Contributions to Zoology, 73 (4) 263-270 (2004)
SPB Academic Publishing bv, The Hague
Alternation of generations in Andricus corruptrix (Schlechtendal): comments on and description of a new sexual form (Hymenoptera: Cynipidae)
¹ ² ² R. Folliot P. Ros-Farré D. Bellido² & J. Pujade-Villar
1 Universite de Rennes I, Faculte Sciences, C.N.R.S. UPRES- A 6026, Equipe Canaux Recepteurs
2 Membranaires, Bat 13-Campus de Beaulieu, 35042-Rennes Cedex - France; Universitat de Barcelona
Facultat deBiologia, Departament de Biologia Animal, Avda Diagonal 645, E-08028-Barcelona
A. A. Keywords: Hymenoptera, Cynipidae, Andricus, corruptrix, improprius, life cycle, new form
Abstract single year. In some species the two generations
ate morphologically different and develop in galls
treated and Life cycle ofA. corruptrix (Schlechtendal, 1870) is of different types on different parts of the host plant.
to discussed. A new sexual corrections previous experiments are The link between the unisexual and bisexual gen- form for A. corruptrix is herein described, while A corruptrix erations of one species can be revealed by care- taxonomical formalarshemi is considered a valid speciesand for fully controlled rearing experiments. In the stat & large reasons is renamed as Andricus improprius n. n.sp, only Andricus for instance, in the two known by the sexual form, and not linked with A. corruptrix. genus Europe,
for ,4. & A. larshemi not Lectotypes impropriusn. stat n, sp. (= generations of many species are known (Pujade-
available) are also designated. Villar but et ai, 2001) many other species are known
only by their unisexual generation and a few other
by their bisexual generation only. Modern molecular Contents biology methods, for example DNA sequencing,
can help to suspect a specific link between two
Introduction 263 previously unlinked bisexual and unisexual forms. Material and methods 264 remain the Rearing experiments necessary method Results 265 to determine the two alternating generations of a The bisexual form ofAndricus corruptrix
(Schlechtendal) 265 species.
Andricus corruptrix (Trotter) new bisexual form 265 In most species the galls of the two alternating
Taxonomic comments on Andricus corruptrix f. generations develop on the same oak species. In larshemi 1958 268 D. van L. & D.-M., the case of Andricus kollari the gall of the uni- Andricus improprius Bellido & Pujade-Villar sexual generation develops on Quercus robur, & 269 Q. n. stat. n. sp and First studies of alterna- Discussion 269 petraea Q. pubescens.
269 tion of Acknowledgements generations in the Andricus kollari group
References 269 were made by Beijerinck (1902), who found that
Andricus circulans, a sexual form galling another
oak the sexual species, Quercus cerris, was gen- Introduction eration ofAndricus kollari. Such a situation is called
heteroecism. These experiments were considered
In is an ex- most oak the reproduction as doubtful gallwasps by many hymenopterologists of his time,
or ample of cyclical parthenogenesis heterogony. until Marsden-Jones (1953) and Folliot (1964)
The lifecycle involves alternation of a bisexual gen- confirmed their results.
eration and a partenogenetic (unisexual or asexual) The Andricus contains several genus groups of generation. The two are strictly alter- generation neighbouring oak gallwasps (Stone & Cook, 1998).
The is completed within a of nating. cycle commonly One them, called the kollari group (Bellido et - 264 R. Folliot et at. Generation alternation in the hymenopterid Andricus corruptrix
ah, 2003), contains the species listed in table 1. One of us (R. F.) found in Rennes (Brittany, France)
unisexual those named Docters van Leeuwen (1956) and Docters van similar galls to A. corruptrix
Leeuwen & Dekhuijzen-Maasland (1958) studied in Docters van Leeuwenpictures. Experiments with
the unisexual adults undertaken in order the lifecycle of two other species of this group, were to
it Andricus lignicolus and Andricus corruptrix. They obtain sexual generation and to compare with
collected galls of the unisexual form of these two the sexual generation obtained by Docters van
and also Leeuwen species on Q. robur they described, on Q. and other known sexual generations of
of the kollari cerris, the sexual form of A. corruptrix and A. species group.
