Description of Five New Species of Inquiline Oak Gall Wasps of the Genus Synergus Hartig (Hymenoptera, Cynipidae: Synergini) with Partially Smooth Mesopleurae From Mexico

Abstract

Here we describe five new species of Synergus Hartig with partially smooth mesopleurae that were obtained from oak cynipid galls on different species of Quercus L. in Mexico: Synergus cibriani sp. nov., Synergus gilletti sp. nov., Synergus longimalaris sp. nov., Synergus longiscapus sp. nov. and Synergus striatifrons sp. nov. To date, this trait has been observed in at least 18 American species, 16 recorded in North America (United States of America) and only 2 in Central America (Panama): Synergus elegans Nieves-Aldrey and Medianero and Synergus laticephalus Nieves-Aldrey and Medianero. The type material and additional material of Synergus punctatus Gillette from the United States have been examined and compared with S. gilletti sp. nov. The type material and additional material of S. elegans from Panama and Synergus citriformis (Ashmead) from the United States have also been examined and compared. This study records Synergus elegans in Mexico for the first time and classifies it as a syn. nov. of S. citriformis. Descriptions and diagnoses for the new species and a key to their identification and differentiation from their closest related species are given. Their distribution and host associations are also commented on.

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© 2017 Academia Sinica, Taiwan
Open Access
Description of Five New Species of Inquiline Oak Gall Wasps of the
Genus Synergus Hartig (Hymenoptera, Cynipidae: Synergini) with
Partially Smooth Mesopleurae From Mexico
Irene Lobato-Vila1,* and Juli Pujade-Villar1
1Universitat de Barcelona, Facultat de Biologia, Departament de Biologia Evolutiva, Ecologia i Ciències Ambientals, Avda. Diagonal 645,
08028-Barcelona, Spain
(Received 5 September 2017; Accepted 14 November 2017; Published 19 December 2017; Communicated by Chiun-Cheng Ko)
urn:lsid:zoobank.org:pub:DAB4D865-5183-4318-ADD6-80A7B7E09F66
Irene Lobato-Vila and Juli Pujade-Villar (2017) Here we describe ve new species of Synergus Hartig with
partially smooth mesopleurae that were obtained from oak cynipid galls on different species of Quercus L.
in Mexico: Synergus cibriani sp. nov., Synergus gilletti sp. nov., Synergus longimalaris sp. nov., Synergus
longiscapus sp. nov. and Synergus striatifrons sp. nov. To date, this trait has been observed in at least 18
American species, 16 recorded in North America (United States of America) and only 2 in Central America
(Panama): Synergus elegans Nieves-Aldrey and Medianero and Synergus laticephalus Nieves-Aldrey and
Medianero. The type material and additional material of Synergus punctatus Gillette from the United States
have been examined and compared with S. gilletti sp. nov. The type material and additional material of S.
elegans from Panama and Synergus citriformis (Ashmead) from the United States have also been examined
and compared. This study records Synergus elegans in Mexico for the rst time and classies it as a syn. nov. of
S. citriformis. Descriptions and diagnoses for the new species and a key to their identication and differentiation
from their closest related species are given. Their distribution and host associations are also commented on.
Key words: Hymenoptera, Cynipidae, Synergus, New species, Mexico.
*Correspondence:. E-mail: ilobato@ub.edu; jpujade@ub.edu
BACKGROUND
Most of the approximately 1400 described
species of Cynipidae are true gall wasps (Ronquist
1999). However, only about 180 species, classied
into 10 genera, develop as inquilines inside galls
of other cynipids (Pujade-Villar et al. 2003; Nieves-
Aldrey and Medianero 2010; Pénzes et al. 2012;
Bozsó et al. 2014 2015; Schwéger et al. 2015a
b; Ronquist et al. 2015). Unlike true gall wasps,
which can induce the formation of complex galls
in different plants, inquilines are not capable
of inducing galls by themselves but instead
develop as guests inside galls of other cynipids
(except for one species described by Abe et al.
(2011) from Japan, Synergus itoensis Abe, Ide
and Wachi, which can induce galls on Quercus
(Cyclobalanopsis) glauca Thunberg, and also for
Synergus hupingshanensis Liu, Yang and Zhu,
which was inferred to be a gall maker because no
gall maker was reared from the galls it emerged
from in Castanopsis carlesii (Hemsl.) Hayata,
according to Liu et al. (2012)). Inquilinism is a form
of kleptoparasitism, usually considered to represent
a unilateral advantageous relationship that benets
only the inquilines (Askew 1984; Ronquist 1994
1999).
Until recently, all inquilines were included in
the Synergini tribe. However, Ronquist et al. (2015)
internally restructured the Cynipidae family, and
three genera of inquilines (Ceroptres, Periclistus
and Synophromorpha) were separated from the
Zoological Studies 56: 36 (2017)
doi:10.6620/ZS.2017.56-36
1

© 2017 Academia Sinica, Taiwan
tribe Synergini to be included in new tribes: the
tribe Ceroptresini, composed only by the genus
Ceroptres Hartig, and the tribe Diastrophini,
which includes the genera Periclistus Förster and
Synophromorpha Ashmead. Now, the Synergini
tribe only includes genera whose rst metasomal
segment is ring-shaped and usually carinated
(Agastoroxenia Nieves-Aldrey and Medianero;
Lithosaphonecrus Tang, Melika and Bozsó;
Rhoophilus Mayr; Saphonecrus Dalla Torre and
Kieffer; Synophrus Hartig; Synergus Hartig; Ufo
Melika and Pujade-Villar). All the species included
in this tribe are related to galls that develop on
Fagaceae (except for Rhoophilus loewi Mayr),
mainly on Quercus L.
The genus Synergus includes 117 species
(Pénzes et al. 2012; Schwéger et al. 2015a;
Pujade-Villar and Lobato-Vila 2016a b; Pujade-
Villar et al. 2016): 47 recorded in the Palearctic
region (30 from the Western Palearctic and
17 from the Eastern Palearctic), 56 in the
Nearctic region and 14 in the Neotropical region.
Neotropical species have been addressed by
Ritchie and Shorthouse (1987), Díaz and Gallardo
(1998), Nieves-Aldrey (2005) and Nieves-Aldrey
and Medianero (2011); for its part, most species
recorded in the Nearctic region are addressed
by Gillette (1896), but also by Fullaway (1911),
McCracken and Egbert (1922) and Burnett (1976).
Among all known species in the genus
Synergus, only four have been recorded in Mexico
(Weld 1930; Pujade-Villar et al. 2008; Pujade-
Villar and Lobato-Vila 2016a b; Pujade-Villar et al.
2016): Synergus filicornis Cameron (= Synergus
furnessana Weld), Synergus mexicanus Gillette
(= Synergus dugesi Ashmead; = Synergus brevis
(Weld)), Synergus equihuai Pujade-Villar and
Lobato-Vila and Synergus estradae Pujade-Villar
and Lobato-Vila. Synergus licornis was obtained
from galls of Andricus furnessae (Weld) collected
at Terecuato (Michoacán); S. mexicanus, from
tuberous galls of the genus Andricus Hartig from
different Mexican locations (Ashmead 1899;
Pujade-Villar and Melika 2005; Pujade-Villar et al.
2008; Pujade-Villar et al. 2015); S. equihuai, from
inconspicuous oak galls inside Q. rugosa Née
acorns collected from Tepoztlán (Morelos) and S.
estradae, from tuberous galls collected from Q.
obtusata Bonpl. near Oaxtepec (Morelos) (Pujade-
Villar et al. 2016).
Morphological and phylogenetical analyses
of species from the Palearctic support that
Synergus is a monophyletic group (Ács et al.
2010; Schwéger et al. 2015a). Nevertheless, only
Western and Eastern Palearctic Synergus species
were involved in the analyses, and many species
from both Nearctic and Neotropical regions
show important morphological differences with
respect to the Palearctic ones. Thus, the global
monophyletism of Synergus should not yet be
assumed.
It is suggested that Synergus species
from America should be classified into three
morphological groups: 1) species with radial
cell open, like S. mexicanus (closed in most
of currently known species), 2) species with
mesopleurae totally striated (the most part of
currently known species) and 3) species with the
speculum smooth or with mesopleurae almost
totally smooth (mesopleurae always striated, with
striae extended into the speculum, in species from
other biogeographical regions). Discrimination of
species based on the sculpture of mesopleurae
was rst dened by Nieves-Aldrey and Medianero
(2011), and the third group of species mentioned
above is the subject of this study.
Until now, there were known only two species
of Synergus from Central America with partially
smooth mesopleurae: Synergus elegans Nieves-
Aldrey and Medianero and Synergus laticephalus
Nieves-Aldrey and Medianero, both recorded for
the first and only time in Panama (Nieves-Aldrey
and Medianero 2011), which have the speculum
smooth. In North America, at least 16 species
belong to this group, but only Synergus punctatus
Gillette is morphologically closely related to one
of the new Mexican species described here:
Synergus gilletti sp. nov. In this study, we describe
ve new species with partially smooth mesopleurae
obtained from oak cynipid galls from Mexico:
Synergus cibriani sp. nov., Synergus gilletti sp.
nov., Synergus longimalaris sp. nov., Synergus
longiscapus sp. nov. and Synergus striatifrons
sp. nov. All of them lack pronotal carinae (usually
present in Synergus from the Palearctic area)
and all except S. gilletti sp. nov. do not have
frontal carinae (always present in Synergus in
the Palearctic area according to Schwéger et
al. (2015b), short in S. deqingensis Pujade-
Villar, Wang and Chen from China, and absent in
some American species). In addition, S. elegans,
previously only recorded in Panama, becomes a
syn. nov. of Synergus citriformis (Ashmead) from
the United States and is for the rst time recorded
in Mexico, thus increasing its known distribution
range.
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© 2017 Academia Sinica, Taiwan
MATERIALS AND METHODS
Adults of the species addressed in the
present study were obtained from galls developed
on different species of Quercus from both Lobatae
and Quercus sections: Q. affinis Scheidw., Q.
candicans Née, Q. castanea Née, Q. conzattii Trel.,
Q. crassipes Humb. et Bonpl., Q. eduardii Trel. and
Q. emoryi Torr. and Q. mexicana Humb. et Bonpl.
(Lobatae Section); Q. glabrescens Benth., Q.
greggii Trel., Q. laeta Liebm, Q. microphylla Née,
Q. obtusata Humb. et Bonpl. and Q. rugosa Née
(Quercus Section). Galls were preserved inside
breeding-boxes under environmental conditions
until the emergence of all adults.
The type material of S. citriformis, S.
punctatus (both deposited in the NMNH,
Washington D.C., USA) and S. elegans (deposited
in the MNCNM, Madrid, Spain) was requested and
studied. The type material of S. citriformis was
requested because, after checking the pictures of
this species deposited in the online database of
Hymenopteran Holotypes (Smithsonian Institution),
we noticed it has partially smooth mesopleurae
and a very similar morphology to that of S. elegans
(and the original description of S. citriformis was
insufcient to decide if they were different species
or not). The type material of S. elegans was
requested to compare it with that of S. citriformis.
Finally, the type material of S. punctatus was
requested because, according to its original
description and pictures from the online database
Hymenopteran Holotypes, it is morphologically
closely related to some of the specimens obtained
from the Mexican galls studied here and it was
difficult to decide if they were different species
without comparing them. Synergus laticephalus
was not requested because it has some
morphological traits that clearly separate it from
the rest of species addressed in this study; also,
because the quality of its original description and
pictures (Nieves-Aldrey and Medianero 2011)
are good enough to be compared with the new
material under study.
For morphological descriptions, it has been
used the current terminology of morphological
structures as given in the following studies:
Liljeblad and Ronquist (1998) and Melika
(2006) for adult morphological structures,
Ronquist and Nordlander (1989) for forewing
venation terminology and Harris (1979) for
patterns of cuticular sculpture. The following
are measurements and abbreviations used in
this paper: F1-F12, the first and the following
flagellomeres; POL (post-ocellar distance), the
distance between the inner margins of the posterior
ocelli; OOL (ocellar-ocular distance), the distance
from the outer margin of the lateral ocellus to the
inner margin of the compound eye; LOL (lateral-
ocular distance), the distance between lateral and
frontal ocellus; transfacial line, distance between
inner margins of compound eyes measured across
toruli; width of radial cell, measured as the distance
between the upper margin of the forewing and the
Rs vein.
Electron microscope images of adults were
taken using a Scanning Electron Microscope (FEI
Quanta 200 ESEM) at Universitat de Barcelona
(Barcelona, Spain); specimens of the new species
were previously dissected and covered with gold,
except for those of S. cibriani sp. nov. Finally,
pictures of the head in frontal view of S. gilletti sp.
nov. were taken by Roca-Cusachs using a Leica
DFC450 digital microscope camera coupled to a
Leica MZ160A binocular microscope.
Type material of the new species is deposited
in the second author’s collection (JP-V col. in UB,
Universitat de Barcelona, Barcelona, Spain), in
the National Museum of Natural History of the
Smithsonian Institution (NMNH, Washington D.C.,
USA) and in the American Museum of Natural
History (AMNH, New York, USA).
RESULTS
The species are given in alphabetical order.
Synergus cibriani Pujade-Villar and Lobato-Vila
sp. nov.
