JHR 37: 77-111 (2014) rte JOURNAL OF | *0eerrrieved opencoss ural
Steen (G-) Hymenoptera

www.pensoft.net/journals/jhr The iternatonl Scie of Hymenopeariss, RESEARCH

The maxillo-labial complex of Sparasion
(Hymenoptera, Platygastroidea)

Ovidiu Alin Popovici', Istvan Mik6?, Katja C. Seltmann?, Andrew R. Deans?”

| University ‘Al. I. Cuza” Iasi, Faculty of Biology, B-dul Carol [, no. 11, RO— 700506; Romania 2 Pennsylvania
State University, Department of Entomology, 501 ASI Building, University Park, PA 16802 USA 3 Department
of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, New
York 10024, USA

Corresponding author: Jstvdn Miké (istvan.miko@gmail.com)

Academic editor: /. Yoder | Received 25 November 2013 | Accepted 28 February 2014 | Published 28 March 2014

Citation: Popovici OA, Miké I, Seltmann CK, Deans AR (2014) The maxillo-labial complex of Sparasion (Hymenoptera,
Platygastroidea). Journal of Hymenoptera Research 37: 77-111. doi: 10.3897/JHR.37.5206

Abstract

Hymenopterans have evolved a rich array of morphological diversity within the maxillo-labial complex.
Although the character system has been extensively studied and its phylogenetic implications revealed in
large hymenopterans, e.g. in Aculeata, it remains comparatively understudied in parasitoid wasps. Reduc-
tions of character systems due to the small body size in microhymenoptera make it difficult to establish
homology and limits the interoperability of morphological data. We describe here the maxillo-labial com-
plex of an ancestral platygastroid lineage, Sparasion, and provide an ontology-based model of the anatomi-
cal concepts related to the maxillo-labial complex (MLC) of Hymenoptera. The possible functions and
putative evolutionary relevance of some anatomical structures of the MLC in Sparasion are discussed.

Anatomical structures are visualized with Confocal Laser Scanning Microscopy.

Keywords
Mouthparts, Hymenoptera Anatomy Ontology, Confocal laser scanning microscopy, maxillary palpus,

labial palpus, galea, lacinia, prementum, glossa

Introduction

Despite relatively recent efforts (e.g., Austin and Field 1997, Murphy et al. 2007), the
phylogeny of Platygastroidea remains largely unresolved, as such insights from previ-
ously unexplored character systems have the potential to make important contributions
to our understanding of the groups evolution. As Platygastroidea are usually reduced

Copyright Ovidiu Alin Popovici et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

78 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

in body size, the number of informative characters is limited to those anatomical com-
plexes that cannot be simplified without negatively affecting the wasps’ survival. In
the case of Platygastroidea two character systems seem to match this requirement: the
ovipositor apparatus (Austin 1983, Field and Austin 1994, Austin and Field 1997) and
the mouthparts including the maxillo-labial complex. While the former is crucial for
the piercing the chorion of host egg and laying the wasp’s own eggs, the second struc-
ture is vital to the process of feeding. Very little is known about the feeding behavior
and diet of adult platygastroids. Some species, e.g., the phoretic mantid egg parasitoid
Mantibaria, feed also on their hosts’ body fluids (Clausen 1976).

Due to the extreme diversity of Hymenoptera foraging strategies homologizing
mouthpart structures within the order is a challenging task. Other than palpal formulae
(Kieffer 1926, Masner 1976, Kononova and Kozlov 2001) and a relatively superficial
survey of the maxillo-labial complex (MLC) in some genera of platygastroids (Popovici
and Fusu 2006) dataon MLC morphology in platygastroids have never been published.

Sparasion is a fairly speciose genus, with 141 valid species Johnson 1992, Johnson et
al. 2008). These wasps are widespread in Eurasia, Africa, and temperate North America
(Johnson et al. 2008), and, as far as is known, these species are egg-parasitoids of Tettigo-
niinae (Orthoptera: Tettigoniidae) Johnson et al. 2008). Based on the latest phylogenetic
analyses, Sparasion is part of a small lineage that is sister to either the vast majority of
Scelionidae sensu Haliday 1839 (Murphy et al. 2007) or Platygastridae sensu stricto (Austin
and Field 1997). Species in this lineage typically exhibit ancestral morphological states
(Murphy et al. 2007, Austin and Field 1997), which should facilitate homologization of
mouthparts with other hymenopterans. Sparasion are also relatively large species, with less
anatomical simplification, which makes them more suitable for dissection and observation.

Material and methods

We examined 34 specimens of 10 Sparasion species (Appendix 1). All specimens were
stored in 70% ethanol prior to dissection. Card-mounted voucher specimens are de-
posited in the Insect Collection of University “Al.I].Cuza” Faculty of Biology, Iasi, Ro-
mania (OPPC), in the C. A. Triplehorn Insect Collection, Ohio State University, Co-
lumbus, OH, USA (OSUC) and in the Frost Entomological Museum, Pennsylvania
State University, State College, PA, USA (PSUC).

We followed the protocols of Prinsloo (1980) for specimen preparation. The head
was boiled in 10% phenol in lactic acid for 30 minutes and rinsed in distilled water.
The MLC were separated from the cranium using forceps (Dumont #5) and insect
pins (size 2), then were transferred into 10% NaOH and macerated for 30 minutes,
than transferred and rinsed in glacial acetic acid for 10 minutes. Clear and neutralized
MLC specimens were dehydrated using ethanol series (70%, 96%, and 100% alco-
hol; 30 min each) and transferred to a clove oil droplet on concave microscope slide.
The labium was separated from the right and left maxillae and mounted separately in
Canada balsam medium.

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea) 72

MLC specimens were examined under Euromex GE 3045 microscope (400x—
1000x). Line drawings were made using Reichart drawing tube attached to the same
microscope. Photos were taken using a Leica DFC-500 camera mounted on a Leica M
205A stereomicroscope.

CLSM images were taken on glycerin-stored specimens with Zeiss LSM 710 Con-
focal Microscope. For visualizing anatomical structures we used excitation wavelength
of 488 and emission wavelength of 510-680 nm, detected using two channels visual-
ized separately using two pseudocolors (510-580 nm=green; 580-680 nm=red). For
visualizing resilin we used excitation wavelength of 405 nm and emission wavelength
of 510-680 nm detected using one channel visualized with blue pseudocolor.

For Scanning Electron Microscopy (SEM) specimens were dried using hexam-
ethyldisilazane (HMDS, Brown 1993), mounted on double adhesive tape and coated
with gold. SEM images were taken using VEGA T SCAN SEM.

Anatomical concepts used here are defined and aligned with those of Alam 1951,
Beutel and Vilhelmsen 2007, Bugnion 1924, 1925, Cockerell 1924, Crampton 1923,
Crosskey 1951, Dhillon 1966; Duncan 1939, Eickwort 1969, Forel 1874, Gotwald
1969, Krenn et al. 2005, Liu 1925; Matsuda 1965, Matsuda 1957, McGinley 1980,
Michener 1944, Michener 1984, Plant and Paulus 1987, Prentice 1998, Ross 1937,
Ritchie and Peters 1981, Ronquist and Nordlander 1989, Snodgrass 1942, Ulrich 1924,
Vilhelmsen 1996, Winston 1979 in the Hymenoptera Anatomy Ontology (HAO, Yoder
et al. 2010). Anatomical terms in the Results section are linked to the HAO via the URI
table (Appendix 2; see Seltmann et al. 2012 for more information about this approach).

Results

Maxilla (integument)

The cardo (cd: Fig. 1A—B) is triangular. The submedial maxillary process of the hypos-
toma (hys: Fig 1A) inserts submedially on the proximal part of the cardo (caf: Fig. 1B).
The mediodistal cardinal ridge, laterodistal cardinal ridge and basal cardinal ridge are
present (Icr, mer, ber: Fig. 1D) and the inner and outer cardinal processes are absent.
The cardo lays almost parallel to the external surface of the hypostoma and is largely
obscured by it even if the maxillo-labial complex is fully protracted (Figs 1A, cd: 4A).
The conjunctiva connecting the cardo to the rest of the maxillo-labial complex is resilin
rich along the stipitocardinal hinge (sch: Fig. 1C).

‘The stipes is triangular in cross section distally. The posterior stipital wall of the stipes
bears the posterior stipital sclerite (pss: Figs 1A, C, 2B, C), while the partly sclerotised
medial wall and the convex and membranous anterolateral wall (conj: Fig. 1B) bears the
galeo-lacinial complex (Fig. 2A, B, D; gal-lac: Fig. 1B, C). The posterior stipital sclerite
is triangular in posterior view, and is margined by the principal carina of stipes (pcs: Figs
1A-C, 3A), which is less rigid and melanized than other regions of the sclerite and is the
most developed medially and distolaterally. The principal carina of stipes is equipped

80 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

C

Figure 1. Mouthparts of Sparasion sp. A SEM micrograph showing the mouthparts, posterior (exter-

nal) view, distal to the bottom B CLSM volume rendered image showing the maxillo-labial complex,
anterior (internal) view, distal to the bottom (doi: 10.6084/m9.figshare.861065, doi: 10.6084/m9.figsha-
re.861058) C Line drawing showing the maxillo-labial complex, posterior (external), distal to the bottom
D Bright field image showing the cardo of Sparasion, lateral to the left.

