Es descriu de Mèxic una nova espècie de cinípid cecidògen, Andricus fusiformis Pujade-Villar n. sp. Aquesta espècie es coneix a partir
de la seva forma asexual provocant gales a les branques joves de Quercus obtusata. Es donen dades referents a la diagnosi, distribució i
biologia d'aquesta nova espècie. S'il·lustren els caràcters diagnòstics.A new cynipid gallwasp species, Andricus fusiformis Pujade-Villar n. sp., is described from Mexico. This species is known only from
asexual females and induces stem galls in young shoots of Quercus obtusata. Diagnosis, biology and distribution data of this new species is
given. Diagnostic characters are illustrated

Equihua Martínez, Amando

Estrada Venegas, Edith G.

Ferrer-Suay, Mar

García Martiñón, Rosa D.

Pujade i Villar, Juli

Revistes Catalanes amb Accés Obert

93
Butlletí ICHN 78, 2014
GEA, FLORA ET FAUNAGEA, FLORA ET FAUNA
A new Mexican species (Hym., Cynipidae) inducing tuberous 
galls in twigs of oaks (Fagaceae)
Juli Pujade-Villar*, Rosa D. García-Martiñón**, Amando Equihua-Martínez**,
Edith G. Estrada-Venegas** & Mar Ferrer-Suay*
* Universitat de Barcelona. Facultat de Biologia. Departament de Biologia Animal. Avda. Diagonal, 645. 08028 Barcelona, Spain. A/e: jpujade@ub.edu; 
mar.ferrer.suay@gmail.com
** Instituto de Fitosanidad. Colegio de Postgraduados. 56230 Montecillo, Texcoco, Estado de México, México. A/e: delisgama@live.com.mx; equihuaa@
colpos.mx; estradae@colpos.mx
Autor per a la correspondència: Juli Pujade-Villar. A/e; jpujade@ub.edu
Rebut: 15.10.2014; Acceptat: 15.11.2014; Publicat: 29.12.2014
Abstract
A new cynipid gallwasp species, Andricus fusiformis Pujade-Villar n. sp., is described from Mexico. This species is known only from 
asexual females and induces stem galls in young shoots of Quercus obtusata. Diagnosis, biology and distribution data of this new species is 
given. Diagnostic characters are illustrated. 
Key words: Cynipidae, tuberous gall-wasp, Andricus, taxonomy, morphology, distribution, biology. Mexico.
Resum
Una nova especie Mexicana (Hym., Cynipidae) inductor de gales en branques de roures (Fagaceae). 
Es descriu de Mèxic una nova espècie de cinípid cecidògen, Andricus fusiformis Pujade-Villar n. sp. Aquesta espècie es coneix a partir 
de la seva forma asexual provocant gales a les branques joves de Quercus obtusata. Es donen dades referents a la diagnosi, distribució i 
biologia d’aquesta nova espècie. S’il·lustren els caràcters diagnòstics. 
Paraules clau: Cynipidae, gales tuberoses, Andricus, taxonomia, morfologia, distribució, biologia, Mèxic.
Butlletí de la Institució Catalana d’Història Natural, 78: 93-98. 2014 ISSN 2013-3987 (online edition): ISSN: 1133-6889 (print edition)
Introduction
The tuberous galls induced by oak wasps are included 
in the Cynipini tribe (Cynipidae). These are located in the 
branches of oaks and more rarely in the roots. Usually they 
have an irregular appearance and are multilocular. The surfa-
ce is finely wrinkled. The coloration is gray with green tones 
when young (similar to the color of the bark when fresh) and 
woody brown at maturity. Hymenoptera induce these galls 
which remain within it for several years. To obtain the adults 
from the harden tissues of the gall is very difficult, even for 
the emerging adults which sometimes die before emergence 
inside the gall.  . In addition, being so longly exposed, there 
are galls with a high rate of parasitism and inquilism, so the 
number of inducers adults is often scarce, very scarce or even 
nonexistent (when all inducing larvae have been attacked), 
making difficult the precise determination because of  the ab-
sence of the inducer adult.
In the United States, northern Mexico, there are several 
species of different genera involved in the formation of this 
type of galls: Andricus Hartig, Bassettia Ashmead, Callirhytis 
Förster, and Loxaulus Mayr (Weld, 1957, 1959, 1960; Burks, 
1979) ; they are located in branches, roots and / or trunks near 
the ground (Weld, 1921). In the Neotropical region highlig-
hts Odontocynips Kieffer (Pujade-Villar, 2008, Mediator et 
al, 2011) and Zapatella Pujade-Villar & Melika (Pujade-Vi-
llar, unpublished data). In Mexico (Pujade-Villar et al, 2012, 
2013) only Andricus produces true tuberous galls, although 
Loxaulus Mayr may produce thin fusiform swellings on the 
branches of oaks (Pujade-Villar et al, 2014.).