(Fig. 1)
urn:lsid:zoobank.org:act:3DD84BE1-758C-437A-A624-
CCECFCC0E491
Type material: Holotype ♀ deposited in
JP-V col. (UB) with the following labels: ‘MEX:
P201, Los Desmontes (Parástaco, MICH),
19°26'44"N, 102°22'40"W, 1416 m’ (white label),
‘Ex. Disholcaspis sp., Quercus sp., (23.ii.2013)
12.iii.2013, E. Estrada and A. Equihua leg.’ (white
label), ‘Holotype ♀ Synergus cibriani Pujade-
Villar and Lobato-Vila, Lobato-Vila 2017 desig.’
(red label). Paratypes (6
♂
and 6♀): same
data as the Holotype: 4
♂
and 3♀; MEX: P232,
Tlaxco-Ciénaga Larga Highway (km 38) (PUE),
19°51'12"N, 98°06'32"W, 1832 m, Ex. Disholcaspis
sp., Quercus sp. (25.vi.2016) vii.2016, JP-V leg.:
2
♂
and 3♀.
page 3 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
Fig. 1. Synergus cibriani Pujade-Villar and Lobato-Vila sp. nov.: (a) Female head in frontal view, (b) mesosoma in dorsal view, (c)
female head in dorsal view, (d) male head in dorsal view, (e) mesosoma in lateral view, (f) female antenna, (g) rst segments of male
antenna, (h) propodeum, (i) tarsal claw, (j) female metasoma in lateral view, (k) detail of the metasomal micropunctures (female), (l)
detail of the metasomal micropunctures (male).
(a)
(b)
(d)
(e)
(h)
(k) (l)
(i) (j)
(c) (f)
(g)
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© 2017 Academia Sinica, Taiwan
Additional material: MEX: P243, Ejido
Santa María Nativitas (Texcoco, EDO. MÉX),
19°27'41.3"N, 98°47'22.2"W, 2703 m, Ex.
undetermined gall, Q. glabrescens, (27.i.2015)
19.ii.2015, A. Equihua and E. Estrada: 3
♂
and
4♀.
Etymology: Named in honor of Dr. David
Cibrián Tovar (Universidad Autónoma Chapingo,
Mexico), who collected galls from Mexico for our
studies.
Diagnosis: This species is morphologically
related to S. citriformis (Ashmead) (= S. elegans
Nieves-Aldrey and Medianero syn. nov.) and
S. laticephalus Nieves-Aldrey and Medianero.
The new species differs from S. laticephalus
by the width of the head in dorsal view, which
is much shorter (2.7 times wider than long
in S. laticephalus); also, by the presence of
metasomal micropunctures (no micropunctures
in S. laticephalus) and the absence of rugae in
the scutellum (with some transverse rugae in S.
laticephalus). Synergus cibriani sp. nov. mainly
differs from S. citriformis by having the malar
space about 0.8 times as long as the height of the
compound eye (0.6 in S. citriformis), frons and
vertex with some scattered little punctures (no
punctures in S. citriformis), metasomal syntergite
with a posterodorsal patch of micropunctures (no
micropunctures in S. citriformis), and F1 of males
slightly expanded basally and straight apically
(slightly expanded apically in S. citriformis).
Female
Length: Body length 3.0-3.4 mm (n = 4).
Color: Head yellowish to brown; frons and
vertex medially black, chestnut laterally; occiput
black; antennae yellowish to brown; mesosoma
generally black, tegulae pale yellow; metasoma
black dorsally, light chestnut basally; legs yellow,
the base of metacoxae with a more or less
extended brownish to blackish coloration; wings
hyaline, veins light brown to yellow.
Head: in frontal view (Fig. 1a) more or less
quadrangular, about 1.2 times wider than high,
genae slightly expanded behind compound eye.
Face faintly pubescent, lower face and genae
with thin, dense and regular striae irradiating from
clypeus and reaching basal margin of compound
eye and basal margin of antennal toruli; medial
carina inconspicuous. Clypeus indistinct, ventral
margin not projected over mandibles. Malar
space about 0.8 times as long as the height of
the compound eye (4.2:5). Anterior tentorial pits
visible; pleurostomal and epistomal sulcus absent.
Transfacial line longer than the height of the
compound eye (1.3). Toruli situated mid-height of
the compound eye; distance between torulus and
compound eye as long as the diameter of toruli;
distance between toruli shorter than the diameter
of toruli. Frons nely coriaceous to reticulated with
some scattered little punctures; frontal carinae
absent. Head in dorsal view (Fig. 1c) is about two
times wider than long. Vertex finely coriaceous
with some scattered and little piliferous punctures
between the ocelli. POL:OOL:LOL = 2.4:1.6:1.2;
OOL about 1.5 times longer than the diameter
of lateral ocellus. Occiput coriaceous, with some
scattered piliferous punctures.
Antennae (Fig. 1f): 14-segmented (2.7: 1.3:
3.4: 2.6: 2.4: 2.2: 1.9: 1.6: 1.3: 1.2: 1.1: 0.9: 0.9:
1.7); filiform not broadened apically; pubescence
dense and short; placodeal sensilla little visible on
agellar segments F3-F12. Pedicel about 2.2 times
longer than wide; F1 1.3 times longer than F2, F2
slightly longer than F3. Last agellar segment 2.8
times longer than wide and 1.9 times longer than
F11.
Mesosoma: Slightly longer than high in
lateral view (including nucha), with short and not
dense pubescence (Fig. 1e). Ratio of length of
pronotum medially/laterally: 0.26. Pronotal plate
absent. Lateral pronotum sculpture coriaceous to
imbricated; lateral margins of pronotum rounded,
without lateral carina. Mesoscutum (Fig. 1b) about
as long as wide, strongly coriaceous to imbricated;
anterior grooves weak, occupying about 1/4
of mesoescutum length. Notauli complete but
faint in the anterior 1/3, wider and convergent
posteriorly. Median groove weakly impressed,
almost inconspicuous. Parapsidial grooves visible,
reaching the anterior margin of tegulae. Scutellum
(Fig. 1b) rounded, as long as wide, coriaceous
anteriorly, imbricate posteriorly; circumscutellar
carina visible but not projected; scutellar foveae
inconspicuous. Mesopleurae (Fig. 1e) with a few
transversal striae located medially and anteriorly,
smooth dorsally and basally, speculum also
smooth; interspaces alutaceous, little pubescent
basally. Metapleural sulcus well dened, reaching
about 3/4 parts of mesopleurae height. Propodeum
(Fig. 1h) pubescent and with a weak sculpture;
propodeal carinae slightly curved and convergent
basally, central area with a weak sculpture and
pubescent. Nucha sulcated dorsally and laterally.
Legs (Fig. 1i): Tarsal claws with a small basal
tooth.
Wings: Forewings pubescent with short
page 5 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
marginal setae, about as long as mesosoma plus
metasoma. Radial cell closed, 2.4 times longer
than wide; areolet absent. Rs + M inconspicuous,
not reaching basal vein. Basal cell faintly
pubescent.
Metasoma: About as long as head plus
mesosoma, longer than high in lateral view. First
metasomal segment sulcated dorsally and laterally.
Syntergite smooth, anterolateral pubescence
composed of a group of 4-5 setae, with a few weak
micropunctures forming a posterodorsal patch;
not incised dorso-distally (Figs. 1j, k). Following
segments and hypopygium clearly punctured.
Hypopygial spine about as long as wide and with a
few lateral setae; without apical setae.
Male
Similar to female, except for the morpho-
logical traits that follow: body length 2.9-3.1 mm
(n = 4). POL:OOL:LOL = 2.5: 1.6: 1.4 (Fig. 1d).
Antennae 15-segmented (2.4: 1.3: 3.7: 2.7: 2.5:
2.2: 2.0: 1.8: 1.6: 1.4: 1.3: 1.2: 1.1: 1.0: 1.4); F1
slightly curved, medial excavation weak, slightly
expanded basally, almost straight apically (Fig. 1g).
Syntergite punctuation slightly extended laterally
but not forming a complete band (Fig. 1l).
Distribution: State of Michoacán (Mexico).
Biology: The studied material was obtained
from soft galls developed on some undetermined
species of Quercus and from Q. glabrescens
Benth. (Quercus section), related to the genera
Disholcaspis and to one undetermined gall,
respectively.
Synergus citriformis (Ashmead, 1885)
Ceroptres citriformis Ashmead, 1885. Trans. Amer. Ent. Soc.,
12: 300
Synergus citriformis (Ashmead) Weld, 1922. Proc. U. S. Nat.
Mus., 61 (18): 26
Synergus elegans Nieves-Aldrey and Medianero, 2011.
Zootaxa, 2774: 4 syn. nov.
Type material of Ceroptres citriformis
Ashmead, 1885 (examined): Lectotype♀ deposited
in the NMNH (Washington D.C., USA) with the
following labels: ‘Jacksnville, Fla’ (white label) /
‘Type No 14538 U.S.N.M.’ (red label) / ‘Collection
Ashmead’ (white label) / ‘Synergus citriformis
♀ Ashmead, Type’ (white label, handwritten) /
‘USNMENT 00779983’ (white label, QR code)
/ ‘Lectotype ♀ Ceroptres citriformis Ashmead,
1885 IL-V desig-2017 (red label) / ‘Synergus
citriformis (Ashmead) Weld, 1922 (= S. elegans)
IL-V det. 2017’ (white label). Paralecototypes (1
♂
and 3♀) deposited in the NMNH (Washington
D.C., USA) with the following labels: ‘Jacksnville,
Fla’ (white label) / ‘Paratype No 14538 U.S.N.M.’
(red label) / ‘Collection Ashmead’ (white label) /
‘Paralectotype ♀ Ceroptres citriformis Ashmead,
1885 IL-V desig-2017’ (red label) / ‘Synergus
citriformis (Ashmead) Weld, 1922 (= S. elegans)
IL-V det. 2017’ (white label) (2♀); ‘Jacksnville, Fla’
(white label) / ‘Collection Ashmead’ (white label)
/ ‘Synergus (Ceroptres) citriformis Ashm.’ (white
label, handwritten) / ‘Paralectotype ♀ Ceroptres
citriformis Ashmead, 1885 IL-V desig-2017’
(red label) / ‘Synergus citriformis (Ashmead)
Weld, 1922 (= S. elegans) IL-V det. 2017’ (white
label) (1♀); ‘Jacksnville, Fla’ (white label) /
‘Collection Ashmead’ (white label) / ‘Paralectotype
♂
Ceroptres citriformis Ashmead, 1885 IL-V
desig-2017’ (red label) / ‘Synergus citriformis
(Ashmead) Weld, 1922 (= S. elegans) IL-V det.
2017’ (white label) (1
♂
).
Type material of Synergus elegans Nieves-
Aldrey and Medianero, 2011 (examined):
Paratypes (2♀) deposited in the MNCNM (Madrid,
Spain) with the following labels: ‘PANAMÁ, Prov.
Chiriquí Boquete, El Salto, 1431 m, 8°47'32.8"N,
82°27'37.9"W, Quercus salicifolia S-ex-A1, 7-V-
2008, E. Medianero leg.’ (white label) / ‘CYNIPIDAE
Synergus elegans sp. nov., Nieves-Aldrey and
Medianero det.’ (white label) / ‘Paratipo’ (red
label) / ‘MNCN Cat. Tipos N° 2095’ (red label)
/ ‘MNCN_Ent N° Cat. 67238’ (white label) /
‘Synergus citriformis (Ashmead) Weld, 1922
det. Lobato-Vila 2017’ (1♀); ‘PANAMÁ, Prov.
Chiriquí Boquete, El Salto, 1431 m, 8°47'32.8"N,
82°27'37.9"W, Quercus salicifolia S-ex-A1, 7-V-
2008, E. Medianero leg.’ (white label) / ‘CYNIPIDAE
Synergus elegans sp. nov., Nieves-Aldrey and
Medianero det.’ (white label) / ‘Paratipo’ (red label)
/ ‘MNCN Cat. Tipos N° 2095’ (red label) / ‘MNCN_
Ent N° Cat. 67240’ (white label) / ‘Synergus
citriformis (Ashmead) Weld, 1922 det. Lobato-Vila
2017’ (1♀).
Additional material: Non-type material
of Synergus elegans (1♀) deposited in the
MNCNM (Madrid, Spain) with the following
labels (examined): ‘PANAMÁ, Prov. Chiriquí
Renacimiento, 1270 m, 8°49'58.7"N,
82°44'44.5"W, Quercus lancifolia S-ex-A55,
22-I-2009, E. Medianero leg.’ (white label)
/ ‘CYNIPIDAE Synergus elegans sp. nov.,
Nieves-Aldrey and Medianero det.’ (white label)
/ ‘MNCN_Ent N° Cat. 67515’ (white label) /
‘Synergus citriformis (Ashmead) Weld, 1922 det.
page 6 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
Lobato-Vila 2017’. New material from Mexico
(147
♂
and 236♀) obtained from the following
localities: AGUASCALIENTES: MEX: P155, A.
N. Protegida Los Alamitos (San José de Gracia,
AGS), 22°10'26.4"N, 102°35'11.9"W, 2598 m, Ex.
Amphibolips hidalgoensis, Q. microphylla, (01.
iii.2012) 20.iii.2012, E. Estrada and A. Equihua
leg.: 1♀. FEDERAL DISTRICT: MEX: P228, Santa
Fe (D.F.), 19°21'14"N, 99°16'24.22"W, 2569,
Ex. Andricus protuberans, Q. laeta, (21.vi.2013)
ix.2013, JP-V leg.: 4
♂
and 3♀; MEX: P224, Santa
Fe (D.F.), 19°21'14"N, 99°16'24.22"W, 2569, Ex.