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea) 81

distolaterally with the marginal fringe of the stipes (mfs: Figs 1A—C, 2B, C) composed
of occasionally branched spines. The extent of the principal carina of the stipes on the
distolateral margin is variable in different Sparasion species (compare pcs of Fig. 1A with
that of Fig. 1B) but it always overlaps the proximomedial surface of the mandible (md:
Fig. 1A). The median part of principal carina of the stipes is divided into the proximo-
median stipital flange (pmf: Figs 1B, C, 2A) and the distomedial stipital flange (dmf:
Figs 1C, 2A, B, C). The proximal part of the distomedial stipital flange posteriorly over-
laps the distal part of the proximomedial stipital flange (pmf, dmf: Fig. 1C). The stipes
articulates with the postmentum (psm: Figs 1C, 2C) and the distal prementum (pmn:
Fig. 1A, C) along the proximomedial stipital flange and with the proximal prementum
via the distomedial stipital flange. The medial stipital groove (msg: Figs 1A, 2B) extends
medially along the proximomedial stipital flange and distomedial stipital flange and
accommodates the first sclerite of the maxillary palp (mpalp: Fig. 1A, C, 3D, E) when
it is adpressed against the stipes. The posterior stipital sclerite is glabrous, except a for
few, elongate, mechanosensory hairs (msh: Figs 1C, 2B) just proximal of the base of
maxillary palp and along the distolateral margin of the stipes abutting the hypostoma.
The distal part of the posterior stipital sclerite is equipped with numerous campaniform
sensilla (cps: Fig. 2C), which are visible only with transmitted light.

The galeo-lacinial complex has four sclerites and two marginal lobes. ‘The proxi-
mal, inverted T-shaped lacinial lever (bls: Fig. 2A, B) and the distal, narrow basal galeal
sclerite (bgs: Fig. 2B, D) are situated on the median wall of the stipes and articulate
with the posterior stipital sclerite along the distomedial stipital flange. The lacinial bar
and lacinial comb are absent. ‘The proximolateral galeal sclerite (pgs: Fig. 2A) is situ-
ated in the middle on the lateral wall of stipes and is connected proximally with the
lacinial lever and distally with the basal galeal sclerite. The number of mechanosensory
hairs in the proximolateral galeal setiferous patch (prs: Fig. 2A) is variable in different
Sparasion species (Table 1) and overlaps the distolateral galeal sclerite (dgs: Fig. 2A, B,
D), which traverses the galeo-lacinial complex and is represented on both its medial
and lateral walls (the complex is unilayered at the sclerite). The proximolateral and dis-
tolateral galeal sclerites are connected to each other by the lateral galeal crease. The api-
comedial galeal plate is absent. The number of setae in the distolateral galeal setiferous
patch (dgp: Fig. 2A) is variable in different Sparasion species (Table 1). The proximal
galeal brush is absent. The single coeloconic sensillum of galea (cfs: Fig. 2B, D) is lo-
cated distally on the median surface of the distolateral galeal sclerite. The single lacinial
lobe (IIb: Fig. 2A, B, D) extends anteroproximal whereas the galeal lobe (glb: Fig. 2A,
B, D) distal to the proximal galeal sclerites. The lacinial lobe and the proximal part
of the galeal lobe are covered with short acanthae (ach: Fig. 2D), which comprise the
spiculate patch of the lacinia and the spiculate patch of galea respectively. The galeal
comb, galeal lamina and galeal fringe are absent. The velum (vlm: Fig. 2D) is fringed
distally in some species (vlm: Fig. 2D). The stipital sclerite is absent from Sparasion.

The maxillary palp (1mp: Fig. 2B, D) is connected at the distal apex of the poste-
rior stipital sclerite trough the ventral dististipital process (vdp: Fig. 2B) adjacent to the
basal galeal sclerite. The maxillary palp is composed of five maxillary sclerites among

82 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

pencil AAS Pg La aN . ~<a

Figure 2. Maxillo-labial complex of Sparasion. A, B CLSM volume rendered images showing the max-
illa: A. anterolateral (internal lateral) view (doi: 10.6084/m9.figshare.861060), distal to the left B postero-
medial (external medial) view (doi: 10.6084/m9.figshare.861057), distal to the left C Brightfield image
showing the maxillo-labial complex, posterior (external) view, distal to the bottom D SEM micrograph
showing the maxilla, posteromedial (external-medial) view, distal to the top.

Table |.

Sparasion
sp-1 | sp.2 | sp.3 | sp.4 | sp.5 |sp.6| sp.7 | sp.8 |sp.9
The number of setae in the proximal galeal brush 5-7 5-7 | 6

The number of setae in the distal galeal setiferous patch | 18-19) 18-20 | 18-20] 11-14] 18-20] 17 |27-34| 18-20] 22
The number of styloconic sensilla on the glossa 14-15] 15-16]10-12}10-12} 17 | 23 |18-19/15-16] 10

Characters

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea) 83

which the second sclerite of the maxillary palp is the shortest, and the fifth sclerite of
the maxillary palp is always the longest (Smp: Figs LA,C, 3D, E). The relative width of
the maxillary palpal sclerites varies between Sparasion species and in some cases even
between different sexes (Table 1). Two different setal types can be differentiated on
the maxillary palp, based on their gross morphology. The type 1 seta (ss1: Fig. 3B) is a
uniporous sensillum whereas the type 2 seta (ss2: Fig. 3B) is a longer mechanosensory
hair. A type 1 seta is present on all but the first sclerite of the maxillary palp, which is
glabrous in Sparasion sp. 4 and sp. 9 and bears only 1—2 type 2 setae in the rest of the
species. Type 1 setae are evenly distributed on the third sclerite of the maxillary palp,
on the fourth sclerite of the maxillary palp and on the fifth sclerite of the maxillary
palp. Type 1 setae are located in 1-4 whorls of setae on the second sclerite of the max-
illary palp. Type 2 setae occur only on the third, fourth sclerite of the maxillary palp
and on the fifth sclerite of the maxillary palp. The number of type 2 setae is positively
correlated with the width of the third sclerite of the maxillary palp and the fourth scle-
rite of the maxillary palp: type 2 setae are absent from the third sclerite of the maxillary
palp if it is not increased in width relative to the second maxillary palpal sclerite.

Maxilla (skeletal muscles)

The cranio-cardinal muscle (cr-cd: Figs 1B, 4A, B) arises medially of the posterior site of
origins of the tentorium and inserts on the cardo just laterally of the cranial fossa of the
cardo (caf: Fig. 1B). The tentorio-cardinal muscle (tnt-cd: Fig. 1B) arises ventrally on the
tentorium just laterally of the site of origin of the tentorio-stipital muscle (tnt-sti: Fig.
4A) and inserts laterally on the stipitocardinal hinge. The tentorio-stipital muscle arises
ventrally from the tentorium, medially of the site of origin of the tentorio-cardinal
muscle and inserts on the median margin of the posterior stipital sclerite just posterior
of the lacinial lever. The cranio-lacinial muscle is absent. ‘The stipito-lacinial muscle (sti-
lac: Fig. 2A) arises from along the lateral margin of the posterior stipital sclerite and
inserts apically on the lacinial lever. The single stipito-palpal muscle (sti-mp1: Figs 2A,
3A) and the stipito-galeal muscle (sti-gal: Fig. 3A) arise medially from the stipito-lacinial
muscle, subsequently. The stipito-galeal muscle inserts on the basal galeal sclerite. The
first intrinsic muscle of the maxillary palp, third intrinsic muscle of the maxillary palp and
fourth intrinsic muscle of the maxillary palp are present, the second intrinsic muscle of the
maxillary palp was not observed. No muscle was attached to the fifth maxillary sclerite.

Labium, distal hypopharynx (integument)

The postmentum is composed of a single interstipital sclerite that is elongate, rectangular,
and is articulated with the posterior stipital sclerites proximolaterally at the proximomedi-
al stipital flange (psm, pmf: Figs 1B, C, 2C). The median postmental ridge (mpr: Figs 1A,
B, 2C) is present and the postmental-premental hinge (pph: Figs 1C, 2C) is resilin rich.

84 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

Figure 3. Maxilla and labium of Sparasion. A SEM micrograph showing the skeletomuscular system of
the maxilla, posterior (internal) view, distal to the top B SEM micrograph showing the maxillary palp,
distal to the top C Bright field image showing the labium, lateral view, distal to the left D, E Bright field
image showing the maxillary palps, distal to the left.

The prementum is connected to the stipes via a conjunctiva (stipito-premental con-
junctiva) extending along the proximal margin of the premental arms (prm: Figs 1B, 3C,
4A, B, 5C, 6A—D) and the lateral margin of the prementum proximally of the arm. The

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea) 85

100 ym

Figure 4. CLSM volume rendered images showing the skeletomuscular system of the labium of Spara-
sion, medial view, distal to the left.

premental carinae (pmc: Fig. 2C and corresponding premental ditches (pmd: Figs 2C,
4A, B, 5C, D, 6A, D) converge proximally and separate the ventral premental face from
(vpf: Figs 2C, 3C, 4A, B, 5C, D, 6A, D) the lateral premental face (pma: Figs 2C, 4A, B,

86 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

oo
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7
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u:
ag

fete Pee oe

48 Pa

os

a . y 7
4 =a ‘
“ath, ; ae '. +e Py £
Z a

Wen AA Le

Figure 5. Labium and distal hypopharynx of Sparasion. A SEM micrograph showing the labium and
distal hypopharynx, anterolateral view, distal to the left B SEM micrograph showing the glossal annuli,
anterior view, distal to the left C, D CLSM volume rendered images showing the labium with retracted
glossa and paraglossae, distal to the left C lateral view (doi: 10.6084/m9.figshare.861064) D medial view
(doi: 10.6084/m9.figshare.861061).