The Mexican species that cause tumberous galls have been 
recently revised in Pujade-Villar et al, (2012, 2013). Diffe-
rent species produce very similar or the same galls,  if the 
adult is not obtained, they cannot be identified. In previous 
studies, multiple tuberous deformations are pending of iden-
tification, precisely because no adults were obtained. One of 
them is described in this study which produced, unlike all 
known Mexican species, thick spindle galls on branches of 
Q. obtusata. After having collected them on multiple occasi-
ons finally in 2014 we obtained the first adults.
Material and methods
Cynipid galls were collected in Mexico D.F from Quercus 
obtusata Humb. & Bonpl. which belongs to the Quercus sec-
tion. The galls were preserved in aerated receptacles waiting 
for adults emergence. The reared adults were preserved in 
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J. PUJADE-VILLAR ET AL.
70 % ethanol until the morphological study was done at the 
Barcelona University. We also examined the reared parasi-
toids and some galls without any emergences were dissected 
for inner examination. 
This paper follows the current terminology of morpho-
logical structures (Liljeblad & Ronquist 1998; Melika 2006). 
Abbreviations for forewing venation follow Ronquist & Nor-
dlander (1989), cuticular surface terminology follows  Harris 
(1979). Measurements and abbreviations used include: F1–
F12, first and subsequent flagellomeres; POL (post-ocellar 
distance) is the distance between the inner margins of the 
posterior ocelli; OOL (ocellar-ocular distance) is the distance 
from the outer edge of a posterior ocellus to the inner margin 
of the compound eye; LOL (lateral-frontal ocellar distance) 
is the distance between lateral and frontal ocelli. The width 
of the forewing radial cell was measured from the margin of 
the wing to the Rs vein.
SEM images of the new species were taken with the field-
emission gun environmental scanning electron microscope 
(FEI Quanta 200 ESEM) which was used for high-resolution 
imaging without gold-coating , in order to preserve the speci-
mens. The wasp habitus and the forewing of the adult was 
photographed with a digital camera-21C INFINITYX at-
tached to a compound microscope Zeiss Discovery V8; the 
program DeltaPix View AZ-Pro was used to combine the se-
ries of images obtained in one single image.
The type material of the new species is deposited in the J. 
Pujade-Villar (JP-V) collection at the University of Barce-
lona (UB, Spain). 
Results
Andricus fusiformis Pujade-Villar n. sp.  (Figs 1–3)
Type material
HOLOTYPE ♀ with the following labels: “R106: MEX, San 
Juan Coajomulco (Jocotitlan, México), Ex. Q. obtusata, (19.
vi.2014) 23.vi.2014, (leg. Delia)” (black label); Holotype of 
Andricus fusiformis Pujade-Villar n. sp. Desig. JP-V 2014” 
(red label). PARATYPE: “R115: MEX, San Juan Coajo-
mulco (Jocotitlan, México), Ex. Q. obtusata, (12.vii.2014) 
14.vii.2014: 1 ♀ (Leg. Delia)” (black label); Paratype of An-
dricus fusiformis Pujade-Villar n. sp. desig. JP-V 2014” (red 
label). Type material deposited in JP-V collection.
Additional material (2♀)
1 ♀ with the same data of Holotype (extracted) and 1♀ with 
the same data of Paratype (extracted).
Diagnosis
This new species belongs to the Andricus tuberous galls 
group, which has metasomal terga completely pubescent and 
forewings ciliated in margin as A. furnaceus Kinsey, 1920; 
A. guanajuatensis Pujade-Villar, 2013; A. tumeralis Pujade-
Villar, 2009; A. montezumus Beutenmüller 1913 and A. dug-
esi Beutenmüller, 1917. Nevertheless, the new species differs 
from all these species by the mesoscutum sculpture. Andri-
cus fusiformis Pujade-Villar n. sp. is the only species without 
linear elements in mesoscutum, punctuated (with fine trans-
verse and parallel carinae in A. guanajuatensis or, more or 
less strongly rugose in the rest of species mentioned). Also 
it has the metasoma punctuated, the lateral pubescence in a 
wide band and ventral spine of hypopygium long (6.0-7.0 
times as long as broad).
Description
Asexual female
Length
Female: 3.5–3.6 mm (n=2). 
Coloration
(Fig. 2d) black. Head brown, darker in lower face. Meso-
soma black, with brown patches. Metasoma black, red-brown 
in the anterior third; hypopigyum light. Legs dark, brown-
black, coxae black. Vein of forewings brown.
Head 
(Figs 1a–b) strongly coriaceous with dense white setae; 
transversally oval in frontal view, 1.3 wider than high; as 
broad as mesosoma; gena coriaceous with come carinae pos-
teriorly, strongly broadened behind eye, 0.6 times as broad 
as cross diameter of eye, measuring along transfacial line; 
malar space 0.3 times as long as the height of the eye, co-
riaceous, without striae radiating from clypeus, malar sul-
cus absent; transfacial line similar to compound eye height. 