Amphibolips sp., Q. crassipes, (21.vi.2013) 1-10.
xii.2013, JP-V leg.: 1
♂
and 7♀. GUANAJUATO:
MEX: P012, Sierra de Santa Rosa (Santa Rosa
de Lima, GTO), 21°07'44.1"N, 101°13'02.0"W,
2559 m, Ex. Disholcaspis sp., Q. castanea, (31.
v.2012) 12.vii.2012, E. Estrada and A. Equihua
leg.: 2
♂
. HIDALGO: MEX: 296, Barrio Tlacpac
(Acaxochitlán, HGO), 20°09'16.4"N, 98°12'25.5"W,
2321 m, Ex. Amphibolips hidalgoensis, Q. rugosa,
(08.vi.2010) 13.vii.2010, E. Estrada and A.
Equihua leg.: 7
♂
and 4♀; MEX: 290, El Arenal
(Acaxochitlán, HGO), 20°13'29.9"N, 98°54'25.0"W,
2210 m, Ex. Amphibolips hidalgoensis, Q.
rugosa, (10.vi.2010) 23.vi.2010, E. Estrada
and A. Equihua leg.: 3
♂
; MEX: 293, Huasca
(HGO), 20°12'05.3"N, 98°35'16.6"W, 2181 m,
Ex. Amphibolips hidalgoensis, Q. candicans, (08.
vi.2010) 15-30.vi.2010, JP-V leg.: 2
♂
; MEX: 283,
Los Romeros (Santiago de Tulantepec de Lugo
Guerrero, HGO), 20°01'45.9"N, 98°24'50.1"W,
2411 m, Ex. Amphibolips hidalgoensis, Q.
mexicana, (03.vi.2010) 17.vii.2010, E. Estrada
and A. Equihua leg.: 4
♂
and 16♀; MEX: P050,
Santiago de Tulantepec de Lugo Guerrero
(HGO), 20°05'07.8"N, 98°17'50.0"W, 2510 m, Ex.
Andricus nr protuberans, Q. greggii, (03.vi.2010)
23.vi.2010, E. Estrada and A. Equihua leg.: 3
♂
and 11♀; MEX: 2784 (UACh Col.), Acaxochitlán
(HGO), 20°10'17.59"N, 98°10'58.59"W, 2154 m,
Ex. Amphibolips damphi, Q. affinis, (19.iii.2014)
v.2014, D. Cibrián leg.: 1
♂
and 6♀. MICHOACÁN:
MEX: P203, Near Amealco (MICH), 20°10'30"N,
100°13'52"W, 2620m, Ex. Undetermined gall, Q.
mexicana, (14.ii.2013) 05.iii-01.iv.2013, E. Estrada
and A. Equihua leg.: 3
♂
and 7♀; MEX: P205,
Los Desmontes (Parástaco, MICH), 19°26'44"N,
102°22'40"W, 1.416 m, Ex. Disholcaspis sp.,
Quercus sp., (23.ii.2013) 03.iv-01.v.2013, E.
Estrada and A. Equihua leg.: 1
♂
and 1♀.
MORELOS: MEX: 278, Felipe Neri (Tlalnepantla,
MOR), 19°02'11.2"N, 98°56'38.8"W, 2512 m, Ex.
Amphibolips sp., Q. rugosa, (29.v.2012) viii.2012,
E. Estrada and A. Equihua leg.: 13♀; MEX:
P008, Tepoztlán (Tepoztlán, MOR), 18°59'04.2"N,
99°05'35.8"W, 2099 m, Ex. Disholcaspis sp., Q.
rugosa, (29.v.2012) 12.vi.2012, E. Estrada and A.
Equihua leg.: 1
♂
and 2♀; MEX: P019, Tepoztlán
(Tepoztlán, MOR), 18°59'04.2"N, 99°05'35.8"W,
2099 m, Ex. Disholcaspis nr taumalipensis, Q.
mexicana, (20.iv.2012) 12.vi.2012, E. Estrada
and A. Equihua leg.: 3
♂
and 2♀. PUEBLA:
MEX: 165, Ometépetl (Tetela de Ocampo,
PUE), 19°50'16.2"N, 97°43'22.9"W, 1687 m, Ex.
Amphibolips nr nevadensis, Quercus sp., (29.
iv.2013) v.2013 (Alondra leg.): 1
♂
and 3♀. STATE
OF MEXICO: MEX: P120, Ixtapaluca (EDO. MÉX),
19°19'07"N, 98°52'56"W, Ex. Amphibolips sp.,
Quercus sp., (30.v.2012) vi.2012, E. Estrada and
A. Equihua leg.: 4♀; MEX: 274, P. N. Zoquiapan
y Anexas (Ixtapaluca, EDO. MÉX), 19°20'06.8"N,
98°41'30.6"W, 3046 m, Ex. Amphibolips
hidalgoensis, Q. crassipes, (30.v.2012) viii.2012,
E. Estrada and A. Equihua leg.: 41
♂
and 39♀;
MEX: 275, P. N. Zoquiapan y Anexas (Ixtapaluca,
EDO. MÉX), 19°20'06.8"N, 98°41'30.6"W, 3046 m,
Ex. Amphibolips hidalgoensis, Q. crassipes, (30.
v.2012) viii.2012, E. Estrada and A. Equihua
leg.: 3
♂
and 5♀; MEX: 276, P. N. Zoquiapan y
Anexas (Ixtapaluca, EDO. MÉX), 19°20'06.8"N,
98°41'30.6"W, 3046 m, Ex. Amphibolips
hidalgoensis, Q. crassipes, (30.v.2012) viii.2012,
E. Estrada and A. Equihua leg.: 1
♂
; MEX: 280,
P. N. Zoquiapan y Anexas (Ixtapaluca, EDO.
MÉX), 19°20'06.8"N, 98°41'30.6"W, 3046 m, Ex.
Amphibolips hidalgoensis, Q. crassipes, (30.
v.2012) viii.2012, E. Estrada and A. Equihua leg.:
47
♂
and 64♀; MEX: 281, P. N. Zoquiapan y
Anexas (Ixtapaluca, EDO. MÉX), 19°20'06.8"N,
98°41'30.6"W, 3046 m, Ex. Amphibolips
hidalgoensis, Q. crassipes, (30.v.2012) viii.2012, E.
Estrada and A. Equihua leg.: 5
♂
and 12♀; MEX:
196, Plateros-Arenales (San Felipe del Progreso,
EDO. MÉX), 19°42'46.8"N, 99°57'05.6"W, 2576 m,
Ex. Andricus group tectucnarum, Q. candicans,
(05.ii.2014) 16.ii.2014, Delia leg.: 2♀; MEX: 235,
Plateros-Arenales (San Felipe del Progreso, EDO.
MÉX), 19°42'46.8"N, 99°57'05.6"W, 2576 m, Ex.
Dros nr picatum, Q. laeta, (18.i.2014) 05.ii.2014,
Delia leg.: 1♀; MEX: 255, San Juan Coajomulco
(EDO. MÉX), 19°45'17.9"N, 99°58'10.2", 2655 m,
W, Ex. Andricus group georgei, Q. obtusata, (18.
i.2014) 16.ii-2.iv.2014, Delia leg.: 1
♂
and 7♀.
TLAXCALA: MEX: 291, San Marcos (Papalotla de
Xicohténcatl, TLAX), 19°19'55.4"N, 98°18'13.4"W,
2342 m, Ex. Amphibolips hidalgoensis, Q .
crassipes, (03.vi.2010) 04.vii.2010, E. Estrada
page 7 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
and A. Equihua leg.: 3
♂
and 7♀; MEX: P079,
San Marcos (Papalotla de Xicohténcatl, TLAX),
19°19'55.4"N, 98°18'13.4"W, 2342 m, Ex.
Amphibolips sp., Q. crassipes, (03.vi.2010)
05.ix.2010, E. Estrada and A. Equihua leg.:
6♀; MEX: P083, San Marcos (Papalotla de
Xicohténcatl, TLAX), 19°19'55.4"N, 98°18'13.4"W,
2342 m, Ex. Amphibolips sp., Q. crassipes, (03.
vi.2010) 12.vii.2010, E. Estrada and A. Equihua
leg.: 2
♂
and 5♀; MEX: P091, San Marcos
(Papalotla de Xicohtncatl, TLAX), 19°19'55.4"N,
98°18'13.4"W, 2342 m, Ex. Amphibolips sp., Q.
crassipes, (03.vi.2010) 12.vii.2010, E. Estrada and
A. Equihua leg.: 1♀; MEX: P167, Calpulalpan-
Tlaxcala Federal Highway (Calpulalpan, TLAX),
19°35'24.6"N, 98°33'24.1"W, 2572 m, Ex. Andricus
group georgei, Q. microphylla, (31.viii.2012)
10.ix.2012, A. Equihua and E. Estrada: 1♀;
MEX: P077, San Diego Recoba (Hueyotlipan,
TLAX), 19°29'05.1"N, 98°22'24.7"W, 2632 m, Ex.
Undetermined gall, Q. crassipes, (03.vi.2010)
15.vi.2010, E. Estrada and A. Equihua leg.: 1
♂
.
VERACRUZ: MEX: P180, Perote-Puebla Federal
Highway (VER), 19°32'30"N, 97°16'23"W, 2410
m, Ex. Antron sp., Quercus sp. (31.viii.2012)
10.ix.2012, E. Estrada and A. Equihua leg.: 1♀.
ZACATECAS: MEX: 285, Las Guacamayas (Monte
Escobedo, ZAC), 21°42'21"N 103°02'39"W,
1631 m, Ex. Amphibolips zacatequensis, Q.
eduardii, (25.v.2012) Before x.2012, JP-V leg.: 1
♂
;
MEX: 277, Monte Escobedo (ZAC), 22°25'06.32"N,
103°28'38.82"W, 2031 m, Ex. Amphibolips
zacatequensis, Q. eduardii, (31.v.2012) vi.2012,
C. Carrillo leg.: 2♀; MEX: P065, Monte Escobedo
(ZAC), 22°25'06.32"N, 103°28'38.82"W, 2031 m,
Ex. Amphibolips sp., Q. eduardii, (31.v.2010)
vi.2010, C. Carrillo leg.: 1♀; MEX: 020, Sierra
de Nochistlán (Jérez, ZAC), 21°20'19.3"N,
102°58'50.2"W, Ex. Undetermined gall, Q. eduardii,
(10.ii.2011) 16.ii.2011, A. Equihua and E. Estrada
leg.: 2♀; MEX: P123, La Cuchilla (Concepción del
Oro, ZAC), 24°37'00.3"N, 101°24'48.8"W, 2048 m,
Ex. Amphibolips sp., Q. conzattii, (25.05.2012)
13.vii.2012, E. Estrada and A. Equihua leg.: 1
♂
.
Comments: According to Ashmead (1885:
300), this species was described based on
numerous specimens. We studied 4 females and
1 male deposited in the NMNH from Ashmead’s
collection, of which 1
♂
and 1♀ have identical
labels to Lectotype’s but the ‘type’ label is
missing. So, we assume these two specimens
also belong to the typical series. Based on the
original description, there are probably more
paralectotypes in Ashmead’s collection without a
‘type’ label.
Synergus citriformis and S. elegans have
the head nely reticulated, without frontal carinae
and 2.0 wider than long from the dorsal view;
antennae 14-segmented and F1 1.3 times longer
than F2; mesoscutum and scutellum finely
coriaceous to imbricated, notauli inconspicuous
in the anterior 1/3, mesopleurae partially smooth
(the horizontal sculpture not extended into the
speculum), scutellum laterally and postero-dorsally
margined (circumscutellar carina clearly visible)
and scutellar foveae inconspicuous or absent;
metasoma not incised dorso-distally and no
micropunctured (or at least with some scarce and
inconspicuous posterodorsal micropunctures in
some specimens). Since the rest of morphological
traits are coincident and that S. citriformis was
described from numerous specimens bred from a
gall related to Amphibolips sp. (Ashmead 1885)
like most of S. elegans, we conclude these are the
same species. So, we propose S. elegans as syn.
nov. of S. citriformis.
The type material of S. citriformis from the
USA and some specimens from Mexico show
a lighter body tonality than thus of S. elegans.
Moreover, some of the Mexican specimens show
morphological variations related to the ratio of F1/
F2 (F1 usually 1.3 times longer than F2 in this
species; ratio longer in some specimens) and the
length of POL and OOL (POL usually 1.5 times
longer than OOL in this species; POL 1.1 times
longer in some specimens). However, since these
subtle variations are already contemplated in the
original description of S. elegans (Nieves-Aldrey
and Medianero 2011), we don’t doubt they are the
same species.
According to Nieves-Aldrey and Medianero
(2011), S. citriformis (= S. elegans) and S.
laticephalus form a distinctive group mainly
characterized by having partially smooth
mesopleurae (with striae not extended into
the speculum), absence of frontal carinae,
scutellum laterally and postero-dorsally margined,
mesoscutum finely coriaceous to alutaceous or
finely imbricate, and metasoma not dorso-distally
incised and not micropunctured. All Mexican
species described here which are related to
these species have a posterodorsal patch of
micropunctures in the metasomal syntergite (see
also the diagnosis of the new Mexican species).