’ lignicolus, respectively named ‘larshemi and ‘van- heurni'. char- Wiebes-Rijks (1978) gave diagnostic
acters to separate the three known sexual forms (A. Material and methods
kollari, A. lignicolus and.A. corruptrix). These three
in of it in species, now widespread a great part Europe, At first, was observed, a glass jar that A. cor-
are not native in The Netherlands and were intro- females could in buds ruptrix lay eggs Q. cerris
duced with of their together one hosts, Q. cerris. (one egg was found after dissection in the heart of
This introduction enabled A. species of the kollari a bud).
Eastern colo- Different three of them group, once restricted to Europe, to experiments were made,
nize Askew & with unisexual In new zones (Quinlan, 1974; Neill, on Q. cerris one female each.
1993). one experiment the oak branch died too early after
Docters After Beijerinck and van Leeuwen, the six months. In another nothing was obtained. The
status of the above three heterogonic species was third experiment was successful and is described thought to be the following: hereunder in more details.
One reared in Rennes female of A. corruptrix on
Unisexual form Sexual form th the 25 of June, 1997 was taken to Puilboreau A. kollari (Hartig, 1843) circulans Mayr, 1870 (Baillac), near to La Rochelle, a more southern place A. lignicolus (Hartig, 1840) vanheurni D. van L. & in France where, despite intense monitoring, A. D.-M., 1958 th corruptrix has never been found. There, on the 27 A. corruptrix (Schlechtendal, larshemi D. van L. & D.-M., sleeve 1870) 1958 [not available] of June, 1997, it was placed inside a on
Quercus cerris.
Table I. Alternation of in A. kollari generations group
Unisexual form Sexual form Sexualformform hosthost tree Author closing cycle
A. kollari A. circulans¹circulans' Q. cerris Beijerinck (1902)
A. suber data) hispanica I Q. Pujade-villar (unpl. data)
3 A. lignicolus A. vanheurni³vanheurni\ Q. cerris D.v.LD.V.L&& D-M (1958)
2 A. cerris Present corruptrix (=A. ambiguus²)ambiguus ) Q. paper
A. Unknown44 cerris? — infectorius Q. cerris? -
A. Unknown cerris? — amblycerus Q. -
A. A. aries Unknown Q. cerris? -—
A. — caliciformis Unknown Q. cerris? -
5s Unknown A. improprius Q. cerris -—
(1) to et not According Pujade-Villar, 1992 and Pujade-Villar at. 2001; known but described; before mentioned as A. kollari in the
Iberian Peninsula,
(2) Non valid species according Bellido, Melika & Pujade-Villar (submitted ).
|3) The new in this do have the 15 of the International Code of sexual, paper, not a specific name, following article Zoological
Nomenclature.
(4) A. burgundus has been recorded as the sexual generation ofA. infectorius tinctoriusnostrus, according to Melika et at, 2000), but recent papers argue against that (Stone & Cook, 1998; Cook et al, 1999).
,5) stat and Andricus n. n. sp. lo corruptrix forma larshemi (in this paper). Contributions to Zoology, 73 (4) - 2004 265
The sleeve left th was from 27 June, 1997 until The bisexual form of Andricus corruptrix 20"' April, 1998, to avoid oviposition by A. kollari (Schlechtendal) and A. this last is lignicolus (although species very rare in the and later, which Sexual adults zone) predation by birds, obtained from the experiment were
attack in the oak buds. with A. larshemi from frequently growing galls compared types Doctors van We obtained and insects of the sexual galls gen- Leeuwencollection, sexual adults ofA. kollari (=cir- eration which will be treated further in more de- culans) and A. lignicolus (=vanheurni) experimen-
tails. obtained and tally by Folliot descriptions present
Our with both uni- the specimens were compared in literature (Wiebes-Rijks, 1978). They are
and sexual Doctors sexual material obtained by van clearly different from all these forms.