POL:OOL:LOL=8:3:3, diameter of lateral ocellus 2.0; 2.5 
times as broad as long in dorsal view; lower face and frons 
without carinae; clypeus impressed, alutaceous, rounded 
ventrally, medially not incised, anterior tentorial pits distinct; 
Figure 1. Andricus fusiformis n. sp.: (a) mesosoma in dorsal view and 
detail without setae, (b) head and mesosoma in lateral view and (c) an-
tenna.
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A NEW MEXICAN SPECIES (HYM., CYNIPIDAE) INDUCING TUBEROUS GALLS IN TWIGS OF OAKS
epistomal sulcus and clypeo-pleurostomal line distinctly 
impressed; striate radiating for clypeus present, but short and 
scarce, not reaching the base of the compound eyes and the 
malar space.  
Antenna 
(Fig. 2c) longer than length of head + mesosoma (42:28), 
with 12 flagellomeres; F1 slightly broader distally, subequal 
to F2; subsequent flagellomeres progressively shortened, F12 
slightly longer than F11. Antennal formula: 6: 4(x2.5): 9(x3): 
8.5: 8: 7: 6.5: 6: 5: 4: 4: 4: 3: 4. Placodeal sensilla on F5–F12. 
Mesosoma 
(Figs 1, 2c) around 1.6 times longer than high, convex in 
lateral view. Sides of pronotum coriaceous with weak and 
parallel carinae in the lower 2/3 latero-posterior. Mesoscu-
tum pubescent, broader than long in dorsal view; coriaceous, 
without linear elements, with piliferous points denser in the 
posterior half; notauli complete, superficial anteriorly, nar-
row, reaching tegulae level, weakly converging posteriorly, 
median mesoscutal line absent; anterior parallel lines differ-
entiated, alutaceous, present until tegulae level; parapsidal 
lines present, smooth, beyond tegulae level. Mesopleuron 
coriaceous, pubescent, with delicate and dense carinae, aluta-
celus and shiny without carinae posteriorly. Metapleuron sul-
cus diferenciated reaching mesopleuron in 2/3 height. Mes-
oscutellum longer than broad, weak rugose and coriaceous; 
scutellar foveae differentiated, small, oval, superficial, with 
alutaceous bottom, widley separated and not delimited pos-
teriorly by a carina. Lateral parts of propodeum uniformly 
alutaceous, densely pubescent; lateral propodeal carinae 
curved, central propodeal area nearly smooth and glabrous. 
Metascutellum subrectangular, alutaceous, strongly incised 
ventrally. Ventral bar of metanotal trough alutaceous; metan-
otal trough alutaceous, with sparse setae. 
Figure 2. Andricus fusiformis n. sp.: (a) head in dorsal view, (b) head in frontal view, (c) propodeum, (d) tarsal claw, (e) metasoma in lateral view, 
and (f) ventral spine of hypopygium.
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Forewing 
(Fig. 3e) translucent, 1.2 times longer than body, pubes-
cent, with cilia on margins; radial cell opened, around 4.0 
times as long as broad; areolet present; Rs+M heading to-
wards the middle of the basal vein, nearly reaching it. 
Legs 
(Fig. 2d) base of tarsal claws with strong tooth. 
Metasoma 
(Figs 2e–f) slightly shorter than head+mesosoma, longer 
than high. 2nd metasomal tergite with a lateral patch of white 
setae, not punctuate dorsaly, all subsequent tergites uniformly 
and entirely punctate. Lateral pilosity forming a band. Promi-
nent part of ventral spine of hypopygium needle-like, 6.0–7.0 
times longer than broad, with sparse setae laterally which not 
form apical tuft in the apex.
Gall 
(Figs 3a-c) galls are developing on thin branches of Q. ob-
tusata. It is a plurilocular gall, woody, more or less fusiform 
(35-45 mm in length and 12-18 mm in diameter), composed 
by different subunits more or less fused. Coloration gray with 
green tones; adults emerge before the gall becomes hard and 
Figure 3. Andricus fusiformis n. sp.: (a) fresh gall, (b) gall cracked observing a larval chamber (see the arrow) and detail of larval chamber (the ar-
row indicates the petiole of the larval chamber), (c) fresh gall opened, (d) habitus and (e) forewing. 
J. PUJADE-VILLAR ET AL.
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take a woody brown color. The surface is glabrous with some 
longitudinal grooves on the mature gall. The inner larval 
chambers are located in the internal tissues; when the gall is 
cracked we can observe the larval chambers inside the crack. 
The single larval chamber is globular (2.6 mm. of diameter), 
dirty yellow, with a hard woody wall (0.3 mm think), which 
is connected with the tissue of the gall by a basal peduncle; at 
the top there is a small umbilicus.
Host plant
Quercus obtusata Humb. & Bonpl. (section Quercus of 
Quercus, white oaks), distributed in many states of Mexico at 
620–2580 m a.s.l. (Valencia, 2004). Galls collected in branch-
es of Q. crassifolia Humb and Q. rugosa Née in the same 
area of type material are similar but  no adults have  emerged; 
we cannot assure these galls are the same of collected in Q. 
obtusata, although all of them belong to the Quercus section.