Distribution: This is the first record of this
species in Mexico. This species was previously
only known from Panama (Nieves-Aldrey and
Medianero 2011) and Florida, USA (Ashmead
page 8 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
1885; Pénzes et al. 2012). The new material
presented here was collected from different states
of Mexico (Aguascalientes, Federal District,
Guanajuato, Hidalgo, Michoacán, Morelos,
Puebla, State of Mexico, Tlaxcala, Veracruz and
Zacatecas).
Biology: The original material described
by Ashmead (1885) was bred from a gall
related to Amphibolips citriformis Ashmead. The
Panamanian specimens studied by Nieves-Aldrey
and Medianero (2011) were obtained from a wide
variety of cynipid galls from different genera,
being very common in galls of the Panamanian
species of Amphibolips and frequent in galls of
species of Disholcaspis, Cynips and other cynipid
genera on different species of Quercus both from
Quercus and Lobatae sections (Nieves-Aldrey
and Medianero 2011). The studied material from
Mexico presented here was also obtained from a
wide variety of cynipid galls, including Amphibolips,
Disholcaspis, Andricus, Dros and Antron’s galls.
These galls were also collected from different
species of Quercus belonging to both Quercus and
Lobatae sections.
Synergus gilletti Pujade-Villar and Lobato-Vila
sp. nov.
(Figs. 2, 3, 8b)
urn:lsid:zoobank.org:act:407A7BC3-CBF9-485B-86B1-
0472D7C442CD
Type material: Holotype ♀ deposited in
JP-V col. (UB) with the following labels: ‘MEX:
101, Charco de la Rana (ZAC), 24°14'23.6"N,
103°25'08.4"W, 1960 m’ (white label); ‘Ex.
Andricus sp., Q. laeta, (19.xii.11) 28.i.12, O.
Cabral, R. Treto, L.G. Landa and C. Carrillo leg.’
(white label); ‘Holotype♀ Synergus gilletti Pujade-
Villar and Lobato-Vila, Lobato-Vila 2017 desig.’ (red
label). Paratypes (35
♂
and 8♀): same data as the
Holotype: 1
♂
; MEX: 068, Sierra el Alicante (Monte
Escobedo, ZAC), 22°30'41.6"N, 103°21'57.2"W,
2360 m, Ex. Atrusca sp., Q. laeta, (19.x.2011)
xi.2011, O. Cabral, R. Treto, L.G. Landa and C.
Carrillo leg.: 34
♂
and 8♀ (5
♂
and 1♀ deposited
in the NMNH; 5
♂
and 1♀ deposited in the
AMNH).
Additional material: 3
♂
and 3♀ with
the same data as MEX: 068 dissected and
covered with gold; MEX: 112, La Cumbre (ZAC),
22°19'00.89"N, 103°38'53.43"W, 2315 m, Ex.
Atrusca sp., Q. rugosa, (05.xii.2011) xii.2011, O.
Cabral, R. Treto, L.G. Landa and C. Carrillo leg:
12
♂
.
Etymology: Named in honor to Clarence
Preston Gillette (1859, Ionia County,
Michigan-1941, Fort Collins, Colorado), the
American entomologist who described the closest
related species Synergus punctatus.
Diagnosis: Species morphologically closely
related to Synergus punctatus Gillette of which
is distinguished by the following combination of
morphological traits: in males, head black except
for a reddish-brown to chestnut colored halo
around the mouth (head yellow except for a black
spot in the vertex in S. punctatus); in females,
colored halo around the mouth less conspicuous
and reddish-brown to chestnut (yellow and clearly
defined in S. punctatus) (Figs. 8a, b); F1/F2
ratio in males about 1.15 (about 1.4 times in S.
punctatus); notauli complete and visible along their
entire length (inconspicuous in the anterior third of
the scutum in S. punctatus); syntergite punctuation
not forming a complete band, less widespread and
shorter (forming a complete band, occupying the
distal half of the syntergite and reaching the ventral
part of metasoma in S. punctatus).
Female
Length: Body length 1.5-2.2 mm (n = 14).
Color: Head black, except for face surfaces
surrounding the oral foveae (sometimes in a
small area), which are reddish-brown to chestnut
(Fig. 8b); frons and vertex black, occiput black;
antennae testaceous; mesosoma black, tegulae
yellowish; metasoma black, reddish basally; legs
yellow, including coxae; forewings hyaline, light
veins.
Head: in frontal view (Fig. 2a) quadrangular
with genae convergent, 1.2 times wider than high,
genae not expanded behind compound eyes.
Face faintly pubescent, lower face and gena
with striae irradiating from clypeus and reaching
basal margin of compound eye and basal margin
of antennal toruli; medial carina also present.
Clypeus indistinct, ventral margin slightly projected
over mandibles. Malar space around 0.5 times as
long as the height of the compound eye (10:18).
Anterior tentorial pits visible; pleurostomal and
epistomal sulcus absent. Transfacial line about
as long as the height of the compound eye.
Toruli situated mid-height of the compound eye;
distance between torulus and compound eye
shorter than diameter of toruli; distance between
toruli slightly shorter than the diameter of torulus.
Frons finely coriaceous to reticulated without
punctures; frontal carinae weak, but visible, not
page 9 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
branched and not reaching lateral ocelli (Figs. 2a,
c). Head in dorsal view (Fig. 2c) is about two times
wider than long. Vertex nely coriaceous, without
punctures. POL:OOL:LOL = 8:3.5:3; OOL as long
as the diameter of lateral ocellus. Occiput (Fig. 2b)
coriaceous, with neither rugosities nor punctures.
Antennae (Fig. 2f): 14-segmented (7: 4: 9: 8:
8: 8: 7: 7: 5: 5: 5: 5: 4: 7.5); liform, not broadened
apically; pubescence dense and short; placodeal
sensilla little visible only on agellar segments F7-
F12. Pedicel 1.3 times longer than wide; F1 slightly
longer than F2, F2 as long as F3. Last flagellar
segment 3.0 times longer than wide and 1.9 times
longer than F11.
Mesosoma: About 1.2 times longer than high
in lateral view (including nucha), with short and
not dense pubescence (Fig. 3b). Ratio of length
of pronotum medially/laterally: 0.36. Pronotal
plate absent; pronotal foveae supercial, long and
separated. Lateral pronotum sculpture imbricated,
with some weak carinae medially and above
pronotal foveae; lateral margins of pronotum
rounded, without lateral carina. Mesoescutum
(Fig. 3a) 1.3 times wider than long, imbricated to
weakly but densely transversely carinated, the
interspaces coriaceous; anterior grooves weak,
occupying almost 1/3 of mesoescutum length.
Notauli complete, wider posteriorly and slightly
convergent. Median groove almost indistinct.
Parapsidial grooves almost indistinct, slightly
surpassing tegulae. Scutellum (Fig. 3a) rounded,
almost as long as wide, wrinkled, circumscutellar
carina not visible; scutellar foveae superficial,
shallowly impressed, smooth, more or less oval-
triangular-shaped, posterior margins almost
indistinct and separated from each other by a
Fig. 2. Synergus gilletti Pujade-Villar and Lobato-Vila sp. nov.: (a) Female head in frontal view, (b) female head in ventral view, (c)
female head in dorsal view, (d) male head in dorsal view, (e) tarsal claw, (f) female antenna, (g) F1 and F2 of male antenna.
(a)
(b)
(c) (f)
(g)
(d)
(e)
page 10 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
narrow septum. Mesopleurae (Fig. 3b) transversely
striated, speculum smooth; interspaces smooth,
little pubescent basally. Metapleural sulcus well
defined, reaching 4/5 parts of mesopleurae
height. Propodeum (Fig. 3c) pubescent and weak
sculptured, alutaceous with some transversal
rugae; propodeal carinae straight and almost
parallel, central area alutaceous and pubescent.
Nucha sulcated dorsally and laterally.
Legs (Fig. 2e): Tarsal claws with a small basal
tooth.
Wings: Forewings pubescent with marginal
setae, longer than mesosoma plus metasoma.
Radial cell closed, 2.5 times longer than wide;
areolet indistinct but visible, small. Rs + M vein
visible in almost 3/4 parts, not reaching basal vein.
Basal cell pubescent.
Metasoma (Fig. 3d): Shorter than head
plus mesosoma length, slightly longer than
high in lateral view. First metasomal segment
sulcated dorsally and laterally. Syntergite smooth,
anterolateral pubescence composed of a group of
4-5 setae, with a band of micropunctures extending
a little more than 1/3 of the metasomal length; not
incised dorso-distally. Hypopygial spine almost as
long as wide and with a few lateral setae; apical
setae not projected beyond the tip of the spine.
Male
Similar to female, except for the following
morphological traits: body length 1.1-1.7 mm
Fig. 3. Synergus gilletti Pujade-Villar and Lobato-Vila sp. nov.: (a) Mesosoma in dorsal view, (b) mesosoma in lateral view, (c)
propodeum, (d) female metasoma in lateral view.
(a)
(c)
(b)
(d)
page 11 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
(n = 43). Transfacial line slightly shorter than the
height of the compound eye (0.9). POL: OOL: LOL
= 7: 5: 4 (Fig. 2d). Antennae 15-segmented (7: 5:
10.5: 9: 9: 9: 7: 6: 6: 5: 5: 5: 5: 5: 6); F1 curved,
medially excavated, more expanded distally (Fig.
2g). Metasoma shorter than females, and almost
as long as mesosoma.
Distribution: State of Zacatecas (Mexico).
Biology: The material studied was obtained
from soft galls developed on Q. laeta Liebm. and
Q. rugosa Née (Quercus section) related to the
genera Atrusca and Andricus.
Synergus longimalaris Pujade-Villar and
Lobato-Vila sp. nov.
(Fig. 4)
urn:lsid:zoobank.org:act:5DB91102-5D22-4144-93AF-
96DE2962C23B
Type material: Holotype ♀ deposited in JP-V
col (UB) with the following labels: ‘MEX: P160,
Sierra de Quila (Tecolotlán, JAL), 20°16'42.6"N,
104°05'26.7"W, 1916 m’ (white label), ‘Ex.
Amphibolips sp., Q. conzattii., (04.xi.2011)
25.xi.2011, E. Estrada and A. Equihua leg.’ (white
label), ‘Holotype♀ Synergus longimalaris Pujade-
Villar and Lobato-Vila, Lobato-Vila 2017 desig.’ (red
label). Paratypes: same data as the Holotype: 7
♂
and 17♀ (1
♂
and 4♀ deposited in the NMNH;
1
♂
and 4♀ deposited in the AMNH).
Additional material: 2♀ with the same data
as the Holotype dissected and covered with gold.
Etymology: Adjective related to its long malar
space.
Diagnosis: This species is morphologically
related to S. cibriani, S. citriformis (= S. elegans)
and S. laticephalus. The new species differ from
S. laticephalus by the width of the head in dorsal
view, which is much shorter (2.7 times wider
than long in S. laticephalus), the presence of
metasomal micropunctures (no micropunctures
in S. laticephalus) and the absence of rugae in
the scutellum (with some transverse rugae in
S. laticephalus). Synergus longimalaris sp. nov.
mainly differs from S. cibriani and S. citriformis by
having POL as long as OOL (POL usually 1.5 times
longer than OOL in these species), F1 1.5 times
longer than F2 (F1 usually 1.3 times longer in
these species), notauli complete and visible along
their entire length (inconspicuous in the anterior
1/3 in these species) and head coloration yellowish
to brown, except for a dark spot in the area
delimited by ocelli, in males (with more dark areas
in these species). It also differs from S. cibriani by
having OOL 2.2 times longer than the diameter of
the lateral ocellus (1.5 times longer in S. cibriani)
and mesoscutum nely coriaceous to alutaceous,
with some little punctures (strongly coriaceous
to imbricated, without punctures, in S. cibriani).
Finally, it also differs from S. citriformis by having
the malar space about 0.8 times as long as the
height of the compound eye (0.6 in S. citriformis),
both frons and mesoscutum coriaceous with some
little punctures (nely sculptured, no punctures, in
S. citriformis) and metasoma with micropunctures
(no micropunctures in S. citriformis).
Female
Length: Body length 2.3-3.4 mm (n = 18).
Color: Head yellowish to brown; frons and
vertex medially black, orange laterally; occiput
black; antennae yellowish; mesosoma black,
tegulae pale yellow; metasoma black, chestnut
basally; legs yellowish, metatarsi darker, the
base of metacoxae with a more or less extended
brownish or blackish coloration; forewings hyaline,
yellow veins.
Head: in frontal view (Fig. 4a) quadrangular,
1.3 times wider than high, genae slightly expanded
behind compound eye. Face faintly pubescent,
lower face and genae with thin, dense and regular
striae irradiating from clypeus and reaching basal
margin of compound eye and basal margin of
antennal toruli; medial carina inconspicuous.
Clypeus indistinct, ventral margin not projected
over mandibles. Malar space almost 0.8 times as
long as the height of the compound eye (19:25).
Anterior tentorial pits visible; pleurostomal and
epistomal sulcus absent. Transfacial line longer
than the height of the compound eye (1.3). Toruli
situated under mid-height of the compound eye;
distance between torulus and compound eye
longer than diameter of toruli; distance between
toruli shorter than the diameter of toruli. Frons
finely coriaceous with some scattered little
punctures; frontal carinae absent. Head in dorsal
view (Fig. 4c) is about two times wider than long.
