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JHR 44: 69-78 (2015) JOURNAL OF *0errviewed opevaccetsjoural
doi: 10.3897/JHR.44.4722 ME Hymenopter a

http://jhr.pensoft.net The international Society of Hymenoptersts. RESEARCH

A new species of Labania Hedavist (Braconidae,
Doryctinae) from Costa Rica, reared from
aerial root galls of Ficus obtusifolia Kunth

Sergey A. Belokobylskij'*, Miguel Artavia Solis’,
Paul E. Hanson’, Alejandro Zaldivar-Riverén*

| Zoological Institute of the Russian Academy of Sciences, Universitetskaya naberezhnaya 1, St Petersburg
199034, Russia 2. Museum and Institute of Zoology of the Polish Academy of Sciences, Wilcza 64, 00-679
Warsaw, Poland 3 Escuela de Biologia, Universidad de Costa Rica, San Pedro de Montes de Oca, Costa Rica
4 Coleccién Nacional de Insectos, Instituto de Biologia, Universidad Nacional Auténoma de Mexico, 3er.
circuito exterior s/n, Ca. Universitaria, Copilco, Coyoacdn, A. P 70-233, C. P 04510, D. FE, México

Corresponding author: Alejandro Zaldivar-Riverén (azaldivar@ib.unam.mx)

Academic editor: G. Broad | Received 13 February 2015 | Accepted 14 April 2015 | Published 11 June 2015
http://zoobank. org/33C017A3-8 LEO-49D2-A373-3DD0170E2A4C

Citation: Belokobylskij SA, Solis MA, Hanson PE, Zaldivar-Riverén A (2015) A new species of Labania Hedgqvist
(Braconidae, Doryctinae) from Costa Rica, reared from aerial root galls of Ficus obtusifolia Kunth. Journal of Hymenoptera

Research 44: 69-78. doi: 10.3897/JHR.44.4722

Abstract

A new species of the Doryctinae genus Labania Hedqvist, L. ficophaga sp. n. from Costa Rica is described.
This new species was reared from aerial root galls of Ficus obtusifolia Kunth. An updated key to the five
described species of Labania and digital pictures of L. ficophaga sp. n. and L. minuta Marsh are provided.

Keywords

Ichneumonoidea, Neotropics, gall-association, Hymenoptera

Introduction

The genus Labania Hedgvist is a small doryctine group restricted to the Neotropical
region and is currently composed of four species. This genus was erected by Hedqvist
(1963) based on a single species from Honduras, L. straminea Hedgqvist. In the de-
scription, Labania was placed within the morphologically heterogeneous subfamily
Hormiinae, but it was subsequently transferred to the Doryctinae in a tribal reclassi-

Copyright Sergey A. Belokobylskij 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.

70 Sergey A. Belokobylskij et al. / Journal of Hymenoptera Research 44: 69-78 (2015)

fication of this subfamily (Belokobylskij 1992). To date, only three additional species
of Labania have been described from Costa Rica (Marsh 2002), L. hansoni Marsh, L.
minuta Marsh and L. prolata Marsh, with the first two being reared from aerial root
galls on Ficus citrifolia Mill. and leaf galls on F. colubrinae Strandl., respectively.

In his tribal reclassification of the Doryctinae, Belokobylskij (1992) placed Laba-
nia as the single member of the tribe Labaniini based on several putative apomorphic
features. More recently, however, three molecular phylogenetic studies consistently
recovered a clade exclusively composed of genera whose species are associated with
galls from different plant families, including Labania (Zaldivar-Riveron et al. 2007,
2008, 2014). In the most recent of these studies (Zaldivar-Riverén et al. 2014), the
authors described three new gall-associated doryctine genera whose species with rear-
ing records were all associated with Ficus species. One of them, Ficobolus Belokob-
ylskij & Zaldivar-Riverén, was recovered as sister to Labania; the latter genus was
represented in that study by an undescribed species from Costa Rica.

In this work, this new species of Labania, which was reared from aerial roots on
F. obtusifolia Kunth is described, and observations that suggest that this new species is
phytophagous are provided. An updated key (adapted from Marsh 2002) to the five

described species of Labania is given.

Material and methods

All specimens of the new species of Labania described here were reared from a single tree
of F. obtusifolia (see details below). Specimens belonging to two of the remaining species
of the genus were also examined, as well as digital photographs of the holotype of ZL.
straminea, deposited at National Museum of Natural History (Washington, D.C., USA).
The type material of the new species is deposited in the following collections: Coleccién
Nacional de Insectos, Instituto de Biologia, Universidad Nacional Auténoma de México
(CNIN IB-UNAM); Museo de Zoologia, University of Costa Rica, San José, Costa Rica
(MZCR); Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia
(ZISP); and National Museum of Natural History, Washington, DC (NMNH).