Leeuwen and Dekhuijzen-Maasland in their experi- We compared also unisexual females used by
ments, in Museum Amsterdam Docters in with deposited Zoologisch van Leeuwen his experiments our
4 unisexual fe- A. (Netherlands). Unfortunately, only corruptrix female and we found they belonged
2 males of Andricus and sexual females to the same As the corruptrix species. unisexual females ap- of A. larshemi are left of the material used in the be identical pear to and the obtained sexual forms
Leeuwen. experiments of Doctors van Our speci- different, one of the two experiments must have
mens were also with material of the uni- been contaminated compared by another species. Docters van
sexual and sexual froms of A. and A. & lignicolus Leeuwen Dekhuijzen-Maasland were success- kollari, obtained Folliot. ful in their to experimentally by Roger attempt produce unisexual form galls
Unfortunately no ofA. vanheurni were found from in other types sexual insects but experiments, they
material collec- among the Doctors van Leeuwen obtained sexual galls underarguable conditions since tions. 3 unisexual females of A. Only lignicolus nets were kept on the host for only two weeks used in the left experiments were (Hogenes, pers. (Docters van Leeuwen & Dekhuijzen-Maasland,
com.). 1958). In our opinion the contaminant could not be
A. SEM were made the second author. or A. kollari since their pictures by lignicolus sexual gen-
The made low and with- pictures were at voltage erations are different, but an undetected species of out from the A. kollari which has coating to preserve specimens damage. group been confused with
We follow the of other the current terminology morpho- species. Consequently, name A. larshemi
logical structures as given in Gibson (1985), points to a valid species the unisexual generation Ronquist and Nordlandcr (1989), and Fergusson of which is unknown.
Abbreviations for fore venation fol- To sum the sexual form (1995). wing up, of A. corruptrix is
low and Nordlander The measure- not the form larshemi described Ronquist (1989). by Docters van
ments and used herein include: FI - Leeuwen but abbreviations a new form, which we describe here-
FI2, 1st and subsequent flagellomeres; POD (post- under.
ocellar distance), the distance between the inner
margins of the posterior ocelli; OOD (oculo-ocel-
lar of distance) for distance between inner margin Andricus corruptrix (Trotter) new bisexual form
eye and lateral ocelli; COD (central-ocellar distance)
for distance between lateral and central ocelli. Studied material:Experimental material,2 males and 5 females
ofthe previously described experiment (VI-1997/ IV-1998); 2
males and 2 females; material deposited in Barcelona University;
2 females (also paratypes) deposited in MNHN Results (Paris).
1.6-1.8 Length: mm (for and 1.8-2 mm In found inside the males) (for April, 1998, 8 isolated galls were
males females). sleeve, from which we reared 5 alive females, 2 Colour: All the black. and body Wing veins dark brown. one dead parasitoid. This parasitoid is not a cynipid Female head la& Without and from the branch used in the, (Figs. lb). pubescence; Parasitoid, may come in dorsal view around 2.5 times wider experiment. One gall remained closed. than long.
Genae coriaceous, not broadenedbehind compound 266 R. Folliot et al. - Generationalternation in the hymenopterid Andricus corruptrix
Fig. I. A. corruptrix bisexual form: a) head of in dorsal view; b) head in frontal view; c) lateral view; d) branch with some galls; e)
isolated gall.
OOD 1.5 times lateral ocel- eyes. POD twice OOD;
lus diameter and more or less equal to COD (ratio
POD:OOD:COD:lateral ocellus diameter is
45:23:23:15). Coriaceous sculpture. Clypeus con-
spicuous and suboval in shape. Face with only some
very short and weak irradiating striae around clypeus,
never reaching neither antenna! toruli nor compound
Transfacial line less eye margin. more or equal to
eye height. Diameter of toruli around twice the Fig. 2. Antenna ofA. corruptrix bisexual form: a) female (scale than dis- distance between them and slightly larger bar 200 micrometres); b) male (scale bar 500 micrometres)
and Male head tance between toruli eye margin.
similar to female.
Female antenna (fig. 2a) with 13 segments, 0.8 segments, the third one curved, dorsally flattened,
times body length or slightly longer; pedicel 2 times proximally excavate and distally expanded; num-
longer than wide; FI is 1.2 times longer than F2 ber of sensilla in each segments higher (6-8) than
and 1.6 times pedicel length; following flagellomeres in females.
in gradually decreasing length, last flagellomeres Female mesosoma (figs. 3b, 3d, 3f). Without pu-
conspicuously longer than wide and last more or bescence except for some hairy zones in the pro-
less twice its width. Male antenna (fig. 2b) with 14 podeum, with coriaceous sculpture. Notauli com- Contributions to Zoology, 73 (4) - 2004 267
Fig 3. Scutum in dorsal view (a, b), scutellum in dorsal view (c, d) and propodeum(e,f) of the bisexual form ofAndricus improprius
& A. form new status n. sp. (a, c, e) and corruptrix new (b, d, f)
plete and deep in all its length, always reaching pletely striated with some striation also in pronotum.