Vertex nely coriaceous to alutaceous with neither
punctures nor striae. POL:OOL:LOL = 10:10:4;
OOL 2.2 times longer than the diameter of lateral
ocellus. Occiput (Fig. 4b) finely coriaceous, with
neither punctures nor striae.
Antennae (Fig. 4e): 14-segmented (15: 8:
19: 12: 11: 10: 8: 7.5: 6: 6: 6: 5: 4.5: 8); liform not
broadened apically; pubescence dense and short;
placodeal sensilla little visible only on flagellar
segments F5-F12. Pedicel about two times longer
page 12 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
Fig. 4. Synergus longimalaris Pujade-Villar and Lobato-Vila sp. nov.: (a) Female head in frontal view, (b) female head in ventral view, (c)
female head in dorsal view, (d) mesosoma in lateral view, (e) female antenna, (f) mesosoma in dorsal view, (g) propodeum, (h) tarsal
claw, (i) female metasoma in lateral view, (j) detail of the metasomal micropunctures.
(a)
(b)
(f)
(i)
(j)
(g) (h)
(c) (e)
(d)
page 13 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
than wide; F1 about 1.5 times longer than F2, F2
slightly longer than F3. Last agellar segment 2.7
times longer than wide and 1.8 times longer than
F11.
Mesosoma: About 1.2 times longer than
high in lateral view (including nucha), with short
and not dense pubescence (Fig. 4d). Ratio
of length of pronotum medially/laterally: 0.2.
Pronotal plate absent. Lateral pronotum sculpture
strongly coriaceous; lateral margins of pronotum
rounded, without lateral carina. Mesoescutum
(Fig. 4f) about 1.3 times wider than long, finely
coriaceous to reticulated, with piliferous punctures
over all its surface; anterior grooves weak but
visible, occupying almost 1/4 of mesoescutum
length. Notauli complete and visible along their
entire length, wider and convergent posteriorly.
Median groove very shallowly impressed but
visible, wider posteriorly, reaching the middle of
the mesoescutum. Parapsidial grooves weak but
visible, surpassing the anterior margin of tegulae.
Scutellum (Fig. 4f) rounded, slightly longer than
wide, coriaceous to alutaceous, with some
punctures; circumscutellar carina visible, forming a
slightly projected margin; scutellar foveae shallowly
impressed, almost inconspicuous; triangular,
separated by a wide septum. Mesopleurae (Fig.
4d) transversely striated, speculum smooth;
interspaces alutaceous, little pubescent basally.
Metapleural sulcus well defined, reaching about
3/4 parts of mesopleurae height. Propodeum (Fig.
4g) pubescent and almost smooth; propodeal
carinae straight, just slightly convergent basally,
central area with a weak sculpture and pubescent.
Nucha sulcated dorsally and laterally.
Legs (Fig. 4h): Tarsal claws with a basal
tooth.
Wings: Forewings pubescent with short
marginal setae, slightly shorter than mesosoma
plus metasoma. Radial cell closed, 2.4 times
longer than wide; areolet indistinct but visible. Rs
+ M inconspicuous, not reaching basal vein. Basal
cell faintly pubescent.
Metasoma: About as long as head plus
mesosoma, longer than high in lateral view (Fig.
4i). First metasomal segment sulcated dorsally
and laterally. Syntergite smooth, anterolateral
pubescence composed of a group of 4-5 setae,
with a few weak micropunctures forming a
posterodorsal patch (Fig. 4j); not incised dorso-
distally. Hypopygial spine almost as long as wide
and with a few lateral setae; without apical setae.
Male
Similar to female, except for the following
morphological traits: body length 2.6-3.4 mm (n =
6). Antennae 15-segmented (13: 7: 17: 12: 11: 10:
8.5: 7.5: 7: 6: 5: 5: 4.5: 4: 6.5), F1 1.4 times longer
than F2; F1 slightly curved, medial excavation
weak, faintly expanded apically and basally. Head
coloration yellowish, except for a dark to black spot
in the area delimited by ocelli; metacoxae entirely
yellow, metatarsi yellow.
Distribution: State of Jalisco (Mexico).
Biology: The material studied was obtained
from galls developed on Q. conzattii Trel. (Lobatae
section) related to the genus Amphibolips.
Synergus longiscapus Pujade-Villar and
Lobato-Vila sp. nov.
(Fig. 5)
urn:lsid:zoobank.org:act:F5EEAC91-B75E-4090-8341-
60D1D91B4D65
Type material: Holotype ♀ deposited in
JP-V col. (UB) with the following labels: ‘MEX:
300, Santa Fe (D.F.), 19°21'14"N, 99°16'24.22"W,
2569 m’ (white label), ‘Ex. Disholcaspis sp., Q.
laeta, (20.vi.2016) vii.2016, JP-V leg.’ (white label),
‘Holotype♀ Synergus longiscapus Pujade-Villar
and Lobato-Vila, Lobato-Vila 2017 desig.’ (red
label). Paratypes: same data as the Holotype: 10
♂
and 36♀ (1
♂
and 5♀ deposited in the NMNH;
1
♂
and 5♀ deposited in the AMNH).
Additional material: MEX: 060, La Cuchilla
(ZAC), 22°18'50.3"N 103°37'14.6"W, 2505 m,
Ex. Disholcaspis nr regina, Q. eduardii c.f., (14.
x.2011) 02.xi.2011, O. Cabral, R. Treto, L.G. Landa
and C. Carrillo leg.: 1♀; MEX: 206, Plateros-
Arenales (San Felipe del Progreso, EDO. MÉX),
19°42'31.5"N, 99°57'03.3"W, 2564 m, Ex. Andricus
tectucnarum or georgei groups, Q. candicans, (21.
i.2014) 19.ii.2014, Delia leg.: 4
♂
and 1♀; MEX:
P016, Monte Escobedo (ZAC), 22°18'07.9"N,
103°33'45.5"W, 2190 m, Ex. Andricus sphaericus,
Q. rugosa, (25.v.2012) vi.2012, JP-V leg.: 2♀;
MEX: P116, Cerro del Potosí (Los Galeana,
NL), 24°52'18.9"N, 100°13'57.5"W, 3716 m, Ex.
Disholcaspis c.f., Q. emoryi, (08.v.2012) 12-26.
v.2012, E. Estrada and A. Equihua leg.: 2
♂
and
2♀; MEX: P187, Perote-Puebla Federal Highway
(VER), 19°32'30"N, 97°16'23"W, 2410 m, Ex.
Disholcaspis sp., Quercus sp., (28.ix.2012)
15.x.2012, E. Estrada and A. Equihua leg.: 1
♂
and 1♀; MEX: P228, Santa Fe (D.F.), 19°21'14"N,
99°16'24.22"W, 2569 m, Ex. Andricus protuberans,
page 14 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
Fig. 5. Synergus longiscapus Pujade-Villar and Lobato-Vila sp. nov.: (a) Female head in frontal view, (b) female head in ventral view, (c)
female head in dorsal view, (d) male head in dorsal view, (e) tarsal claw, (f) female antenna, (g) male antenna, (h) mesosoma in dorsal
view, (i) mesosoma in lateral view, (j) propodeum, (k) female metasoma in lateral view, (l) detail of the metasomal micropunctures.
(a)
(b)
(c)
(d)
(e)
(i)
(h)
(j) (k) (l)
(f) (g)
page 15 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
Q. laeta, (21.vi.2013) ix.2013, JP-V leg.: 3♀;
MEX: 304, Ejido de San Miguel Tlaixpan (Texcoco,
EDO. MÉX), 19°26'29.3"N, 98°46'02.6"W, 2995 m,
Ex. Andricus sphaericus, Q. rugosa, (01.vii.2016)
vii.2016, A. Equihua and E. Estrada: 2♀; MEX:
308, Ejido Santa María Nativitas (Texcoco, EDO.
MÉXICO), 19°27'41.3"N, 98°47'22.2"W, 2703 m,
Ex. Andricus nievesaldreyi, Q. rugosa, (13.
i.2015) 19.ii.2015, J. Acuña leg.: 1
♂
; MEX: P238,
Ejido de San Miguel Tlaixpan (Texcoco, EDO.
MÉX), 19°26'29.3"N, 98°46'02.6"W, 2995 m, Ex.
undetermined gall, Q. glabrescens, (08.ii.2016)
03.iv.2016, A. Equihua and E. Estrada: 1♀ and
1
♂
; MEX: P239, Ejido de San Miguel Tlaixpan
(Texcoco, Estado de México), 19°26'29.3"N,
98°46'02.6"W, 2995 m, Ex. undetermined gall, Q.
glabrescens, (08.ii.2016) 03.iv.2016, A. Equihua
and E. Estrada: 1♀; MEX: P240, Ejido Santa
María Nativitas (Texcoco, EDO. MÉXICO),
19°27'41.3"N, 98°47'22.2"W, 2703 m, Andricus
nr cylindratum, Q. glabrescens, (27.i.2015)
09.iii.2015, A. Equihua and E. Estrada: 1♀; MEX:
P242, Ejido Santa María Nativitas (Texcoco,
EDO. MÉXICO), 19°27'41.3"N, 98°47'22.2"W,
2703 m, Andricus nr cylindratum, Q. rugosa, (13.
i.2015) 09.iii.2015, J. Acuña leg.: 1♀; 1
♂
and 2♀
dissected and covered with gold with the same
data as MEX: 206.
Etymology: Adjective related to its long
scapus.
Diagnosis: This species is morphologically
related to S. cibriani, S. citriformis (= S. elegans),
S. laticephalus and S. longimalaris. The new
species differs from all these species and from
S. striatifrons sp. nov. by having a long scapus,
slightly longer than F1 (shorter than F1 in all the
species addressed here). It also differs from S.
laticephalus by the width of the head in dorsal
view, which is much shorter (2.7 times wider
than long in S. laticephalus), the presence of
metasomal micropunctures (no micropunctures
in S. laticephalus) and the absence of rugae in
the scutellum (with some transverse rugae in S.
laticephalus), and from S. cibriani, S. citriformis
and S. longimalaris, by having F1 and F2 subequal
(F1 clearly longer than F2 in all these species)
and the transfacial line about as long as the height
of the compound eye (1.3 times longer in these
species). Synergus longiscapus sp. nov. also
differs from S. cibriani and S. citriformis by having
the scutum with some piliferous punctures (no
punctures in these species) and head coloration
yellowish to brown, except for a dark spot in the
area delimited by ocelli, in males (males’ head
with more dark areas in these species); from S.
cibriani and S. longimalaris, by having the malar
space about 0.6 times as long as the height of
the compound eye (0.8 in these species); from
S. cibriani, by having OOL 2.7 times longer than
the diameter of the lateral ocellus in males (1.5
times longer in S. cibriani in both sexes); from S.
citriformis, by having the frons finely coriaceous
with some scattered little punctures and the
metasomal syntergite with a posterodorsal patch
of micropunctures (no punctures in S. citriformis);
finally, from S. longimalaris by having POL 1.6
times longer than OOL (POL as long as OOL in
S. longimalaris) and notauli inconspicuous in the
anterior 1/3 (visible along their entire length in S.
longimalaris).
Female
Length: Body length 1.4-2.9 mm (n = 10).
Color: Head yellowish to light brown; frons
and vertex black medially, yellowish to light brown
laterally; occiput black; antennae yellowish to light
brown; mesosoma black, tegulae pale yellow;
metasoma black dorsally, light chestnut basally;
legs light yellow, metatarsi usually darker, the base
of metacoxae with brownish or blackish coloration
more or less extended; wings hyaline, veins yellow
to light brown.
Head: in frontal view (Fig. 5a) more or less
rounded, about 1.2 times wider than high, genae
slightly expanded behind compound eye. Face
faintly pubescent, lower face and genae with thin,
dense and regular striae irradiating from clypeus
and reaching basal margin of compound eye and
basal margin of antennal toruli; medial carina
inconspicuous. Clypeus indistinct, ventral margin
slightly projected over mandibles. Malar space 0.6
times as long as the height of the compound eye
(12:19). Anterior tentorial pits visible; pleurostomal
and epistomal sulcus absent. Transfacial line just
slightly longer than the height of the compound
eye (1.1). Toruli situated mid-height of the
compound eye; distance between torulus and
compound eye shorter than diameter of toruli;
distance between toruli shorter than the diameter
of toruli. Frons nely coriaceous to alutaceous with
some scattered little punctures; frontal carinae
absent. Head in dorsal view (Fig. 5c) is about 1.8
times wider than long. Vertex finely coriaceous
to alutaceous with neither punctures nor striae.
POL:OOL:LOL = 8:5:4; OOL about 2.0 times
longer than the diameter of lateral ocellus. Occiput
(Fig. 5b) nely coriaceous, with neither punctures
page 16 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
nor striae.
Antennae (Fig. 5f): 14-segmented (12: 6:
10.5: 10: 9: 8: 7: 6.5: 5: 5: 5: 4.5: 4.5: 8.5); liform
not broadened apically; pubescence dense and
short; placodeal sensilla little visible only on
flagellar segments F7-F12. Pedicel about two
times longer than wide; F1 and F2 subequal, F2
slightly longer than F3. Last agellar segment 2.8
times longer than wide and 1.9 times longer than
F11.