Distribution
Currently known only from Mexico State. Galls were col-
lected in San Juan Coajomulco (Jocotitlan).
Biology
Only the asexual (parthenogenetic) females are known. 
Galls develop in dry season and adults emerge in July. No 
parasitoids  emerged, only Synergus on 24.vii.2014.
Etymology
The species name refers to the shape of gall.
Discussion
Gallwasps from different genera that induce woody stem 
swelling-like galls on oaks -Andricus, Bassettia, Callirhy-
tis, Holocynips Loxaulus and Odontocynips- are common in 
North and Central America. However, galls of different spe-
cies and genera in stems, branches and twigs are quite dif-
ferent in size and shape. For example, Bassettia and Loxau-
lus induce galls in twigs without visible twig enlargements. 
Many Callirhytis species induce small spindle-shaped or 
club-shaped galls in twigs. Gall shape depends on the posi-
tion of the twig: if terminal, the gall is club-shaped; if the 
gall is in the middle of the twig, then it is usually spindle-
shaped. There is a smaller group of species belonging to An-
dricus, Callirhytis, Dryocosmus, Eumayria, Holocynips and 
Odontocynips, which induce large, tuberous galls on stems 
and twigs, commonly located in the tree crown or at the base 
of young sprouts (near or slightly underground surface), so-
called «subterranean galls» (Weld, 1921). Tuberous galls are 
a phenotype extended among gallwasps, which have evolved 
independently several times in different phylogenetic unrela-
ted groups. In the Neotropical area a tuberous gall produced 
by Zapatella (Pujade-Villar unpublished data) has been de-
tected when these are usually restricted to Odontocynips in 
Panama and Costa Rica.
In Mexico 12 Andricus species produce tuberous galls 
(Table 1). Most of them, when the host is mentioned, are as-
sociated to white oaks. Only one species is found in red oak: 
A. duguesi. Nevertheless, 6 species belong to an unknown 
host (Table 1).
The species here described differs from all Andricus spe-
cies with tuberous galls by morphology (see diagnosis) and 
by galls shape because it is the only species with fusiform 
tuberous galls in branches in Mexico. The North-American 
species Andricus wheeleri (Beutenmüller, 1907) has a simi-
lar morphology of A. fusiformis gall, nevertheless, the adults 
are completely different; in Andricus wheeleri the metasoma 
is scarcely pubescent on tergite II and the mesoscutum sculp-
ture is strongly rugose (all metasoma terga pubescent and 
mesoscutum sculpture punctuated without linear elements in 
A. fuciformis).
Acknowledgements
We thank Christian Equihua for reviewing the English.
Table 1. Andricus species producing tuberous galls, their host (including Quercus section) and state where these species have been collected.
Andricus tuberous galls species Quercus species Section State
A. bonanseai Mayr sp ¿? unkown
A. carrilloi Pujade-Villar Q. rugosa Née Quercus Zacatecas
 Q. obtusata Humb. & Bolpl. Quercus Morelos
A. duguesi Beutenmüller Q. castanea Née Lobatae Guanajuato , Puebla
A. durangensis Beutenmüller sp ¿? Durango
A. furnaceus Kinsey, 1920 sp ¿? San Luis Potosí
A. guanajuatensis Pujade-Villar Q. castanea Née Quercus Guanajuato
 Q. obtusata Humb. & Bolpl. Quercus Querétaro
A. fusciformis n. sp. Q. obtusata Humb. & Bolpl. Quercus Mexico
A. montezumus Beutenmüller sp ¿? unkown
A. peredurus Kinsey sp ¿? San Luis Potosí
A. santafe Pujade-Villar Q. laeta Liebm Quercus Mexico DF
A. tumefaciens Pujade-Villar Q. chihuahuensis Quercus Zacatecas
A. tumeralis Pujade-Villar sp ¿? Guanajuato
A NEW MEXICAN SPECIES (HYM., CYNIPIDAE) INDUCING TUBEROUS GALLS IN TWIGS OF OAKS
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J. PUJADE-VILLAR ET AL.