and wid- Scutellum pronotal margin, converging posteriorly subquadrate, only slightly wider than long, notauli width least third of with reticulated ening (posterior at one closely sculpture, not marginated distance area and between them), delimiting an with laterally not lobed posteriorly. Scutellar foveae
with regular closely reticulate sculpture (not longitudi- (Fig. 3d) oval, a transversal disposition, smooth,
Median mesoscutal im* and not nally striated) (Fig. 3b). shiny pubescent inside, not delimited poste-
absent. almost com- a carina and pression Mesopleuron (Fig. 1c) riorly by separated from each other by 268 R. Folliot et al. - Generation alternation in the hymenopterid Andricus corruptrix
carina. outside. The surface of a median Lateral carinae of propodeum (Fig. and apical part projected
without 3lj thin, ofan uniform thickness, bowed outwards, the gall is brown, nearly smooth, striation,
closed The internal area shiny and smooth, not in its and with a short and deciduous pubescence. superior part (a carina close to the superior part of tops are pointed, not rounded, and the gall wall is
thin. Adults leave the a propodeal area in A. larshemi). Wings hyaline. Fore- very gall through circular, wing margin with short and scattered setae in its big and lateral opening below the top. anterior margin; radial cell 4-4.5 times longer than broad; 2r vein angled; areolet usually conspicuous.
claws with basal lobe and Taxonomic f. Tarsal of legs an acute comments on Andricus corruptrix
with forming a close angle; anterior tibiae provided larshemi D. van L. & D.-M., 1958 short and applied hairs. Male mesosoma similar to
“ ” female. The insects named larshemi by Docters van
Female metasoma. Slightly shorter than head plus Leeuwen & Dekhuijzen-Maaland (1958) do not mesosoma together; without pubescence except at represent the sexual form of A. corruptrix but cor-
rd the basis of the 3 abdominal tergite. Third abdomi- respond to another valid species only known by its nal tergite covering between Vi and Ya of metasoma sexual form. Consequently, what is the status of in “ dorsal vision. Ventral spine of hypopigium slightly the name larshemi”?. longer than broad, with sparse setae, which do not As evidences in this study show the life cycle of form an apical tuft. Male metasoma smaller than Andricus corruptrix was erroneously described by mesosoma. Docters van Leeuwen (1956: 255). The sexual form
A. forma larshemi was given the name corruptrix
from other Diagnosis. It can be easily separated in Docters van Leeuwen & Dekhuijzen-Maasland sexual forms of the A. kollari group by the follow- (1958: 102, 104) and not in Docters van Leeuwen ing combinations of characters (special emphasis (1956) as it’s stated in Wiebes-Rijks (1978: 140).
A. “ ” is given to the separation of A. larshemi and According ICZN, the name larshemi is not avail- corruptrix bisexual form because they are the most able for two reasons: because it’s described as a similar forms). Upper part of propodeum not closed form of a previous species (art. 45.6.1) and because
from by a complete carina, while in A. larshemi it is it’s described galls and not adults (arts. 1.3.6 conspicuously closed (figs. 3e & 3f), mesoscutal i 13.1.1). sculpture between notauli regularly reticulated, not Actually, Docters van Leeuwen & Dekhuijzen- elongated like in A. larshemi (figs. 3c & 3d) and Maasland (1958: 102) stated that sexual adults of
“ ” scutellar foveae not delimited posteriorly while they larshemi form are indistinguishable from bisexual
A. arc delimited in larshemi (figs. 3c & 3d), sculp- adults ofAndricus kollari. Moreover Eady & Quin-
between notauli reticulated in lan differences between sexual ture regularly (like (1963: 49) gave
like A. circulans) but never elongated in A. larshemi females of“larshemi” and circulans since males ,
(figs. 3a & 3b), scutellar sculpture weaker than in were unknown to them (Eady & Quinlan, 1963:
circu- A. lignicolus (= vanheurni) or A. kollari (= 52). These authors never explicitly mentioned that
“ ” lars), and regularly reticulated and not coarse. larshemi belong to a taxonomical category other
than ‘form’, and thus the name continues as not
Distribution. Only known in experimental condi- available. Later, Wiebes-Rijks (1978) gave diag-
“ ” tions from France, where the unisexual form has nostic characters for both sexes of larshemi but probably been introduced recently. continued considering them as a form of A. cor-
ruptrix, and thus the name can’t be used according
ICZN (art. 15.2, 45.6.1 i 46.6.3). Gall. Similar to other sexual galls of the A. kollari