Mesosoma: Slightly longer than high in
lateral view (including nucha), with short and not
dense pubescence (Fig. 5i). Ratio of length of
pronotum medially/laterally: 0.36. Pronotal plate
absent. Lateral pronotal sculpture coriaceous to
reticulated; lateral margins of pronotum rounded,
without lateral carina. Mesoscutum (Fig. 5h) about
1.2 times wider than long, finely coriaceous, with
some weak piliferous punctures over all its surface;
anterior grooves weak, occupying about 1/4 of
mesoescutum length. Notauli complete but faint in
the anterior 1/3, wider and convergent posteriorly.
Median groove almost indistinct but visible.
Parapsidial grooves weak, reaching the anterior
margin of tegulae. Scutellum (Fig. 5h) rounded,
almost as long as wide, nely coriaceous, weakly
imbricated posteriorly, circumscutellar carina visible
but not projected; scutellar foveae inconspicuous.
Mesopleurae (Fig. 5i) with medial and anterior
areas transversely striated, smooth basally,
speculum also smooth; interspaces alutaceous,
little pubescent basally. Metapleural sulcus well
dened, reaching about 3/4 parts of mesopleurae
height. Propodeum (Fig. 5j) pubescent and with a
weak sculpture; propodeal carinae almost straight,
slightly convergent basally, central area with a
weak sculpture and pubescent. Nucha sulcated
dorsally and laterally.
Legs (Fig. 5e): Tarsal claws with a small basal
tooth.
Wings: Forewings pubescent with short
marginal setae, longer than mesosoma plus
metasoma. Radial cell closed, 2.4 times longer
than wide; areolet absent. Rs + M inconspicuous,
not reaching basal vein. Basal cell faintly
pubescent.
Metasoma: Slightly shorter than head plus
mesosoma, longer than high in lateral view (Fig.
5k). First metasomal segment sulcated dorsally
and laterally. Syntergite smooth, anterolateral
pubescence composed of a group of 4-5 setae,
with a few weak micropunctures forming a
posterodorsal patch (Fig. 5l); not incised dorso-
distally. Hypopygial spine as long as wide and with
a few lateral setae; without apical setae.
Male
Similar to female, except for the morphological
traits that follow: body length 1.4-2.0 mm (n = 7).
POL:OOL:LOL = 6.5: 4: 3; OOL 2.7 times longer
than the diameter of lateral ocellus (Fig. 5d).
Antennae (Fig. 5g) 15-segmented (9: 5: 9: 8: 7:
6: 6: 5: 5: 4.5: 4: 4: 4: 3.5: 5); F1 slightly longer
than F2, F1 slightly curved, medial excavation
weak, faintly expanded apically and basally. Head
coloration yellowish, except for a black spot in the
area delimited by ocelli; mesopleurae sometimes
brownish to black; metacoxae usually entirely
yellow or with dark coloration less extended; tarsi
yellow.
Distribution: Federal District (D.F.), Nuevo
León, State of Mexico, Veracruz and Zacatecas
(Mexico).
Biology: The studied material was obtained
from soft galls developed on Q. candicans Née, Q.
eduardii c.f. and Q. emoryi Torr. (Lobatae section);
Q. glabrescens Benth., Q. laeta Liebm. and Q.
rugosa Née (Quercus section), related to the
genera Andricus and Disholcaspis, as well as to
some undetermined galls.
Synergus punctatus Gillette
(Figs. 6, 8a)
Synergus punctata Gillette, 1896: 90, 94 (
♂
and ♀)
Type material: Lectotype ♀ deposited in
NMNH (Washington D.C., USA) with the following
labels: ‘Colo. 841’ (white label) / ‘Biol. 116’ (white
label) / ‘Type’ (white label, handwritten) / Acc n°
71950, USNM to be loaned only to ColoAgCol’
(white label) / ‘Synergus punctata Gill.’ (white
label bordered with two red lines, handwritten)
/ ‘USNM 00802265’ (white label, QR code) /
‘Lectotype of Synergus punctatus ♀ Gillette, 1896,
JP-V desig-2017’ (red label). Paralecototypes
(3
♂
and 6♀) deposited in NMNH (Washington
D.C., USA) with the following labels: ‘Colo. 841’
(white label) / ‘Biol. 116’ (white label) / ‘Type’
(white label, handwritten) / ‘Collection Ashmead’
(white label) / ‘Synergus punctata Gill.’ (red label,
handwritten) / ‘USNM 00960582’ (white label,
QR code) / ‘Paralectotype of Synergus punctatus
♀ Gillette, 1896, JP-V desig-2017’ (red label)
(1♀); ‘Colo. 1864’ (white label) / ‘Type’ (white
label, handwritten) / ‘Collection Ashmead’ (white
label) / ‘USNM 00960208’ (white label, QR code)
page 17 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
/ ‘Paralectotype of Synergus punctatus
♂
Gillette,
1896, JP-V desig-2017’ (red label) (1
♂
); ‘Colo.
841’ (white label) / ‘Biol. 116’ (white label) / ‘Type’
(white label, handwritten) / ‘Collection Ashmead’
(white label) / ‘USNM 00960641’ (white label, QR
code) / ‘Paralectotype of Synergus punctatus
♂
Gillette, 1896, JP-V desig-2017’ (red label) (1
♂
);
‘Colo. 843’ (white label) / ‘Biol. 117’ (white label)
/ ‘Type’ (white label, handwritten) / ‘Collection
Ashmead’ (white label) / ‘Synergus punctata ♀
Gill.’ (white label, handwritten) / ‘USNM 00960471’
(white label, QR code) / ‘Paralectotype of Synergus
punctatus ♀ Gillette, 1896, JP-V desig-2017’ (red
label) (1♀); ‘Colo. 843’ (white label) / ‘Biol. 117’
(white label) / ‘Type’ (white label, handwritten) / Acc
n° 71950, USNM to be loaned only to ColoAgCol’
(white label) / ‘USNM 00960234’ (white label,
QR code) / ‘Paralectotype of Synergus punctatus
♀ Gillette, 1896, JP-V desig-2017’ (red label)
(1♀); ‘Colo. 1864’ (white label) / ‘Type’ (white
label, handwritten) / Acc n° 71950, USNM to be
loaned only to ColoAgCol’ (white label) / ‘USNM
00960481’ (white label, QR code) / ‘Paralectotype
of Synergus punctatus ♀ Gillette, 1896, JP-V
desig-2017’ (red label) (1♀); ‘Colo. 1864’ (white
label) / ‘Type’ (white label, handwritten) / Acc n°
71950, USNM to be loaned only to ColoAgCol’
(white label) / ‘USNM 00960708’ (white label, QR
code) / ‘Paralectotype of Synergus punctatus ♀
Gillette, 1896, JP-V desig-2017’ (red label) (1♀);
‘Colo. 1722’ (white label) / ‘Type’ (white label,
handwritten) / ‘Coll. CF Baker’ (white label), ‘USNM
00960567’ (white label, QR code) / ‘Paralectotype
of Synergus punctatus
♂
Gillette, 1896, JP-V
desig-2017’ (red label) (1
♂
); ‘Colo. 1719’ (white
label) / ‘Type’ (white label, handwritten) / ‘Coll.
CF Baker’ (white label), ‘USNM 00960892’ (white
label, QR code) / ‘Paralectotype of Synergus
punctatus
♂
Gillette, 1896, JP-V desig-2017’ (red
label) (1♀).
Additional material: Col. CF Baker (1
♂
and
5♀) with the following QR codes: 00960262 (♀),
00960464 (♀), 00960565 (♀), 00960598 (♀),
00960742 (♀) and 00960874 (
♂
); Col. Ashmead
(1♀) with the following QR code: 00960652.
Comments: According to Gillette (1896:
94), this species was described based on 24
females and 22 males. We located 4 males and 10
females in the NMNH from Ashmead and Baker’s
collections; 3
♂
and 6♀ have similar labels to the
ones of the Lectotype and a handwritten label with
the word ‘type’. So, we assume these specimens
belong to the typical series. Baker’s material
mentioned in ‘Additional material’ probably belongs
to the typical series too, but the ‘type’ label is
missing.
Synergus punctatus has partially smooth
mesopleurae (striae are not extended into the
speculum) as all the species mentioned in this
study. Synergus punctatus is closely related to
S. gilletti because females of both species have
head and body coloration entirely black, except
for a light halo around the mouth; also, both sexes
of these two species have frontal carinae, the
mesoscutum with weak transverse carinae, the
scutellum wrinkled with the circumscutellar carina
not visible and the metasomal punctuation forming
a wide band, amongst others (see the differences
between these species in the diagnosis of S.
gilletti). On the contrary, S. punctatus mainly differs
from S. cibriani, S. citriformis (= S. elegans), S.
laticephalus, S. longimalaris and S. striatifrons
sp. nov. by having frontal carinae (absent in
all these species), the mesosoma with a few
weak transverse carinae (coriaceous or weakly
imbricated in all these species), the circumscutellar
carina not visible (visible in all these species), a
wide band of micropunctures in the metasomal
syntergite (no punctures or only forming a
posterodorsal patch in all these species), and body
coloration entirely black in females, except for a
yellow and well defined halo around the mouth
(other coloration in all these species).
Short description: Synergus punctatus is
characterized by the following combination of
morphological traits: in females, body and head
black, except for a yellow and well-defined halo
around the mouth (Fig. 8a); in males, head yellow
except for a black spot in the vertex. Frons and
vertex coriaceous, without punctures (Figs. 6a, b,
c). Frontal carinae weak but visible, incomplete
and not branched (Figs. 6b, c). Female antennae
14-segmented, F1 just slightly longer than F2
(Fig. 6f); in males, F1 about 1.4 times longer than
F2, curved, medially excavated, more expanded
distally (Fig. 6b). Mesoescutum (Fig. 6d) weakly
but densely transversely carinated; notauli
inconspicuous in the anterior third of the scutum;
mesopleurae (Fig. 6g) transversely striated,
speculum smooth. Scutellum (Fig. 6d) wrinkled;
circumscutellar carina not visible. Metasomal
syntergite punctuation forming a band, occupying
the distal half of the syntergite and reaching the
ventral part of metasoma; not incised dorso-distally
(Fig. 6h). Tarsal claws with a small basal tooth (Fig.
6e). Radial cell closed, 2.5 times longer than wide.
Distribution: USA: Manitou, Colorado (Gillette,
1896); Stanford, California (McCracken and Egbert
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© 2017 Academia Sinica, Taiwan
Fig. 6. Synergus punctatus Gillette, 1896: (a) Female head in frontal view, (b) male head and antennae in dorsal view, (c) female head
in dorsal view, (d) mesosoma in dorsal view, (e) tarsal claw, (f) female antenna, (g) mesosoma in lateral view, (h) female metasoma in
lateral view.
(a)
(b)
(g) (h)
(c)
(e)
(f)
(d)
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© 2017 Academia Sinica, Taiwan
1922).
Biology: According to Gillette (1896) and
McCracken and Egbert (1922), this species is
related to the galls of Disholcaspis rubens (Gillette),
Disholcaspis eldoradensis (Beutenmüller) and
Biorhiza eburnea (Bassett) (= Dryophanta glabra
Gillette), always reared from oak species from
Quercus section: Q. arizonica, Q. dumosa, Q.
gambelii, Q. garryana, Q. lobata, Q. oblongifolia, Q.
reticulata, Q. submollis, Q. toumeyi, Q. turbinella
and Q. undulata.
Synergus striatifrons Pujade-Villar and Lobato-
Vila sp. nov.
(Fig. 7)
urn:lsid:zoobank.org:act:96C806A3-E7F3-45A1-9AA2-
5892F549CCBA
Type material: Holotype ♀ deposited in JP-V
col. (UB) with the following labels: ‘MEX: 292, San
Marcos Contla (Papalotla de Xicohténcatl, TLAX),
19°11'54.2"N, 98°11'58.7"W, 2342 m’ (white label),
‘Ex. Amphibolips hidalgoensis, Q. crassipes, (03.
vi.2010) 04.vii.2010, A. Equihua and E. Estrada’
(white label), ‘HOLOTYPE ♀ Synergus striatifrons
Pujade-Villar and Lobato-Vila, Lobato-Vila 2017
desig.’ (red label). Paratypes: same data as the
Holotype: 2
♂
and 49♀ (10♀ deposited in the
NMNH; 10♀ deposited in the AMNH).
Additional material: Same data as holotype:
20♀; MEX: 286, Laguna de Gamboa (Monte
Escobedo, ZAC), 22°16'57.04"N, 103°32'58.38"W,
2197 m, Ex. Amphibolips zacatequensis, Q.
eduardii, (27.iv.2011) vi.2011, C. Carrillo leg.: 2
♂
and 74♀; MEX: 287, Monte Escobedo (ZAC),
22°18'23.49"N, 103°32'47.56"W, 2188 m, Ex.
Amphibolips zacatequensis, Q. eduardii, (22.
iv.2011) 05.vii.2011, C. Carrillo leg.: 1♀; MEX:
299, Fracc. Bosques de Santa Fe (Cuajimalpa,
D.F.), 19°21'09.78"N, 99°15'55.02"W, 2.543 m,
Ex. Amphibolips sp., Q. crassipes, (10.vii.2014)
vii.2014, D. Cibrián leg.: 2
♂
and 20♀; 2♀
dissected and covered with gold with the same
data as MEX: 286 and MEX: 287, respectively.
Etymology: Name in apposition related to
weak striae present in its frons.