5C, D, 6A, D). The ventral premental face is flat, diamond-shaped and is equipped with
campaniform sensillae of the prementum (cps: Fig. 2C) and distally-oriented mecha-
nosensory hairs (msh: Fig. 2C). The number and pattern of both the campaniform sen-
silla and mechanosensory hairs and the length of the mechanosensory hairs are variable
between different species and sexes (Table 1). The lateral face of the prementum is mostly
overlapped ventrally by the distomedial stipital flange (Fig. 1A). The labial palpal excision

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea) 87

100 um

i 7 Ve oo Oe ema el

Figure 6. Labium of Sparasion. A CLSM volume rendered images showing the labium with protracted
glossa and paraglossae, lateral view, distal to the left (doi: 10.6084/m9.figshare.861063) B CLSM vol-
ume rendered images showing the labium, glossa removed, lateral view, distal to the left (doi: 10.6084/
m9.figshare.861062, doi: 10.6084/m9.figshare.861066) C CLSM volume rendered images showing the
labium, glossa removed, posterior view, distal to the left (doi: 10.6084/m9.figshare.861056, doi: 10.6084/
m9.figshare.861067) D CLSM volume rendered images showing the labium with protracted glossa and
paraglossae, medial view, distal to the left (doi: 10.6084/m9.figshare.861059) E, F Bright field images

showing the labium, posterior view, distal to the top.

88 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

(pin: Figs 2C, 3C) accommodating the base of the labial palp is distinct on the disto-
lateral corner of the prementum. The labial palp is composed of three sclerites that are
equipped with both the uniporous sensilla (type 1) and mechanosensory hairs (type 2).

The glossa (glo: Figs 1C, 3C, 4A, B, 5C, GA) is separated anteroproximally from
the paraglossae (pgl: Figs 5C, 6A, C) and the distal hypopharynx by the basal glos-
sal invagination (inv: Figs 4A, 5D, 6D) adjacent to the salivarium (svr: 6B, C, F).
The anterior glossal sclerites (ags: Figs 3C, 4A, B, 5C, 6A, D) are embedded into the
distal wall of the invagination and are connected medially to the ventral wall of the
salivarium via a ligament (lig: Figs 4A, B, 5D, 6D). The ventrolateral basiglossal arms
are distinct (vba: Fig. GA, F) and each anterior glossal sclerite is equipped with a sin-
gle campaniform sensillum of the glossa (csg: Fig. 6F). The posterior glossal plate is
composed of a single, median sclerite (pgp: Figs 4A, B, 5D, 6E, F). The flat posterior
surface of the glossa (pwg: Figs 3C, 4A, B, 5D, 6D) is glabrous with distinct ventral
glossal lines. The distal glossal edge (dge: Figs 3C, 4A, B, 5A, D, 6D) is thicker than
the proximal region of the posterior surface of the glossa. The apical glossal hairs are
usually present (agh: Fig. 6A). The apical glossal setae (sts: Figs 5D, 6D, E) are stylo-
conic sensilla with variable number in different species (Table 1).

The arched anterior surface of the glossa (awg: Figs 4A, B, 5A, D, 6D) is evenly
covered with glossal annuli (gla: Fig. 5A), which are composed of transverse rows of
spatulate and anteriorly curving annular hairs, each with a dentate distal margin (anh:
Fig. 5B).

The bilobed paraglossae arise proximolaterally of the basal glossal invagination
and encircle the proximal part of the glossa. The elongate, triangular basiparaglossal
sclerite (pls: Figs 3C, 5C, GA—D, F) corresponds to a posterior lobe of the paraglos-
sa whereas the larger anterior lobe of the paraglossa bears the distal, less sclerotised
paraglossal acroglossal button (pgb: Fig. 6B). Proximally the paraglossa is continuous
with the wall of the distal hypopharynx (dhy: Fig. 5A, C) and the paraglossal sclerite
is connected to the distal hypopharyngeal sclerite via an elongate ligament. ‘The ba-
siparaglossal brush (bgb: Figs 4B, 6A, B, F) is distinct but the paraglossal sclerite and
paraglossal annuli are absent.

The distal hypopharynx is supported laterally by the premental arms and the hy-
popharyngeal rod (hypr: Fig. 3C), which is continuous with the distal hypopharyn-
geal sclerite via the hypopharyngeal button (hyb: Figs 3C, 5C, 6A—D). The hy-
popharyngeal pecten (hpc: Figs 4B, 5C, 6A, B) is distinct and abuts the basiparaglos-
sal brush when the ligula is retracted (hpc: Figs 5C). The hypopharyngeal buttons
are connected to each other medially trough the ventral wall of the salivarium, that
is either sclerotised (svr: Figs 3C, 6B; salivarial sclerite) or is a resilin rich conjunc-
tiva (svr: Fig. GF). In the former case the hypopharyngeal buttons and the ventral,
sclerotised wall of the salivarium compose the distal hypopharyngeal sclerite (dhs:
Figs 4B, GC) encompassing ventrally the salivary duct (svd: Figs 3C, 4A). The distal
hypopharyngeal invagination (dhi: Figs 4A, 5C, D) corresponds to the dorsal bend
on the hypopharyngeal button.

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea) 89

Labium, distal hypopharynx (skeletal muscles)

The postmento-premental muscle is absent. ‘The posterior tentorio-premental muscle (tntp-
pmn: Fig. 4A, B) arises from the tentorium anterolaterally from the site of origin of the
anterior tentorio-premental muscle (tnta-pmn: Fig. 4A, B) and inserts on the proximal
part of the hypopharyngeal button. The anterior tentorio-premental muscle arises from
the tentorium medially of the site of origin of the posterior tentorio-premental muscle
and inserts proximomedially on the postmental-premental hinge. The premento-para-
glossal muscle (pmn-pgs: Fig. 4A, B) arises medially from the internal edge corresponding
to the premental ditch and from the intima of the median premental face anterior to
the edge and inserts on the posterior glossal plate. The dorsal premento-salivarial muscle
(pmnd-slv: Fig. 4A, B) arises distomedially from the premental arm and inserts dorsally
on the salivarium. The ventral premento-salivary sclerite muscle (pmnv-slv: Figs 4A, B)
arises from the proximal premental area and inserts proximomedially on the distal hy-
popharyngeal sclerite. The premento-palpal muscle arises anterolaterally from the site of
origin of the premento-paraglossal muscle. The first intrinsic muscle of the labial palp and
the second intrinsic muscle of the labial palp are present. The premento-glossal muscle is
apparently absent.

Discussion

The maxillo-labial complex of Sparasion

The term maxillo-labial complex (MLC) is used by Snodgrass (1935). However, many
entomologists prefer to use the term labio-maxillary complex, e.g. Duncan (1939),
Labandeira (1997), Jervis (1998), Vilhelmsen (1996), Krenn (2007). The reason for
using the latter seems to be that it is more easily pronounced (Duncan 1939). In this
paper the term maxillo-labial complex is preferred because it corresponds with the
anatomical position of the component pieces.

The hymenopteran labium, unlike that of other holometabolan insects, is so tight-
ly connected to the maxillae by the labiomaxillary hinge that they protract and retract
together as a single unit (Duncan 1939, Snodgrass 1942, Vilhelmsen 1996, Krenn
2007). While the labiomaxillar hinge connects their proximal parts, the distal, more
sclerotised regions of the maxillae and the labium still are able to move with some free-
dom in basal Hymenoptera (Vilhelmsen 1996, Beutel and Vilhelmsen 2007) and ina
few apocritan taxa (e.g. in Gasteruption, Evania (Evanioidea) (Fig. 8) and Gryon (Plat-
ygastroidea) Miko, pers. obs.). However, in Sparasion and some other apocritan taxa
(Ibalia (Cynipoidea) Ronquist & Nordlander, 1989, Vespula (Vespoidea) Duncan,
1939) where the posterior wall of the labium and maxillaarealmost exclusively sclerotised,
the freedom of the independent movement between the appendages is restricted:
they can not be separated from each other and sometimes — e.g. in Sparasion

90 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

— not even from the hypostoma. The development of rigid connections between
sclerites that are flexibly connected in basal Hymenoptera seems to be an apocritan
evolutionary trend, which is arguably related to their more sclerotised body (Vilhelm-
sen 2000a, 2000b, Vilhelmsen et al. 2010).

Although the MLC of Sparasion is reduced in size and highly sclerotized, we were
able to homologize most of its anatomical structures to those of other Hymenoptera
(Appendix 2, Figs 7-9). We hypothesize that the platygastroid MLC is an anatomical
system with enough phenotypic diversity to serve as a source of morphological charac-
ters for phylogenetic analyses and species diagnosis.

Maxilla

One consequence of the more heavily sclerotised and less moveable MLC is that the
cardo remains relatively simple in Spavasion, retaining its main function: providing
rigid attachment of the MLC to the cranium. The inner and outer processes of the
cardo (Winston 1979) that articulate with the stipes are missing from Sparasion,
similarly to other Proctotrupomorpha (e.g., /balia and Pelecinus pers. obs.). Because
the stipito-cardinal articulation is absent, the cardo seemingly interacts with the
posterior stipital sclerite only by the resilin rich stipito-cardinal hinge. Similarly to
Sparasion, the cardo is not visible externally in these taxa and even if the MLC is in
a protracted position, the cardo is obscured by the proximal portion of the principal
carina of the stipes.

Different levels of sclerotization can be observed on the intercardinal and interstipi-
tal areas in different hymenopteran taxa. Only the intercardinal area contains a sclerite
in basal Hymenoptera (Ross 1937), while both intercardinal or interstipital sclerites
may be present in Apocrita. The intercardinal and interstipital areas are separated from
each other by the labial suture (Snodgrass 1935, Prentice 1998). The median area of the
MLC posterior to the suture contains an anterior and a posterior sclerite in some holo-
metabolan insects (i.e. Coleoptera), the submentum and the mentum. ‘This condition
never occurs in Hymenoptera (see references listed in materials and methods section).
Even in those Aculeata where two medial sclerites are in the postmental area, one is al-
ways anterior to the labial suture. Based on the their position relative to the stipites and
cardines, the intercardinal postmentum of basal Hymenoptera might be homologous
with the postmentum in other insects and the interstipital postmentum of Apocrita is
most likely not homologous to these structures. The postmental area is fully membra-
nous in many Apocrita (Evania, psm: Fig. 8; Vespula, Duncan 1939; Stenobracon, Alam
1951). Only the intercardinal sclerite is present in /balia (Ronquist and Nordlander
1989) leading Ritchie and Peters (1981) to homologize this structure with the mentum
of other holometabolans.