A New Mexican species (Hym., Cynipidae) inducing tuberous galls in twigs of oaks (Fagaceae)

Hemeroteca Cientifica Catalana

93Butlletí ICHN 78, 2014GEA, FLORA ET FAUNAGEA, FLORA ET FAUNAA new Mexican species (Hym., Cynipidae) inducing tuberous galls in twigs of oaks (Fagaceae)Juli Pujade-Villar*, Rosa D. García-Martiñón**, Amando Equihua-Martínez**,Edith G. Estrada-Venegas** & Mar Ferrer-Suay** Universitat de Barcelona. Facultat de Biologia. Departament de Biologia Animal. Avda. Diagonal, 645. 08028 Barcelona, Spain. A/e: jpujade@ub.edu; mar.ferrer.suay@gmail.com** Instituto de Fitosanidad. Colegio de Postgraduados. 56230 Montecillo, Texcoco, Estado de México, México. A/e: delisgama@live.com.mx; equihuaa@colpos.mx; estradae@colpos.mxAutor per a la correspondència: Juli Pujade-Villar. A/e; jpujade@ub.eduRebut: 15.10.2014; Acceptat: 15.11.2014; Publicat: 29.12.2014AbstractA new cynipid gallwasp species, Andricus fusiformis Pujade-Villar n. sp., is described from Mexico. This species is known only from asexual females and induces stem galls in young shoots of Quercus obtusata. Diagnosis, biology and distribution data of this new species is given. Diagnostic characters are illustrated. Key words: Cynipidae, tuberous gall-wasp, Andricus, taxonomy, morphology, distribution, biology. Mexico.ResumUna nova especie Mexicana (Hym., Cynipidae) inductor de gales en branques de roures (Fagaceae). Es descriu de Mèxic una nova espècie de cinípid cecidògen, Andricus fusiformis Pujade-Villar n. sp. Aquesta espècie es coneix a partir de la seva forma asexual provocant gales a les branques joves de Quercus obtusata. Es donen dades referents a la diagnosi, distribució i biologia d’aquesta nova espècie. S’il·lustren els caràcters diagnòstics. Paraules clau: Cynipidae, gales tuberoses, Andricus, taxonomia, morfologia, distribució, biologia, Mèxic.Butlletí de la Institució Catalana d’Història Natural, 78: 93-98. 2014 ISSN 2013-3987 (online edition): ISSN: 1133-6889 (print edition)IntroductionThe tuberous galls induced by oak wasps are included in the Cynipini tribe (Cynipidae). These are located in the branches of oaks and more rarely in the roots. Usually they have an irregular appearance and are multilocular. The surfa-ce is finely wrinkled. The coloration is gray with green tones when young (similar to the color of the bark when fresh) and woody brown at maturity. Hymenoptera induce these galls which remain within it for several years. To obtain the adults from the harden tissues of the gall is very difficult, even for the emerging adults which sometimes die before emergence inside the gall.  . In addition, being so longly exposed, there are galls with a high rate of parasitism and inquilism, so the number of inducers adults is often scarce, very scarce or even nonexistent (when all inducing larvae have been attacked), making difficult the precise determination because of  the ab-sence of the inducer adult.In the United States, northern Mexico, there are several species of different genera involved in the formation of this type of galls: Andricus Hartig, Bassettia Ashmead, Callirhytis Förster, and Loxaulus Mayr (Weld, 1957, 1959, 1960; Burks, 1979) ; they are located in branches, roots and / or trunks near the ground (Weld, 1921). In the Neotropical region highlig-hts Odontocynips Kieffer (Pujade-Villar, 2008, Mediator et al, 2011) and Zapatella Pujade-Villar & Melika (Pujade-Vi-llar, unpublished data). In Mexico (Pujade-Villar et al, 2012, 2013) only Andricus produces true tuberous galls, although Loxaulus Mayr may produce thin fusiform swellings on the branches of oaks (Pujade-Villar et al, 2014.).The Mexican species that cause tumberous galls have been recently revised in Pujade-Villar et al, (2012, 2013). Diffe-rent species produce very similar or the same galls,  if the adult is not obtained, they cannot be identified. In previous studies, multiple tuberous deformations are pending of iden-tification, precisely because no adults were obtained. One of them is described in this study which produced, unlike all known Mexican species, thick spindle galls on branches of Q. obtusata. After having collected them on multiple occasi-ons finally in 2014 we obtained the first adults.Material and methodsCynipid galls were collected in Mexico D.F from Quercus obtusata Humb. & Bonpl. which belongs to the Quercus sec-tion. The galls were preserved in aerated receptacles waiting for adults emergence. The reared adults were preserved in 94Butlletí ICHN 78, 2014GEA, FLORA ET FAUNAJ. PUJADE-VILLAR ET AL.70 % ethanol until the morphological study was done at the Barcelona University. We also examined the reared parasi-toids and some galls without any emergences were dissected for inner examination. This paper follows the current terminology of