Diagnosis: This species is morphologically
related to S. cibriani, S. citriformis (= S. elegans), S.
laticephalus, S. longimalaris and S. longiscapus.
The new species differs from all these species
by having the frons covered by multiple weak
striae extending from lower face and toruli (no
striae in these species). It also differs from S.
laticephalus by the width of the head in dorsal
view, which is much shorter (2.7 times wider
than long in S. laticephalus), the presence of
metasomal micropunctures (no micropunctures
in S. laticephalus) and the absence of rugae in
the scutellum (with some transverse rugae in S.
laticephalus), and from S. cibriani, S. citriformis,
S. longimalaris and S. longiscapus by having
OOL 3.0 times longer than the diameter of the
lateral ocellus (much shorter in all these species,
except in males of S. longiscapus), scutellar
foveae shallow, but visible (inconspicuous in
these species) and metasoma clearly longer than
head plus mesosoma (about as long as head plus
mesosoma in these species). Synergus striatifrons
sp. nov. also differs from S. cibriani, S. longimalaris
and S. longiscapus by the absence of punctures
in the frons (frons with a few punctures in these
species); from S. cibriani, S. citriformis and S.
longiscapus, by having POL about as long as OOL
(POL about 1.5 times longer than OOL in these
species) and F1 1.5 times longer than F2 (shorter
in these species); from S. cibriani and S. citriformis,
by having the mesoscutum with a few piliferous
punctures (no punctures in these species); from S.
citriformis and S. longiscapus, by having the malar
space 0.8 as long as the height of the compound
eye (0.6 in these species); from S. longimalaris
and S. longiscapus, by having frons, vertex and
occiput black in both sexes (head coloration
yellowish to brown, except for a dark spot in the
area delimited by ocelli in males of these species);
from S. citriformis, by the presence of metasomal
micropunctures forming a posterodorsal patch
(no punctures in S. citriformis); finally, from S.
longimalaris by having notauli inconspicuous in
the anterior 1/3 (visible along its entire length in S.
longimalaris).
Female
Length: Body length 2.9-4.3 mm (n = 19).
Color: Head black, except for the face and
genae, which are yellowish orange; frons and
vertex black, occiput black; antennae dark brown;
mesosoma black, tegulae yellowish; metasoma
black, reddish to chestnut basally; legs testaceous
to yellow, metatarsi darker, basal half of metacoxae
black; forewings hyaline, brownish veins.
Head: in frontal view (Fig. 7a) quadrangular,
1.2 times wider than high, genae slightly expanded
behind compound eye. Face faintly pubescent,
lower face and genae with thin, dense and regular
striae irradiating from clypeus and reaching
basal margin of compound eye and basal margin
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© 2017 Academia Sinica, Taiwan
Fig. 7. Synergus striatifrons Pujade-Villar and Lobato-Vila sp. nov.: (a) Female head in frontal view, (b) female head in ventral view, (c)
female head in dorsal view, (d) male head in dorsal view, (e) tarsal claw, (f) female antenna, (g) last segments of female antenna, (h) F1
and F2 segments of male antenna, (i) mesosoma in dorsal view, (j) mesosoma in lateral view, (k) propodeum, (l) female metasoma in
lateral view, (m) detail of the metasomal micropunctures.
(a)
(b)
(i)
(k) (l) (m)
(j)
(c)
(d)
(e)
(f)
(h)
(g)
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© 2017 Academia Sinica, Taiwan
of antennal toruli; medial carina also present.
Clypeus indistinct, ventral margin not projected
over mandibles. Malar space about 0.8 times as
long as the height of the compound eye (11:14).
Anterior tentorial pits visible; pleurostomal and
epistomal sulcus absent. Transfacial line longer
than the height of the compound eye (1.3). Toruli
situated under mid-height of the compound eye;
distance between torulus and compound eye about
as long as the diameter of toruli; distance between
toruli shorter than the diameter of torulus. Frons
finely coriaceous without punctures; absence of
true frontal carinae, however, frons covered by
multiple weak striae extending from lower face
and toruli. Head in dorsal view (Fig. 7c) is about
1.8 times wider than long. Vertex nely coriaceous
to imbricated, without punctures. POL:OOL:LOL
= 10:9:5; OOL about 3.0 times longer than the
diameter of lateral ocellus. Occiput (Fig. 7b)
coriaceous, with a few inconspicuous piliferous
punctures and a few weak striae.
Antenna (Figs. 7f, g): 14-segmented (17: 8:
25: 18: 15: 14: 13: 11: 9.5: 8: 8: 6: 6: 10); liform
not broadened apically; pubescence dense and
short; placodeal sensilla little visible only on
flagellar segments F3-F12. Pedicel about 1.5
times longer than wide; F1 about 1.4 times longer
than F2, F2 longer than F3 (Fig. 7f). Last agellar
segment 2.5 times longer than wide and 1.7 times
longer than F11 (Fig. 7g).
Mesosoma: About 1.2 times longer than high
in lateral view (including nucha), with short and
not dense pubescence (Fig. 7j). Ratio of length
of pronotum medially/laterally: 0.3. Pronotal
plate absent. Lateral pronotum sculpture finely
coriaceous; lateral margins of pronotum rounded,
without a lateral carina. Mesoscutum (Fig. 7i) 1.3
times wider than long, finely coriaceous, without
carinae and with a few piliferous punctures
between notauli and parapsidial grooves;
anterior grooves weak, occupying almost 1/3 of
mesoescutum length. Notauli indistinct in the
anterior 1/3, wider and not strongly convergent
posteriorly. Median groove almost indistinct but
visible. Parapsidial grooves weak, reaching the
anterior margin of tegulae. Scutellum (Fig. 7i)
rounded, almost as long as wide, nely coriaceous,
circumscutelar carina visible, forming a clear and
projected margin; scutellar foveae weak, posterior
margins indistinct, shallow, covered with some
striae and separated from each other by a narrow
septum. Mesopleurae (Fig. 7j) transversely striated
medially, smooth and shiny dorsally and basally;
interspaces alutaceous, little pubescent basally.
Metapleural sulcus well dened, reaching just over
3/4 parts of mesopleurae height. Propodeum (Fig.
7k) pubescent and weak sculptured, alutaceous
with some rugae; propodeal carinae slightly
convergent basally, central area alutaceous and
pubescent. Nucha sulcated dorsally and laterally.
Legs (Fig. 7e): Tarsal claws with a small basal
tooth.
Wings: Forewings pubescent with marginal
setae, as long as mesosoma plus metasoma.
Radial cell closed, 2.3 times longer than wide;
areolet indistinct but visible. Rs + M inconspicuous,
not reaching basal vein. Basal cell faintly
pubescent.
Metasoma: Slightly longer than head plus
mesosoma, clearly longer than high in lateral
view (Fig. 7l). First metasomal segment sulcated
dorsally and laterally; however, striae are
incomplete in some specimens, in which case
the anterior part is smooth. Syntergite smooth,
anterolateral pubescence composed of a group
of 6-7 setae, with a few micropunctures forming a
posterodorsal patch (Fig. 7m); not incised dorso-
distally. Hypopygial spine short, or almost as long
as wide, with a few lateral setae; apical setae not
projected beyond the tip of the spine.
Male
Similar to female, except for the following
morphological traits: body length 2.8-2.9 mm
(n = 2). Transfacial line as long as the height of the
compound eye. POL: OOL: LOL = 10: 5: 4 (Fig.
7d). Antennae 15-segmented (12: 5: 18: 12: 10:
10: 8: 7: 7: 6: 5: 5: 5: 5.5: 7); F1 curved, slightly
excavated medially, faintly expanded apically and
basally (Fig. 7h).
Distribution: Federal District (D.F.), Tlaxcala
and Zacatecas (Mexico).
Biology: The studied material was obtained
from soft galls developed on Q. eduardii Trel.
and Q. crassipes Humb. et Bonpl. (Lobatae
section) related to the genus Amphibolips (A.
zacatequensis and A. hidalgoensis).
For the identication of the Synergus species
from Mexico with mesopleurae partially smooth
and their closest related species, we propose
the following key
1. Frontal carinae present, not branched (Figs. 2a, c, d; 6b,
c). Mesoescutum with weak transverse carinae, scutellum
wrinkled (Figs. 3a; 6d). Circumscutellar carina not visible
(Figs. 3a, b; 6d). Syntergite punctuation occupying
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© 2017 Academia Sinica, Taiwan
the distal half (Fig. 6h) or the distal 1/3 (Fig. 3d) of the
syntergite. In females, body and head entirely black, except
for a more or less extended pale halo around the mouth
(Figs. 8a, b); males with the same color pattern as females
or not ................................................................................ 2
- Frontal carinae absent (Figs. 1c, d; 4a, c; 5a, c, d; 7a, c,
d). Mesoescutum and scutellum nely coriaceous to nely
imbricated, with neither carinae nor wrinkles (except in S.
laticephalus, which presents some weak wrinkles in the
scutellum) (Figs. 1b; 4f; 5h; 7i). Circumscutellar carina
visible (Figs. 1b; 4d, f; 5h, i; 7i, j). Syntergite punctuation
absent or only forming a posterodorsal patch (Figs. 1k, l; 4j;
5l; 7m). Body coloration of both females and males lighter;
generally dark dorsally and light ventrally ......................... 3
2. Colored halo around the mouth yellow and clearly dened
(Fig. 8a). Notauli inconspicuous in the anterior 1/3 of
mesoescutum (Fig. 6d). Syntergite punctuation forming
a band, occupying the distal half of the syntergite and
reaching the ventral part of metasoma (Fig. 6h). In males,
F1 about 1.4 times longer than F2 (Fig. 6b) and head
yellow except for a black spot in the vertex .........................
............................................................. S. punctatus Gillette
- Colored halo around the mouth less conspicuous and
reddish-brown to chestnut (Fig. 8b). Notauli complete
and visible along their entire length (Fig. 3a). Syntergite
punctuation not forming a complete band, less widespread
and short (Fig. 3d). In males, F1 just slightly longer than F2
(Fig. 2g) and head with the same color pattern as females
.................................................................. S. gilletti sp. nov.
3. Head strongly transverse in dorsal view, more than 2.5
times wider than long. Scutellum with some wrinkles.
Syntergite not punctured posteriorly ....................................
..................... S. laticephalus Nieves-Aldrey and Medianero
- Head in dorsal view less than 2.5 as wide as long.
Scutellum finely coriaceous to finely imbricated, not
wrinkled. Syntergite punctured posteriorly or not ............. 4
4. Frons and mesoescutum finely coriaceous to reticulated,
with neither punctures nor striae. Malar space 0.6 times
as long as the height of the compound eye. Metasomal
syntergite smooth, not punctured (or at least with scarce
and inconspicuous posterodorsal punctures) ......................
S. citriformis (Ashmead) (= S. elegans Nieves-Aldrey and
Medianero syn. nov.)
- Frons nely coriaceous to reticulated, with some scattered
piliferous punctures or with weak striae (Figs. 1a, c, d; 4a, c;
5a, c, d; 7a, c, d); mesoscutum with some weak punctures,
except in S. cibriani. Malar space 0.8 as long as the height
of the compound eye, except in S. longiscapus. Metasomal
syntergite with punctures forming a posterodorsal patch
(Figs. 1k, l; 4j; 5l; 7m) ....................................................... 5
5. Frons covered by multiple weak striae extending from lower
face and toruli, not forming true frontal carinae; without
punctures (Figs. 7a, c, d). Scutellar foveae shallow and not
delimited posteriorly, but visible (Fig. 7i). Metasoma longer
than head plus mesosoma. In both sexes, head black,
except for the face and genae, which are orange ...............
........................................................... S. striatifrons sp. nov.
- Frons without striae, but with some scattered little
punctures (Figs. 1a, c, d; 4a, c; 5a, c, d). Scutellar foveae
inconspicuous (Figs. 1b; 4f). Metasoma about as long
as head plus mesosoma. In females, head yellowish to
brown; frons and vertex medially black, orange to chestnut
laterally; occiput black. In males, head coloration yellowish
to brown, except for a dark spot in the area delimited by
ocelli (except in S. cibriani) ............................................... 6
6. Scapus long, longer than F1 in females (Fig. 5f) and about
as long as F1 in males (Fig. 5g). F1 as long as F2. Malar
space 0.6 as long as the height of the compound eye (Fig.
5a). OOL 2.0 times the diameter of lateral ocelli in females
(Fig. 5c) and 2.7 times in males (Fig. 5d) ............................
........................................................ S. longiscapus sp. nov.
- Scapus not as long, always shorter than F1 both in males
and females (Figs. 1f, g; 4e). F1 clearly longer than F2
(Figs. 1f, g; 4e). Malar space 0.8 as long as the height of
Fig. 8. Detail of the pale halo around the mouth: (a) Synergus punctatus female (picture extracted from the online database of
Hymenopteran Holotypes (Smithsonian Institution): http://bit.ly/2q3gV9G), (b) Synergus gilletti female.
(a) (b)
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the compound eye (Figs. 1a; 4a). OOL and diameter of
lateral ocelli ratio not different between males and females,
always much less than 2.7 (Fig. 1c, d; 4c) ....................... 7
7. F1 1.5 times longer than F2 (Fig. 4e). POL as long as
OOL (Fig. 4c). OOL 2.2 times the diameter of lateral ocelli.