The terminology employed in this work for the morphological features and meas-
urements follows Belokobylskij and Maeté (2009). The wing venation nomenclature
follows Belokobylskij and Maeté (2009) and Sharkey and Wharton (1997) in paren-
theses. Photographs were taken with a Leica IC 3D digital camera mounted on a Leica”
MZ16 microscope and using the Leica Application Suite® imaging system.

Other examined material

Labania straminea. Holotype: female, Honduras, La Ceiba. Labania minuta: 5 females,
5 males. Costa Rica, Turriatlico, Turrialba, ex leaf-ball gall of Ficus colubrinae, 1X-2013,
K. Nishida col. Labania hansoni: 2 females, 1 male. “Costa Rica: DNA Hym- 46”.

A new species of Labania Hedgvist (Braconidae, Doryctinae)... |

Results

Taxonomy

Labania ficophaga Belokobylskij & Zaldivar-Riverén, sp. n.
http://zoobank.org/7A4A0738-1F19-46C8-B8AA-1C9788785478
Figs 1; 2A—B

Diagnosis. ‘This new species is morphologically similar to L. hansoni Marsh; however,
L. ficophaga can be distinguished from the latter species by the following features: 1)
greater diameter of eye 2.0—2.2 times wider than temple (1.4—1.7 times in L. hansoni);
2) eye covered by short and sparse setae (long and very dense setae in L. hansoni); 3)
malar space 0.35—0.4 times height of eye (0.4—0.5 times in L. ansoni); 4) propodeum
mainly rugose-striate, smooth only in basolateral areas (mainly smooth, rugose only
medially in L. hansoni); 5) discoidal (first discal) cell distinctly sessile anteriorly (peti-
olate anteriorly in L. ansoni); 6) dorsal carinae of the first metasomal tergite almost
complete (incomplete in L. hansoni); 7) vertex and frons light reddish brown or brown-
ish yellow (reddish brown to black in L. /ansoni); 8) basal antennal segments black
or almost black (yellow or brownish yellow in L. hansoni); 9) hind femur and tibia of
female dark reddish brown to reddish brown (yellow or brownish yellow in L. hansoni).

Labania ficophaga is also morphologically similar to L. minuta, but differs from
this species by having: 1) a larger body length, from 3.6 to 4.1 mm (1.0 to 1.8 mm
in L. minuta; Fig. 3A); 2) antenna 23—26-segmented (less than 20 antennomeres in
L. minuta; Fig. 3A); 3) notauli joining at the end of mesoscutum in a wide longitu-
dinally striate-rugose area (joining at the end of mesoscutum in narrow rugose area
in L. minuta; Fig. 3B); 4) head light reddish brown or yellowish brownish (face and
temple honey yellow to brown, frons and vertex black in L. minuta; Figs 3A—C); and
5) entirely smooth second metasomal tergite (with mediobasal longitudinal striae in
L. minuta; Fig. 3E).

Description. Female. Body length 3.6—4.1 mm; fore wing length 3.1—3.2 mm.

Head: width 1.8—1.9 times its median length, 1.15—1.20 times width of mesoscu-
tum. Head behind eyes (dorsal view) distinctly roundly narrowed. Transverse diameter
of eye 2.0—2.2 times wider than temple. Ocelli medium-sized, arranged in triangle
with base 1.2—1.3 times its sides. POL 0.8—1.0 times Od, 0.30—0.35 times OOL.
Frons excavation deep, not wide, not extending beyond antennal sockets, with shallow
or very shallow median furrow. Eye without emargination opposite antennal socket,
1.3 times as high as broad. Malar space 0.35—0.40 times height of eye, 0.8—1.0 times
basal width of mandible. Face slightly convex, its width 0.9 times height of eye and
0.9-1.0 times height of face and clypeus combined. Clypeus convex, about twice as
wide as high. Width of hypoclypeal depression almost equal to distance from edge of
depression to eye, 0.5—0.6 times width of face. Hypostomal flange narrow. Occipital
carina not fused with hypostomal carina and obliterated ventrally. Palpi short, maxil-

lary palpus 5-segmented, labial palpus 2-segmented.