morpho-logical structures (Liljeblad & Ronquist 1998; Melika 2006). Abbreviations for forewing venation follow Ronquist & Nor-dlander (1989), cuticular surface terminology follows  Harris (1979). Measurements and abbreviations used include: F1–F12, first and subsequent flagellomeres; POL (post-ocellar distance) is the distance between the inner margins of the posterior ocelli; OOL (ocellar-ocular distance) is the distance from the outer edge of a posterior ocellus to the inner margin of the compound eye; LOL (lateral-frontal ocellar distance) is the distance between lateral and frontal ocelli. The width of the forewing radial cell was measured from the margin of the wing to the Rs vein.SEM images of the new species were taken with the field-emission gun environmental scanning electron microscope (FEI Quanta 200 ESEM) which was used for high-resolution imaging without gold-coating , in order to preserve the speci-mens. The wasp habitus and the forewing of the adult was photographed with a digital camera-21C INFINITYX at-tached to a compound microscope Zeiss Discovery V8; the program DeltaPix View AZ-Pro was used to combine the se-ries of images obtained in one single image.The type material of the new species is deposited in the J. Pujade-Villar (JP-V) collection at the University of Barce-lona (UB, Spain). ResultsAndricus fusiformis Pujade-Villar n. sp.  (Figs 1–3)Type materialHOLOTYPE ♀ with the following labels: “R106: MEX, San Juan Coajomulco (Jocotitlan, México), Ex. Q. obtusata, (19.vi.2014) 23.vi.2014, (leg. Delia)” (black label); Holotype of Andricus fusiformis Pujade-Villar n. sp. Desig. JP-V 2014” (red label). PARATYPE: “R115: MEX, San Juan Coajo-mulco (Jocotitlan, México), Ex. Q. obtusata, (12.vii.2014) 14.vii.2014: 1 ♀ (Leg. Delia)” (black label); Paratype of An-dricus fusiformis Pujade-Villar n. sp. desig. JP-V 2014” (red label). Type material deposited in JP-V collection.Additional material (2♀)1 ♀ with the same data of Holotype (extracted) and 1♀ with the same data of Paratype (extracted).DiagnosisThis new species belongs to the Andricus tuberous galls group, which has metasomal terga completely pubescent and forewings ciliated in margin as A. furnaceus Kinsey, 1920; A. guanajuatensis Pujade-Villar, 2013; A. tumeralis Pujade-Villar, 2009; A. montezumus Beutenmüller 1913 and A. dug-esi Beutenmüller, 1917. Nevertheless, the new species differs from all these species by the mesoscutum sculpture. Andri-cus fusiformis Pujade-Villar n. sp. is the only species without linear elements in mesoscutum, punctuated (with fine trans-verse and parallel carinae in A. guanajuatensis or, more or less strongly rugose in the rest of species mentioned). Also it has the metasoma punctuated, the lateral pubescence in a wide band and ventral spine of hypopygium long (6.0-7.0 times as long as broad).DescriptionAsexual femaleLengthFemale: 3.5–3.6 mm (n=2). Coloration(Fig. 2d) black. Head brown, darker in lower face. Meso-soma black, with brown patches. Metasoma black, red-brown in the anterior third; hypopigyum light. Legs dark, brown-black, coxae black. Vein of forewings brown.Head (Figs 1a–b) strongly coriaceous with dense white setae; transversally oval in frontal view, 1.3 wider than high; as broad as mesosoma; gena coriaceous with come carinae pos-teriorly, strongly broadened behind eye, 0.6 times as broad as cross diameter of eye, measuring along transfacial line; malar space 0.3 times as long as the height of the eye, co-riaceous, without striae radiating from clypeus, malar sul-cus absent; transfacial line similar to compound eye height. POL:OOL:LOL=8:3:3, diameter of lateral ocellus 2.0; 2.5 times as broad as long in dorsal view; lower face and frons without carinae; clypeus impressed, alutaceous, rounded ventrally, medially not incised, anterior tentorial pits distinct; Figure 1. Andricus fusiformis n. sp.: (a) mesosoma in dorsal view and detail without setae, (b) head and mesosoma in lateral view and (c) an-tenna.95Butlletí ICHN 78, 2014GEA, FLORA ET FAUNAA NEW MEXICAN SPECIES (HYM., CYNIPIDAE) INDUCING TUBEROUS GALLS IN TWIGS OF OAKSepistomal sulcus and clypeo-pleurostomal line distinctly impressed; striate radiating for clypeus present, but short and scarce, not reaching the base of the compound eyes and the malar space.  