Notauli visible along their entire length (Fig. 4f). In females,
frons and vertex medially black, orange to chestnut
laterally; in males, head coloration yellowish to brown,
except for a dark spot in the area delimited by ocelli ...........
........................................................ S. longimalaris sp. nov.
- F1 1.3 times longer than F2 (Figs. 1f, g). POL 1.5 times
longer than OOL (Figs. 1c, d). OOL 1.5 times the diameter
of the lateral ocellus. Notauli inconspicuous in the anterior
1/3 (Fig. 1b). Frons and vertex medially black, orange to
chestnut laterally both in males and females .......................
................................................................ S. cibriani sp. nov.
DISCUSSION
Morphological limits between genera
Synergus and Saphonecrus are imprecise.
According to the results presented in Schwéger et
al. (2015a) about Palearctic species of Synergus,
the only consistent morphological trait found to
distinguish Saphonecrus from Synergus is the
presence (Synergus) or absence (Saphonecrus)
of lateral frontal carinae. Six of the species
addressed in our study (S. cibriani, S. citriformis
(= S. elegans), S. laticephalus, S. longimalaris,
S. longiscapus and S. striatifrons) do not have
frontal carinae; nevertheless, all of them have the
radial cell of forewings closed (according to the
same study, always open in Saphonecrus and
usually closed in Synergus) and female antenna
with 14 segments (usually with 13 segments in
Saphonecrus and 14 segments in Synergus),
so we do not doubt that they all belong to the
genus Synergus. This is yet more evidence
that Saphonecrus would be a polyphyletic or
paraphyletic genus closely related to Synergus, as
several authors have been suggested (Pénzes et
al. 2012; Bozsó et al. 2014 2015).
Neither the new species described in
our study nor their morphologically closest
related species (S. citriformis (= S. elegans), S.
laticephalus and S. punctatus) have pronotal
carinae. This trait is always present in the core or
main group of Palearctic Synergus species and
absent or weak in Synergus plagiotrochi Nieves-
Aldrey and Pujade-Villar, S. variabilis Mayr, S.
flavipes Hartig, S. acsi Melika and Pujade-Villar,
and S. consobrinus Giraud in Houard, which
are considered the most basal clades within
the Palearctic Synergus close to the genera
Saphonecrus and Synophrus (Ács et al. 2010;
Nieves-Aldrey and Medianero 2011). Like all the
Neotropical species studied by Nieves-Aldrey and
Medianero (2011), Synergus species described
from Mexico to date (see Pujade-Villar et al.
2016) and several species from the USA (like S.
punctatus and S. citriformis (= S. elegans)) lack
pronotal carinae. As Nieves-Aldrey and Medianero
(2011) suggested, the absence of lateral pronotal
carinae could be a plesiomorphic trait state in the
Palearctic Synergus shared with the Neotropical
species; so, it might be shared with some Nearctic
species too.
Before this study, there were 18 recorded
species with partially smooth mesopleurae in
America. After the publication of this study, the
total number of species with partially smooth
mesopleurae reaches 22 (since S. elegans is now
synonymous with S. citriformis). This trait seems
to be quite frequent among American Synergus
species from both Nearctic and Neotropical
regions; in contrast, it has not been observed
among Eastern nor Western Palearctic species,
which always have totally striated mesopleurae
(see Pujade-Villar et al. 2014; Schwéger et al.
2015a).
Both the absence of frontal carinae and
pronotal carinae, as well as the absence of striae
covering all the mesopleuron in some American
species, constitute further evidence that both
Nearctic and Neotropical species of Synergus
show important morphological differences with
respect to the Palearctic ones. Thus, it will
be necessary to include species from these
biogeographical regions in future analyses to
discuss the global monophyletism of the genus
Synergus.
In this study, only one species described
north of Mexico that belongs to this group (S.
punctatus) has been included in the key due to
its great morphological resemblance to one of the
newly described species: S. gilletti. According to
their original descriptions, the rest of the species
with partially smooth mesopleurae described
north of Mexico (Synergus agrifoliae Ashmead
(= Synergus maculatus Fullaway, = Synergus
obscurus McCracken and Egbert); Synergus
campanula Osten Sacken; Synergus castanopsidis
(Beutenmüller); Synergus confertus McCracken
and Egbert; Synergus flavens McCracken and
Egbert; Synergus laeviventris (Osten Sacken);
Synergus mendax Walsh; Synergus multiplicatus
Fullaway; Synergus niger Fullaway; Synergus
pacificus McCracken and Egbert; Synergus
profusus McCracken and Egbert; Synergus stelluli
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© 2017 Academia Sinica, Taiwan
Burnett; Synergus varicolor Fullaway; Synergus
variegatus McCracken and Egbert and Synergus
walshii Gillette (= Synergus albipes (Walsh))
can be easily separated from all of the species
addressed in this study by the following traits:
Synergus agrifoliae differs from S. cibriani,
S. citriformis (= S. elegans), S. laticephalus, S.
longimalaris, S. longiscapus and S. striatifrons by
having the mesoscutum finely carinated and the
scutellum rugose (mesoscutum and scutellum
nely coriaceous in these species). It differs from
S. gilletti and S. punctatus by the presence of
punctures in the head (no punctures in these
species) and by the absence of metasomal
micropunctures extending laterally (with
micropunctures extending 1/3-1/2 of metasomal
length in these species).
Synergus campanula differs from all the
species addressed in this study by having the
mesoescutum finely and densely carinated, the
scutellum densely and deeply rugose and the
metasoma as long as broad (mesoscutum and
scutellum nely coriaceous and metasoma longer
than broad in all the species addressed here). It
also differs from S. gilletti and S. punctatus by the
absence of metasomal micropunctures extending
laterally (with micropunctures extending 1/3-1/2 of
metasomal length in these species).
Synergus castanopsidis differs from all the
species addressed in this study by having the
metasoma dorsally incised (not incised in none
of the species addressed here). It also differs
from S. cibriani, S. citriformis (= S. elegans),
S. laticephalus, S. longimalaris, S. longiscapus
and S. striatifrons by having frontal carinae, the
mesoescutum with weak carinae and the scutellum
rugose (no frontal carinae and mesoscutum and
scutellum nely coriaceous in these species).
Synergus confertus, S. avens, S. pacicus,
S. profusus and S. varicolor mainly differ from
S. cibriani, S. citriformis (= S. elegans), S.
laticephalus, S. longimalaris, S. longiscapus and
S. striatifrons by having the mesoscutum carinated
and the scutellum rugose (mesoscutum and
scutellum nely coriaceous in these species), and
from S. gilletti and S. punctatus by the absence of
metasomal micropunctures extending laterally (with
micropunctures extending 1/3-1/2 of metasomal
length in these species).
Synergus mendax differs from all the species
addressed in this study by having the mesoscutum
with coarse transverse waving carinae and
antennae nearly as long as the body (mesoscutum
finely coriaceous and antennae not as long in all
the species addressed here).
Synergus multiplicatus differs from all the
species addressed in this study by having the
head rugose-punctate, and Synergus stelluli,
by having the vertex wrinkled in the ocellar
area (head coriaceous without rugae in all the
species addressed here); moreover, S. stelluli
has frontal carinae (absent in most of the species
mentioned in this study) and a postero-dorsal
patch of micropunctures in the metasoma (with
micropunctures extending laterally in S. gilletti and
S. punctatus).
Synergus niger mainly differs from S. cibriani,
S. citriformis (= S. elegans), S. laticephalus, S.
longimalaris, S. longiscapus and S. striatifrons
by having the scutellum deeply rugose and the
scutellar foveae large (scutellum nely coriaceous
and with inconspicuous foveae in all these
species), and from S. gilletti and S. punctatus
by having the mesoscutum smooth and shining,
coriaceous-reticulate (nely carinated, not shining,
in these species).
Synergus variegatus mainly differs from
S. cibriani, S. elegans, S. laticephalus, S.
longimalaris, S. longiscapus and S. striatifrons
by having the mesoscutum finely transversely
carinated and the scutellum rugose (mesoscutum
and scutellum nely coriaceous in these species),
and from S. gilletti and S. punctatus by its lighter
body coloration (head and body mainly black
in these two species); moreover, this species
has notauli incomplete (more or less visible but
complete in all the species addressed in this
study).
Finally, Synergus walshii and S. laeviventris
differ from all the species addressed in this
study by having the mesoscutum finely rugose-
punctate (with a different sculpture in all the
species addressed here) and from S. gilletti
and S. punctatus by the absence of metasomal
micropunctures extending laterally (with
micropunctures extending 1/3-1/2 of metasomal
length in these species). Synergus walshii also has
the metasoma as long as broad (longer than broad
in all the species addressed in this study).
Mexico is an area of Quercus diversication,
since between 135 and 150 of the 450 worldwide
estimated species of this genus are in this country
(Pujade-Villar et al. 2009) and 86 of them are
considered endemic (Nixon 1998; Zavala 1998).
Of them, eight oaks with gall wasps belong to the
Lobatae section, 23 to the Quercus section and
two to the Protobalanus section.
It is generally assumed that there is a positive
page 25 of 28Zoological Studies 56: 36 (2017)

© 2017 Academia Sinica, Taiwan
correlation between the number of Quercus
species and their associated host gall-inducers
and inquiline fauna. According to this, the number
of inquiline species from Mexico is poorly known,
given that only nine species, all of them belonging
to the genus Synergus, are currently known in front
of the 135-150 estimated species of Quercus. The
main reason for this is the little attention paid to
this fauna by different researchers (Pujade-Villar et
al. 2008). In this sense, for example, Alfred Kinsey
(Hoboken, Nova Jersey 1894 - Bloomington,
Indiana 1956) described more than 130 gall wasps
in a few years after several biological expeditions
in Mexico and Central America (1931-32 and 1935-
36), but none of them were inquilines (Pujade-Villar
et al. 2009). In contrast, 11 inquiline species have
been described from Panama (1 Agastoroxenia
and 10 Synergus) in front of nine species of
Quercus; 75 species of inquiline gall wasps are
known from North America (17 Ceroptres, 54
Synergus and 4 Saphonecrus) in front of 90
species of Quercus; nally, 45 species of inquiline
gall wasps are known from Europe (2 Ceroptres,
34 Synergus, 6 Saphonecus and 3 Synophrus) in
front of 14 species of Quercus (Tutin et al. 1993;
Melika 2006; Nixon 2008; Nieves-Aldrey and
Medianero 2011; Pénzes et al. 2012).
Mexico harbors a huge biodiversity
of inquilines (Lobato-Vila and Pujade-Villar
unpublished data), so the species addressed in
this study represent only a fraction of the actual
Mexican inquiline biodiversity. In this sense, much
more samplings and use of sequencing DNA
techniques (see techniques used by Schwéger
et al. (2015b) with cynipids from the Eastern
Palearctic) will be required to achieve a deeper
understanding of this fauna in this country,
and especially because Mexico includes two
biogeographic areas: Nearctic and Neotropical.
Acknowledgments: This work and the five new
species names have been registered with ZooBank
under: urn:lsid:zoobank.org:pub:DAB4D865-5183-
4318-ADD6-80A7B7E09F66. We want to express
our thanks to Matt Bufngton (NMNH, Washington
D.C., USA) for sending us the type material and
additional material for Synergus punctatus and
Synergus citriformis; to Mercedes Paris (MNCNM,
Madrid, Spain) and Marcos Roca-Cusachs
(Universitat de Barcelona, Barcelona, Spain), for
sending and delivering us the type material and
additional material for Synergus elegans; also
to Marcos Roca-Cusachs, for taking the digital
picture of the head in frontal view of Synergus
gilletti; to Olivia Cabral-Gamboa and Rosa Treto-
Pereyra (Universidad Autónoma de Zacatecas),
Luis G. Landa-Orozco and Carlos Carrillo-Sánchez
(Centro de Biodiversidad, Monte Escobedo,
Zacatecas), David Cibrián (Universidad Autónoma
de Chapingo, Texcoco), Edith G. Estrada-Venegas,
Armando Equihua-Martínez and Rosa D. García-
Martiñón (Colegio de Postgraduados, Texcoco),
and Alondra G. Pérez-García (Facultad de
Ingeniería Agrohidráulica-Ingeniería Agroforestal,
Tetela de Ocampo, Puebla), for the collection of
the galls from which we obtained the material
mentioned in this work; to Jesús Alberto Acuña-
Soto (Fitosanidad, Entomología y Acarología,
Colegio de Postgraduados, Campus Montecillo,
México), for the determination of Quercus species;
finally, to Miguel A. Alonso-Zarazaga (MNCNM,
Madrid, Spain), for helping us to apply the correct
zoological nomenclature and etymology for the
new species’ names.
Authors’ contributions: Both authors analyzed all
the specimens addressed in this study, identified
the new species, decided the new synonymy,
structured the content of the manuscript and took
the electron microscope images. IL-V wrote the
manuscript, described the species and made the
morphological key. Both authors participated in
revising the manuscript. Both authors read and
approved the nal manuscript.
Competing interests: IL-V and JP-V declare that
they have no conict of interest.
Availability of data and materials: This work
and the five new species name have been
registered with ZooBank under: urn:lsid:zoobank.
org:pub:DAB4D865-5183-4318-ADD6-
80A7B7E09F66. Materials addressed in this study
are deposited in collections of several research
institutions, as stated in the manuscript.
Consent for publication: Not applicable.
Ethics approval consent to participate: Not
applicable.
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