Only the interstipital postmentum (Prentice 1998) is present in Sparasion. This
sclerite is not connected with muscles, nor is it articulated with any other sclerites. It’s
only function might be to separate the posterior stipital sclerites from each other.

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea)

Figure 7. CLSM volume rendered images showing the MLC of Athalia rosae. A maxilla, medial view,
doi: 10.6084/m9.figshare.956279 B MLC, anterior view, distal to the left, doi: 10.6084/m9.figsha
te.956279

The development of the principal carina arguably correlates with the presence of
rigid connection between the stipes and neighboring sclerites. In those taxa, where the

stipes articulates with the hypostoma, but not with the labium, the principal carina is

present only laterally (Evania, pcs: Fig. 8).

92 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

Figure 8. CLSM volume rendered image showing the mouthparts of Evania sp, posterior view, distal to
the bottom, doi: 10.6084/m9.figshare.956280

Both the medial and lateral portions of the carina are well developed in Sparasion,
having two unique characteristics that were not found in other Hymenoptera (see ref-
erences listed in the materials and methods section):

1. ‘The median portion of the carina is divided into two sections that overlap each other
and accommodate the proximal and distal part of the lateral premental margin.

2. ‘The presence of an “apical fringe”, a branched flattened evagination along the
distolateral margin of the carina. While the first specialization might be the con-
sequence of the articulation between the posterior stipital sclerite and the pre-
mentum, the second is most probably related to the unique movement of the
mandible in Sparasion. While pivotal axis of the mandible is anteroposterior in
most Hymenoptera taxa (dorsoventral in a prognathous head) it is directed medi-

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea)

stip-mp1

Figure 9. CLSM volume rendered image showing the maxilla of Orthogonalys pulchella, posteromedial
view, distal to the top-left, doi: 10.6084/m9.figshare.956281

olaterally in Sparasion and the mandibles are moved parallel to the body axis (Mik6
et al. 2007). Due to this unique, oblique position of the mandible, the distolateral,
fringed margin of the principal carina is positioned inside the internal concavity
of the mandible and seemingly acts as a cleaning “brush”. Similar movement of

the mandible has been reported from many other Hymenoptera taxa, including

94 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

two platygastrid genera, Tyrannoscelio and Encyrtocelio. It would be worthwhile
to study the MLC of these taxa and to clarify whether the fringed lateral portion
of the principal carina is unique for Sparasion or whether it can be found in other
genera with modified mandible movement.

The presence of the campaniform sensillum of the stipes was considered as one of
the synapomorphies of the clade composed of Platygastroidea and Cynipoidea (Shar-
key et al. 2012). Although the sensillum is present in Sparasion and some other plat-
ygastrid taxa (Sharkey et al. 2012), it is absent from T7iteleia, Calliscelio, Macroteleia,
Apegus, Duta, Psilanteris, Anteris (Popovici and Fusu 2006), which implies that this
character might be useful for generic classification in Platygastroidea.

Prentice (1998) reported the presence of two apical sclerites on the galea, the apico-
lateral and apicomedial stipital plates. We were able to locate only one apical sclerite, the
distolateral galeal sclerite on the galeo-lacinial complex of Sparasion. This sclerite traverses
the complex and bears both the distolateral galeal setiferous patch and the medial coelo-
conic sensillum that define the apicolateral and apicomedial stipital plates respectively.

Appendage segments and annuli are ring-like and repetitive sclerites of the legs,
antenna, labial palps and maxillary palps. While appendage segments have muscles at-
taching to them, annuli do not. Annuli are traditionally differentiated from appendage
segments by names with the sufhx “-mere” e.g. flagellomere and tarsomere. Among
the five maxillary palpal sclerites of Sparasion the fifth one does not have muscle at-
tachments. We observed a similar condition in Orthogonalys (Trigonalidae), where the
fifth and sixth sclerites lack any muscle attachment (Fig. 9). If we apply the annuli vs.
segment terminology system to these taxa, we should call the first four sclerites “palpal
segments” and fifth sclerite in Sparasion and the fifth and sixth sclerites in Orthogonalys
“palpomeres”. To avoid misinterpretations of the segment identity of the palpi, and
keeping a simple and easily applicable terminology we prefer to use palpal sclerites for
the ring-like sclerites of the palpi.

Sparasion has five maxillary palpal sclerites that is the highest in Platygastroidea
(Masner 1976). Since the presence of five maxillary palpal sclerites is a possible syna-
pomorphy for Proctotrupomorpha (Sharkey et al. 2012), and is also shared by other,
putatively basal lineages of Platygastroidea (i.e. Archaeoteleia, Nixonia and Plauman-
nion, Johnson et al. 2008) it is most probable that this condition is plesiomorphic for
Platygastroidea.

The maxillary palp is located medially on the lateral margin of the posterior stipital
sclerite in basal Hymenoptera distantly from the apical end of the sclerite bearing the
base of the galea (Fig. 7B., Vilhelmsen 1996, Beutel and Vilhelmsen 2007). In Sparasion
the palp is connected to the apical vertex of the posterior stipital sclerite, adjacent to the
base of the galeo-lacinial complex. This distal position makes it difficult to interpret the
presence of minute anatomical structures such as the palpifer (“flexible and transparent,
very difficult to detect structure” at the base of the maxillary palp, Prentice 1989).

The 2™4, 3 and 4" maxillary palpal sclerites are distinctly wider than the more
proximal or distal maxillary palpal sclerites in some Sparasion species (Figures 3D, E).

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea) 95

The number and position of these modified sclerites are sexually dimorphic in some
Sparasion species and are seemingly useful for species groups definitions. A widened
4th palpal sclerite was found in most species of the plesiomorphic platygastrid genus
Nixonia, and representatives of Plaumannion, Archaeoteleia, Sceliomorpha and Neuro-
scelio (Johnson and Masner 2006, Johnson et al. 2008); the presence of enlarged 3"
maxillary palpal sclerites have been reported from Sparasion (Johnson et al. 2008) and
enlarged 2™ and 3" sclerites from the putatively most primitive extant platygastroid
genus, Huddlestonium (Masner et al. 2007).

Although the position of enlarged palpal sclerites in the maxillary palp has often
been used for the classification of different Hymenoptera taxa (i.e. Evaniidae, Deans
and Huben 2003; Aculeata, Bohart and Menke 1976), our knowledge about their
possible function is rather incomplete. Albeit our study confirms Prentice’s (1998) ob-
servation that enlarged sclerite size corresponds to larger muscle mass, the role of this
modification in the mechanics of palpal movement is still unrevealed.

Although the presence of only two stipito-palpal muscles is considered as the Hy-
menoptera groundplan (stia-mp1, stip-mp1: Fig. 9; Beutel and Vilhelmsen 2007),
similarly to Sparasion, only one maxillary plapal muscle has been reported from many
Hymenoptera taxa (Snodgrass 1942, Matsuda 1957, Prentice 1998). The presence of
resilin-rich conjunctiva at the base of the maxillary palpus in Sparasion supports Pren-
tice’s hypothesis that the expansion of the maxillary palpus is facilitated by a resilin
rich region between the posterior stipital sclerite and the palpus in taxa with a single
stipito-palpal muscle.

The terms, galea and lacinia refer usually to the evaginated distal, sclerotised regions
of the maxillar cuticle that are adjacent to the sites of insertions for the stipito-lacinial,
cranio-lacinal and the stipito-galeal muscles in insects. Although these muscles are pre-
sent in most hymenopterans (sti-gal, sti-lac: Fig. 7A, and more proximal and more distal
lobes can almost always be differentiated, the proximal limits of the galea and lacinia are
difficult to define and thus these structures are difficult to homologize to that of other
insects where the galea and lacinia are well sclerotised. Therefore we preferred to use
galeal lobe and lacinial lobe instead of galea and lacinia in the present paper.

Two distinct setiferous areas can be defined on the lateral area of the galeal lobe
in Athalia: a proximal row of setae traversing the galea (prs: Fig. 7B) and a more distal
setiferous patch (dsp: Fig. 7B). The presence of these two setiferous areas seems to be
consistent within Hymenoptera (Prentice 1998) and can be used for the delimitation
of areas of the galeo-lacinial complex.

In Sparasion, similarly to some other, more derived Hymenoptera taxa these set-
iferous areas correspond to two sclerites, the proximolateral and distolateral sclerites
of the galea (Prentice 1998, Ronquist and Nordlander 1989). ‘Two distal sclerites are
located on the medial wall of the galeo-lacinial complex in Azhalia (pgs, dgs: Fig. 7A),
with some campaniform sensillae (cfs: Fig. 7A). Based on the presence of the sensilla
on the medial wall of the distal galeal sclerite of Sparasion it is possible that the distal
galeal sclerite, or at least its medial part (the sclerite traverses the galeo-lacinial com-
plex) is homologous with one of the apical galeal sclerites of Athalia.

96 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

Although the velum is well developed in numerous other Hymenoptera (Prentice
1998), we are not aware of any case in the order where the lobe is fringed apically as it
was found in some Sparasion species (vlm: Fig. 2D).

The galeal comb and the galeal lamina are important characters that have been
used in aculeate systematics (Prentice 1998, “comb of galea” in Michener 1944, “Bor-
stenkamm” in Ulrich 1924). Although neither of these structures are present in Spara-
sion, they are present in some other scelionines (Encyrtoscelio, Teleas, Trimorus Popo-
vici pers. obs.).