22 Sergey A. Belokobylskij et al. / Journal of Hymenoptera Research 44: 69-78 (2015)

Figure |. Labania ficophaga sp. n. (female, paratype): A habitus, lateral view B head and palpi, ventro-

latero-posterior view C head and mesosoma, dorsal view D head and mesosoma, lateral view E metasoma,

dorsal view F fore wing G hind wing.

Antenna: thickened, weakly setiform, covered by dense and dark setae, 23—26-seg-
mented. Scape 1.5—1.6 times longer than its maximum width, almost twice as long
as pedicel. First flagellar segment 4.0—4.5 times longer than its apical width, 1.2-1.3
times longer than second segment. Penultimate segment 2.2—2.3 times longer than its
width, 0.7—0.8 times as long as apical segment; the latter pointed apically.

Mesosoma: length 1.6-1.7 times its height. Median lobe of mesoscutum dis-
tinctly and highly convex (lateral view), slightly protruding forward and rounded

A new species of Labania Hedqvist (Braconidae, Doryctinae)... 73

anteriorly (dorsal view). Oblique submedian furrow on side of pronotum narrow
and densely crenulated. Notauli complete, deep, narrow and densely crenulated,
joining at the end of mesoscutum in a longitudinally striate-rugose area. Prescutel-
lar depression long, deep, with several more or less high carinae, densely and dis-
tinctly reticulate-striate between carinae, 0.25—0.35 times as long as scutellum.
Scutellum 1.2 times longer than maximum anterior width. Scuto-scuttelar suture
absent. Precoxal sulcus almost absent or very weak, short. Metanotal tooth low
(lateral view). Metapleural flange quite short and wide, almost indistinct. Propodeal
lateral tubercles large and wide.

Wings: fore wing 2.7—3.0 times longer than its maximum width. Pterostigma
2.9-3.5 times longer than width. Metacarp (R1) 1.2—1.3 times longer than pter-
ostigma. Radial (marginal) cell long, open apically after strong desclerotisation of
apical part of third radial abscissa (3RSb). First (r) and second (3RSa) radial abscis-
sae forming an obtuse angle. Second radial abscissa (3RSa) 4.0—4.3 times longer
than first abscissa (r), 0.35—0.40 times as long as the straight third abscissa (3RSb).
First radiomedial vein (2RS) mainly or completely absent, rarely present and short,
or (exceptionally) with elongate anterior part of abscissa. Second radiomedial (sec-
ond submarginal) cell fused with first radiomedial (first submarginal) cell, slightly
narrowed apically. First medial abscissa ((RS+M)a) distinctly sinuate. Discoidal (first
discal) cell sessile. Recurrent vein (1m-cu) 0.3—0.4 times as long as basal vein (1M);
these veins parallel. Distance from nervulus (1cu-a) to basal (1M) vein 0.7—0.8 times
nervulus (1cu-a) length. Brachial (first subdiscal) cell long, wide, opened apico-pos-
teriorly after strong desclerotisation of second abscissa of longitudinal anal vein (2-
1A). Brachial vein (2cu-a) long and mainly pigmented. Hind wing 4.0—4.1 times
longer than its maximum width. Hamuli 3-4. Medial (basal) cell apically widely
open after strong desclerotisation of second abscissa of costal vein (SC+R). Nervellus
(cu-a) absent; submedial (subbasal) cell widely open apically. Recurrent vein (m-cu)
distinct or fine, straight or slightly curved, more or less inclivous, interstitial, dis-
tinctly or sometimes faintly pigmented.

Legs: hind coxa without basoventral corner and tubercle, 1.4—1.5 times longer than
maximum width, about 0.8 times as long as propodeum. Hind femur 2.8—3.1 times
longer than width. Hind tarsus 0.8—0.9 times as long as hind tibia. Basitarsus of hind
tarsus 0.55—0.60 times as long as second to fifth segments combined. Second segment
of hind tarsus 0.5 times as long as basitarsus, 0.9 times as long as fifth segment (with-
out pretarsus). Claws large.

Metasoma: 1.0—1.1 times longer than head and mesosoma combined. First ter-
gite distinctly and almost linearly widened from base to apex, with slightly impressed
spiracular tubercles in basal third, with distinct, almost complete and slightly con-
vergent posterior dorsal carinae. Maximum width of first tergite about 2.0 times its
minimum width; length 0.9-1.0 times its apical width, 1.4-1.5 times length of pro-
podeum. Suture between second and third tergites quite shallow and almost complete.
Median length of second and third tergites 1.1—1.2 times basal width of second tergite,
0.8—0.9 times maximum width of these tergites. Ovipositor short, its sheath about as

74 Sergey A. Belokobylskij et al. / Journal of Hymenoptera Research 44: 69-78 (2015)

rp is { wig R
= se i Hs 4 + YA ‘ "hs ' iy ie fe
3 4 : ht x hai : Prt oe

- Sasa ae th atl Wace & |:
rks an SSR OSS ees sae i ak Lei wi zt

Figure 2. Labania ficophaga sp. n. (male, paratype): A habitus, lateral view B hind femur C Aerial root

galls on F. obtusifolia Kunth where specimens of L. ficophaga were reared.

long as first tergite of metasoma, 0.40—0.45 times as long as mesosoma, 0.18—0.22
times as long as fore wing.