Antenna (Fig. 2c) longer than length of head + mesosoma (42:28), with 12 flagellomeres; F1 slightly broader distally, subequal to F2; subsequent flagellomeres progressively shortened, F12 slightly longer than F11. Antennal formula: 6: 4(x2.5): 9(x3): 8.5: 8: 7: 6.5: 6: 5: 4: 4: 4: 3: 4. Placodeal sensilla on F5–F12. Mesosoma (Figs 1, 2c) around 1.6 times longer than high, convex in lateral view. Sides of pronotum coriaceous with weak and parallel carinae in the lower 2/3 latero-posterior. Mesoscu-tum pubescent, broader than long in dorsal view; coriaceous, without linear elements, with piliferous points denser in the posterior half; notauli complete, superficial anteriorly, nar-row, reaching tegulae level, weakly converging posteriorly, median mesoscutal line absent; anterior parallel lines differ-entiated, alutaceous, present until tegulae level; parapsidal lines present, smooth, beyond tegulae level. Mesopleuron coriaceous, pubescent, with delicate and dense carinae, aluta-celus and shiny without carinae posteriorly. Metapleuron sul-cus diferenciated reaching mesopleuron in 2/3 height. Mes-oscutellum longer than broad, weak rugose and coriaceous; scutellar foveae differentiated, small, oval, superficial, with alutaceous bottom, widley separated and not delimited pos-teriorly by a carina. Lateral parts of propodeum uniformly alutaceous, densely pubescent; lateral propodeal carinae curved, central propodeal area nearly smooth and glabrous. Metascutellum subrectangular, alutaceous, strongly incised ventrally. Ventral bar of metanotal trough alutaceous; metan-otal trough alutaceous, with sparse setae. Figure 2. Andricus fusiformis n. sp.: (a) head in dorsal view, (b) head in frontal view, (c) propodeum, (d) tarsal claw, (e) metasoma in lateral view, and (f) ventral spine of hypopygium.96Butlletí ICHN 78, 2014GEA, FLORA ET FAUNAForewing (Fig. 3e) translucent, 1.2 times longer than body, pubes-cent, with cilia on margins; radial cell opened, around 4.0 times as long as broad; areolet present; Rs+M heading to-wards the middle of the basal vein, nearly reaching it. Legs (Fig. 2d) base of tarsal claws with strong tooth. Metasoma (Figs 2e–f) slightly shorter than head+mesosoma, longer than high. 2nd metasomal tergite with a lateral patch of white setae, not punctuate dorsaly, all subsequent tergites uniformly and entirely punctate. Lateral pilosity forming a band. Promi-nent part of ventral spine of hypopygium needle-like, 6.0–7.0 times longer than broad, with sparse setae laterally which not form apical tuft in the apex.Gall (Figs 3a-c) galls are developing on thin branches of Q. ob-tusata. It is a plurilocular gall, woody, more or less fusiform (35-45 mm in length and 12-18 mm in diameter), composed by different subunits more or less fused. Coloration gray with green tones; adults emerge before the gall becomes hard and Figure 3. Andricus fusiformis n. sp.: (a) fresh gall, (b) gall cracked observing a larval chamber (see the arrow) and detail of larval chamber (the ar-row indicates the petiole of the larval chamber), (c) fresh gall opened, (d) habitus and (e) forewing. J. PUJADE-VILLAR ET AL.97Butlletí ICHN 78, 2014GEA, FLORA ET FAUNAtake a woody brown color. The surface is glabrous with some longitudinal grooves on the mature gall. The inner larval chambers are located in the internal tissues; when the gall is cracked we can observe the larval chambers inside the crack. The single larval chamber is globular (2.6 mm. of diameter), dirty yellow, with a hard woody wall (0.3 mm think), which is connected with the tissue of the gall by a basal peduncle; at the top there is a small umbilicus.Host plantQuercus obtusata Humb. & Bonpl. (section Quercus of Quercus, white oaks), distributed in many states of Mexico at 620–2580 m a.s.l. (Valencia, 2004). Galls collected in branch-es of Q. crassifolia Humb and Q. rugosa Née in the same area of type material are similar but  no adults have  emerged; we cannot assure these galls are the same of collected in Q. obtusata, although all of them belong to the Quercus section.DistributionCurrently known only from Mexico State. Galls were col-lected in San Juan Coajomulco (Jocotitlan).BiologyOnly the asexual (parthenogenetic) females are known. Galls develop in dry season and adults emerge in July. No parasitoids  emerged, only Synergus on 24.vii.2014.EtymologyThe species name refers to the shape of gall.DiscussionGallwasps from different genera that induce woody stem swelling-like galls on oaks -Andricus, Bassettia, Callirhy-tis, Holocynips Loxaulus and Odontocynips- are common in North and Central America. However, galls of different spe-cies and genera in stems, branches and twigs are quite dif-ferent in size and shape. For example, Bassettia and Loxau-lus induce galls in twigs without visible twig enlargements. Many Callirhytis species induce small spindle-shaped or club-shaped galls in twigs. Gall shape depends on the posi-tion of the twig: if terminal, the gall is club-shaped; if the gall is in the middle of the twig, then it is usually spindle-shaped. There is a smaller group of species belonging to An-dricus, Callirhytis, Dryocosmus, Eumayria, Holocynips and Odontocynips, which induce large, tuberous galls on stems and twigs, commonly located in the tree crown or at the base of young