Labium

The prementum is articulated with the stipes via the premento-stipital articulation
that is composed of the premental ditch, premental carinae, and the principal carina
of the stipes. Duncan (1939) reported a similar connection between the stipes and the
prementum and considered its importance in the simultaneous movement of these
two structures. Although we are unaware of other hymenopterans with a complete pre-
mento-stipital articulation, the premental carinae are present in some Evaniidae (pmc:
Fig. 8) and in some platygastroid genera (illustrations in Popovici and Fusu 2006).

Vilhelmsen (1996) proposed the presence of acanthae on the hypopharyngeal rod
as a possible synapomorphy for Apocrita. In Sparasion and in some other apocritans
(e.g. Vespula, Duncan 1939; in Apoidea, =spiculate patch of the hypopharynx in Pren-
tice 1989, Evania pers. obs.) two regions of the hypopharyngeal rod, the hypopharyn-
geal pecten (hpc: Fig. 10) and the basiparaglossal brush (bgb: Fig. 10) are extensively
covered with acanthae. Contraction of the posterior tentorio-premental muscle might
protract, whereas contraction of the ventral premento-salivarial muscle retract the hy-
popharyngeal buttons. The distal hypopharyngeal invagination, which is apparently
unique for Platygastroidea, is defined by the hypopharyngeal button. Protraction and
retraction of the button, therefore, might actually control the orientation of the invagi-
nation. We were not able to locate an infrabuccal pouch (Fig. 10) in Sparasion between
the transverse line connecting the tips of the lateral arms of the prementum and the
sitophore in basal Hymenoptera (Vilhelmsen 1996).

The maximum number of labial palpal sclerites in Apocrita is four (some Formici-
dae (Gotwald, 1969), Braconidae (Belokobylskij, 2006), Vespinae (Duncan, 1939)).
In Sparasion, the labial palp is composed of three sclerites that are moved directly by
muscles. In those Hymenoptera where the labial palp is composed of four sclerites
(Vilhelmsen and Beutel 2007, Gotwald 1969, Belokobylskij and Chen 2006, Duncan
1939) the apical sclerite has no muscles and is considered to be a secondary subdivi-
sion of the apical segment. The presence of four labial palp sclerites is considered a
hymenopteran synapomorphy (Vilhelmsen and Beutel 2007) and the ground plan of
almost all other apocritan superfamilies (Sharkey et al. 2012). The ground plan of a
three-segmented labial palp of Platygastroidea is probably the result of the secondary
loss of the apical palpal sclerite.

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea) 97

NE , — :
2p Jp tabuccal
= pouch

labrumy y
f
y

Figure 10. CLSM volume rendered image showing the ventral region of the head of Evania sp., medial
view, distal to the bottom, doi: 10.6084/m9.figshare.956282

Duncan (1939) hypothesized that the glossal annuli have rasping function or are
involved in retaining liquids in Vespinae. Glossal annuli with elongated and well de-
veloped spatulate annular hairs are present in most Apocrita (Duncan 1939, Prentice
1998, Ronquist and Nordlander 1989, Dangerfield et al. 2001, Popovici and Fusu
2006) while are present only in a few basal Hymenoptera taxa (Cephoidea, Megalo-
dontoidea and Orussioidea, Vilhelmsen 1996).

Along the proximal margin of distal edge of glossa in Sparasion, a row of 9-23
apical glossal setae can be observed. ‘These setae are styloconic sensilla, which are con-
sidered to be bimodal contact chemo-mechanosensillae (Shields 2010). Although they
have never been mentioned elsewhere, these styloconic setae are present in other Hy-
menoptera (Evania, sts: Figs 8, 10, Gasteruption, Ibalia, Miko pers. obs.) and might
play crucial role in the chemo- and mechanosensation in Apocrita.

98 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

The ventral glossal lines have been reported only from a few Hymenoptera (Sal-
man 1929, Prentice 1998, Evania, Fig. 8). The lines radiate from the posterior glossal
plate and extend toward the apical glossal hairs and therefore might be related to the
presence of these styloconic sensilla. We have found the lines present in taxa whith
apical styloconic sensilla.

Acknowledgements

We wish to thank Mariana Popovici and Gordon Ramel for gift specimens, Irina Gostin
and Raileanu for the SEM pictures, Eva Johannes (NCSU-NSCORT, Department of
Plant Biology) for her assistance in Confocal Laser Scanning Microscopy, Lubomir Mas-
ner, Lucian Fusu, Mircea Mitroiu, and Lars Vilhelmsen for their valuable comments, and
Norman EF Johnson and Luciana Musetti for maintaining the sites “The genera of Platy-
gastroidea (Hymenoptera) of the World” and “Hymenoptera On-line Database”. The ma-
terial is based on work supported, in part, by a grant of the Romanian National Authority
for Scientific Research, CNCS-UEFISCDI, project number PN-II-RU-TE-2012-0057
to O. Popovici. This research is also based on work supported by the National Science
Foundation (grant numbers DBI-0850223, DEB-0842289, DEB-0956049, and EF-
0905606); and the National Evolutionary Synthesis Center and benefited from discus-
sions initiated through the Phenotype Research Coordination Network. Any opinions,
findings, and conclusions or recommendations expressed in this material are those of the
authors and do not necessarily reflect the views of the National Science Foundation.

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102

Appendix |

Specimens examined.

Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)

Taxon Riies o Data labels
specimens
Anteris sp. 1 ROMANIA: Botosani, 10.viii.2004, leg. O. Popovici
Anteris sp. iL ROMANIA: Iasi, 26.vi.2004, leg. O. Popovici
Apegus sp. 1 ROMANIA: Iasi, 30.vi.2004, leg. O. Popovici
Athalia rosae 1 Wildtype culture, 2005, M. Hatakeyama
Belyta sp. 2 ROMANIA: Iasi, 10.ix.2005, leg. O. Popovici
Calliscelio sp. 1 ROMANIA: Bacau, 3 1.viii.2002 , leg. O. Popovici
Calliscelio sp. 1 ROMANIA: Holboca (Iasi), 30.06.2002, leg. I. Moglan
Duta sp. 1 ROMANIA: Bacau, 17.vii.2004, leg. O. Popovici
ae 5 Costa Rica, 21.11.2005 15/M/16/016 1500-site Costa Rica Heredia, 9km NE Vara
ents Blanca, 10°14'N (INBIO)
Gasteruption sp. 2 USA WV, Hardy Co., 11—22.v.2006, MT, D. Smith
lbalia sp 1 USA: Virginia, Fairfax Co., Great Falls, 18-30.v. 2007, leg. D. Smith
lbalia sp 1 USA: Motgomery Co., 21.v. 2010, leg. N. E. Woodley
lbalia sp. 1 USA: Virginia, Fairfax Co., Great Falls, 3 1.v.-13.vi.2007, leg. D. Smith
Macroteleia sp. 1 ROMANIA: Iasi, 17.viii.2004, leg. O. Popovici
Macroteleia sp. 1 ROMANIA: Iasi, 13.viii.2000, leg. M. Mitroiu
Macroteleia sp. 1 ROMANIA: Iasi, 25.vii.2005, leg. L. Fusu
Orthogonalys 4 USA, WV, Hardy Co. 3mi NE Mathias, 38°55'N, 78°49'W, 2-15.vii.2004, Ma-
pulchella laise trap. D. Smith
Paramesius sp. 1 ROMANIA: Iasi, 10.ix.2005, leg. O. Popovici
ee 1 USA: VA: Fairfax Co., Great Falls, 27.vii-4.viii. 2006, leg. D. Smith
polyturator
taste 1 USA: VA: Fairfax Co., Turkey Run, 5—24.viii.2006, leg. D. Smith
polyturator
Proctotrupes sp. 1 GREECE: Kerkini Nat. Res., 15—17.vii.2007, leg. G. Ramel
Proctotrupes sp. 1 ROMANIA: Iasi, 7.x.2004, leg. O. Popovici
Proctotrupes sp. 1 ROMANIA: Vorona, 21.v.2007, leg. C. Lisenchi
Psilanteris sp. 1 ROMANIA: Bacau, 17.vii.2004, leg. O. Popovici
Ropronia sp 2 USA: Virginia, Fairfax Co., 18—30.v.2007, leg. D. Smith
Scelio sp. 3 ROMANIA: Iasi, 24.viii.2004, leg. O. Popovici
ean 7 ROMANIA: Iasi county, Barnova forest, meadow in glade; 12.vii.2007
? P. N46°59.617'; E27°35.452', Leg. Popovici O. & Popovici Mariana YPT
Oe a 1 ROMANIA: Iasi county, Barnova forest, meadow in glade; 12.vii.2007
? P. N46°59.617'; E27°35.452', Leg. Popovici O. & Popovici Mariana YPT
Se ey My 5 ROMANIA: Iasi county, Botanical Garden; N47°11.167'; E27°32.983';
Ee 26.vii.2007; Leg. Popovici O. YPT
‘See 3 3 ROMANIA: Constanta county; sand dune natural reservation from Agigea; 27—
ae a oe 29.vii.2008; N44°5.184'; E28°38.517'; Leg. Popovici O. & Popovici Mariana YPT
Ge oe, 10 ROMANIA: Constanta county; sand dune natural reservation from Agigea; 27—
? P. 29.vii.2008; N44°5.184'; E28°38.517'; Leg. Popovici O. & Popovici Mariana YPT
eine ts 1 ROMANIA: Tulcea county; forest border, 9 km S of Babadag; 6—7.vii.2009;
@ P- N44°48.817'; E28°42.41'; Leg. Popovici O. & Fusu L. YPT
; ROMANIA: Tulcea county; forest border, 9 km S of Babadag; 6—7.vii.2009;
Spamasion sp.6 1

N44°48.817'; E28°42.41'; Leg. Popovici O. & Fusu L. YPT

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea) 103

Taxon Nie e Data labels
specimens
Saran 1 ROMANIA: Tulcea county; forest border, 9 km S of Babadag; 6—7.vii.2009;
N44°48.817'; E28°42.41'; Leg. Popovici O. & Fusu L. YPT
Sparasion sp.7 2 GREECE: Kerkini Lake, N.P. Lithotopos, 1—7.viii.2006 leg. G. Ramel
Sparasion sp.8 2 JAPAN: Kitahira, Otsu-shi Shiga-ken, 19—22.vii.2008, leg. T. Yoshida.
Sad UGANDA: Kibale, N.P. Kanyawara Biol. Station, 14~21.xi.2010, leg. S. Katusabe
parasion sp.9 1 & Co.
Sparasion sp. 1 USA: NC, Raleigh 03.VII.2009 Winkler, Benoit
Sparasion sp. is USA: West Virginia, Hardy Co. VIII-1 1-28-06, D. Smith
Triteleia sp. 1 ROMANIA: Iasi, 8.ix.2004, leg. O. Popovici
Vanhornia sp. 3 USA: VA: Loudoun Co., 1.vi-17.vii.2000, leg. D. Smith
Appendix 2

Anatomical terms used, cross-referenced to an ontological (formal) definition (Hymenoptera Anatomy

Ontology; URI = Uniform Resource Identifier).