Sculpture and pubescence: head entirely smooth, sometimes frons near antennal
sockets with several curved striae. Mesoscutum smooth, with rugulosity in short and
small medioposterior area. Scutellum mainly smooth, finely striate laterally. Meso-
pleuron mostly smooth. Propodeum widely rugose-striate, with all areas distinctly
delineated by carinae; basolateral areas short, semi-rounded, mainly smooth; areola
wide, about as long as maximum width; basal carina 0.2—0.3 times as long as basal
fork of areola. Hind coxa and femur smooth. First tergite mainly striate with rugosi-
ties, finely reticulate to almost smooth between dorsal carinae. Remaining tergites
entirely smooth. Vertex with numerous, dense and semi-erect pale setae. Mesoscu-
tum mainly glabrous, with dense and erect pale setae arranged in almost single lines
along notauli and laterally. Mesopleuron medially rather widely glabrous. Hind tibia
dorsally with dense, relatively long and semi-erect pale setae, its length 0.5—0.8 times
maximum width of hind tibia.

Colour: head entirely light reddish brown or sometimes yellowish brownish, ocel-
lar triangle often black. Antenna black, dark reddish brown basally, two basal segments
yellowish brown. Palpi pale brown. Mesosoma almost reddish black, with light brown
on narrow stripe along lower margin of pronotal sides and along area of precoxal sulcus.
Metasoma mainly dark reddish brown, dorsal basal and apical parts black. Fore and mid-

A new species of Labania Hedgvist (Braconidae, Doryctinae)... 75

dle legs light reddish brown to yellowish brown. Hind leg mainly dark reddish brown,
tibia faintly paler and light reddish brown basally and apically, hind tarsus entirely light
reddish brown. Ovipositor sheath almost black. Fore and hind wings distinctly and even-
ly darkened, veins dark brown. Pterostigma black, slightly paler apically.

Male. Body length 2.9-3.9 mm; fore wing length 2.5—2.7 mm. Antenna slender,
filiform or narrow basally and weakly thickened apically, 20—22-segmented. First fla-
gellar segment 4.5—4.8 times longer than its apical width. Hind femur strongly wid-
ened, 1.9—2.1 times longer than width, dark brown or dark reddish brown. Hind tibia
almost entirely light reddish brown. Metasoma 0.9-1.0 times as long as head and
mesosoma combined. Length of first tergite 1.1 times its apical width, 1.2—1.5 times
length of propodeum. Otherwise similar to female.

Distribution. Known only from Costa Rica.

Type material. Holotype: female, “Costa Rica: San Jose, Reserva el Rodeo, root
galls on Ficus obtusifolia, 1V.2011, Col. Miguel Artavia (IB-UNAM). Paratypes: 30
females (9 in alcohol), 16 males (4 in alcohol), the same label as holotype (IB-UNAM,
MZCR, ZISP, NMNH); 1 female, same data as holotype, DNA voucher number
CNIN1078 (IB-UNAM), GenBank accession numbers KJ586713 (COI), KJ586690
(wingless) and KJ586780 (28S) (Zaldivar-Riverén et al. 2014) (IB-UNAM).

Biology. All examined specimens of L. ficophaga were reared from aerial root galls
ona single F. obtusifolia tree. Each gall (Fig. 2C) was approximately 1.5 cm in diameter
and contained multiple larval chambers. Galls were frequently grouped together form-
ing a mass that measured up to 6 cm in diameter and 12 cm in length. A photograph
of these galls is shown in Zaldivar-Riverén et al. (2014; Fig. 1B). They are similar to
the galls induced by L. hansoni Marsh on F. citrifolia, as reported by Marsh (2002).
It should be noted that the identification of the F. citrifolia requires verification; it is
possible that the correct identity is F. eximia Schott.

Galls of L. ficophaga were tracked on the host tree during a two years period (2011-—
2012), during which time the tree was visited every month. All wasps, including four
additional species that are probably parasitoids, emerged during April and May. No
wasps emerged during any other month. New galls began appearing in May and June
and became full-sized in August.