“ First author larshemi” different taxo- small bud giving a group. A gall (fig. Id), measuring be- nomic status 2 and 2.5 from (A. corruptrix ssp larshemi) was tween mm basis to top, on Q. cerris,
Ambrus (1974: 26), but his name can’t be used either, isolated (unlike those ofA. circulars or A. larshemi), because he described and 1CZN with its basal part hidden from view by bud-scales only galls, according 73 - Contributions to Zoology, (4) 2004 269
infra terms can’t be used after 1960 but is in Baillac again, specifics Brittany very rare (France) where
and several it (art. 15.2) gall descriptions are not considered since decades has been found only once. valid for taxonomic 1930 2 nd purposes after (art. 1.3.6). A. kollari, in Baillac emerges (in the year) usu-
“ ” Therefore larshemi remains available after from At the not ally July. any rate, sexual adults ob-
Ambrus’ tained do paper. not belong neither to A. lignicolus (=van-
Melika consider A. Finally, et al. (2000: 269) heurni) nor to kollari (=circulans), as we have
“ ” larshemi as a different species from data shown discussed above. in this but The of both study, it’s not characterised morphologi- presence sexes originating from one
and thus “larshemi” unisexual female cally, can’t be available either may seem strange as the norm in
(art. 1.3.6). Andricus is usually to find sexual adults originat-
At from one unisexual female of last we consider the previously named form ing only one sex (all
“ males all larshemi' neither linked to Andricus or females). But in the allied species A. , corruptrix
Andricus kollari, Folliot (1964: 493) observed a similar situ- nor to Andricus kollari and lignicolus as ation in several instances, one or a few sexual a valid species only known by its sexual form but usually
females in the of a male uni- for taxonomic reasons we rename this species as progeny producing sexual female small Andricus and eventually when numbers impropius n. stat. & n. sp.
are females At concerned, more than males. any
rate both sexes described here are morphologically Andricus improprius Bellido & Pujade-Villar n. stat. similar and different from the other three known & n. sp. sexual forms in the group. Therefore we consider
the ndw sexual form as the valid form alternant Andricus corruptrix forma larshemi Docters van Leeuwen & with Dekhuijzen-Maasland (1958) [not available] Andricus corruptrix (Trotter).
Andricus To corruptrix (=larshemi ) Eady & Quinlan (1963) [not sum all the known up, life-cycles of the spe- available] cies of A. kollari in group are presented table I. Andricus corruptrix forma larshemi Wiebes-Rijks (1978) [not Sexual ofother of the generation species group may available] have O. cerris as host as most known species but, Andricuscorruptrix ssp larshemi Ambrus (1974) [not available] the host tree of these sexual still Andricus larshemi Melika, Csoka & Pujade-Villar (2000) [not generations is
unknown and therefor here indicated with available] a ques-
tion mark.
Studied material(deposited in Zoologisch Museum Amsterdam).
Lectotype 9 herein designatedwith following labels: Andricus corruptrix Schlechtendal forma larshemi 0*9 D. v. L. el J. Acknowledgements M.D.-M.Leersum, 10.v. 1955” (handwriting); “Lectotype” (red label); ‘Andricus Bellido & det. 2000” improprius, Pujade-Villar We thank Willem Hogenes (Zoologisch Museum Amsterdam, (white label). Paralectotype 9 with the same labels except for for material Netherlands) loaning and for his help in search of a "Paralectotype” label (red). Docters van Leeuwen experimental material, and Dr. Karla
Schneider (Institut fuerZoologie der Marthin-Luther-Universitaet, characters for adults of A. Halle, Diagnostic improprius Germany) for loan oftype material ofA. corruptrix. Finally
remain Dr. are given in Wiebes-Rijks (1978) and in this study we grateful to M.A. Alonso Zarazaga (Museo de Ciencias Naturales, Madrid, Spain) for comments concerning (fig. 3a, c, e), galls are described in Docters van 1CZN, to Dr. G. Melika (Systematic ParasitoidLab, Hungary) Leeuwen & Dekhuijzen-Maasland (1958). for additional material of A. corruptrix and A. ambiguus for
and comparison to ourfriend Albert Maso for pictures ofAndricus
in corruptrix galls this paper,
Discussion
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This last as in Askew Neill MP. 1993. species can emerge as early May RR, Parasitoids and inquilines of - 270 R. Folliot et al. Generation alternation in the hymenopterid Andricus corruptrix
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