Abbreviation |Term Ontological definiton URI
racine The process that corresponds to a single http://purl.obolibrary.org/obo/
epidermal cell. HAO_0002119
wh rnmurnar [eeDngon deal an HRS publ ob
Shee, featke soen's | HAO_0002205
glossal hair by a carina.
head anterior glossal aU eon a agae one ee http://purl.obolibrary.org/obo/
8 sclerite : J HAO_0000112
anterior glossal plate.
fs anterior surface |The area of the glossa that is between the distal | http://purl.obolibrary.org/obo/
8 of the glossa glossal edge and the anterior glossal plate. HAO_0002208
anterior ‘ ;
tentorio- De Pe ey arises ane, none ang http://purl.obolibrary.org/obo/
tnta-pmn a ty cranium and inserts on the posterior margin of HAO 0001064
P the prementum. =
muscle
: anterior stipito- ecap poig ape nes ee Soa eda http://purl.obolibrary.org/obo/
stia-mp1 ietneice [On the proximal part of the stipes and inserts HAO 0000909
ees anteroproximally on the first maxillary sclerite. E:
The area that extends between anterior margin
: anterolateral wall | of the median wall of the stipes and the lateral | http://purl.obolibrary.org/obo/
ety P p:i/Pp y-Org
J of the stipes margin of the posterior wall of the stipes and }|HAO_0002215
the base of the mandible and the labrum.
agh i ae gigs The spines that are on the distal glossal edge. eae “ Re ee
, The row of setae of the glossa that is adjacent | http://purl.obolibrary.org/obo/
ie to the distal glossal edge. HAO_0002210
These taser em appr btn
wi R HAO_0002142
the coeloconic sensillum of galea.
basal cardinal ak Pa ; http://purl.obolibrary.org/obo/
ber fide The cardinal ridge that is median. HAO. 0002083
b basal galeal The sclerite that receives the site of insertion of | http://purl.obolibrary.org/obo/
8s sclerite the stipito-galeal muscle. HAO_0002143
The anterior invagination of the labium that is
: basal glossal adjacent with the distal end of the salivarium _|http://purl.obolibrary.org/obo/
a & ) p://P yOrg

invagination

and separates the glossa proximally from the | HAO_0002231

distal hypopharynx and the paraglossae.

104 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)
The anteroproximal area of the paraglossa that
beb basiparaglossal _ | is covered with long acanthae and corresponds |http://purl.obolibrary.org/obo/
8 brush to the ligament connecting the acroglossal HAO_0002199
button with the basiparaglossal sclerite.
The sclerite that is located medially on the
ie basiparaglossal _| paraglossa and articulates with the anterior http://purl.obolibrary.org/obo/
P sclerite glossal plate and is continuous with the HAO_0002201
hypopharyngeal rod..
campaniform |The aporous sensillum without a hairlike http://purl.obolibrary.org/obo/
a sensillum cuticular component. HAO_0001973
coeloconic The coeloconic sensillum that is located on the eis RGaR
cfs sensillum of medial surface of the galeo-lacinial complex FED POUEh SHON UEatyOrB Ope,
: ae HAO_0002141
galea distal to the base of the lacinial lobe. a
campaniform |The campaniform sensillum of the anterior ;
csg sensillum of surface of the glossa that is located proximal to HetpHt Dut QUO NDAAN ORB ORO!
x HAO_0002212
glossa the glossal annuli.
pee anor The campaniform sensillum that is on the http://purl.obolibrary.org/obo/
cps sensillum of the P Pye yO%8
isfémentam ventral premental face. HAO_0002244
cardinal articular | The condyle that is located on the cranium Aesiantkebolits ihe
condyle of the | and articulates with the cranial fossa of the aan pak gts G end
: HAO_0002074
cranium cardo.
eedinal lever The process that receives the site of attachment | http://purl.obolibrary.org/obo/
of the cranio-cardinal muscle. HAO_0002075
The sclerite that is articulated with the
cranium at the cranio-cardinal articulation, is h
: ttp://purl.obolibrary.org/obo/
cd connected to the stipes distolaterally via the
ae : ; i : HAO_0000187
stipitocardinal hinge and receives the site of
attachment of the cranio-cardinal muscle.
P meti The area of the integument that is weakly http://purl.obolibrary.org/obo/
Caria ai sclerotized, with thin exocuticle. HAO_0000221
at cranial fossa of cp ames mae an nee http://purl.obolibrary.org/obo/
the cardo Bae a) HAO_0002219
The maxillar muscle that arises medially from
ei cranio-cardinal | the occiput dorsally of the occipital foramen __|http://purl.obolibrary.org/obo/
muscle and inserts on the proximolateral part of the | HAO_0001592
cardo.
cranio-lacinial Tne ee ENE e eee ofihe http://purl.obolibrary.org/obo/
muscle facinial lobe, HAO_0001593
d The transverse edge that extends distally on http://purl.obolibrary.org/obo/
Be the glossa. HAO_0002206
dhi ibe TBE aOR OF ae eset ee yas http://purl.obolibrary.org/obo/
i hypopharyngeal | that id adjacent to the concavities of the HAO 000221
Es wea ai a! 5
invagination hypopharyngeal rods.
The area that is located on the anterior
surface of the hypopharyngeal wall and is
distal ep ases come Ys Es ee http://purl.obolibrary.org/obo/
dhy RE 2s pouch or the distal margin of the sitophore, HAO. 0001575
Slane are distally by the salivarial orifice and laterally =
by the lateral parts of the prementum and the
hypopharyngeal rods.
distal The sclerite that receives the site of insertions
dhe Ry popharpneeal of the dorsal and ventral premento-salivarial | http://purl.obolibrary.org/obo/

sclerite

muscles and is continuous with the

HAO_0002228

hypopharyngeal rod.

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea)

The sclerite that is located distolaterally on the

dis distolateral galeal etal walla the-ealenelacimial complevand http://purl.obolibrary.org/obo/
sclerite f HAO_0002127
bears the apicolateral galeal setae.
d distolateral galeal | The setiferous patch that is located http://purl.obolibrary.org/obo/
SP setiferous patch | distolaterally on the galeo-lacinial complex HAO_0002128
distomedial ape ued ee Oe Era ag or http://purl.obolibrary.org/obo/
dmf ae the stipes that is overlapped by the premental
stipital flange HAO_0002217
carina and overlaps the lateral premental face.
aes dorsal premento-| The salivarial muscle that inserts http://purl.obolibrary.org/obo/
Pete salivarial muscle | proximodorsally on the salivary duct. HAO_0000274
fifth sclerite of | The sclerite that is ringlike and is connected east intl shall beawe terse:
5mp the maxillary distally to the fourth sclerite of the maxillary Hi #6 a 0 0222 0 y0rs
palp palp via conjunctiva. “,
first intrinsic The muscle that arises from the first sclerite of ;
http://purl.obolibrary.org/obo/
mp1-mp2 muscle of the —_| the maxillary palp and inserts on the second HAO 0002114
maxillary palp _|sclerite of the maxillary palp. =
first intrinsic The muscle that arises from the first sclerite
http://purl.obolibrary.org/obo/
muscle of the —_|of the labial palp and inserts on the second HAO. 0002237
labial palp sclerite of the labial palp. =
ll first sclerite of fs ue i in pie ne . coir’ http://purl.obolibrary.org/obo/
P thelabial palps |e che, Se ee ee HAO_0002194
conjunctiva and muscle.
first sclerite of _ | The sclerite that is ringlike and is connected
; ane: .__._|http://purl.obolibrary.org/obo/
Imp the maxillary distolaterally to the posterior stipital sclerite via HAO 0002109
palp conjunctiva and muscle. a
fourth intrinsic |The muscle that arises from the third sclerite
http://purl.obolibrary.org/obo/
muscle of the | of the maxillary palp and inserts on the fourth HAO 0002222
maxillary palp _|sclerite of the maxillary palp. =
fourth sclerite —_ | The sclerite that is ringlike and is connected
http://purl.obolibrary.org/obo/
4mp of the maxillary | distally to the third sclerite of the maxillary HAO 0002113
palp palp via conjunctiva. te
The row of setae that is located on the medial
wall of the galeo-lacinial complex proximal to Ras age
the coeloconic sensilla of galea. id
The row of setae that extends along the margin
galeal fringe of the galeo-lacinial complex, distal to the " eas : vate ate
lacinial lobe. ve
galeal lamina Le ae enn ae and ArepH put LOUs lb Apr re ope
; HAO_0002136
margined by the galeal comb. =
The lobe that is located on the stipes at the :
glb distal part of the posterior alta ey eae
distolateral to the lacinia. 3
The area of the stipes that is delimited
galeo-lacinial P rgamomegialy p vane SUPIEP remnictiral http://purl.obolibrary.org/obo/
gal-lac eee conjunctiva, proximolaterally by the stipito- HAO 0002126
ee mandibular conjunctiva and posteroproximally r
by the margin of the posterior stipital sclerite.
The lobe of the labium that is limited
I | posteroproximally by the prementum, http://purl.obolibrary.org/obo/
5° Bayes anteroproximally by the fold traversing the HAO_0000376
salivary orifice and laterally by the paraglossae.
la ierianeie The anatomical cluster of the glossa that is http://purl.obolibrary.org/obo/
8 5 composed of annular hairs. HAO_0002204
hyb hypopharyngeal he sara of ae Hyper eS ioe sr http://purl.obolibrary.org/obo/
y ee receives the site of insertion of the posterior fry 4) Q92234

tentorio-premental muscle.