Two lines of evidence suggest that L. ficophaga is the gall inducer. First, it is larger
than the other four species emerging from these galls (parasitoids are expected to be
slightly smaller, unless they feed on gall tissue). Second, of the five species of hyme-
nopterans reared from these galls (no other insects were reared), L. ficophaga was the
only species that was present in all the samples and in most samples it was also the
most numerous species. For example, eight galls collected in April 2011 yielded 389
Labania, 89 Eurytomidae (probably Phylloxeroxenus Ashmead) species #1, and 28 Eu-
rytomidae (probably Phylloxeroxenus Ashmead) species #2. Two galls collected in April
2012 yielded 45 Labania and 4 Eurytomidae species #2. One gall collected in May
2012 yielded 10 Labania, 17 Torymus Dalman (Torymidae), 1 Eurytomidae species
#1, and 2 Platygastridae.

76 Sergey A. Belokobylskij et al. / Journal of Hymenoptera Research 44: 69-78 (2015)

a

Ul

Figure 3. Labania minuta Marsh (female A—C, E; male D): A habitus, lateral view B head and mesosoma,

dorsal view C head and mesosoma, lateral view D hind femur E metasoma, dorsal view.

Updated key to species of Labania (after Marsh 2002)

1 Body length 1.0—1.8 mm (Fig. 3A). Antenna with less than 20 segments .....
PN, Ra Re POO ee cc Bere Brae Ie L. minuta Marsh (Fig. 3A—E).
= Body length 2.5—4.1 mm (Fig. 1A). Antenna with more than 20 segments..... 2
2 Second metasomal tergite completely smooth (Fig. 1E). Mesosoma and
metasoma mainly or entirely black or dark reddish brown (Figs 1A, C, D, E).
Wringssdistinetly: darkened (Picgmlincs Eos Cy) ayia) a Gece alee esan theca. 3
- Second metasomal tergite longitudinally striate basally. Mesosoma and meta-
soma at least partly brown. Wings mainly subhyaline oe eee 4
3 Transverse diameter of eye 1.4—1.7 times wider than temple. Eye covered
with long and very dense setae. Malar space 0.4—0.5 times height of eye.
Propodeum mainly smooth, rugose only medially. Discoidal (first discal) cell

A new species of Labania Hedqvist (Braconidae, Doryctinae)... res

petiolate. Dorsal carinae of first metasomal tergite incomplete. Vertex and
frons:darksreddish browt-to: black.14..0i.s cee on et L. hansoni Marsh
= Transverse diameter of eye 2.0—2.2 times wider than temple (Fig. 1C). Eye
covered with short and sparse setae. Malar space 0.35—0.40 times height of
eye (Fig. 1D). Propodeum mainly rugose-striate, smooth only in basolateral
areas (Fig. 1C). Discoidal (first discal) cell sessile (Fig. 1F). Dorsal carinae of
first metasomal tergite almost complete (Fig. 1E). Vertex and frons light red-
dish brown or brownish yellow... cee eeeeseeseeeeeeeeees L. ficophaga sp. n.
4 Ovipositor extending beyond apex of mesosoma by length of hind basitarsus.
First metasomal tergite parallel-sided, its length greater than apical width.
Bodytlenethis3 03 Samii. ts thats veut sade bien ituteieot cal L. prolata Marsh
aa Ovipositor usually not extending beyond apex of mesosoma, rarely only very
weakly and much less than length of hind basitarsus. First metasomal tergite
distinctly widened apically, its length about equal to posterior width. Body
Jerse lr) SAO iiriina Pte. ston end tvteesduasnee aesetesosroeeed eeuts L. straminea Hedqvist

Acknowledgements

We thank Laura Sanchez for initiating the collection of galls, José Francisco DiStefano
for logistical support with the field trips, Cristina Mayorga Martinez and Guillermina
Ortega for mounting most of the examined specimens, Susana Guzman for taking the
digital pictures, and Elijah Talamas (Washington, D.C.) for images of the type species
of the genus. This study was supported by a grant given by the Russian Foundation
for Basic Research (grant No. 13—04—00026), programa PAEP-Posgrado en Ciencias
Biolégicas, UNAM (convocatoria 2014), and the Coordinacién de la Investigacién
Cientifica (CTIC-UNAM; convocatoria intercambio académico 2013) to SAB, and by
grants given by DGAPA PAPITT-UNAM (Convocatoria 2013) and CONACyT (Red
Tematica del Codigo de Barras de la Vida, México) to AZR.

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