sprouts (near or slightly underground surface), so-called «subterranean galls» (Weld, 1921). Tuberous galls are a phenotype extended among gallwasps, which have evolved independently several times in different phylogenetic unrela-ted groups. In the Neotropical area a tuberous gall produced by Zapatella (Pujade-Villar unpublished data) has been de-tected when these are usually restricted to Odontocynips in Panama and Costa Rica.In Mexico 12 Andricus species produce tuberous galls (Table 1). Most of them, when the host is mentioned, are as-sociated to white oaks. Only one species is found in red oak: A. duguesi. Nevertheless, 6 species belong to an unknown host (Table 1).The species here described differs from all Andricus spe-cies with tuberous galls by morphology (see diagnosis) and by galls shape because it is the only species with fusiform tuberous galls in branches in Mexico. The North-American species Andricus wheeleri (Beutenmüller, 1907) has a simi-lar morphology of A. fusiformis gall, nevertheless, the adults are completely different; in Andricus wheeleri the metasoma is scarcely pubescent on tergite II and the mesoscutum sculp-ture is strongly rugose (all metasoma terga pubescent and mesoscutum sculpture punctuated without linear elements in A. fuciformis).AcknowledgementsWe thank Christian Equihua for reviewing the English.Table 1. Andricus species producing tuberous galls, their host (including Quercus section) and state where these species have been collected.Andricus tuberous galls species Quercus species Section StateA. bonanseai Mayr sp ¿? unkownA. carrilloi Pujade-Villar Q. rugosa Née Quercus Zacatecas Q. obtusata Humb. & Bolpl. Quercus MorelosA. duguesi Beutenmüller Q. castanea Née Lobatae Guanajuato , PueblaA. durangensis Beutenmüller sp ¿? DurangoA. furnaceus Kinsey, 1920 sp ¿? San Luis PotosíA. guanajuatensis Pujade-Villar Q. castanea Née Quercus Guanajuato Q. obtusata Humb. & Bolpl. Quercus QuerétaroA. fusciformis n. sp. Q. obtusata Humb. & Bolpl. Quercus MexicoA. montezumus Beutenmüller sp ¿? unkownA. peredurus Kinsey sp ¿? San Luis PotosíA. santafe Pujade-Villar Q. laeta Liebm Quercus Mexico DFA. tumefaciens Pujade-Villar Q. chihuahuensis Quercus ZacatecasA. tumeralis Pujade-Villar sp ¿? GuanajuatoA NEW MEXICAN SPECIES (HYM., CYNIPIDAE) INDUCING TUBEROUS GALLS IN TWIGS OF OAKS98Butlletí ICHN 78, 2014GEA, FLORA ET FAUNAReferencesBURKS, B.D. 1979. Superfamily Cynipoidea. In: Krombein, K.V., Hurd, P.D., Jr., Smith, D.R. & Burks, B.D. (Eds), Catalog of Hy-menoptera in America North of Mexico. Vol. 1. Symphyta and Apocrita. Smithsonian Institution Press,  Washington, DC. P. 1045-1107.LILJEBLAD, J. & RONQUIST, F. 1998. A phylogenetic analysis of higher-level gall wasp relationships (Hymenoptera: Cynipidae). Systematic Entomology, 23: 229-252. MEDIANERO, E., NIEVES-ALDREY, J .L. & PUJADE-VILLAR, J. 2011. The genus Odontocynips Kieffer, 1910 (Hymenoptera, Cynipidae, Cynipini) in Panama, with redescription of Cynips championi Cameron, 1883. Graellsia, 67: 35-46.MELIKA, G. 2006. Gall Wasps of Ukraine. Cynipidae. Vestnik zoo-logii, supplement 21 (1–2), 1–300, 301–644. PUJADE-VILLAR, J. 2008. Description of Odontocynips hansoni n. sp., from Costa Rica (Hymenoptera. Cynipidae). Dugesiana, 15: 79-85. PUJADE-VILLAR, J. & PARETAS-MARTÍNEZ, J. 2012. A new species of woody tuberous oak galls from Mexico (Hymenoptera: Cynipidae) and notes with related species. Dugesiana, 19 (2): 79-85.PUJADE-VILLAR, J., PÉREZ-GARCÍA, A. G., EQUIHUA-MAR-TÍNEZ, A.,  ESTRADA-VENEGAS, E. G., CIBRIÁN-TOVAR, D., BARRERA-RUÍZ, U. M. & FERRER-SUAY, M. 2013. Re-view of Andricus species (Hym., Cynipidae) producing woody tuberous oak galls in Mexico and bordering areas of United States. Dugesiana, 20 (2): 183-208. PUJADE-VILLAR, J., CIBRIÁN-TOVAR, D., EQUIHUA-MAR-TÍNEZ, A.,  ESTRADA-VENEGAS, E. G., BARRERA-RUÍZ, U. M. & MELIKA, G. 2014. First Record of Loxaulus Mayr from Mexico, with Descriptions of Two New Species (Hymenoptera: Cynipidae, Cynipini). Southwestern Entomologist, 39 (2):343-354.RONQUIST, F. & NORDLANDER, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement, 33: 1-60.WELD, L.H. 1921. American gallflies of the family Cynipidae producing subterranean galls on oak. Proceedings of the United States National Museum, 59: 187-246.WELD, L. H. 1957. Cynipid galls of the Pacific Slope. Ann Arbor, Michigan, Privately printed.WELD, L. H. 1959. Cynipid galls of the Eastern United States. Ann Arbor, Michigan, Privately printed.WELD, L. H. 1960. Cynipid galls of the Southwest (Hymenoptera, Cynipoidea) an aid to their identification. Ann Arbor, Michigan, Privately printed.J. PUJADE-VILLAR ET AL.

A New Mexican species (Hym., Cynipidae) inducing tuberous galls in twigs of oaks (Fagaceae)

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