105

106 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)
The anterolateral area of the distal
i‘ hypopharyngeal AYROP EEL aUe acl scene Wa Me http://purl.obolibrary.org/obo/
pc proximal part of the hypopharyngeal rod
pecten ; . |HAO_0002214
proximal to the hypopharyngeal button and is
covered with acanthae.
The ligament that connects the proximolateral hipdipurzabolipanrcre obo
hypr margin of the prementum with the proximal HAO 0000408
part of the ligula. 2
The area that extends on the posterior (ventral)
TuareMn oleae ra foramen along ne pee of http://purl.obolibrary.org/obo/
hys attachments of the conjunctiva connecting the HAO 0000411
cranium with the maxillae and is delimited =
laterally by the pleurostomal fossa.
The anatomical system that forms the covering
; layer of the animal, ectodermal in origin and __|http://purl.obolibrary.org/obo/
bead a composed of epidermal cells producing the HAO_0000421
cuticle.
The area that is located between the cardines
intercardinal ane pe i P tena Dane sa Sea gu http://purl.obolibrary.org/obo/
‘ anatomical line that extends between the HAO 0002145
of proximal and distal ends of the left and right ra
cardines.
The area of the postmental area that is limited
laterally by the median margins of the stipites, h
rt ttp://purl.obolibrary.org/obo/
interstipital area | proximally by the anterior margin of the HAO 0002146
intercardinal area and distally by the proximal =
margin of the prementum.
interstipital sie cae beathatls era he inteetip i http://purl.obolibrary.org/obo/
ite area and is connected to the prementum via [i744 9992223
PE conjunctiva. 7
invagination The area where the cuticle is invaginated. De eas er
The anatomical structure that is distal to the
; ; http://purl.obolibrary.org/obo/
labial palp proximal margin of the first sclerite of the HAO_0000450
labial palp.
labial palpal be ala as sts cei seen! ai ie http://purl.obolibrary.org/obo/
pin CRS ad see um and receives the base of the la HAO_0002153
The appendage that is encircled by the area
that is proximally delimited by the lateral
margins of the cardo and the posterior stipital | http://purl.obolibrary.org/obo/
sclerite laterally, and the anatomical line that is } HAO_0000453
tangential to the salivary duct and traverses the
salivary orifice anteriorly.
lacinial b The sclerite that is located on the lateral wall of | http://purl.obolibrary.org/obo/
salami the lacinial lobe. HAO_0002117
eine b The row of setae on the lacinial lobe that is http://purl.obolibrary.org/obo/
acimar com | marginal. HAO_0002124
The sclerite that is located on the medial :
bls lacinial lever stipital wall and receives the site of insertion of eps! purl ove ipratsere obo!
ee NC HAO_0002093
the stipito-lacinial muscle.
The lobe that extends proximally on the distal
a margin of the medial stipital wall, is adjacent | http://purl.obolibrary.org/obo/
Ilb lacinial lobe to the basal lacinial sclerite, overlaps the HAO_0000457
proximal part of the galea.
The area of the prementum, that lays parallel ;
ound lateral premental sad Bommesieditia pomiensavattonhe medial http://purl.obolibrary.org/obo/

face

stipital vall.

HAO_0002152

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea)

The cardinal ridge that arises distally from

laterodistal oa apa 2 ee http://purl.obolibrary.org/obo/
Icr bandinal idee eae ridge and is oriented HAO__0002084
; ; Beak oid mere http://purl.obolibrary.org/obo/
ligament The area of the cuticle that is resilin rich. HAO_0002229
The anatomical cluster that is composed of the | http://purl.obolibrary.org/obo/
glossa and paraglossae. HAO_0000496
The evagination that is mostly membranous. SS aaa ee eas
The sclerite that is connected to the cranium
d along the anterior margin of the oral foramen |http://purl.obolibrary.org/obo/
uP via the anterior and posterior cranio- HAO_0000506
mandibular articulations.
marginal fringe OG some teat ey 5 composes. ore http://purl.obolibrary.org/obo/
mfs spines on the distal margin of the distolateral
of the stipes Abe ; HAO_0002216
portion of the principal carina of the stipes.
The appendage that is encircled by the area
oe ae fos pete Py oe Syeesene http://purl.obolibrary.org/obo/
maxilla, maxillae | posteriorly, the median margin of the mandible
HAO_0000513
laterally, the labrum anterolaterally and the
labium medially.
The anatomical structure that is distal to the Remon nesiecbakits aoe,
| maxillary palp | proximal margin of the first sclerite of the Pe Ar cae os
pests HAO_0000515
maxillary palp.
; http://purl.obolibrary.org/obo/
The sclerite that is part of the maxillary palp. HAO_0002183
th The articular process that bears the cardinal _| http://purl.obolibrary.org/obo/
eee orn’ | condyle of the cranium. HAO_0002073
ypostoma
The anatomical cluster that is composed of
the maxillae and the labium and is connected
maxillo-labial —_| by conjunctivae laterally to the cranium along |http://purl.obolibrary.org/obo/
complex the hypostoma, to the mandible along the HAO_0000452
proximomedial margin of the mandible and
proximally to the hypopharynx.
h mechanosensory | The aporous sensillum that has a hair-like http://purl.obolibrary.org/obo/
oe hair cuticular component. HAO_0001039
Abe medial stipital ee nae Een ade a al http://purl.obolibrary.org/obo/
groove eaisrones HAO_0002106
fea spall The area that is medial and lays parallel with | http://purl.obolibrary.org/obo/
lei the lateral wall of the prementum. HAO_0002092
median te :
conjunctiva of a es ee ea: res - i eae oS http://purl.obolibrary.org/obo/
the anterior Bey eet gee Pe ase to AOLO002203
glossal plate ;
median The ridge that is limited laterally by the stipital | http://purl.obolibrary.org/obo/
paren 8 ¥ DY Pp p:i/P 8g
P postmental ridge | articular surfaces of the postmentum. HAO_0002225
mediodistal ee ares rae ae re http://purl.obolibrary.org/obo/
oa cardinglridvels [Rn wee ee ge HAO_0002085
distomedially.
The lobe that is connected to the distal margin
of the prementum posteroproximally, to the
pel premental hypopharynx proximolaterally http://purl.obolibrary.org/obo/

HAO_0000686

and anteroproximally, to the glossa
proximomedially and bears the basiparaglossal
brush and the paraglossal sclerite.

107

108 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)
bataplossa! The sclerite that is located distally on the http://purl.obolibrary.org/obo/
pgb acroglossal ;
bie paraglossa. HAO_0002232
paraglossal The anatomical cluster of the paraglossa that is | http://purl.obolibrary.org/obo/
annuli composed of connected annular hairs. HAO_0002233
paraglossal The anteroproximal sclerite of the paraglossa__| http://purl.obolibrary.org/obo/
sclerite that bears the basiparaglossal brush. HAO_0002200
The sclerite that is connected to the distal
posterior glossal | margin of the prementum and receives the http://purl.obolibrary.org/obo/
PSP plate site of insertion of the premento-paraglossal | HAO_0000747
muscles.
The sclerite that is located on the posterior
stipital wall, articulates with the cardo
Re posterior stipital |and with the labial palp, is connected by http://purl.obolibrary.org/obo/
P sclerite conjunctiva distolaterally to the galeo-lacinial |HAO_0002097
complex, proximally to the hypostoma and the
cardo, proximolaterally to the mandible.
The stipito-palpal muscle that arises medially
posterior stipito- |on the proximal part of the stipes and inserts | http://purl.obolibrary.org/obo/
stip-mp 1 8
palpal muscle _| posteroproximally on the first maxillary palpal | HAO_0001814
segment.
. ane gieaiat ls ite sy pedal Eee Bene a! http://purl.obolibrary.org/obo/
posterior wall —_| stipital wall and limited proximolaterally by
; a Ae) HAO_0002097
the margin of the posterior stipital sclerite.
posterior surface a i t io contac aera ne http://purl.obolibrary.org/obo/
Pwe of the glossa ia meee spt Ly cane ih eal aa HAO_0002207
eS as The tentorio-labial muscle that arises from
tentorio- http://purl.obolibrary.org/obo/
tntp-pmn the cranium and inserts distally on the labium
premental HAO_0000264
adjacent to the level of the salivary orifice.
muscle
The area that is limited distally by the posterior
tmental area |™2"sin of the prementum and laterally by http://purl.obolibrary.org/obo/
Postmentar atc | the median margins of the cardines and the HAO_0002144
stipites.
h postmental- The conjunctiva that is between the http://purl.obolibrary.org/obo/
PP premental hinge | postmentum and prementum. HAO_0002226
ponerse ial mac ar une ie Fm Tangalanga
Pp“ P HAO_0000803
muscle margin of the prementum.
be wae The anatomical cluster that is composed of the | http://purl.obolibrary.org/obo/
r pone gure sclerites that are on the postmental area. HAO_0000785
a premental arms receives the ste oforign of the dows [Rttpel(pus.obolibrary.org/obo
WIPE HAO_0002155
premento-salivarial muscle.
The flange that is adjacent with the border
eh eaten between the ventral and lateral premental faces | http://purl.obolibrary.org/obo/
P P and that overlaps externally the median part of | HAO_0002157
the posterior stipital sclerite.
The scrobe of the prementum that is adjacent
iid premenealatiten to and extends lateral to the premental carina | http://purl.obolibrary.org/obo/

HAO_0002227

and accommodates the medial part of the
posterior stipital sclerite.

premento-glossal
muscle

The labial muscle that arises on the ventral
part of the prementum, laterally to the ventral |http://purl.obolibrary.org/obo/
premento-salivarial muscle and inserts on the |HAO_0000377

anterior glossal plate.

