric. Monogr ., 2.

Richards, O. W. 1956. Hymenoptera introduction and key to fami¬
lies, in Handbooks for the identification of British Insects ,
6 (1) : 1-94, London, Royal Entomological Society.

Rosen, H. von. 1956. Erne phytophage Eurytoma in Mittel.-und
Nordschweden (Hym. Chalcid.). Opusc. ent. Lund., 21 : 16-20.

Thomson, C. G. 1875. Hymenoptera Scandinaviae, 4 (1), Lund.

Walker, F. 1832. Monographia Chalciditum. Ent. Mag., 1 : 12-29.

Walker, F. 1846. List of the specimens of Hymenopterous insects in
the collection of the British Museum, Part 1, Chalcidites,
London.

Walker, F. 1871. Notes on Chalcidiae, Parts 1 and 2. London.

Westwood, J. O. 1839. Synopsis of the genera of British insects, in
An introduction to the modern classification of insects; fov/nded
on the natural habits and corresponding organization of the
different families. London.

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