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea)

premento-palpal
muscle

The labial muscle that arises from the
prementum and inserts on the first sclerite of

the labial palp.

http://purl.obolibrary.org/obo/
HAO_0000314

The labial muscle that arises from the ventral

premento- part of the prementum, anterior to the origin : ¢
pmn-pgs paraglossal of the premento-glossal muscle and ventral AEDS Pu OUe DIANE ORB SG
ciate j . HAO_0000687
muscle premento-salivarial muscle and inserts just
distally of the distal margin of the prementum.
The sclerite that is median, is connected via
conjunctiva along its proximolateral margins
to the stipites, is articulated with the labial
palps, is continuous along its distal margin http://purl.obolibrary.org/obo/
\ er pe with the ligula and distolateral margins with |HAO_0000804
the distal hypopharynx and receives the site
of attachments of the extrinsic labial palp
muscles.
principal carina |The flange that extends along the margin of __|http://purl.obolibrary.org/obo/
Re of the stipes the posterior stipital wall. HAO_0002099
proximal galeal | The row of setae that is on the distal margin of | http://purl.obolibrary.org/obo/
brush the proximolateral galeal sclerite. HAO_0002135
é The sclerite that is located on the lateral wall ‘
: proximolateral BF die glen dncdialientiplecandibes ele http://purl.obolibrary.org/obo/
P8& galeal sclerite ei) s HAO_0002130
proximolateral galeal setiferous patch.
proximolateral | The setiferous patch that is located on the j .
prs galeal setiferous | lateral wall of the galeo-lacinial complex Dsl Put Ope Mp rary Ore! O00)
: , ‘ HAO_0002129
patch proximally of the distolateral setiferous patch.
The medial part of the principal carina of
f proximomedial jee ae i ern by Pa http://purl.obolibrary.org/obo/
pm Sdpial flange’ §[c te ces Cue Sanna and Over apes FAO: 0002218
lateral premental face and the proximal part of
the prementum.
The area that is at the proximal end of the
ae salivary duct and corresponds to the site of http://purl.obolibrary.org/obo/
ok Ticks insertion of the dorsal and ventral premento- | HAO_0000906
salivarial muscles.
svd salivary duct The duct that leads from the salivary gland. Bp Vip sl Obs IeiatiorBiobe,

HAO_0002236

salivarial sclerite

second intrinsic
muscle of the
maxillary palp

The sclerite that is located in the ventral wall
of the salivarium and corresponds to the site

of insertion of the ventral premento-salivarial
muscle.

http://purl.obolibrary.org/obo/
HAO_0001682

The area of the integument that is strongly
sclerotised, with thick exocuticle and is
surrounded by conjunctivae.

http://purl.obolibrary.org/obo/
HAO_0000909

The muscle that arises from the first sclerite
of the maxillary palp and inserts on the third
sclerite of the maxillary palp.

http://purl.obolibrary.org/obo/
HAO_0002115

second intrinsic
muscle of the

labial palp

second sclerite of

the labial palp

second sclerite
of the maxillary

palp

The muscle that arises from the second sclerite
of the labial palp and inserts on the third
sclerite of the labial palp.

http://purl.obolibrary.org/obo/
HAO_0002238

The sclerite that is ringlike and is connected
distally to the first sclerite of the labial palp via
conjunctiva and muscle.

http://purl.obolibrary.org/obo/
HAO_0002195

The sclerite that is ringlike and is connected
distolaterally to the first sclerite of the
maxillary palp via conjunctiva and muscle.

http://purl.obolibrary.org/obo/
HAO_0002111

109

110 Ovidiu Alin Popovici et al. / Journal of Hymenoptera Research 37: 77-111 (2014)
The sensillum that is multicellular and consists
‘ of trichogen, tormogen, and sense cells and http://purl.obolibrary.org/obo/
Si the cuticle secreted by and adjacent to the HAO_0000935
trichogen cell.
Realise The muscle that is attached at either end to http://purl.obolibrary.org/obo/
the cuticle. HAO_0001922
spiculate patch |The area on the galeo-lacinial complex distal to | http://purl.obolibrary.org/obo/
P Pp g Pp p:i/P yOrg
of galea the lacinial lobe that is covered with acanthae. | HAO_0002139
spiculate patch |The area on the lacinial lobe that is covered _| http://purl.obolibrary.org/obo/
of the lacinia with acanthae. HAO_0002138
Bont The process that lacks non-sclerotised ring at | http://purl.obolibrary.org/obo/
7 the base. HAO_0000949
The appendage that is connected
posteroproximally to the hypostoma,
: oe, una ay ane aye Spree te http://purl.obolibrary.org/obo/
stipes, stipites | mandible and medioproximally to the labium HAO 0000958
and the hypopharynx via conjunctiva, is =
connected to the cranium via muscles and that
bears the maxillary palp.
rustlers The sclerite that is on the medial stipital wall, | http://purl.obolibrary.org/obo/
Stipitar sent’ | bears the medial stipital process. HAO_0002096
The maxillar muscle that arises from the
stipito-galeal Pees a Sete of a eee edit ee http://purl.obolibrary.org/obo/
sti-gal ae site of origin of the stipito-lacinial muscle and HAO 0001661
inserts on the median wall of the galeo-lacinial .
complex distal to the stipito-lacinial muscle.
The maxillar muscle that arises along the
Pttic stipito-lacinial | lateral margin of the posterior stipital sclerite | http://purl.obolibrary.org/obo/
muscle and inserts proximally on the median wall of |HAO_0001660
the stipes just proximal to the lacinal lobe.
; stipito-palpal The ee pee chat ae Don the http://purl.obolibrary.org/obo/
sti-mp1 aes posterior stipital sclerite and inserts on the first HAO 0002110
sclerite of the maxillary palp. =
eapitos The conjunctiva that extends along the
fic aden posterior (dorsal) margin of the premental arm |http://purl.obolibrary.org/obo/
z pnjaneche and the proximal margin of the medial stipital | HAO_0002125
wall.
stipitocardin e membranous area linking the cardo an ttp://purl.obolibrary.org/obo
h p dinal |Th b linking the cardo and ___|http://purl.obolibrary.org/obo/
_ hinge stipes. HAO_0002076
styloconic ; http://purl.obolibrary.org/obo/
sts cae The seta that is on a process. u RON onoD ot ae
tentorio-cardinal ae pally pee ches Alls pon ne http://purl.obolibrary.org/obo/
tnt-cd anterior region of the cranium and inserts
muscle dye. ai HAO_0001638
adjacent to the cardino-stipital hinge.
The maxillar muscle that arises on the
. tentorio-stipita osteroventral part of the anterior tentoria ttp://purl.obolibrary.org/obo
peta pital |p al part of th | http://purl.obolibrary.org/obo/

muscle

tentorium

arm and inserts on the median wall of the
stipes.

The apodeme that has its sites of origin
marked by the anterior and posterior tentorial
pits and gular sulci.

HAO_0001002

http://purl.obolibrary.org/obo/
HAO_0001003

third intrinsic
muscle of the
maxillary palp

third sclerite of
the labial palp

The muscle that arises from the second sclerite
of the maxillary palp and inserts on the third
sclerite of the maxillary palp.

The sclerite that is ringlike and is connected
distally to the second sclerite of the labial palp

via conjunctiva and muscle.

http://purl.obolibrary.org/obo/
HAO_0002116

http://purl.obolibrary.org/obo/
HAO_0002196

The maxillo-labial complex of Sparasion (Hymenoptera, Platygastroidea) 111

third sclerite of |The sclerite that is ringlike and is connected
3mp the maxillary distally to the second segment of the maxillary
palp palp via conjunctiva and muscle.

http://purl.obolibrary.org/obo/
HAO_0002112

uniporous T
ssl sensilla, Type
1 seta

he sensillum whose cuticular component has |http://purl.obolibrary.org/obo/
one cuticular pore. HAO_0002221

The flange that is transparent, and extends
vlm along the anterodistal margin of the galeo-
lacinial complex distal to the lacinial lobe.

http://purl.obolibrary.org/obo/
HAO_0002140

The projection that is located distally on the urbe foueRobalibeaipensiabor

vdp posterior stipital sclerite and encircles the base HAO_0002102
of the maxillary palp.
ventral glossal | The carinae that radiates from the posterior _| http://purl.obolibrary.org/obo/
lines glossal plate towards the apical glossal setae. HAO_0002230
£ ventral The area of the prementum that is delimited | http://purl.obolibrary.org/obo/
vp premental face _ | laterally by the lateral premental face. HAO_0002156
ventral The salivarial muscle that arises from the
| premento- proximal end of the ventral part of the http://purl.obolibrary.org/obo/
ae at salivary sclerite | prementum and inserts on the salivarial HAO_0001072
muscle sclerite.
The projection that is located proximolaterall
tha ventrolateral a Le Raa Se Naik es dias eam Y |http://purl.obolibrary.org/obo/
basiglossal arm or ae HAO_0002202
with the basiparaglossal sclerite.
Appendix 3

Volume rendered CLSM media files on figshare.com
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doi: 10.6084/m9.figshare.861057
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